Synopsis of Phylum Chordata

Identifying Characteristics of the Phylum

-most advanced phylum of animal kingdom
-includes fish, amphibian, reptiles, birds and mammals; some of the largest or most massive animals
-Notochord; flexible rodlike structure; extends the length of the body; in larva and/or adult
-Dorsal tubular nerve cord; (in invert ; nerve cord is ventral & paired)
-Pharyngeal (gill) slits; first evolved as a filter feeding apparatus
-endostyle or thyroid gland; specific kind of tissue found only in chordates
-Post-anal tail; in aquatic chordates it provides motility in terrestrial chordates it is used for balance
- ventral heart (inverts have dorsal heart)
-endoskeleton; most members have an internal skeleton of cartilage and/or bone

Subphylum Urochordata (tunicates, sea squirts)

-all marine; widely distributed in all marine waters
-most are bag-like sessile suspension feeders as adults, often colonial
-tadpole-like larva has typical chordata features
-adults have tough, nonliving, tunic covering body; secreted by mantle; forms 2 siphons
-filter feeders: incurrent siphon  pharynx (branchial sac) slits atrium excurrent siphon
-pharynx also serves as a respiratory organ
-simple open circulatory system with small ventral heart and 2 major blood vessels
-nerve with ganglia and plexus of nerve fibers
-all tunicates are hermaphrodites with single ovary and testis; free-swimming larva
-Human Impacts of Tunicates: antiviral, antitumor

Suphylum Cephalochordata (lancelets)

-closest living relatives to vertebrates
-slender, translucent, laterally compressed, fishlike or eel-like body
-instead of tunic, outer body is covered by soft epithelium; 2 folds of skin = metapleural folds
-springy notochord for support supports body while swimming or burrowing
-with well developed “V” myotomes (=myomeres)
-are filter feeders: mouth surrounded by oral hood with tentacles (=oral cirri) an a wheel organ
-pharynx strains food from water; water passes through pharynx into atrium and out the atriopore
-all are dioecious with males and females

Subphylum Vertebrata
-internal jointed skeleton of bone or cartilage
-complex skin; multilayered: epidermis, dermis
-digestive system uses muscular contractions rather than cilia to move food through
-increasingly efficient closed circulatory system with pumping heart (2,3, or 4 chambered)
-most complex and best developed nervous system of all animals; more emphasis on brain & senses
-Improved efficiency of excretory system; paired kidneys (most cephalochordates had none)
-almost all are dioecious and reproduce only sexually

Vertebrates: The Fish Classes

-fish are the most diverse, most abundant and successful group of living vertebrates
-all fish are aquatic & and highly adapted for aquatic life
-skin: epidermis usually secretes slimy mucus; dermis produces scales in most fish
-highly flexible “backbone” of cartilage or bone is the main support for swimming muscles
This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions 1
 -most of a fish’s body mass is myomeres (=myotomes)
-most fish have gills for getting O2 from water; some fish can also breath through their skin; a few
fish can breath air
-circulation is tied to gas exchange through gills; 2 chambered heart and a single circuit of bloodflow
-brains are relatively small and simple; cerebrum (higher centers) very small; cerebellum
(coordination of movement) relatively large; brain stem (automatic activities)relatively large
-probably the most important sense in fish is lateral line system = “distance touch”
-sound is an important means of communication in fish, especially deepwater fish
-kidneys remove wastes (Nitrogen wastes); gills also play role in excretion and osmoregulation
-most fish are dioecious; most with external fertilization (oviparous); a few bear live young

Class: Agnatha (Jawless Fish)

-oldest known vertebrates; most ancient & primitive vertebrate group
-not technically “vertebrates” since they have no vertebrae – just a cartilage rod for support
-only living vertebrate group with no jaws; also lack paired fins
-three main groups of agnatha: ostracoderms - all extinct; hagfish; lampreys
-Human Impacts: bane to some commercial fishermen using gill or set nets; collected for “leather” to
make golf bags and boots; in 1950’s lampreys destroyed great lakes fisheries

Class: Chondrichthyes (Sharks and Rays)

-evolution of jaws was one of the major events in the history of vertebrates
-Body Form; either fusiform (spindle shaped) or flattened
-skin is very tough & leathery muscles of shark pull on skin rather than pulling on the skeleton
-bony scales reduced to small, hard, knife-like (placoid) dermal scales embedded in skin
-all members of the group have a skeleton made mostly of cartilage
-paired appendages: pectoral and pelvic fins; but pectoral fins are rigid, not flexible
-most sharks are predators with powerful jaws; the teeth and (dermal) scales of sharks identical
-digestive system has new structures eg. liver, gall bladder, pancreas; spiral valve to improve absorption
-gills are inside 5 pairs of gill slits similar to agnatha; a pair of spiracles behind the eyes
-sharks retain urea to help maintain internal fluids isosmotic to sea water; rectal gland assists kidney
-most sharks, but only a few kinds of other fish posess a cloaca
-all chondrichthyes have internal fertilization; many bear live young
-Human Impacts: shark attacks. shark fishing, medicinal/pharmaceuticals

Class Osteichthyes (Bony Fish)

-most successful vertebrate class; more species than all other kinds of vertebrates combined
-most bony fish are designed for active swimming but with an amazing diversity of body form
-most bony fish have thin, overlapping dermal scales in dermis that grow throughout life
-most bony fish can control their color to some degree due to chromatophores
-freely moveable pectoral and pelvic fins for better maneuvering
-most bony fish today have swim bladder to control buoyancy
-most modern fish are carnivores; small, numerous, sharp teeth are used to seize prey
-much more efficient gills; often have “gill rakers”; covered by operculum
-most bony fish with external fertilization; a few bear live young (eg. guppies)
-A few fish make nests and show fairly elaborate mating behaviors and parental care
-some fish migrate between fresh and saltwater for spawning
-Human Impacts: pets, research, food

Class Amphibia
-modern amphibians still retain a unique blend of aquatic and terrestrial characteristics
This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions 2
 -most with thin moist, glandular skin without scales; often with many glands
-stronger, skeleton, mostly of bone, with toes; supports body weight & movement on land
-most amphibians are predators (carnivores); eat mostly insects
-most have long flexible tongues for capturing prey
-some amphibians have teeth to hold prey; food swallowed whole, not chewed
-amphibians can take in oxygen in four ways: lungs, through skin, mouth, gills
-circulatory system has 3 chambered heart & two complete circuits of blood flow
-amphibian brain is about same size as fish relative to body size
-Senses: lateral line; vision is dominant sense in many amphibians; smell has also become more
important; hearing – amphibians have both middle and inner ear
-skin and kidneys are the main way salts and water are gained or lost
-all amphibians have poison glands in their skin; some toxins are lethal
-dioecious; no sexual dimorphism; mating is controlled by season; external fertilization
-most frogs undergo metamorphosis into adult in a year or less
-during winter most temperate frogs hibernate in mud at bottoms of pools and streams
-Ecology: Frogs are critical links between predators and the bottom of the food chain
-Human Interactions: food, education, research, poisonings, as environmental indicators

Class Reptilia
-reptiles include lizards, snakes, turtles
-reptiles were the 1st vertebrates no longer tied to water, even for reproduction
-complete independence from water due to development of amniotic egg
-another major innovation of reptiles is a thick, tough, dry, waterproof skin; the skin of reptiles
contains scales but unlike fish scales reptile scales are in the epidermis, not under the
epidermis  also reptile scales are made of keratin, a waxy protein, not enamel and dentin
-more powerful muscles than amphibians; limbs are stronger and more flexible for walking
-most reptiles are carnivores
-tongue is muscular and mobile; in some tongue serves as touch receptor
-most reptiles have teeth
-in some salivary glands are modified into poison glands
-stomach often has pebbles to help grind food (=gastroliths) common find at dinosaur sites
-lungs are more efficient, more folding, more surface area; air is sucked into lungs, not gulped
-like amphibians, most with three chambered hearts; but partial septum separates the ventricle
-some reptiles in past were warmblooded
-nervous system similar to mammals in basic structure, only smaller
-vision is most important sense organ eyes usually with 2 moveable eyelids
-also have well developed sense of smell; Jakobson’s organ assists in sense of smell/taste
-some snakes have IR sensors
-more efficient (metanephric) kidneys
-venomous snakes use their poisonous fangs for protection as well as for subduing prey
-dioecious; copulatory organs; all reptiles have internal fertilization
-almost all reptiles go through early development within an amniotic egg
-many reptiles have well developed abilities to regenerate missing body parts
-Humans Impacts: snakebites, medical research, pharmaceuticals, farmed reptiles – semi-
domesticated, reptiles as food, world trade in live reptiles, invasive species

Class: Aves (Birds)

-birds clearly evolved from dinosaurs
-in spite of the great diversity of birds they are amazingly similar in structure; entire anatomy is
designed around flight
This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions 3
 -bird skin is thin, light and flexible, most of body is covered by feathers
-wings and body covered by feathers; light & strong and tough, feathers are molted regularly
-chromatophores impart colored pigments during feather development
-the skeleton is exceptionally light and delicate yet sturdy
-since birds lose the use of their forelimbs their beaks are used as tools; neck is extremely flexible
-breast muscles are the flight muscles
-beaks of birds are highly adapted for their feeding type
-birds are voracious feeders due to high metabolic rate
-crop stores food to provide a continuous supply of energy during flight
-modern birds have no teeth, grinding is done in gizzard; some birds “eat” pebbles to aid this process
-some birds of prey form pellets of undigested material (bones and fur) and regurgitate them
-birds & mammals are warm blooded (homeothermic)
-bird lungs are relatively small; instead of microscopic sacs bird lungs contain air capillaries & system of
air sacs in body; these air sacs also serve as an air conditioning system
-most birds have a larynx but use syrinx to generate sounds
-have 4 chambered heart & 2 completely separate circuits: pulmonary & systemic
-brain is same relative size as mammals
-predatory birds have eyes in front of head; other birds have eyes that look out to sides
-some of the most obvious and characteristic features of birds are the nests they make
-some with elaborate courtship, nesting, mating, and parental behavior
-Bird Ecology: pollination, disperse seeds, pest control
-Human Interactions: meat and eggs, introduced pests, domestication, bird watching, hunting, bycatch,
research, wildlife photography, art

Class Mammalia
-today, is one of most successful group of vertebrates
-most massive of all animals today or that ever existed is a mammal; blue whale
-skin is thicker and more complex; many different glands; sensory structures
-body covered with complex layer of skin with hair (fur); is periodically molted
-other keratinized (horny) structures of mammals: bristles, spines, vibrissae, horns
-mammals have a great variety of skin glands: sweat, scent, oil, mammary, wax
-great variation in structure of skeleton based on method of locomotion and lifestyle
-mammals are warmblooded and much more active than reptiles
-teeth represent the greatest evolutionary diversification of the mammalian skeleton
-the digestive system may also be modified in various ways determined by their diet
-all mammals have very efficient lungs and breath air
-mammals have 4 chambered heart with 2 completely separate circuits of blood flow
-relatively large, highly developed brain  disproportionately larger per body wt
-vision and hearing well developed in most mammals
-dioecious, internal fertilization, all but one small group of mammals are viviparous
-nurse young with milk  mammary glands
-3 patterns of reproduction: egg laying, marsupials, placental mammals
-Mammal Ecology: pollination & plant dispersal
-Human Impacts: domestication, pets, service animals, hunting, fur & game farming, zoos education,
research, food and crop loss, sickness & disease, illegal trade in mammal products, bycatch, pollution,
tourism, wildlife photography, art, entertainment

This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions 4
 Synopsis of Phylum Echinodermata

Identifying Characteristics of the phylum

-means “prickly skin”; include: starfish, sea cucumbers, basket stars, brittle stars, sea lilies, etc
-has an extremely abundant and diverse fossil record; much more diverse fossil record than species
existing today
-all marine; found in all oceans at all depths; some of the most abundant of all marine animals
-almost all are bottom dwellers
-only major invertebrate phylum with affinities for vertebrates
-most with pentamerous (=pentaradial) radial symmetry
-outer surface covered by epidermis; below epidermis is thick dermis made of connective tissues
-dermis secretes skeletal pieces (= ossicles) = endoskeleton
-echinoderms can vary rigidity of dermis = “catch collagen”
-water vascular system; madreporite leads to stone canal, joins ring canal around the mouth, radial
canals extend into each arm, lateral canals branch off , lead to ampullae, connected to tube feet
-echinoderms are particle feeders, scavengers or predators; no parasitic species
-simple, usually complete digestive tract; stomach has 2 chambers: cardiac & pyloric; digestive
enzymes are secreted into stomach by pyloric caecae
-respiration by dermal branchae (or papulae)
-no brain or centralized processing area; circumoral ring and radial nerves branching from it
-few specialized sense organs; have some simple tactile, chemical and photoreceptors and statocysts
-in many starfish the body surface bears small jaw-like pedicellariae
-sexes typically separate  dioecious; external fertilization; planktonic bipinnaria larva
-some can also reproduce asexually by fragmentation; excellent powers of regeneration

Class: Asteroidea (sea stars, starfish)

-inhabit all seas except low salinity areas
-bottom dwellers; mostly found on hard rocky surfaces; many live in deep ocean
-body composed usually of 5 rays (arms) projecting from a central disc
-mouth in center of oral surface
-wide furrows project from mouth into each arm = ambulacral grooves
-aboral surface with madreporite toward one side & numerous pedicellariae
-movement mainly by tube feet
-many sea stars are scavengers; a few are suspension feeders; most asteroids are carnivores
-many starfish regularly reproduce asexually
-most are dioecious; gonads in small area at base of each arm; most produce free swimming larvae

Class: Ophiuroidea (brittle stars, basket stars, serpent stars)

-the most active of the phylum
-found in all types of marine benthic habitats; mainly benthic; tend to be secretive
-long thin arms sharply set off from disc; no ambulacral groove; visceral organs confined to central disc
-locomotion by snake-like arm movements; muscles are much more important in this group; ossicles of
arms are arranged into flexible columns (called “vertebrae”) connected by muscle strands
-brittle stars are carnivores, scavengers, deposit feeders or filter feeders
-mouth on oral side has 5 jawlike plates; incomplete digestive tract
-no dermal branchiae’ brittle stars have internal sacs called bursae for respiration
-gametes discharged through the genital slits to the outside; some brood their young in the bursae
-brittle stars can spontaneously cast off arms; the pieces can regenerate into whole brittle stars

This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions 1
Class Echinoidea (sea urchins, heart urchins, sand dollars & sea biscuits)
-widely distributed in all seas; all are benthic; seem to prefer hard substrates
-compact body enclosed within a test (or shell) of closely fitting ossicles sutured firmly together
-no arms, but 5 ambulacral areas on test through which very long tube feet extend; with long spines
-mouth with Aristotle’s lantern; used to scrape and chew algae from rocks
-use very long tube feet and prehensile spines to move
-most sea urchins are grazers; scrape algae from substrates with teeth

Class: Holothuroidea (Sea Cucumbers)

-rule the deep ocean benthos  make up 90% of biomass on deep ocean floor
-like sea urchins have no arms
-tend toward bilateral symmetry; with mouth and anus are on opposite ends
-body has a leathery appearance; ossicles reduced to microscopic plates embedded in body wall
-mouth is surrounded by 10-30 tentacles (modified tube feet)
-large fluid filled coelomic cavity serves as a hydrostatic skeleton
-mainly deposit feeders and suspension feeders; use tentacles to collect food and deliver it to mouth
mouth opens into a muscular pharynx; then to esophagus and stomach, then a long, looping intestine
leads to anus which opens into cloaca
-most have a respiratory tree for respiration & excretion
-many sea cucumber are capable of evisceration; the organs are later regenerated
-some also have tubules of Cuvier that can be aimed and shot out of the anus for protection
-most are dioecious; a few are hermaphrodites; some brood their young inside coelom

Class: Crinoidea (sea lilies, feather stars)

-an ancient group; many fossil species
-some are stalked sessile animals; others are free living and motile
-most live at depths of 100 M or more
-flower shaped body; body disc, = calyx, is covered in leathery skin
-upper surface of calyx bears mouth and anus; arms have pinnules giving feather-like appearance
-no madreporite, spines or pedicellariae
-crinoids are suspension feeders
-very slow metabolism
-can probably live for 1000’s of years
-dioecious; either brood eggs or release them

Ecological Role of Echinoderms

-a wide variety of other animals make their homes in or on echinoderms
-sea stars are often the top predators in some benthic communities
-though unpalatable to most organisms to some they are the preferred meal: eg. some fish with strong
teeth, eg. sea otters
- “crown of thorns” starfish destroys Pacific coral reefs; feed on coral polyps; sometimes attack in “herds”
-sea urchins destroy kelp forest

Economic/Human Impacts
-echinoderms never attack humans; don’t transmit any diseases
-handling poisonous forms can kill
- predatory starfish can devastate commercial clam or oyster beds
- in China and Pacific Islands sea cucumbers are eaten as a delicacy
-roe (gonads & eggs) are sold, raw or roasted, as a delicacy in Japan and in sushi restaurants
-echinoderms have been widely used in developmental research
This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions 2
 Major difference between reptiles and mammals
REPTILE
Epidermis covered with scales
Most with three chambered heart
Most with no diaphragm
Low metabolic rate
Oviparous (egg laying)
little care of young
Growth continues through life
Skull with small brain case,
several bones forming jaw
Teeth continually replaced with
simple cheek teeth
Anterior brain (cerebrum) relatively
small
Skull with one occipital condyle
Single middle ear bone
(columella=stapes)
Bio 1413, Ziser, 2000
MAMMAL
Epidermis covered with hair
All with four chambered heart
Diaphragm for respiration
High metabolic rate; homeotherms
Viviparous (young develop within
mother) placenta, live birth
& care of young with milk
Growth limited after adulthood
Skull with expanded brain case,
single bone forming jaw
Two sets of teeth only (deciduous &
permanent) with complex cheek
teeth
Anterior brain (cerebrum) larger and often
convoluted
Skull with two occipital condyles
Three middle ear bones
(malleus, incus, stapes)
Bony palate usually incomplete Bony palate completely separates nasal
passages from mouth

Ribs present on all vertebrae Free ribs confined to thoracic vertebrae

Pelvic bones separate Pelvic bones fused

Sprawling gait with limbs emerging Upright stance wth limbs directly beneath
horizontally from body body
 Animal Welfare
“Animal Rights” vs “Animal Welfare”

Animal Rights

 based on the idea that animals have inherent

legal and moral rights just as humans do.

animals can’t have “rights”

 implies ability of animals to reason with humans

and agree on mutually accepted principles

 implies lives of all animals, including humans,

are equal

but this implies it is unethical to use animals as pets or

for any other purpose

eg. food, clothing, recreation, education, research

eg. pets = form of slavery

eg. killing rats or roaches is murder

Animal Welfare

 any use of animals should be motivated by

humanitarian goals

 we are obligated to minimize pain

Frontiers in Biology: Animal Welfare; Ziser, 2008 1
  we are required to show accountability for our
actions

there are many animal welfare movements

need to define “animal”

warm blooded vs cold blooded

vertebrates vs invertebrates

does a sponge or an earthworm deserve the same

consideration as a primate?

if so, why draw the line at animals

what about protozoa, fungi, plants, bacteria?

Frontiers in Biology: Animal Welfare; Ziser, 2008 2

 Avoiding contact with animal products is
virtually impossible

Many uses of animal products are hidden:

eg. medicines, film, rubber, ceramics, plastics,
paint, perfumes, glue, explosives, cosmetics, shaving
cream all contain materials from slaughter houses

eg. cellophane  made with animal fats

eg. freon animal fats used to make it

eg. makers of synthetic fibers use tallow based products to

control static cling

eg. animal based lubricants are used in jet engines

 all flying miles are animal based

eg. used in corrosion inhibitors for oil pipelines

eg. cars manufacture alone:

galvanized steel body, fan belts, gaskets, anti freeze,
hydraulic brake fluid, battery, steering wheel, dashboard,
tires

animal fats and hides are even used in asphalt on the roads
the car drives on.

Extent of animal products “exposure”:

foods ~75%

clothing ~10-20%

soaps & cosmetics ~5-10%

The animal by-products industry brings in $2 Billion/yr

Frontiers in Biology: Animal Welfare; Ziser, 2008 3

 eg. 1000 lb steer:
432 lbs retail beef
568 lbs by products
27 lbs: variety meats; hearts, livers, brain,
tongue, kidneys
358 lbs hide, hair, bones, horns, hoofs, glands
and organs
46 lbs blood
183 lbs fat

eg. hide: $50-$75/hide

 US sends 90% of hides overseas for fabrication then

back to US for product sale

hide: clothing, insulation, ointment base, binder for

plaster and asphalt

hair: toothbrush bristles, mattresses, air filters,

upholstery covers

hair from inside of cows ear

 “camel’s hair” paint brushes

eg. hides and connective tissues, cartilage, blood, bones:

glue in plywood, paper matches, textiles, cardboard,
window shades

eg. bones, horns, hoofs: gelatin for photofilm and

pharmaceuticals (gelatin capsules)

cattle horns: imitation tortoise shell

hooves: white  imitation ivory;

black  potassium cyanide  used to extract
gold from ore

eg. bones: electrical bushings, dice, chessmen, crochet needles,

piano keys, buttons, knife handles,

bone charcoal is used as refining material to purify

Frontiers in Biology: Animal Welfare; Ziser, 2008 4
 steel, filter sugar cane, manufacture high grade
steel ball bearings

eg. blood: dried and used in cattle, turkey and hog feeds; pet
food, fertilizers, clotting factors are extracted for
pharmaceuticals

eg. glands: >130 different medicines and pharmaceuticals

eg. tallow and lard:

(tallow hydrolysis glycerine and crude fatty acids stearic & oleic acids)
glue, agricultural chemicals, candles, cosmetics, detergents,
drugs, metal castings, paints, inks, paper, shaving cream

eg. if animal has gall stones

 in orient $1000-$2000/lb of gallstones
used as aphrodisiac

Frontiers in Biology: Animal Welfare; Ziser, 2008 5

 Animals as Food
by far the largest use of animals (~99%)

 >10 Billion animals are killed each year for food

including:
9.5 billion chickens turkeys & ducks

42 million cattle

97 million pigs

4 million sheep

also,

an additional 125 million animals are hunted

with estimates that two animals are

wounded/injured for every one killed

Frontiers in Biology: Animal Welfare; Ziser, 2008 6

 Furs
farmed animals provide 85% of furskin production per
year worldwide

80 million animals are killed in the US each year

includes 50 million raised in captivity

and 30 million hunted

mink and fox are the most common furbearing animals

raised on farms
eg. mink pelt production in the US was 2.6 million pelts in
2002

the US is the world’s largest volume producer of

furskins derived from wild animals

there are ~150,000 licensed trappers in the US

annual (97-98) US harvest by trappers:

otter 29,000
coyote 159,000
nutria 398,000
mink 190,000
red fox 164,000
muskrat 2,183,000
beaver 429,000
raccoon 2,896,000
other 613,000

TOTAL 7,062,000

Frontiers in Biology: Animal Welfare; Ziser, 2008 7

 Pets
>100 M mammals are kept as pets

300,000-500,000 from “puppy mills”

Benefits
can improve physical and mental well being

provide companionship

especially effective for lonely and depressed

but:
 up to 20 Million cats and dogs are abandoned
each year to starve or be put to sleep

Frontiers in Biology: Animal Welfare; Ziser, 2008 8

 Animal Use in Education
three most commonly dissected vertebrates:

frogs
cats
fetal pigs

9M frogs shipped to US schools (1969)

~1/2 live, 1/2 preserved

this is the only animal experience of most students

41% of K-12 teachers report no time spent on live

animals

only 26% spent >5 hrs/semester on live animals

65% spent >5 hrs/semester on preserved animals

Frontiers in Biology: Animal Welfare; Ziser, 2008 9

 Animal Use in Research
probably one of the most contentious issues with
animal welfare advocates

Biomedical research is a major enterprise

US spent $9 B in 1984:

70 M vs 10 Billion killed for food (1.4%)

90% of research animals are rodents bred specifically

for research

for every dog or cat used in research

~100 are killed at shelters and pounds

each year cosmetic and drug companies use millions of

mice, rats and rabbits to determine the safety of
chemical they plan to use in their products

 20% of all animal use in science(‘87)

about half of the biomedical research carried out in US

would not have been possible without lab animals

about 2/3rd’s of projects that led to Nobel Prizes in

Physiology and Medicine used animal
experimentation

What kinds of research?

animals are used for both basic and applied research

Frontiers in Biology: Animal Welfare; Ziser, 2008 10
 some argue that they should only be used for
applied research but you can’t separate the two

diagnostics
1-5% of all lab animals are used to diagnose disease
eg. TB, diptheria, anthrax, burcellosis, etc

disease models:
eg. inbred mice for Hodgkins lymphoma
eg. primates for HIV

organ transplantation:
eg. tissue typing techniques, immunosuppression
drugs

bionics research

development of anticancer drugs

determine treatment regimens

minimizing side effects

psychoactive drugs:
eg. lithium  calming effect on pigs

surgical procedures:
eg. balloon angioplasty

extraction of medical products

eg. hormones, blood for culture media

production of antisera, antibodies, & vaccines:

eg. diptheria, whooping cough, tetanus, polio
eg. smallpox vaccine from skin of calves or sheep
eg. rabbits as antisera factories

antibiotic testing

Frontiers in Biology: Animal Welfare; Ziser, 2008 11

 large scale screening of potential drugs and antibiotics

toxicity testing

food and water safety

search for new drugs

in terms of specific animals, the most commonly used

animals are preferred because:
easy to handle & maintain
inexpensive to maintain
most appropriate model for a particular kind of research
generally docile
breed quickly
easily acquired from pounds and shelters or purchased from
internet
easily bred in labs

examples of specific animals used in research:

Mamals:
mice & rats: 95% of all animal research are done on mice and
rats.

used in virtually every kind of scientific investigations

cows: narcolepsy, reproductive physiology, vaccine testing,

infectious disease research, heart studies

pigs: very important animal model for human physiological

studies; cardiovascular research, blood dynamics, nutritional
deficiencies, alcoholism & drug abuse, general metabolism,
digestive related disorders, respiratory disease, diabetes,
kidney and bladder disease, organ toxicity studies,
dermatology, neurological studies, burn studies, cystic
fibrosis research
Frontiers in Biology: Animal Welfare; Ziser, 2008 12
 sheep: pregnancy related research, multiple sclerosis, medical
implant studies, burn and injury evaluation, smoke
inhalation

70% of sheep used in research experience pain

goats: studies in cartilage repair, respiratory physiology, medical

diagnostics, gene therapy, anesthetics research, used to
produce antibodies, and to produce genetically engineered
products

dogs: heart and lung research, transplantation experiments,

cancer research, microbiology, genetics, orthopedics,
surgeries, vet medicine, toxicity studies of drugs, additives
and industrial chemicals

cats: neurological research, spinal cord injury, used to study

vision, sleep and hearing problems, Parkinsons disease,
cancer, genetic disorders, HIV/AIDS research

50% of cats used in these experiments suffer pain and

distress

rabbits: toxicity testing for cosmetics and household products;

especially Draize eye & skin irritancy tests
 very painful
 rabbits often locked in full body restraints

also used as models for eye diseases, skin, heart and

immune system studies, asthma research, cystic fibrosis
studies, diabetes and used to produce antibodies for
research and diagnosis

guinea pigs: toxicity & safety testing, effects of cigarette

smoke, alcohol and drugs, spinal cord injury investigations,
TB research, kidney functionk, osteoarthritis research,
nutrition and genetics studies, reproductive biology and
study of infectious diseases

12% of guinea pigs are subjected to pain and distress

Frontiers in Biology: Animal Welfare; Ziser, 2008 13
 hamsters: taste and vision research, cardiopulmonary research,
cancer and muscular dystrophy investigations, studies of
aging, asthma, and biorhythms

20% of hamsters are subjected to pain and distress

birds

those used for research are excluded from animal

welfare regulations
no US stats

but: 100,000 birds/ yr in UK and 650,000 birds/ yr in EU

are used for research

they are the third most common experimental animal

after rodents & fish

fish

some cold-blooded animals are also used for

research but they are exempt from animal
welfare regulation

probably ranges in US from 3.5-7 M/yr

used for biomedical research, toxicity testing, aquaculture, cancer
research

What is the value of animal research:

some of this information cannot be learned any other

way

Frontiers in Biology: Animal Welfare; Ziser, 2008 14

  its unethical to test surgeries or drugs in
humans 1st (=human rights issue)

 can set up controlled experiments that you

cannot do with humans
eg. genetically identical pairs
eg. exact feeding regimes

many surgical and medical procedures used in

research had spinoffs in veterinary sciences
pets, livestock, zoo animals generally live longer, are
healthier and live more comfortably because of animal
experimentation

animal experimentation has helped to preserve

endangered species :
treat illnesses,
eliminate parasites,
promote breeding (eg. artificial insemination, embryo
transfer, capitve breeding)

Are there alternatives?

Criticisms of animal researchers:

a. inadequate self regulation
only recently have standards of care been dramatically
improved; they were slow in coming

biomedical research has always been closely regulated

but really are not many inspections done

b. slow to replace animal models with alternatives

Frontiers in Biology: Animal Welfare; Ziser, 2008 15

 few incentives to change even when alternatives are
available

c. tend to point fingers in other directions

it’s the other groups, not us, who are mistreating animals

Criticisms of animal welfare activists:

a. oversimplistic generalizations, loose thinking

eg. animal testing compared to Nazi legacy of human abuses
for “research”

but:
ironic that animal research was almost banned in Nazi
Germany before the war

b. misstatements, misrepresentation of the

problem
eg. development of polio vaccine cost 2 M monkeys and
didn’t reduce polio rate from 1916 to 1962

but:
polio research only started in 1953
by ‘70’s polio rate dropped to near 0 in US

eg. thalidomide touted as drug that got through animal

testing and still proved dangerous;

also has been stated that many tests were performed

on pregnant animals
but:
actually, didn’t get enough animal testing
no pregnant animals were used in research

eg. some believe that all animals suffer agony at some

Frontiers in Biology: Animal Welfare; Ziser, 2008 16
 stage of research

tout statistic that 80% of experiments are done

without anesthetic

but: most didn’t require any, there was no pain

involved

c. some antivivisectionists tactics result in more

pain and mistreatment than the research they
oppose

eg. “freeing” lab animals

most will be hunted and killed by wild animals

eg. one group was charged with animal abuse for keeping
over 200 dogs on a 1 acre enclosure to prevent their
use in medical research

d. the “animal rights” movement has:

driven up the cost of research

more money spent on tighter security and to repair

damaged facilities

may slow development of therapies and

treatments
reduces the amount of research being done

some research must be started over when facility is

damaged or animals released

other methods are often cheaper and require less

paperwork: scientists use them when they can

Frontiers in Biology: Animal Welfare; Ziser, 2008 17

animals are used only when it is the best way to get
the appropriate information
eg. some aspects of the causation, treatment or prevention
of blindness cannot be studied in bacteria, fungi or
plants  need complex animals

eg. high blood pressure cannot be studied in invertebrates

still, there is an effort to find alternatives when

possible
eg. may be best to use alternatives for toxicity studies,
or do nonanimal tests first then use fewer animals for trials

most researchers hold non-animal experiments cannot

completely replace animals:

adjuncts not alternatives

Laws

animal experiments are subjected to a wide variety of

overlapping laws, regulations, and guidelines

Animal Welfare Act (1966)

standards for handling, housing, transporting, feeding, vet
care, use of pain relieving drugs
in dogs, cats primates, rabbits, hamsters, etc
investigators must consider use of alternatives
allows for unannounced inspections of facilities

Health Research Extension Act (1985)

human care and use of animals
procedures established to insure compliance

Frontiers in Biology: Animal Welfare; Ziser, 2008 18

Animal Facilities must be accredited by Institutional
Animal Care and Use Committee

There are also state and local laws against animal

cruelty

Pain

probably one of the biggest concern is causing pain to

animals

most animals are capable of experiencing pain

 generally scientists acknowledge and accept that

all warmblooded animals and most
coldblooded vertebrates (frogs, fish, etc)
experience pain

even though experiencing pain, many animals may not

show any external signs of pain

animals that show distress in nature might attract a predator

eg. recent evidence has shown that even fish have pain receptors
and experience pain when caught on fishing line

one simple test for pain:

“a stimulus is said to be painful if it is consistently

terminated or escaped by subject”

animals tend to begin to escape pain sensations at

Frontiers in Biology: Animal Welfare; Ziser, 2008 19
 about the same intensity that humans begin to
report pain

Most animals experience only minimal pain in research

settings:

eg. Animal Welfare Enforcement Report (1988)

94% of all lab animals are not exposed to
painful procedures or given drugs to relieve any pain

6% are exposed to painful procedures which

are usually not severe or long lasting

eg. Biomedical Research Study (1989)

58% experienced no pain, received no pain medication
35% received anaesthesia  little or no pain
7% experienced significant pain

eg. there are safeguards to insure animals for research are well
cared for:
 unhealthy animals can lead to eroneous results
 animal research is expensive; can only afford high
quality research
 pain can invalidate an experiment because stress
induces physiological changes in virtually all body
systems

some kinds of research subject animals to considerable

pain:

eg. orthodontic research

eg. car crash studies

originally used human cadavers, but their use was
banned

eg. oral radiation research

Frontiers in Biology: Animal Welfare; Ziser, 2008 20
 subject animals to enough radiation to cause death

eg. tumor therapies

there are also cases of:

inadequate use of anaesthesia and

inadequate care of laboratory animals

Alternatives to Animal Testing

2006: beginning to develop the first realistic software

models of human and animal organs

can show thousands of molecular interactions

can manipulate physiological processes

modelers agree that still, some degree of animal

testing will be needed; but new models can
definitely reduce the need and extent of animal
testing; some have doubts due to complexities
of living sysstems

Frontiers in Biology: Animal Welfare; Ziser, 2008 21

 Additional perspective on animal welfare:

 up to 20 Million cats and dogs are abandoned

each year to starve or be put to sleep

 many more pets and farm animals are

neutered, some without anaesthetic, than are
subjected to experimental surgery for research

 habitat destruction kills many millions more of

animals, and whole species are lost, yet this is
NOT the major issue with “animal rights”
advocates

Frontiers in Biology: Animal Welfare; Ziser, 2008 22

 Human Evolution single man (“Adam”) who lived sometime
between 120,000 to 156,000 years ago
modern humans are a relatively new species on the however, some Cameroonians and one African American
planet tested have Y chromosomes not related to the above
male ancestor; their Y-chromosome date to between
oldest Homo sapiens fossil are ~200,000 yrs old 180,000 to 581,000 years ago

about 100 B people have lived on earth so far or about 100,000 the human genome is 99.9% identical worldwide
generations
the human genome is also ~98% the same as the
today we can trace human history using the males’
genome of chimps (our closest living relatives)
“Y-chromosome” and mitochondrial DNA
the Y chromosome is passed to the males of every generation ! how did we get to that point?

DNA in the mitochondria and passed the males & females of Primates
every generation

humans are in the order Primates and evolved from

One misconception surrounding mitochondrial Eve is that since all women alive today
descended in a direct unbroken female line from her, she must have been the only
some early member of this group
woman alive at the time. However, nuclear DNA studies indicate that the size of the
ancient human population never dropped below tens of thousands. Other women earliest primates appeared ~65 MY ago during time of great
living during Eve's time have descendants alive today, but at some point in the past
each of their lines of descent did not produce a female, thereby breaking the diversification of mammal after the extinction of the dinosaurs
mitochondrial DNA lines of descent.

primates lived in trees (= arboreal) which selected

results of mitochondrial genetic studies indicate for a specific group of traits important in human
that all living humans are related to a single evolution:
woman (“Eve”) who lived ~150,000 years ago
in east Africa a. 5 digits sometimes with opposable thumb
! better gripping and dexterity
results of Y-chromosomal inheritance studies
indicate that all living humans are related to a b. nails rather than claws

Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 1 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 2

! greater tactile sensitivity of fingertips

more use of hands and arms for manipulation
c. visual acuity
may have moved to grasslands early
could form sharp images
had color vision ! seems to have eaten roots and seeds rather
decreasing reliance on chemical senses than soft vegetation of plants

d. greater maternal care modern Apes & Chimps

longer, stronger mother-child relationships most are larger than other monkeys
larger brain in proportion to size
long arms & short legs ! brachiation
e. tendency toward upright posture
4 modern genera
at least could sit upright
eg. gorillas, chimps

Hominidae
Humans and Human Ancestors (Hominins)
primates are divided into 13 “families” which include
uniquely “human” traits
the gibbons, marmosets, old world monkeys,
tailed monkeys and the Hominidae which include
– not found in other living primates:
the “great apes”: gorillas, chimps and us.
used to be “the family of man” more recent genetic analysis a. habitually erect posture ! truly bipedal
showed it should also include closely related primates b. lower limbs longer than forelimbs
c. non-opposable larger toes
apes first appeared ~20 million years ago (miocene)
d. reduced canines
e. bony chin
a time when woodland savannas were replacing
f. prominent nose
forests and spreading across Africa, Europe
g. relatively hairless
and N. America
h. much larger brain led to:
! abstract thinking
earliest fossils were smaller than a chimp ! speech and language
! use of complex implements
tended toward bipedalism
Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 3 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 4
 ! belief in supernatural and life after death
! importance of cultural evolution Ardipithecus was probably in direct line of
descent to us
the evolutionary branch leading to humans probably
began ~7 million years ago if so, they are separated from us today by
over 300,000 generations
molecular evidence indicates that Great Apes diverged from
human line ~10 MY ago (DNA is ~97% similar) 2. Australopithecus sp. [4.2 –2.3 MY ago]

chimps diverged ~8 MY ago (DNA is ~98% similar) several species known (eg. “Lucy” & Taung Child);

some not in our direct line of ancestry

it apparently wasn’t a clean break

! both ancestral lines apparently continued to interbreed 4’ tall, ~50 lbs

for ~1.2 MY after the split (based on DNA analysis)
the first bipedal primate (the only one of the time)
! chimps share 98% of our genes
walked upright (bipedal), ground walker
! 40 million molecular changes from them to us
only 250,000 seem most responsible for the lived in open grasslands (eg. Lucy & footprints)
physical and mental differences between us
and them(AAS 9-1-05) ! better vision in grasslands

!frees hands for weapons and harvesting

the earliest fossils of human ancestors appear in the
fossil record ~4.4 Million years ago still had relatively long arms

1. Ardipithecus ramidus (“Ardi”) [4.4 MY] still probably spent lots of times in trees

lived in the woodlands of E Africa no dramatic increase in brain size;

~4’ tall, weighed ~110 lbs
had long arms, short legs with opposeable toes slightly larger brain than modern monkeys and apes
spent most of their time in trees; ate, slept and raised (relative to size)
young in trees
but could easily walk on 2 legs and carry food in arms, relatively small canines and incisors compared to
cruised the trees on all 4’s
probably several distinct species of the genus apes
Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 5 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 6

! tooth size decreased

some species began using primitive tools
diet changed from largely vegetarian to more
improving tool use in human line probably led to the carnivorous
development of speech
! large increase in relative brain size
making tools uses the same areas of the brain as
speech higher protein diet fed brain development

during this time our ancestors acquired one of brain uses 20x’s more energy than muscle
today’s uniquely human parasites, pubic lice ! larger brain needed more high quality food
genetic analysis indicates the pubic lice evolved from gorilla
body lice about 3 M years ago at exactly this time when our genus
(Homo) first appears, fossil and genetic
this also indicates that our ancestors of the time may have evidence indicates that:
spent considerable time with gorillas then
! a single gene was duplicated that gave brain cells
abt 2.5 MY ago Australopithecus split into 2-3 (neurons) the ability to form more complex
shapes and connections allowing the exchange of
major evolutionary lines: more information with a larger number of
neighboring cells
A. robustus, A. boisei & Homo habilis
! Infant skulls became more flexible allowing them to
3. Homo habilis [2.5 – 1.6 MY] accommodate larger brains

earliest “human” fossils (same genus as us) H. habilis was also a tool user

existed same time as Australopithecus and shares many of same 4. Homo erectus (eg. Peking Man) [1.8 MY – 30,000 yrs ago]
traits with following changes:
the fossil called H. habilis is so similar to Australopithecus that it
! more delicately built is questioned by some to not actually be in our genus at all

! females ~ 1/2 as large as males by 1.5 MY a group of hominins had evolved that
are indisputably in our genus
males probably had harems as do apes

Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 7 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 8
 distinctly different from Australopithecus and
much more similar to us:
! skeleton similar to ours
! close to our size, ~5’ tall, ~100 lbs;
! males and females similar sizes
! similar stride
more efficient walk than Australopithecus
wider ranging than Australopithecus
first hominids to migrate out of Africa
fossils found in Africa, Asia & Europe
the main difference between H. erectus and us is
the skull
! their skull is thick and massive
!large jaws and teeth, no chin
show another dramatic increase in brain size
brain size is correlated with intelligence
it evolved 100’s of x’s more quickly than most traits evolve
this occurred after bipedalism and tool use
strong selective pressure for brain development
were efficient hunters
Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11
before fire use early hominins ate tough, raw foods that
required lots of chewing and digesting to break food
down
! cooking made all foods more easily digestible
more nutrition for less effort
enhanced brain development
teeth got smaller
!1.2 MY ago: genetic evidence indicates ancestral
“fur” was shed; we became “naked apes”
a naked body allowed us to sweat to keep cooler in a warm
environment
fewer fleas and other ectoparasites
H. erectus probably had limited speech ability
fossil evidence suggests that areas of the brain responsible
for language existed over 500,000 years ago
spoken language developed before written language
5. Homo neanderthalensis [600,000 – 30,000]
(Neanderthal man)
mainly in Europe and near east; earliest fossils from England and
Germany
more heavily built, very strong
protruding face, low skull, heavy brow
Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11
earliest use of projectile weapons
plenty of game, more grasslands in their time
killed elephants, rhinos, antelopes, bears, hippos and
giant babboons
sometimes stampeded them into marshes or over cliffs
new analyses indicate that the the anatomy and
physiology of humans became adapted toward
distance running and spear throwing about 2 M
years ago
running was perhaps an adaptation for hunting
running prey to exhaustion
a well conditioned human can run 5 miles or more
as good as wild dogs, zebras, anteloes and
wildebeasts
it would also allow us to compete with dogs and
hyenas for widely dispersed carcasses
humans are the only primates who can launch a
spear or rock overhand with speed, force, and
precision
all other primates throw underarm with poor aim
1st ancestor to tame fire; all before “ran from
it’
9 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 10
brain size equivalent to ours
had language
communication and culture became more important
than physical evolution
used tools
cared for invalids
buried their dead with weapons, food & flowers
! belief in afterlife
lived with H. sapiens (us) for over 100,000 years
occasionally interacted with us
even bred with us a few times
6. Homo floresiensis (the “hobbit”) [95,000 - 13,000]
recently discovered in a cave on a remote island in
Indonesia
lived with pygmy elephants & Komodo dragons
may have evolved as separate line from H. erectus
~ 3.3’ tall, weighed ~55 lbs
upright, bipedal, large feet!
longer arms than modern humans
11 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 12
 small brained !most recent analysis indicted by genetic evidence for evolution
of human body louse species that only lives in clothing, not
used fire and made tools on skin or fur from human head louse. a reanalysis
indicates it might have been much earlier
hunted large game ! required communication and planning
~100,000 years ago earliest known use of jewelry and
modern humans didn’t reach the island until ~11,000 years
ago so we don’t think the hobbits interacted with
cosmetics
modern humans

7. Homo sapiens (Cro Magnons) [200,000 – present] ~60,000 yrs ago earliest needles appear

our species apparently first appeared ~200,000 indicating that people had begun to sew and create clothing

years ago
~35,000 yrs ago: earliest cave paintings
H. erectus, H. neanderthalensis, H. foresiensis & H. sapiens and
possibly 1 or 2 more species coexisted for 1000’s of years ~60,000 years ago perhaps ~1000 people
moved out of Africa and into western Asia
H. erectus & neanderthals disappeared abruptly about
34,000 yrs ago
(some evidence indicates that humans first left Africa ~120,000
yrs ago but this group quickly died out and left no
replaced by Cro Magnon (except H. floresiensis)
descendents)

physically indistinguishable from us ancestors of this group eventually replaced all

earlier kinds of humans including the
less massively built than Neanderthals
neanderthals
made far better tools: knives, awls, chisels,
engravers, etc all “non-Africans” are descendants of this small
band
could make spears, harpoons, fishhooks, needles
they reached Australia ~50,000 years ago where
170,000 yrs ago: humans and our “cousins” began
they became genetically isolated from other
wearing clothing
humans and became the aborigines

Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 13 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 14

~40,000 years ago humans pushed to central Social Evolution

Asia and to Japan and Siberia
after Homo sapiens emerged about 200,000 years
genetic evidence suggests that humans moved earlier, it took another 140,000 years before
into the Americas 20,000-15,000 years ago any sign of modern civilisation emerged

during this time groups in some areas became up until this time biological factors controlled our
relatively isolated as agriculture and village life evolutionary development
caused our species to become more settled
about 65,000 years ago the biological structure of our
! genetic differences between races brain stopped changing
arose during this time
we began to interact more as groups; families,
eg. black skin evolved as protection against UV radiation of sun clans, villages
in warm tropical areas

[Biologically speaking, Race is a genetically distinct variation human interactions allowed us to learn and do
within a particular species of animal or plant. What we consider
things we could not do alone
“racial differences” among humans do not fit this criterion;
genetically the differences between human races are not
significantly different from the genetic variation among individuals] we could teach these things to our kids; pass on
our knowledge
by end of Paleolithic (~10,000BCE):
very few examples of this in other primates and other
!only a few places had not yet been settled by animals
humans
from then on ‘social evolution’ supersceeded biological
evolution in learning to interact with the world and
!human population was ~ 600,000,000
solve problems

Agricultural Revolution (10,000BCE)

Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 15 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 16
 Some Milestones in Human Biological & Cultural Evolution
65 MY ago: earliest primates appear (diversification after dinosaur extinctions)

a pivotal moment in our social development came with 20 MY ago: earliest examples of Hominidae (great apes)

the agricultural revolution 7 MY ago: beginning of Hominid line that lead to us

4.4 MY ago: earliest fossils of this line (“Ardi”)

led to a more settled existence 4.2 MY ago: first bipedal primate Australopithecus
2.5 MY ago: Homo habilis; earliest fossils of genus Homo

less time spent on survival 1.8 MY ago: first stone projectile weapons
1.5 MY ago: Homo erectus; dramatic increase in brain size, 1st migration out of Africa
more time spent on higher brain functions 1.2 MY ago: shed fur to become “naked ape”
1 MY ago: first evidence of controlled fire use
600,000 yrs ago: first neanderthals, a very closely related european species - could even interbreed
with H. sapiens that appeared later
500,000 yrs ago: first evidence of construction of shelters
200,000 yr ago: first modern humans, Homo sapiens, appear
140,000 yrs ago: earliest evidence of long-distance trading
100,000 yrs ago: earliest known use of jewelry, body paint and water containers
72,000 yrs ago: humans began wearing clothes – probably animal skins
60,000 yrs ago: oldest known needle – animal skins gave way to manufactured clothing
60,000 yrs ago: second migration out of Africa; all non-africans are descendants of this small group
50,000 yrs ago: oldest indication of ritualistic burial
35,000 yrs ago: oldest cave art
20,000 yrs ago: “racial” differences evolved as small groups of humans became more isolated and
more settled
10,000 yrs ago: origin of agriculture, 1st known lunar calendar; the entire human population was
about 600,000,000
8,000 yrs ago; plough invented
5,000 yrs ago: earliest writing, simple sewage systems to dispose of wastewater
2,600 yrs ago: oldest written legal code
2,300 yrs ago: birth of science (Aristotle)
250 yrs ago: the Industrial Revolution began
450 yrs ago: origin of the germ theory of disease

Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 17 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 18

The Scars of Human Evolution distributed along all of vertebral column

all organs are piled on top of each other
Bipedalism ! hernias

various evolutionary changes, esp bipedalism, resulted 5. the weight of our upper body is borne by
in a “terrific mechanical imbalance” in body pelvis and its junction with vertebral
column =sacroiliac
we are born with a backbone that has the ancestral ! lower back pain
curve of a 4 legged creature
6. also, wider distance between rib cage and
pelvis
! less protection for lower abdominal
organs
old curve
retained
7. also, harder to pump blood from feet back up
to the heart which is 4 ft off ground
! varicose veins, hemorrhoids
as we become bipedal
8. much bigger burden on our feet
1. stress points shifted from more evenly ! fallen arches
distributed 4 legged posture bunions
callouses
2. vertebrae had to become more moveable to
facilitate all these new curves Head
! weakens the back ! sore back
humans are the only living animals that are
3. the whole pelvis was tilted upward habitually bipedal
! narrow birth canal
head is balanced on top of vertebral column
4. rather than viscera hanging evenly and weight foramen magnum moved to inferior location
Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 19 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 20
 Relics of Human Evolution
face became flatter
eg. birds lost their teeth 70 million years ago but still have the
gene to produce them
don’t need strong neck muscles as does a 4 footed
animal ! brow ridges became small 1. The Appendix

Brain in plant-eating vertebrates the appendix is large and contains

lots of microorganisms to help in the digestion of food

1. in skull, brain case has expanded greatly modern humans eat mostly meat and no longer have a need for
! larger face, smaller snout such structure

2. while face bones have decreased in size 2. Wisdom Teeth

our teeth have remained large as our face lost its snout the mouth had less room for teeth
! impacted wisdom teeth
however there may have been an early advantage since our
3. big head, small birth canal early ancestors probably frequently lost jaw teeth and the
wisdom teeth would serve as useful replacements
! difficult & painful births
3. Tailbone
small fused vertebrae are only a vestige of the tail of our primate
ancestors.

Occasionally today a baby is born with a tail

there are > 100 medical reports of babies born

with tails,
sometimes tiny vestigial structures,
some have vertebrae and can even move

If that had happened in the dark ages the child and his mom
would have been burned as witches

4. Extrinsic ear muscles

Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 21 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 22

Atavisms
three muscles allowed ancestors to move ears as in dogs
and rabbits.
less clear possible remnants of human evolution
we still have them and some learn to use them
1. large canines (possibly)
5. Arrector Pili Muscles some have large canines
Darwin viewed them as an atavism
small bands of muscles attached to our hair follicles fossil evidence indicates that our canines began shrinking as
soon as we diverged from chimp ancestors
in other mammals it helps them to fluff up their fur against the
cold 2. extra nipples or breasts (yes when on
in humans, who have shed most of their fur, they just cause milkline)
goose bumps most mammal have multiple pairs of nipples
most primates have only 1 pair
1 in 20 people have at least 1 extra nipple
they usually occur along the “milk lines”

3. hand walking (no)

five Turkish siblings walk on all 4’s
gait resembled apes
not an atavism; is brain damage affecting their balance

4. undeveloped thumb (unclear)

some are born with short thumbs lacking some of the usual
muscles
a few also have long slender fingers
in some it leads to lack of fully opposable thumb

5. lack of sweat glands (probably not)

some people lack sweat glands
but most cases seem to involved mutations that block sweat
gland formation altogether whereas in apes there are
sweat glands ins the palms and soles

6. polydactyly (probably)
extra fingers or toes are one of the most common
Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 23 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 24
 congenital defects in humans and other animals
seems to be controlled by two genes during development
7. syndactyly (yes)
webbed fingers or toes are a common birth defect
probably the result of the self destruct mechanism being
disrupted to cells between are not destroyed
development is very similar to development of fish fins
except for the destruction of the web later in
development
8. Branchial cleft cyst (perhaps)
in 4th week of development vertebrate embryos develop
five ridges in the neck region
in fish, these ridges go on to form gills
in humans they form various structures in head or neck
the clefts between ridges normally disappear but sometimes
a fluid filled sac called a branchial cleft cyst remains
9. twinning (unclear)
some think it’s a throwback to large litters of our early
mammalian ancestors
some women have tendency to have nonidentical twins
10. hiccups (perhaps)
may be a primitive reflex inherited from our land
conquering ancestors
these primitive air breathers had both gills and lungs
the pattern of muscle movements in hiccups resemble those
seen when such animals close their glottis to stop
water getting into the lungs and contract the mouth
cavity to pump water over the gills
11. various muscles that were once useful in our
quadrupedal ancestors and are greatly reduced or absent in
many humans today:
Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 25
Do Humans Continue to Evolve?
evolutionary changes in the past 100,000 yrs have
been minor
eg jaws, teeth, size, toes, etc
the brain and other major things remain essentially
unchanged
structurally, the human form is generalist
= highly adaptable
the most dramatic structural changes in past 10,000
years are mainly due to better nutrition and health
humans are not dependent solely on physical structure
for adaptability anymore
! we are tool users and social animals
our tools have evolved greatly in the past few 1000
yrs
! tools are an extension of us
“the speed of man’s development is equal to the speed
with which new tools can be invented and made”
-Lewis (Man & Evol)
also, our social structure is an outgrowth of
Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 27
subclavius muscle – small muscly stretching from rib to
collar bone would have been more useful if we walked
on all 4’s. Missing in some people
palmaris muscle – long narrow muscle from elbow to wrist
may once have been important in hanging and
climbing. Today is missing in 11% of humans
plantaris muscle – useful for other primates for grasping
with their feet. Today missing in 9% of the population
12. 5th toe
apes use all their toes for grasping or clinging to branches.
Humans use mainly the big toe for balance. The
smallest toe is almost nonfunctional now
13. The thirteenth rib
chimps and gorillas have an extra set of ribs. 8% of adults
have this extra set but most of us have only 12 pairs
14. Paranasal sinuses
the sinuses of our early ancestors were lined with odor
receptors that gave a heightened sense of smell.
Humans have retained these sinuses but lost the smell
receptors that lined them
Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 26
technological development
language, knowledge, culture exist outside of the
physical body
tools and society are subjected to different
evolutionary processes
New (2007) research on the human genome indicates
that, genetically, humans are evolving 30-40x’s
faster today than 40,000 years ago.
in terms of physical changes
!those who say humans are no longer evolving
sometimes confuse evolution with
speciation
evolution is a change in gene frequencies
over time
speciation occurs when there is so much change
that members of the new gene pool can no longer
interbreed with the original gene pool
the human gene pool is always changing
! it will never “stagnate”
but there may never be a new human species
! this would require isolation of a population
Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 28
humans haven’t really changed the rules of Natural Is Man becoming less well adapted to nature?
Selection
the physical environment has become far less
! we are as much a product of evolution as severe a selective factor than in any other animal
any other organism
behavior is a strong selective pressure in the
culture and technology might change the kinds of animal kingdom
genes in the human gene pool but they cannot ability to learn
remove the force of evolution to overcome obstacles
parental care and protection
we can now measure evolution at the molecular scale:
this behavioral shift initiates new selective pressures:
eg. new evidence suggests that two key brain-
building genes which were critical in evolution to modern
humans are still evolving and spreading rapidly throughout 1. we are constantly finding more genetic links to
the human population behaviors and psychological factors
dramatic changes in the past coincided with huge leaps in
human intellectual development 2. the conquest of disease and mitigation of the
effects of aging have achieved spectacular
not everyone has these genes but the genes are increasing results
in the human population

one gene, microcephalin, began its spread among human ! new medical procedures may prevent
ancestors about 37,000 years ago weeding out of physically defective
humans
! a period marked by a creative explosion in music,
art, religious expression and tool making
eg. genetic diseases
the other gene, ASPM, arose only about 5,800 yrs ago
such things actually counteract natural selection
!about the time of emergence of major civilizations in
the mid east ! may produce an increase in deleterious
phenotypes

Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 29 Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 30

“are we sowing the seeds of our own species’ doom?”

while modern humans have largely mastered their

environment

and communication and interactions are tending to

normalize our gene pool

still ~50% of human race live in poor conditions,

shortages of food & water, disease is common,
poor health

! still very subject to natural selection

Genetics & Evolution: Human Evolution, Ziser Lecture Notes, 2014.11 31

 Phylum Chordata - Vertebrata
Mammals
relatively small group

5,100 species

~half # of birds; ~1/5th # of fish species

today, is one of most successful group of vertebrates

Cenozoic = age of mammals

occupy every major habitat on earth

most massive of all animals today or that ever existed

blue whale  105’, to 150 tons

blue whale, Balaenoptera musculus,

Mature blue whales typically measure anywhere from 75

feet (23 m) to 100 feet (30.5 m) from head to tail

and can weigh as much as 150 tons (136 metric tons).

The largest blue whale on record is a 110’ female that

weighed 195 tons (177 tonnes).

their bulk is several times greater than the

largest dinosaur

elephants are largest land mammal

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 1
 11’ tall, 14,500 lbs (=6,590 kg)

smallest mammals:

pygmy shrew  ~0.1oz (4 cm, few grams)

Kitti hognosed bat  0.05 oz (1.5 g)

mammals are also the vertebrate group most affected

by human activities:

domestication
food
clothing
beasts of burden
pets
research
education
hunting
alien animals
pleistocene extinctions
modern extinctions

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 2

Origin of Mammals

mammals developed from mammal-like reptiles

(therapsids; from synapsids) ~200 MY ago

mammal-like reptiles share many skeletal features

with mammals
legs held closer to body

not splayed out as in most reptiles

faster more agile

decreased stability due to new gait lead to increased

development of cerebellum for muscle coordination

secondary palate separates nasal and mouth cavities

teeth differentiated into incisors, canines, cheek teeth

many reptiles had hair (none today do)

some were warm blooded

but:
mammals have lower jaw consisting of a single bone with
articulation between jaw and squamosal bone

mammal-like reptiles lack this articulation

mammals first appeared 200 MY ago

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 3
  early Mesozoic
(same time as 1st dinosaurs)

1st mammal:
very closely resembled their reptile ancestors
about size of mouse (or ground shrew)
reptilian skeleton
had sharp teeth
 ate insects, worms, fruits, vegetables
large eyes
probably nocturnal
warm blooded
(many reptiles were warm blooded then)

for 160 MY they lived in the shadow of the dinosaurs

most were small, shrew-like animals, largest were 3’

long.

“suddenly” the dinosaurs disappeared ~65 MY ago

when dinosaurs vanished near beginning of Cenozoic

mammals diversity greatly increased

the first modern (placental) mammals appeared

and diversified
the most successful group, placental mammals, originated
and diversified

mammals were now agile, more intelligent, warm blooded,

well insulated, and suckled their young

moved into habitats vacated by dinosaurs and became the

dominant “large animals” on land and in sea

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 4

  Dawn of Cenozoic = “age of mammals”

may have been 2 separate lines of mammal origin & evolution:

prototheria  egg laying mammals; poor

thermoregulation

theria  all others

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 5

 Skin & Derivatives

skin is thicker and more complex than in other

vertebrate groups (or any other animal)

thicker layers esp dermis (hide=leather)

many different glands

sensory structures

hair

a. Hair

hair (fur) replaced scales as main covering for

better insulation

some facial hairs (=vibrissae) became sensory in

function

today, especially characteristic of mammals

in past, some reptiles had fur and/or feathers

grows from follicle in epidermis and dermis

cells at base of follicle produce hair

when it reaches a certain length,it stops growing

made of keratin (protein)

 same as nails, claws, hooves, feathers
of birds and scales of reptiles and birds
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 6
 most mammals have two kinds of hair:

underhair
dense and soft for insulation

 traps layers of insulating air

in aquatic mammals (fur seal, otter, beaver)

its almost impossible to wet skin

guard hair
coarse and long

protection against wear

coloration

some also have hair modifications:

a. defensive hairs
eg. porcupines, hedgehogs

b. horny or bony plates

eg. armadillo, pangolins

c. some have lost most of their fur

eg. hippos, elephants, porpois, humans

shedding (molting)

in most mammals entire coat is periodically

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 7
 molted
eg. foxes and seals  1x/yr

eg. most mammals have 2 annual molts

spring  replaced by thinner hairs

fall  replaced by thicker hairs

in humans hair is shed and replaced

continuously throughout life

coloration of hair:

camoflage
protective camoflage:

eg. arctic  white

eg. outside arctic  somber colors

disruptive camoflage

eg. leopard spots

eg. tiger stripes
eg. fawn spots

warning
eg. skunk

modified hairs:

bristles of hogs

spines of porcupines
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 8
 vibrissae (whiskers)  tactile, sensory hairs

b. horns and antlers:

horns or antlers are found in only 5 families

of ungulates:
Rhinoceri
cattle, sheep, goats, etc
pronghorns
moose, caribou, elk, deer

[virtually all even toed ungulates with ruminant (4-

chambered) stomachs have horns or antlers]

1. horns
esp cattle, sheep, goats, rhinos, etc

unbranched

horns originated early, >100M yrs ago, in large reptiles

hollow sheaths of keratinized epidermis

(same as hair, scales, feathers, claws, nails, hooves)

surrounds bony core

grow continuously throughout life

not normally shed; do not regenerate of cut off

usually used as a weapon for protection

eg. only pronghorn seasonally sheds its horns

eg. rhino horn has been collected for >1000 yrs

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 9
 2. antlers
esp deer, caribou, moose, elk

antlers originated in mammals ~50M years after horns

entirely bone, no keratinized layer covering it

tend to be large complex and ornate

used mainly for sexual display during mating season

sometimes require a significant investment in resources

to grow them (esp. large amounts of minerals)

eg. moose or elk need 50lbs of Calcium/season to

grow them

eg. antlers of irish elk weighed more than the rest

of its skeleton; 3 M across, 154 lbs

living tissue; sensitive to touch and pain

develop beneath highly vascularized sheath= velvet

velvet dropped off after breeding season

3. Giraff Horns

antler-like but retain skin covering; are not shed

c. Glands

mammals have a great variety of skin glands

the glands confer some of the most important

mammalian traits
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 10
 1. sweat glands (ecrine glands)
important in warmbloodedness; temperature control

esp on hairless regions; eg foot pads

simple, tubular, highly coiled
only mammals have sweat glands
heat regulation
part excretory organ

2. scent glands & apocrine glands

 smell important in most mammal social behaviors

almost all mammals, inc humans

their location and function vary greatly
used for communication:
territory
warning
defense
mating

3. oil (sebaceous) glands

associated with hair follicles
used to keep skin and hair pliable and waterproof

4. mammary glands
parental care; secrete milk

all mammals feed their young milk

all females; rudimentary in males
probably modified sweat glands
in all female mammals
rudimentary in males

Skeleton & Support

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 11
each species of mammal has a typical adult size
 skeleton doesn’t keep growing throughout
life as in amphibians and reptiles

4 limbs for locomotion (=tetrapods)

 pectoral & pelvic appendages

most have fore and hind limbs similar

limbs are up under the body

not sprawled out to the sides

much more efficient movement than other

land animals

many mammals walk more on their toes

 greater speed for both predators and prey

often smaller mammals can move at same speed

as larger mammals
eg. horse vs greyhound

(but larger need more powerful limbs and muscles)

predators tend to have retractable claws

Muscles & Movement

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 12
mammals display a wide variety of movements other
than walking and running

typically require modifications of bones of the

appendages

hopping
provides sudden bursts of speed and quick changes of
direction

at high speeds, the metabolic act of hopping is

much lower than that of running on all 4’s

eg kangaroo

brachiation
tree life

arms longer than legs

eg. primates

burrowing
limbs are short and powerful

eg. badgers, marmots, moles

have very large ears to pick up sounds

flying
only bats

moved into niche largely unoccupied by birds

 night flying
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 13
 for wing, skin is stretched between elongated fingers and
attached to legs and tail

beats up to 20x’s/second

use echolocation to avoid objects and find prey

emit high frequency sound waves that bounce off

objects and return

 can detect distance from objects

bats generally have large ears to pick up sound

a few bats don’t use echolocation

large eyes & good sense of smell

feed on fruits & nectar

some bats migrate up to 500 miles annually

gliding
generally nocturnal

can travel 40-50 M at a time

“flying” squirrels, some marsupials, lemurs

Feeding and Digestion

mammals are warmblooded and much more active

than reptiles

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 14

 require 10-13 x’s the food energy as living
reptiles and amphibians to maintain the same
body mass

teeth

all but a few mammals have teeth

eg. monotremes, anteaters, some whales

earliest mammals were insectivores

teeth were all the same peglike shape

today, teeth represent the greatest evolutionary

diversification of the mammalian skeleton

teeth more than any other physical characteristic

reveal the life habit of a mammal
all major mammal groups can be identified from a single
molar
 often even to species

mammals typically have 2 sets; milk teeth &

permanent teeth
most other vertebrates continuously replace teeth as
needed and their teeth continue to grow throughout
life

rather than all teeth in the mouth being the same,

mammals developed 4 major kinds of teeth
incisors  snipping and biting
canines  piercing and holding
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 15
 premolars  shearing and slicing
molars  crushing and chewing

each set of teeth then became specialized for

different lifestyles and feeding types

Digestive Tract (GI tract)

the digestive system may also be modified in

various ways determined by their diet:

a. herbivores (horses, deer, antelope, cattle, sheep,

goats, many rodents, rabbits and hares)

canines reduced or absent

large flattened grinding teeth (molars)

require lots of plant food for nutrition since

most of it is “indigestible”
eg. elephant = 4 tons eats 300-400 lbs/day

often have symbiotic bacteria and

microorganisms that can produce enzymes
to digest plant material

long large digestive tract

large caecum and stomach

eg. ruminants (cattle, bison, goats, sheep,
deer, giraffe)

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 16

 food is bitten off and swallowed, at first - not
chewed,

food is periodically regurgitated and chewed the

reswallowed

have 4 chambered stomachs with 1 way valves

rumen  bacterial action  cud
reticulum  fermentation
omasum  water soluble foods absorbed
abomasum  pepsin & HCl

rumen: >30B bacteria/oz of rumen material and

1000-100M protists/oz

forage remains in rumen up to 36 hrs

omasum: undigested portion passes to omasum

via reticulum

omasum is a holding chamber  soluble

nutrients are absorbed and large
particles are prevented from proceeding
further

abomasum: after screening and absorption food

passes to abomasums  functions as our
stomach: secretes pepsin and Hcl; also
digests microbes that accompahy food

eg. rabbits and many rodents eat their fecal pellets

giving food a second pass through the digestive
system

b. carnivores (foxes, dogs, weasels, wolverines, cats, etc)

biting and piercing teeth

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 17

 long sharp canines and incisors

powerful claws and limbs

much shorter digestive tract

smaller or no caecum (part of lg intestine)

c. omnivores (pigs, raccoons, many rodents, bears, most

primates including us)

teeth lack extreme adaptations of herbivores

and carnivores

d. insectivores (moles, shrews, anteaters, bats)

eat larval and adult insects

some mammals store food for winter

 collect nuts, seeds, fungi, etc

eg. all tree squirrels, chipmunks, gophers

eg. some mice

the amount of food a mammal must consume is

inversely proportional to its size

generally smaller animals need more food per

gram body weight than do larger animals

eg. a 3 g mouse consumes 5x’s more food per gram body

weight than a 10 kg dog
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 18
 and 30x’s more than a 5000kg elephant

eg. small shrews, bats and mice must spend much more
time hunting and eating than large mamals

eg. a shrew must consume its weight in food each day; it

will starve to death in a few hours if it stops feeding

eg. large carnivores can easily survive on 1 meal every few

days

eg. average (100 ton) blue whale requires ~ 2 tons a krill

(2% body wt) daily for sustenance

Respiration

mammals are warm blooded (endothermic &

homeothermic) and therefore have a relatively
high metabolism and therefore a high oxygen
demand

all mammals have lungs and breath air

whether terrestrial or aquatic

lungs are very efficient, second only to birds

 contain alveoli  blind ended sacs surrounded

by capillaries

 provide much greater surface area for gas

exchange
eg. humans: 760 sq ft (~tennis court)
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 19
mammals also have a muscular diaphragm which
“sucks” air into the lungs
much more efficient than gulping air or expanding chest
cavity

Circulation

mammals have 4 chambered heart

 completely separates the two circuits of blood

flow

smaller mammals with higher metabolism have faster

heartrate
eg. shrews heart beats 760 times/minute (10 x’s ours)

Nervous System

relatively large, highly developed brain

 disproportionately larger per body weight

cerebrum most highly developed in mammals

unique “neocortex” that is folded to increase

surface are

many more interconnections

 more intelligent
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 20
  capable of complex social behaviors

 behaviors based more on “learning”

cerebellum also relatively larger & folded

allows for more complex movements

Senses

a. smell

the sense of smell is particularly well developed in

most mammals

in most olfaction is more important than vision

new (2013) evidence indicates that an improved

sense of smell was the first stimulus for an
enlarged brain

smell is an integral part of feeding and social

activities for mammals

humans are unusual mammals since they have

largely lost their sense of smell
eg. dogs have 100 M – 300 M smell receptors in their
noses; humans have 6 M

a. vision
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 21
 vision and hearing well developed in most
mammals

moveable eyelids

vision especially good in daytime predators

b. hearing

fleshy external ears

hearing adaptations:

eg. bats - echolocation

sonar (echolocation) is the ability to emit sounds then

to detect their return to estimate distances from
objects

in general the higher the frequency of emitted sound,

the better echolocation works

sonar has evolved in several vertebrates; marine

mammals, a few birds and bats

bats seem to have perfected it

bats emit very high frequency sound waves

(human limit = 20,000Hz; bats 30-100,000Hz)

rate=10/sec to 200/sec if prey near

bats have very large ears to receive signals

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 22

 many insects have evolved defensive measures to
avoid bat predation

they have learned to detect the sonar signal of

bats and fly defensive patters to avoid them

some use sonar themselves to confuse bats

eg. whale communication

low frequency “songs” for communication

low frequencies travel well through water

can use explosive sounds to stun or kills fish

toothed whales have highly developed echolocation

not emitted by larynx; originates near blow hole

produce clicks that are focused by “melon” in forehead

return sounds perceived through lower jaw

some fish have adapted to detect and avoid them

c. touch

the sense of touch is also extremely well

developed in mammals

the earliest mammals were mainly nocturnal and

depended on touch for most social activities

eg. star nosed mole - feel their way through their burrows
with their noses

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 23

d. detection of earth’s magnetic field

bats have been found to be able to sense the

earth’s magnetic field like birds.

the only other mammals known to be able to

do this are the naked mole rat and
Siberian hamsters

Excretion

kidneys effectively filter blood to remove waste

products

 usually urea

kidneys also very effective at maintaining salt/water

balance

Protection and Defenses

mammals use a variety of methods to protect

themselves from predators:

some have hairs modified into relatively hard outer

“shell”
eg. armadillo

or sharp spines

eg. porcupine

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 24

others may play dead when approached by danger
eg. opossum

shrews are one of the few venomous mammals

can send a mouse into a coma (wont hurt us)

Reproduction

most mammals are dioecious with internal

fertilization

most mammals have definite mating season

usually winter or spring

usually limited by female estrous (ovulation; in heat)

female advertises receptivity by distinctive visual,

behavioral or pheromonal signals

a few mammals (eg some primates) females show no

obvious signs of ovulation and are receptive year
round
= concealed ovulation

Courtship Behaviors

In many, especially the larger mammals, courtship

begins active competitions between males to
demonstrate their strength and fittness to the
females
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 25
 sometimes it’s bluster; the animal with the loudest longest call
shows he is the most macho

sometimes it’s actual battles; eg elk; only in rare cases is one

hurt or killed, the challenger will usually back off before
then

of all mammals, Bonobo’s whole society revolves

around sex more than any other vertebrate (even
humans)

they use sex as greetings, for solving disputes,

making up after fights and as favors in
exchange for food

“Chimpanzees and Bonobos both evolved from the same

ancestor that gave rise to humans, and yet the Bonobo is
one of the most peaceful, unaggressive species of mammals
living on the earth today.

They have evolved ways to reduce violence that permeate

their entire society. They show us that the evolutionary
dance of violence is not inexorable.”

all but one small group of mammals are viviparous

(=bear live young)

mammals generally produce few young but devote

considerably parental care to insure their survival

it is rare for parental duties to be shared

usually the female does most of the work

nurse young with milk  mammary glands

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 26
some mammals (eg mongoose) give synchronous
birth:
a females in the group give birth on the same day as the
dominant female, regardless of when they became pregnant

subordinate mothers can slip their kids into the communal litter
where they will be safer

3 patterns of reproduction

characteristic of the three major mammal groups:

1. egg laying (arose ~140 MY ago)

monotremes

most ancient kind of mammals

produce thin leathery shell

 no pregnancy (gestation)

after hatching, young are fed milk

2. marsupials (arose ~130 MY ago)

brief gestation

then crawl to pouch and attach to nipple

= embryonic diapause

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 27

 3. Placental mammals (~65 MY ago)

most recent evolutionary strategy

relatively long gestation period

eg. mice  21 d
rabbits  30 d
cats/dogs  60 d
cattle 280 d
elephants  22 mo

Hibernation

eg. black bears

in winter they can lower their body temperature up to 5.5º C
(~15º F) and their total metabolic rate to only 25% of its
normal rate. Heartrate drops from 55bpm to 14 irratic
bpm’s

in spring their metabolism takes several weeks to return to

normal

Migration

migration is much more difficult for mammals than for

birds
walking requires much more energy than swimming or flying

Energy Consumption/kg body wt/km:

swimming: 0.39 kcal (salmon)
flying: 1.45 kcal (gull)
walking: 5.43 kcal (squirrel)

only a few mammal migrate

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 28
 most of these are in North America
eg. caribou

migrate 100-700 miles (160-1100 km) twice/yr

eg. plains bison

eg. seals

northern fur seals  1740 miles (2800 km)

eg. whales

humpback whales have the longest migration of any

mammal; over 5000 miles

migrate to warmer waters to raise their young

gray whales  11,250 miles (18000 km); twice/year

the oil with which they store energy makes them more
buoyant and poor heat conductor

eg. a few bats migrate

Mexican free tailed bats in Austin

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 29

 Classification of Mammals
when mammals began their explosive evolutionary
radiation 65-70 MY ago Eurasia, Greenland and
North America were still joined together and
shared a common fauna

Africa and Australia were isolated and developed

their own fauna

3 major mammal groups identified by the type of

gestation

also, mammals have developed a great variety of

teeth types for eating many different kinds of
foods

 individual teeth, esp molars can often

identify the species of mammal

1. Monotremes

(single hole or opening for reproductive, urinary

and digestive systems)

reptilian structure

horny birdlike beak for mouth

lost all traces of teeth

lay eggs
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 30
 in Australia only
eg. platypus
aquatic
muskrat habitats
false marsupium

eg. echidna
ant eater
has pouch for young

2. Marsupials
eg. possums, kangaroo, koala

very short gestation period

eg. opossum  12 days

abdominal pouch for rearing young

mainly Australia, Tasmania, New Zealand

diversified into niches taken by other kinds of

mammals in the rest of the world:
Tasmanian devil weasel
Tasmanian wolf wolf
Marsupial moles moles
bandicoot rabbit

opossum only marsupial in Americas

3. Placental Mammals (Eutherians)

most successful group

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 31
 increased reproductive investment
 relatively long gestation period
 lots of parental care

placental mammals are subdivided into orders

based on teeth and skeletal characteristics

Some Kinds of Placental Mammals:

a. insect eating mammals

usually small animals

eat mainly invertebrates, esp. insects

all Mesozoic Mammals (age of dinosaurs) were insect eating

earliest placental mammals were insect eating as well

eg. shrews, moles,

b. carnivores
require keen sensory perception

diet mainly of other vertebrates esp. rodents

at least 1 pr of specialized shearing or carnaissial teeth to slice

meat

most are medium sized animals

more highly evolved carnivores hunt in packs to get larger prey

compared to herbivores carnivores have a relatively few

anatomical and physiological specializations
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 32
 skeleton needs flexibility and strength

in running forms, feet are lightly built and only toes touch ground

most cats have retractable claws to keep them sharp

brain is usually relatively large & convolutions more complex

eyes give stereoscopic vision

ears have greater frequency range

olfactory senses well developed

eg. dogs, cats, bears,

c. Paddlers and Swimmers

total number of aquatic mammal species is large: 9 orders

aquatic invasion could never have been an easy option to

competition on land

heat retention is a major problem for aquatic mammals

 thermal conductivity of water is 25x’s that of air

 aquatic mammals grow large (less surface to volume

ratio-less heat loss)

some are carnivores some are herbivores

herbivores (vegetarians)

tend to be less specialized,

browse near shore, none cruise the ocean,

most are large

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 33
 relatively few in numbers

eg. hippos, sea cows

carnivores

often large breeding colonies

simplified dentition

some can dive very deep

eg. sea lion can dive to 200 meters

eg. seals, sea lions, otters, walruses,

d. Whales and Dolphins

largest animals that have ever lived

 blue whale: 30M long 130 tonnes

evolved from an aquatic four legged deer-like creature no larger

than a fox

whales are the most specialized of all mammals

completely adapted to aquatic life except for their need

for air

hind limb is absent

forelimbs are short, webbed
bones are filled with oil for floatation and energy reserve
thick fat layer
no sweat glands
large complex brain
communication skills second only to humans
poor smell, poor vision
acute touch and exceptional hearing
large tidal volume (10x’s humans)
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 34
 can extract 10% of O2 in air (humans use 4%)
average (100 ton) blue whale requires ~ 2 tons a krill
(2% body wt) daily for sustenance

there are freshwater whales in rivers of India, China and So.

America

deep diving cetaeans can last an hour or more on one breath:

have considerably more (up to 10 x’s more )muscle
myoglobin than in most land mammals
these muscles can work much longer anaerobically than in
most land mammals

hearing is probably more important in cetaceans than any of

their other senses

whale sonar is extremely precise & can apparently produce a

mental image as precise as vision can

e. gliders & fliers

animals can move through air in 3 ways:
parachute, glide, or fly

gliding: 3 orders - marsupials, lemurs, flying squirrels

flying: bats only

ancient writers: bats were peculiar birds; Pliney called them

winged mice; only birds with teeth and bore live young that
they suckled with milk

powered flight

echolocation used by insectivores, whales and bats

bats emit and pick up high frequency sounds

most bats 20-80 kHz

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 35

 sound is produced in larynx and emitted through mouth or
nose

returning sound is picked up by ears  ear often large

each bat species has distinctive signal

one nectar eating bat has tongue that extends over 1.5 x’s
its body length (only chameleons can top that)

f. gnawers
includes largest of orders =rodentia (40% of all mammal
species)

most are mouse to rat sized

but some fossil forms were much bigger

all have at least 1 pr of large, curved, continuously growing

incisors
 need to bite continuously to wear them down or
would be unable to close jaws

gap (or diastema) behind incisors

eg rodents, lagomorphs, a few extinct orders

eg. naked mole rats

only mammal that lives in colonies like the social insects
native to Ethiopia, Soimalia and Kenya
3.5”long and weigh 1-3 oz
pinkish or yellowish wrinkled skin look naked
live in well organized colonies of 20-30 individuals
only one pair breeds
the other males and females belong to 2 castes
small working casts: dig the burrows and carry food and
nesting material
larger working cast: spend most time in nest with breeding
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 36
 female; may help defend her and colony
large communal nest
has a large communal nest chamber
with smaller tunnels where animals forage for tubers,
roots and corms
all mole rats care for young but only breeding female
suckles them
once weaned, the juveniles join the owrker caste

g. rooters and browsers

1st great wave of vegetarians

medium to elephant sized creatures

only two groups survive today

feed on tubers
 food is mashed and ground by cheek teeth

eg. elephants
2nd incisor teeth become tusks for food gathering and
display
elephant head can weigh 1 ton

eg. conies & elephants

h. hoofed herbivores
even or odd toed; 2 orders perissodactyla and artiodactyla

gut is more complex than that of insectivores and carnivores

microorganisms in gut digest cellulose

many have horns or antlers for defense

domestication of wild animals is one of the greatest

achievements of mankind
 ~7000 BC
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 37
 eg. horses, rhinos, pigs, deer cattle, antelope,

i. primates
adaptations of primates are mainly anatomical trends related to
behaviors

free and precise hand and forelimb movements

shift from reliance on smell to vision leads to good spatial

perception

cerebral cortex increased in size and complexity

lengthening of prenatal and postnatal life required

prolonged care of dependent young and allowed time
for learning

overall primate evolution shows an increasing dependence on

intelligence as a way of life

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 38

Orders of Mammals

Sirenia (manatees)
first appeared ~50 MY ago
ancestors were terrestrial mammals that also gave rise to
elephants, hyraxes and possibly aardvarks
in past were exploited for food eg 1950’s 7000 were killed each
year in So America for food
graze on aquatic plants
much lower metabolism than other mammals their size
 can go 7 months without eating
reproduction: 1 calf every 2-3 years for 20 years

Seals & Sea lions

evolved ~25 MY ago from 2 different groups; mustelids and
ursids, respectively

Rodentia
40% of all living mammal species are in this order; largest
mammal order

Chiroptera (bats)
20% of all living mammal species are in this order; 2nd largest
mammal order

Insectivora
10% of all mammal species are in this order; 3rd largest mammal
order

Primates
earliest fossils ~65 MY ago

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 39

 Ecological Roles of Mammals
1. Mammals are important at most levels of the
food chain
eg. predatory mammals help to control population sizes of prey
species

eg. salmon-eating bears in northern forests play a significant role

in fertilizing northern forests with fish remains

eg. many mammals are scavengers and omnivores that help to

recycle dead plants and animals

2. Pollination

bat pollinated
mainly in tropics
strong odor
dull color
open only at night

mice, rats, shrews, gerbils

mainly in tropics

visit several of the low-to-the-ground species

“Rodents are attracted by a strong musty odour

they are rewarded with a syrupy sugar which is secreted in

large quantities.

In order to prevent birds and insects from stealing this

nectar, rodent-pollinated flowers are an inconspicuous
brown or black.

the flowers are usually hidden inside the bush at ground

level, where they are more accessible.

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 40

 the center of the flower may be pale white and the tips of
the flowers may be shiny red

 both serve to guide the rodent to the nectar

in the dark.

3. Seed Dispersal

seeds dispersed in edible fruits

attracts birds or mammals
may eat whole fruit or spit out pits
if swallowed seeds resistant to digestive juices
squirrels and birds bury fruits and seeds
nuts stored underground are forgotten

seeds passively carried by animals

hooks or spines to catch in fur or on skin
in mud on feet of birds, etc.
burs, beggars ticks, devils claw, etc.

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 41

 Human Impacts of Mammals
1. Domestication of Mammals

a. Agricultural Animals

by far the largest use of mammals is as food

3.3 billion cattle, pigs, sheep and goats worldwide

cattle: 1.4 Billion (42 M in US; 24% world use)

pigs: 1 Billion (97 M pigs in US)
sheep: 1 Billion (>4 M in US)
goats: 700 Million
rabbits: 450 Million
domestic buffalo: 162 Million

meat and milk, fiber production

694 Million tonnes of milk/yr globally

domestication began about the same time as

origin of agriculture
dogs might have been first animal domesticated

sheep were probably first domesticated farm animals

(~11,000 yrs ago)

cattle: domesticated ~8500 yrs ago; 1200 distinct

breeds

horses: ~5500 yrs ago horses were tamed

b. Pets

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 42

 >105 Million mammals are sold as pets in US
each year

cats 51 M
dogs 50 M (300,000-500,000 from puppy mills)
rabbits 1.4 M
hamsters 600,000
guinea pigs 400,000
gerbils 400,000

can improve physical and mental well being

provide companionship

especially effective for lonely and depressed

but: up to 20 Million cats and dogs are abandoned each
year to starve or be put to sleep

c. Service Animals
horses: 61 Million worldwide

donkeys: 43 Million worldwide

mules: 14 Million worldwide

camels: 19 Million worldwide

llamas & alpacas: 5.5 Million worldwide

seeing eye dogs, search and rescue

military --dolphins

2. Hunting

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 43

 140 Million wild animals are killed in the US/yr:

60 Million wild mammals are hunted & killed in the

US/yr for sport:
deer  3 M
rabbits  27 M
squirrels  32 M
bear, caribou, moose, antelope  250,000

for each animal killed at least 2 are wounded

hunting is also having an effect on marine

mammals:

while large scale whaling has decreased in the

last several decades the consumption of
small whales, dolphins and manatees is on
the rise in poor nations

largely due to the decline in coastal fish catch

and more unintentional kills as bycatch

3. Furskin production

the US is the world’s largest volume producer of

furskins derived from wild animals

about 30 M/yr mammals are hunted for their

fur

provide 85% of furskin production per year

worldwide
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 44
 there are ~150,000 licensed trappers in the US

over 50 million animals are raised in captivity for

their fur in the US each year

mink and fox are the most common furbearing

animals
eg. mink pelt production in the US was 2.6 million
pelts in 2002

4. Zoos

conservation and management of wildlife

breeding programs for endangered and threatened

species

education of general public to value and plight of

wildlife

but many problems in keeping animals in

unnatural captivity

4. Animal By-Products

Many uses of animal products are hidden:

eg. medicines, film, rubber, ceramics, plastics,
paint, perfumes, glue, explosives, cosmetics,
shaving cream all contain materials from slaughter
houses

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 45

 eg. cellophane  made with animal fats

eg. freon animal fats used to make it

eg. makers of synthetic fibers use tallow based products

to control static cling

eg. animal based lubricants are used in jet engines

 all flying miles are animal based

eg. used in corrosion inhibitors for oil pipelines

eg. cars manufacture alone:

galvanized steel body, fan belts, gaskets, anti freeze,
hydraulic brake fluid, battery, steering wheel,
dashboard, tires

eg. animal fats and hides are even used in asphalt on the
roads the car drives on.

The animal by-products industry brings in over $2

Billion/yr
eg. 1000 lb steer:
432 lbs retail beef
568 lbs by products
27 lbs: variety meats; hearts, livers, brain,
tongue, kidneys
358 lbs hide, hair, bones, horns, hoofs, glands
and organs
46 lbs blood
183 lbs fat

eg. hide: $50-$75/hide

 US sends 90% of hides overseas for fabrication then

back to US for product sale

hide: clothing, insulation, ointment base, binder for

plaster and asphalt
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 46
 hair: toothbrush bristles, mattresses, air filters,
upholstery covers

hair from inside of cows ear

 “camel’s hair” paint brushes

eg. hides and connective tissues, cartilage, blood,

bones:

glue in plywood, paper matches, textiles,

cardboard, window shades

eg. bones, horns, hoofs: gelatin for photofilm and

pharmaceuticals (gelatin capsules)

cattle horns: imitation tortoise shell

hooves: white  imitation ivory

black  potassium cyanide  used to
extract gold from ore

eg. bones: electrical bushings, dice, chessmen, crochet

needles, piano keys, buttons, knife handles,

bone charcoal is used as refining material to purify

steel, filter sugar cane, manufacture high grade
steel ball bearings

eg. blood: dried and used in cattle, turkey and hog feeds;
pet food, fertilizers, clotting factors are extracted for
pharmaceuticals

eg. glands: >130 different medicines and pharmaceuticals

eg. tallow and lard:

(tallow hydrolysis glycerine and crude fatty acids stearic & oleic acids)

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 47

 glue, agricultural chemicals, candles, cosmetics,
detergents, drugs, metal castings, paints, inks, paper,
shaving cream

eg. if animal has gall stones

 in orient $1000-$2000/lb of gallstones used as

aphrodisiac

Ambergris one of the most rare and unusual

animal by-products
formed from the undigestible parts of >1 ton of squid eaten
per day by the whale

this undigestible mass occasionally damages the

intestinal lining and accumulates with impacted
feces to coalesces into a dense concretion

may be passed by the whale or may block the intestine

and kill the whale

when released the up to 1 ton “boulders” float in

seawater for decades and “matures”

has a complex, hard to describe, smell

rare and extremely valuable; up to $10,000/lb

has been used for centuries as an ingredient in fine

perfumes, medicines, incense and flavorings and, of
course, as an aphrodisiac

5. Education

most commonly dissected mammals are fetal

pigs and cats
fetal pigs: 500,000
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 48
 cats: >100,000

6. Research

probably one of the most contentious issues with

animal welfare advocates

Biomedical research is a major enterprise

>115 Million mammals are used in research

each year, worldwide

>70 Million/yr in US alone

__US___ Worldwide
/yr: rats & mice 60 M 115 M
guinea pigs 204,809
hamsters 167,571
dogs 87,000 140,016

most from shelters; ~40,000 bred for research

cats 21,637 35,004

most from shelters

primates 62,315

many from breeding colonies

pigs 57,000 175,020

rabbits 554,385 1,003,448
sheep 3,700

90% of research animals are rodents bred

specifically for research
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 49
 What kinds of research?

animals are used for both basic and applied

research

some argue that they should only be used for

applied research but you can’t separate the
two
diagnostics
1-5% of all lab animals are used to diagnose disease
eg. TB, diptheria, anthrax, burcellosis, etc

disease models:
eg. inbred mice for Hodgkins lymphoma
eg. primates for HIV

organ transplantation:
eg. tissue typing techniques, immunosuppression
drugs

bionics research

development of new drugs

determine treatment regimens, treatment regimens,

study of side effects, etc

surgical procedures:
eg. balloon angioplasty

extraction of medical products

eg. hormones, blood for culture media

production of antisera, antibodies, & vaccines:

eg. diptheria, whooping cough, tetanus, polio
eg. smallpox vaccine from skin of calves or sheep
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 50
 eg. rabbits as antisera factories

antibiotic testing

toxicity testing

food and water safety

search for new drugs

examples of specific mammals used in research:

mice & rats: 95% of all animal research are done on mice
and rats.

used in virtually every kind of scientific investigations

cows: narcolepsy, reproductive physiology, vaccine testing,

infectious disease research, heart studies

pigs: very important animal model for human physiological

studies; cardiovascular research, blood dynamics,
nutritional deficiencies, alcoholism & drug abuse,
general metabolism, digestive related disorders,
respiratory disease, diabetes, kidney and bladder
disease, organ toxicity studies, dermatology,
neurological studies, burn studies, cystic fibrosis
research

sheep: pregnancy related research, multiple sclerosis,

medical implant studies, burn and injury evaluation,
smoke inhalation

goats: studies in cartilage repair, respiratory physiology,

medical diagnostics, gene therapy, anesthetics
research, used to produce antibodies, and to produce
genetically engineered products

dogs: heart and lung research, transplantation

experiments, cancer research, microbiology, genetics,
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 51
 orthopedics, surgeries, vet medicine, toxicity studies of
drugs, additives and industrial chemicals

cats: neurological research, spinal cord injury, used to

study vision, sleep and hearing problems, Parkinsons
disease, cancer, genetic disorders, HIV/AIDS research

rabbits: toxicity testing for cosmetics and household

products; also used as models for eye diseases, skin,
heart and immune system studies, asthma research,
cystic fibrosis studies, diabetes and used to produce
antibodies for research and diagnosis

guinea pigs: toxicity & safety testing, effects of cigarette

smoke, alcohol and drugs, spinal cord injury
investigations, TB research, kidney function,
osteoarthritis research, nutrition and genetics studies,
reproductive biology and study of infectious diseases

hamsters: taste and vision research, cardiopulmonary

research, cancer and muscular dystrophy
investigations, studies of aging, asthma, and
biorhythms

7. Food and Crop Loss

rodents and rabbits cause “staggering” amounts of

damage to crops and stored food each year

8. Sickness & Disease

rodents & others carry diseases

eg. bubonic plague, typhus

eg. tularemia: reservoirs; rabbits, muskrats &

other rodents(vector=wood tick)

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 52

 eg. rocky mtn spotted fever: squirrels & dogs (ticks)

eg. lyme disease: deer (ticks)

9. Illegal Trade in mammal products

2006: 510 sp of mammals critically endangered

eg. Rhino horns

used in China to reduce fever & treat heart, liver and skin
disease

some breeds on brink of extinction

1970-1997: horns from 22350 rhinos were imported into

Yemen alone

10. Herbal Medicine

especially in China:
the skulls of fgazelles are ground into powder taken to
improve strength

gallstones of bulls are highly valued as a treatment for

fevers and inflammation

elephant skin is taken for acne

monkey heads are eaten for headaches

11. Bycatch

dolphins bycatch of Tuna fisheries:

115,000 US/yr

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 53

12. Pollution

cattle lots, hog farms

13. Tourism, Wildlife Photography, Art

There is a wildlife refuge in every state and within an hour's

drive of most American cities

More than 35 million people visit refuges annually, generating

nearly $1.7 billion for local economies and supporting
almost 27,000 private sector jobs

14. Entertainment

eg. circuses, rodeos, movies, horse racing, dog

racing, dog fights

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 54

 Animal Welfare
more than any other group of animals, mammals are
most closely associated with “animal welfare”
concerns

the original phrase used was “animal rights” but most

(not all) now agree that the legal connotations of
that phrase are not possible

animals can’t have “rights”

 implies ability of animals to reason with humans

and agree on mutually accepted principles

 implies lives of all animals, including humans,

are equal

 implies that it is unethical to use animals as

pets or for any other purpose
eg. food, clothing, recreation, education, research

eg. pets = form of slavery

eg. killing rats is murder punishable by execution

Animal Welfare requires that:

 any use of animals should be motivated by

humanitarian goals

 we are obligated to minimize pain

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 55
  we are required to show accountability for our
actions

there are many animal welfare movements

need to define “animal”

warm blooded vs cold blooded

vertebrates vs invertebrates

does a sponge or an earthworm deserve the same

consideration as a primate?

if so, why draw the line at animals

what about protozoa, fungi, plants, bacteria?

avoiding all contact with animal products is virtually

impossible

many uses of animal products are hidden:

eg. medicines, film, rubber, ceramics, plastics,
paint, perfumes, glue, explosives, cosmetics, shaving
cream all contain materials from slaughter houses

eg. cellophane  made with animal fats

eg. freon animal fats used to make it

eg. makers of synthetic fibers use tallow based products to

control static cling

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 56

 eg. animal based lubricants are used in jet engines
 all flying miles are animal based

eg. used in corrosion inhibitors for oil pipelines

eg. cars manufacture alone:

galvanized steel body, fan belts, gaskets, anti freeze,
hydraulic brake fluid, battery, steering wheel, dashboard,
tires

animal fats and hides are even used in asphalt on the

roads the car drives on.

Our extent of animal products “exposure”:

foods ~75%

clothing ~10-20%

soaps & cosmetics ~5-10%

scientific research using animals is probably one of

the most contentious issues of “animal welfare”

What is the value of animal research?

a. some of this information cannot be learned any

other way

 its unethical to test surgeries or drugs in

humans 1st (=human rights issue)

 can set up controlled experiments that you

cannot do with humans
eg. genetically identical pairs
eg. exact feeding regimes
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 57
 b. many surgical and medical procedures used in
research had spinoffs in veterinary sciences
pets, livestock, zoo animals generally live longer, are
healthier and live more comfortably because of animal
experimentation

c. animal experimentation has helped to preserve

endangered species :
treat illnesses,
eliminate parasites,
promote breeding (eg. artificial insemination, embryo
transfer, capitve breeding)

Criticisms of animal researchers:

a. inadequate self regulation

standards of care been dramatically improved; they were
slow in coming

biomedical research has always been closely regulated

but really are not many inspections done

b. slow to replace animal models with alternatives

few incentives to change even when alternatives are

available

c. tend to point fingers in other directions

it’s the other groups, not us, who are mistreating animals

Criticisms of animal “rights” activists:

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 58

 a. oversimplistic generalizations, loose thinking
eg. animal testing compared to Nazi legacy of human
abuses for “research”

but:
ironic that animal research was almost banned in Nazi
Germany before the war

b. misstatements, misrepresentation of the

problem
eg. development of polio vaccine cost 2 M monkeys and
didn’t reduce polio rate from 1916 to 1962

but:
polio research only started in 1953
by ‘70’s polio rate dropped to near 0 in US

eg. thalidomide is touted as drug that got through animal

testing and still proved dangerous

also has been stated that many tests were performed

on pregnant animals

but:
actually, didn’t get enough animal testing
no pregnant animals were used in research

eg. some believe that all animals suffer agony at some

stage of research

tout statistic that 80% of experiments are done

without anesthetic

but:
most didn’t require any, there was no pain
involved

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 59

 c. some antivivisectionists tactics result in more
pain and mistreatment than the research they
oppose

eg. “freeing” lab animals

most will be hunted and killed by wild animals

eg. one group was charged with animal abuse for keeping
over 200 dogs on a 1 acre enclosure to prevent their
use in medical research

d. the “animal rights” movement has:

driven up the cost of research
more money spent on tighter security and to repair
damaged facilities

may slow development of therapies and

treatments
reduces the amount of research being done

some research must be started over when facility is

damaged or animals released

Are there alternatives?

other methods are often cheaper and require less

paperwork:

 scientists tend to use them whenever they can

animals are used only when it is the best way

to get the appropriate information

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 60

 eg. some aspects of the causation, treatment or prevention
of blindness cannot be studied in bacteria, fungi or
plants  need complex animals

eg. high blood pressure cannot be studied in invertebrates

still, there is an effort to find alternatives when

possible
eg. many toxicity tests are done using cell or tissue cultures
now

eg. new chemical and mechanical simulations can provide

valuable information about how a tissue or organ
will react to certain medications

eg. we are beginning to develop the first realistic software

models of human and animal organs that can show
thousands of molecular interactions & can manipulate
physiological processes

however, most researchers hold that these non-animal

techniques cannot completely replace animals:

Pain

probably one of the biggest concern is causing pain to

animals

most animals are capable of experiencing pain

 generally scientists acknowledge and accept that all
warmblooded animals and most coldblooded vertebrates
(frogs, fish, etc) experience pain

even though experiencing pain, many animals may not

show any external signs of pain
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 61
 animals that show distress in nature might attract a predator

eg. recent evidence has shown that even fish have pain
receptors and experience pain when caught on fishing line

one simple test for pain:

“a stimulus is said to be painful if it is consistently

terminated or escaped by subject”

animals tend to begin to escape pain sensations at

about the same intensity that humans begin to
report pain

Most animals experience only minimal pain in research

settings:

eg. Animal Welfare Enforcement Report (1988)

94% of all lab animals are not exposed to

painful procedures or given drugs to relieve any pain

6% are exposed to painful procedures which

are usually not severe or long lasting

eg. Biomedical Research Study (1989)

58% experienced no pain, received no pain medication

35% received anaesthesia  little or no pain

7% experienced significant pain

eg. there are safeguards to insure animals for research are

well cared for:

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 62

  unhealthy animals can lead to erroneous results

 animal research is expensive; can only afford high

quality research

 pain can invalidate an experiment because stress

induces physiological changes in virtually all body
systems

some kinds of research subject animals to

considerable pain:

eg. orthodontic research

eg. car crash studies

originally used human cadavers, but their use was
banned

eg. oral radiation research

subject animals to enough radiation to cause death

eg. tumor therapies

there are also cases of:

inadequate use of anaesthesia and

inadequate care of laboratory animals

Additional perspective on animal welfare:

 1000x’s more mammals are killed for food than used for
research

 for every dog or cat used in research ~100 are killed at

shelters and pounds
Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 63
  many more pets and farm animals are neutered, some without
anaesthetic, than are subjected to experimental surgery for
research

 about half of the biomedical research carried out in US

would not have been possible without lab animals

 about 2/3rd’s of projects that led to Nobel Prizes in

Physiology and Medicine used animal experimentation

 habitat destruction kills many millions more animals, and

whole species are lost, yet this is NOT a major issue with
“animal welfare” advocates

Animals: Phylum Chordata-Vertebrata; Mammals; Ziser Lecture Notes, 2015.11 64

 Phylum Chordata - Vertebrata
Birds
of all higher vertebrates, birds are probably best
known

9700 species

 2nd most abundant vertebrate group

 outnumber all other vertebrate groups except fish

smallest bird: bee hummingbird 1.8g (.06oz)

one of smallest warmblooded vertebrates

only slightly larger than a bumblebee

largest living flying bird:

is the wandering albatross with a 12 ft wingspan; weighs about
25 lbs

largest known flying bird:

a condor-like bird, ~6 MY ago

155 lbs (70 kg) with 21’ (6.4M) wingspan

had to run downhill into a headwind to take off

largest bird: elephant bird of Madagascar is most

massive bird that ever lived

2 M tall, 450kg(~1000 lbs)

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 1

 also: tallest was extinct moas of New Zealand
flightless bird, related to emus

12 ft (3.6 M) to 550 lbs (250kg)

birds are found in all habitats:

forests, deserts, mountains, praries, oceans

some live in caves in total darkness

some can dive to 140’ to capture prey

birds are even found at the north and south poles

Origin of Birds

for over 50 MY amphibians and reptiles were the sole

terrestrial vertebrates

before the Permian extinction, 250 MY ago, a branch

of reptiles leading to birds an mammals arose

fossil evidence indicates that true birds evolved

from feathered velociraptors (of Jurassic Park
fame)

birds clearly evolved from dinosaurs

more similar to dinosaurs than dinosaurs are

to turtles, snakes and lizards

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 2

earliest fossil of a true bird Archaeopteryx
(=ancient wing) 150 MY ago

first fossil discovered in 1861 –2 yrs after Darwin’s

origin of species

rare find since delicate bones and feathers

don’t fossilize well

if not for impressions of feathers would be

classified as a small dinosaur

following the rules of taxonomy they should be

in same class (reptilia)

Archaeopteryx (=ancient wing)

~ size of crow
jaws had teeth
clawed fingers
reptilian skeleton
long reptile like tail

but
feathers may not be similar to modern bird feathers
no keel for flight muscles
 probably didn’t fly
bones not thin and hollow as in modern birds
brain comparable to reptile not to larger bird
brain

by cretaceous, fossils clearly indicate birds that could fly

by end of tertiary (3 MY ago) many modern families were already

in existence

some were up to 12 ft tall

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 3
recent genetic analysis indicates that the large
flightless birds such as ostrich, kiwi & emu are the
most ancient and most “dinosaur-like” birds

the most recent birds to evolve are the very

specialized shorebirds, birds of prey, flamingos
and penguins

Origins of Flight

flight has several advantages over other forms of

locomotion:
permits sudden and rapid escape from predators

 easier to find food, water, nesting areas, mates, etc

fast straight line travel from place to place

inaccessible places become accessible; opens up new

niches

 facilitates migrations over long distances

flight had evolved at least 4 different times in history

of life:
insects: 400 MY;devonian
reptiles: 200MY pterosaurs; late jurassic ,
birds: 150MY; coexisted with pterosaurs for~90MY
bats: 54 MY; (Eocene)

birds didn’t evolve feathers & wings “in hopes” of

achieving flight

there had to be an advantage at each state

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 4
 in its evolution

two major theories:

1. arboreal (gliding, trees down):

wings evolved in reptiles that climbed trees to

hunt for insects

 could glide to base of next tree

eg. used today by some woodpeckers

2. cursorial (running ground up):

wings evolved in running reptile

perhaps as stabilizers

eg. archaeopteryx evolved from a running reptile:

has running legs and feet, not perching

Body Form

in spite of the great diversity of birds they are

amazingly similar in structure

birds evolved as flying machines

entire anatomy is designed around flight

small compact body; reduced weight; with all

heavy organs close to center of gravity

center of gravity and propulsion systems are

properly balanced

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 5

  body is light yet strong enough for flight

Skin

bird skin is thin, light and flexible

 the major functions of skin (insulation and protection from the
elements is taken over by the feathers

loosely attached to body

directly attached to bone in several places

skull and beak

wing tips

legs

no sweat glands

single oil gland at base of tail

 preening

skin over most of body is covered by feathers

on legs only: scales instead of feathers

on head and neck in some birds

 combs and wattles

often brightly colored “ornaments” used for dominance or

sexual signaling
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 6
other bare areas:

vulture head
 keeps feathers clean while feeding on
carcass

ostriches & relatives

 unfeathered legs used for cooling after
heavy exercise

arctic birds have NO bare areas

Feathers & Flight

wings and body covered by feathers

today, the single unique trait that identifies all birds

almost weightless but incredibly strong and tough

feathers smooth the surface and streamline the

contour of the body

 make flying more efficient

but origin of feathers was clearly not for flight

 many dinosaurs apparently had feathers

feathers are epidermal structures derived from

reptile scales

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 7

 developing feathers closely resemble developing scales

scales elongate, edges fray

some dinosaurs clearly had feathers

feathers tend to grow in dense tracts with bare

areas between

while growing feather has a blood supply

when fully grown blood supply is sealed off, and

feathers are dead structures

feathers can be moved individually by muscles in

skin (arrector pili)

a feather consists of:

shaft (=rachis)

quill part of shaft below vanes

vanes

barbs & hooked barbules overlapping extensions of vane

preening “zips” barbs and barbules back together

kinds of feathers

contour feathers
most of the visible feathers
 smooth and streamline body surface

flight feathers = contour feathers that extend beyond the

body and used in flight

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 8

 primaries are the most critical in flying

 their loss may prevent flight

down feathers (plumules)

soft tufts without rachis

lack vane, barbs fan out, not hooked together

hidden beneath contour feathers

especially on breast and abdomen of water birds

to conserve heat

filoplumes (decorative feathers & bristles)

hairlike, degenerative feathers; simple shafts or with tuft of

bristles at end

some decorative  displays

eg. the phoenix fowl (a domesticated relative of

chickens) has tail feathers up to 34 ft long

bristles sensory

on head
around eyes
around mouth and nostrils

birds spend much time on feather maintenance:

preening  reconnects barbs & barbules

oiling  waterproofing

bathing

dust baths  to remove ectoparasites

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 9
feathers can be replaced individually as need or as a
group by molting

Molting

feathers are shed regularly = molting

highly orderly process

(except for penquins who molt all at once)

frequency of molt depends on wear and tear

and seasonal factors
most birds molt once/yr

usually late summer after nesting season

feathers must be shed gradually and

symmetrically (matched pairs) to retain ability
to fly

replacements emerge before next pair is shed

 only ducks and geese are grounded during molting

 wing clipping: removing critical flight feathers on one

wing to prevent flight

Coloration:

among vertebrates, only tropical reef fish show

the same intensity and diversity of color

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 10

 a feather is naturally white

coloration due to:

a. pigments

chromatophores impart colored

pigments during feather development

 color deposited in barbules as they

form
melanins: black, brown, dull yellow, dull red

carotenoids: bright yellow, orange, many reds

porphyrin: bright green, some reds

b. structural color

coloration due to refraction or scattering of

light rays

 all blues, most greens and some purples of

animals

eg. blue jays, indigo buntings, bluebirds

eg. there is no “color” in blue jay feathers

eg. Tyndall blue  caused by light scattering

(refraction) from keratin layers

eg. there is no ‘color’ in blue jay feathers

eg. iridescence due to interference patterns of

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 11
 light; light

interacting with bundles of hollow tubules

inside feathers

color may change when viewed at different

angles

eg. starlings & peacocks

color is used for:

camoflage
eg. in many species, juveniles and females are
camouflaged with melanin pigments

eg. arctic birds white in winter, darker in summer

breeding/communication
eg. males breeding plumage often brightly colored

warning
eg. toxins similar to that of poison frogs has been
found in skin and feathers of some brightly
colored New Guinea species of Pitohui

Skeletal System

some of the most important flight adaptations are

found in the skeleton

the skeleton is exceptionally light and delicate yet

sturdy

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 12

 frigate bird: 7’ wingspan skeleton = 4 oz
 less than weight of feathers

vs humans 6’ skeleton (6-7’ armspan) weighs ~10 lbs

bones light and hollow with air sacs

Many bones are fused together to make them light,

but still strong
skull is light, bones fused together, no teeth

many vertebrae are fused together (not neck) for more rigid
support of body

pelvic girdle is also fused and joined to stabilize legs for

landing

anterior skull bones are elongated to form beak

(or bill) covered with hardened skin attached to
skull
modified lips

since birds lose the use of their forelimbs their beaks

are used as tools

long tubular beaks for nectar

sturdy wedge shaped to pry insects from bark

curved overlapping beaks to crack nuts and seeds

long upper beak that curves down over lower to tear flesh

neck is extremely flexible with more vertebrae than

most vertebrates
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 13
 most mammals have 7 vertebrae

birds have 11-25 vertebrae

the rest of the vertebral column is very rigid with

vertebrae fused together for support

fused ribs and interlocking vertebrae provide

additional attachment sites for flight muscles

the major flight muscles attack to large keel on

sternum

collar bones are fused (wishbone) and connected to

shoulder blade for additional support of wings

the pelvic bones are also fused and fused to vertebrae

to provide a stable support for landing and walking

while the limbs of birds are made up of the same

bones found in the limbs of all vertebrates they
are modified in a characteristic way:
the pectoral appendage forms 3 major joints that support the
flying feathers

humerus - first wing segment; upper arm of us

radius & ulna - second wing segment; lower arm of us

carpometacarpus and phalanges - third wing segment of

bird; hand of us

the pelvic appendage has 4 major joints:

femur - only slightly moveable

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 14
  acts as shock absorber for landing

tibiotarsus - upper “leg” of bird; lower leg and foot of us

tarsometatarsus - lower “leg” of bird; foot of us

phalanges - feet of bird; toes of us

5th toe has been lost and first digit rotated 180o

Movement

numerous muscles in neck provide tremendous

agility

forelimbs modified into wings

breast muscles are the flight muscles

flight muscles (breast muscles) often very large

% of body weight
eg. pigeon up to 50%

not so in gliders and soarers

main muscle mass is near a bird’s center of

gravity

wings have to be large enough to generate enough lift

to support birds weight

 direct relationship between body wt and

wing area

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 15

 largest bird that can fly is the great bustard Otis tarda

22 kg (~10 lbs)

arched wing creates concave depression on underside

creates upward suction as wing passes through air

some birds fly in “V” formations:

when flying in flocks birds use each others energy like fish in
shoals

takes advantages of leading birds slipstream; called drafting,

like bicyclists; helps conserve energy;

birds take turns at lead position

Kinds of Flight:

wings are designed to facilitate a particular kind of

flight
launching

gliding and soaring

use up drafts to stay airborne

flapping flight

complex “figure-8” pattern

hovering

maneuvering

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 16

 smaller faster wings

swooping

diving

swimming

Bird Flight:

some birds spend most of their lives in flight

eg. swifts

eg. alpine swifts spend up to 6 consecutive months

alfot, not even resting after migration each year

eg. common swifts feed, communicate and mate in

flight

only lands to sleep and nest

some swifts spend over 200 days each year in flight

while migrating

average ~135,000 miles/yr (217,300 km/yr)

one recorded a nonstop trip of 310,000 miles

(498,00 km)

cruising speeds are usually ~40 km/hr (25 mph)

peregrine falcon can reach 190 km/hr (120 mph)

many birds can hover at 0 mph

highest flying bird recorded:

Alpine Choughs, Pyrrhocorax graculus  8200 M (26,902’)
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 17
Feet

are nearly devoid of muscles

 greater agility

 very resistant to freezing damage

since mostly bone, tendons & tough skin

when perching, toes lock around branch

 prevents bird from falling off while sleeping

Tail

early birds had long reptilian tail

modern birds have replaced tail with up to 1000

tail feathers; each under individual muscular
control

Digestive System

first birds were probably carnivores

 probably fed mainly on insects

head very flexible & versatile

Beak used like a tool or limb:

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 18
 bird gave up “hands” millions of years ago and
now use beak/bill in their place:

eg. catch bugs, shatter seeds, crush shells, drill holes,

dismemeber carcasses, snare fish

eg. attack enemies, build nests, preen, impress mates and

feed young

birds lives revolve around their beaks

beaks of birds are highly adapted for their feeding

type:
eg. crows generalized type has strong, pointed beak

eg. woodpecker straight, hard, chisel-like, creates forces

of 10 g’s when pecking a tree (humans
can only survive 9g’s for a few
seconds); insert sticky tongue into hole
to find insects

eg. hummingbird long tubular, ~20% birds feed on nectar

eg. seagull  basketlike sac below beak

contrary to conventional “wisdom” birds are voracious

feeders due to high metabolic rate

hummingbird has the fastest metabolic rate of

all birds
eg. 12x’s MB of pigeon & 25x’s MB of chicken

hummingbird may eat 100% body wt/day

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 19

crop:

in many birds there is an enlargement at lower

end of the esophagus = crop

stores food to provide a continuous supply of

energy during flight

used to store food for regurgitation to feed young

in pigeons, doves and some parrots: crop not only

stores food but produces “bird milk”

breakdown of epithelial lining

much higher fat content than cow milk

 feed young for a few days after hatching

gizzard:

modern birds have no teeth

grinding is done in gizzard

muscular with hard keratinized plates to help grind

food

some birds “eat” pebbles to aid this process

some birds of prey form pellets of undigested

material (bones and fur) and regurgitate them
eg. owl pellets
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 20
  another way to reduce weight

birds have very efficient digestion

eg. shrike - can completely digest a mouse in 3 hours

eg. thrush - berries pass completely through GI tract in

30 minutes

Respiration

birds & mammals are warm blooded (homeothermic)

 they maintain a constant body temperature

independent of environment

flight is energy intensive; requires a consistently

high metabolism
higher than land mammals (eg. 110º vs 98º F)

have fast heart rate

eg. hummingbirds 1000 bpm (humans 70bpm)

respiratory system is specially adapted to meet this

metabolic demand
 very different from other vertebrates

bird lungs are different than those of mammals:

 bird lungs are relatively small

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 21

  instead of microscopic sacs (=alveoli) that fill
with air with each breath,

bird lungs contain microscopic tubes, open at

both ends called air capillaries
(=parabronchi)

 in addition to lungs, birds have extensive

system of air sacs in body

usually consists of 5 air sacs connected to

lungs

branches throughout the body and enters

larger bones

the air sacs comprise ~80% of the

respiratory system and may completely
surround the heart, liver, kidneys, gonads
and intestine

air flows in only one direction in bird lungs compared

to the two-way flow of mammals

fresh air goes through lungs on inhale & exhale

as it circulates through the air sacs and then
through the lungs
in alveoli of mammals new air mixes with old
in blind ended sacs

in parabronchi fresh air is constantly moving

through tubes
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 22
 doesn’t get diluted with old air

 much more efficient gas transfer

new studies indicate that some reptiles (alligators) also have a
similar one-way flow of air through lungs; as perhaps did
dinosaurs and ancestors of birds from when O2 levels were
~half what they are today

air sacs and lungs often make up 20% of body

volume (humans lungs=5%)

these air sacs also serve as an air conditioning

system

 cool bird during vigorous flight

eg. pigeon produces 27x’s more heat flying than
at rest

bones with air sacs help to lighten weight of bird

even provides a little buoyancy

 hot air rises

The main breathing muscle in mammals is the

diaphragm which contracts to draw air in and
relaxes to push air out of the lungs

birds do not have a diaphragm, instead they use

muscular contractions to expand and compress the
ribcage for inspiration and expiration
much like reptiles
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 23
the “wishbone” of birds also compresses one of the
airsacs to increase ventilation while in flight to
improve the efficiency of air flow

Circulation

similar to mammals:

4 chambered heart

2 completely separate circuits: pulmonary &

systemic

heart is relatively large

very fast heartbeat (humans ~70-75bpm at rest):

eg. turkey 93 bpm

eg. chicken 250 bpm

eg. blackcapped chicadee 500 bpm

 exercise to 1000 bpm

actual blood pressure is similar to mammals of similar size

Nervous System & Senses

brain is same relative size as mammals

while most birds seem to operate mainly on instinct

and preprogrammed behaviors, birds are the first
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 24
 nonhuman vertebrates to be shown to have
instinctive tendencies to make and use tools
eg. New Caledonian crows use twigs to poke food out of crevices
(without being shown)

several other birds such as woodpecker finches and

burrowing owls also routinely use tools but it’s not known if
their skills come from copying other birds or are innate

cerebellum is relatively larger than in mammals

or reptiles

 coordination of flight muscles

a. eyes are perhaps the most important sense organ

birds eyes are disproportinately larger than in

other vertebrates
eg. the eye of the ostrich is ~2” diameter; the largest of any
vertebrate

eg. the eyes of most large birds; eg hawks and eagles are larger
than human eyes

also disproportionately large compare orbits in

skulls

no eye muscles

 all space is filled with eyeball

can’t move eyes to track objects

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 25

 flexible neck compensates

plate-like sclerotic ring strengthens and focuses

eyes

pecten is a thin, greatly folded tissue extending

from the retina toward the lens.

unique structure in birds eyes

supplies nutrients and oxygen throughout the

vitreous humor
this reduces the number of blood vessels in the retina
that obscure the rods and cones.

With fewer blood vessels to scatter light coming into

the eye vision is enhanced

predatory birds such as eagles and hawks

have the largest and most elaborate
pecten of all the birds. and the best vision
of all animals

generally:

predatory birds have eyes in front of head

 stereo vision = depth perception

some birds of prey have 2 foveas (areas of

greatest visual acuity - mammals have
one)
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 26
 equivivalent to a camera having both a
telephoto and a macro lens

visual acuity of hawk is 8x’s that of

humans

best vision in animal kingdom:

 can clearly see crouching rabbit >1 mile away

vegetarian birds have eyes that look out to

sides

 greater field of view

eyes have nictitating membrane (reptile trait)

in some water birds this membrane acts as a

“contact lens” to help birds focus
underwater

a few birds (eg zebra finches) can see UV light

b. hearing is also well developed in birds

they can hear the same range of sounds as we do

yet their ear structure remains reptilian:
have retained single middle ear bone (columella)

do have short outer ear canal

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 27

 but better cochlea (not coiled as in mammals,
but long and straight; banana-shaped)

birds can replace damaged hair cells (we

can’t)

In some their hearing ability improves during

mating season and declines the rest of the
year

night predators such as owls have asymetric ears

(not the feathery “ear” tufts) improving their
ability to locate prey in the dark.

great grey owls, which hunt in the daytime

can locate a mouse beneath the snow with
precision

a few birds (oilbirds & swiftlets) can use

echolocation in the dark to avoid objects

bird ears are not as effective as mammal ears

for sonar

not as good as bats, not used for hunting

c. Sound and Communication

bird songs are some of the most recognizable and

endearing characteristics of birds; used to
find mates, mark territories and other kinds of
communication
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 28
 most birds have a larynx (voice box) but don’t use it to generate
sounds

they do have a syrinx

a cartilaginous chamber at the base of the

trachea with muscle controlled membranes
which they use to make familiar bird songs

membranes on each side can produce separate sounds to

generate chords or harmonies when singing

one species, the club-winged manakin of Ecuador

creates a courtship sound by knocking his
wings together over his back to go with his
song
 no other vertebrate produces this kind of sound, although
stridulation is common in insects

c. smell, we used to think birds lacked sense of smell

now we know they have very refined sense of
smell (some as good as mammals)

used to find food, breeding colonies, their

individual nests and recognize their
partners

vultures & petrels probably have some of the best smell

receptors; songbirds the worst

d. taste not very well developed

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 29

 hundreds of receptors located on upper and
lower jaws
mammals have 10,000’s mainly on tongue

e. touch

the bills of ducks & wading birds contain very

sensitive touch receptors

as they dabble in muddy water or probe for

hidden food they can distinguish between
what’s edible and what’s not

touch also plays an important role in social

relationships that promotes bonding

many spend hours preening each other (as in

primate grooming)

Excretion

kidney is similar to that of reptiles

contains nephrons which filter blood and remove

metabolic wastes

water is conserved by excreting insoluble uric acid as

main nitrogen waste
takes 20x’s more water to get rid of urea than to get rid of uric
acid

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 30

metabolic wastes are passed directly to cloaca
birds have no bladder

water is reabsorbed in cloaca

the white, paste-like uric acid that remains is

eliminated along with feces through the vent

like reptiles, marine birds have salt glands that

empty through nostrils to get rid of excess salt

Life Cycle

some birds live over 70 years (eg. Andean condors)

reproductive strategy for most birds is to have a few

offspring at a time and provide lots of care initially

usually both males and females contribute to care.

Reproduction, Nesting & Egg Laying

birds are dioecious with internal fertilization

to save weight:
in males testes enlarge only during mating season

females only have left ovary; not paired

Courtship

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 31

courtship in birds involves

 marking and defending a territory

 and sometimes elaborate rituals to entice a

female into the territory

selection of territory usually occurs a few weeks

before nesting season

male selects nest location

solitary species defend fairly large area

gregarious species that nest in colonies defend

a very small area

sometimes this seasonal instinct to defend

territory becomes obsessive
eg. robin or cardinal that returns day after day to
struggle futily with its reflection in a window pane

courtship rituals

males are sometimes very colorful during breeding

season, dull rest of time

many develop seasonal ornamentation

eg. inflated skin pouch on throat

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 32

 courtship almost always involves singing to a
potential mate

sometimes also involves elaborate dances

eg. lyrebird - to attract a mate:

male will stand on a small mound of dirt and spreat his

decorative tail feathers up over his head

he then sings both his own songs and mimicks other

bird’s songs

he will even mimic the noise of a nearby car

as he sings he jumps about

eg. frigatebird

male has a throat sac that it can inflate over a period

of 20 minutes into a heart shaped balloon

he then waggles his head from side to side, shakes his

wings and calls the female

a female frigatebird will mate with the male with the

largest and shiniest balloon

during sex the male will “sweetly” put its wings over
the females eyes … to make sure she doesn’t get
distracted by a better offer

eg. Long tailed Manakin (of Costa Rica)

males work in pairs who begin perched on a branch

near the ground

they both call a whistle-like call for females

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 33

 a female lands on the branch indicating she is ready to
be courted

both birds launch into a prolonged acrobatic display

they step daintily and hop, they somersault and leap-

frog, they take turns hovering in the air, all while
calling to the female

as the tempo picks up the males emit a buzzing sound

and the female becomes even more excited

at the critical point the leading male utters a shrill cry

this is the lesser male’s cue to make himself scarce

following a brief dance the male quickly mounts the

female

eg. bowerbirds

19 sp in Australia and New Guinea

no other bird has developed its breeding and mating

behaviors to such a degree

the key to successful mating is construction of a

“bower”  a performance arena used only to
entice a female, not a nest for eggs

the least colorful males produces the most

elaborate bowers

it apparently takes the young males several years

to learn how to build one

each bower consists of a mat of mosses and ferns,

decorated with shells, feathers, pieces of fruit, etc

often includes an ‘avenue’ bordered by walls of

sticks
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 34
 some bowers form a coneshaped hut several feet tall
with a door and a ‘front lawn’ decorated with fruit
and flowers

once the bower is completed the male defends it and

entices females with complex calls and dances

if the female is impressed she will mate then leave to

build a nest and lay eggs

the male promptly begins advertising for a new partner

-------------------

most birds have no transfer organ press cloacas

together
a few birds have erectile penis with external groove to guide
sperm into females cloaca

a few bird species are monogamous for life

eg. swans, geese, eagles, owls

most birds are monogamous while mating

but after mating or rearing their chicks, they

go their own ways
however new genetic data shows that many female
birds once thought monogamous are actually
quite promiscuous bearing offspring from several
fathers

Nests

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 35

some of the most obvious and characteristic features
of birds are the nests they make to lay eggs and
care for their young

nests vary from simple depressions or accumulations

of materials on the ground to large communal
nests for 100’s of birds
some of the most elaborate nests are associated with some
swallows and weaver finches

nests typically take 2 to 7 days to construct

cavity nests in trees can take up to 4 weeks to

excavate

the most elaborate nests can take months

nests vary in complexity based at least partly on their

flying skills

eg. typical nest of smaller bird is cup shaped “basket”

lined with finer material

eg. barn and cliff swallows mold nests of mud from

softened pellets

eg. largest bird nest is that of bald eagle

 to 10’ wide, 20’long and 5,500 lbs

(the weight of almost 3 cars)

 the same nest can be used for decades

most birds void outside of nest

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 36
types of nests:

scrape nests – simple depressions in the ground or litter

eg. penguins, shorebirds, gulls, terns, nighthawks,

vultures

burrow nests – burrows dug in the sides of cliffs

are very effective at protecting eggs and young from
predators

eg. bank swallows, kingfishers, burrowing owls

cavity nests – nests excavated in trees or cacti are used by

numerous birds

eg. woodpeckers, owls, parrots, some waterfowl

platform nests – relatively flat nests that may be on the

ground, in a tree or across the tops of vegetation in shallow
water

eg. western grebe

cupped nest – some of the most commonly seen nests,

usually in branches of trees and shrubs and supported from
below or suspended from branches; there are many
variations in this kind of nest

eg. many “passerines” & hummingbirds

Eggs

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 37

 all birds lay eggs (=oviparous)

not sure why they never evolved to bear live

young (=viviparity)

live bearing did evolve in all other vertebrate groups:

fishes
amphibians - at least twice
reptiles - 100’s of separate times
mammals - once

one argument is that it would be difficult for a

pregnant bird to fly

but: bats can

another idea is that bird’s body temperature is usually

a few degrees higher than the body temp of
mammals and that temperature is lethal to the
eggs

birds typically incubate their eggs at ~37º C

(98.6º F)

all bird eggs have hard shells

with lots of microscopic pores

shell is soft when formed & hardens before

being layed

egg size & shape

largest: known bird egg is from extinct Elephant bird
(Aepyornis) of Madasgascar

13” long, 9.5” dia; 2 gallon volume

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 38

 smallest: some hummingbird species <1/4th “

abnormal eggs:

a. runt egg

yolk smaller than normal

 parasite or debris in oviduct triggers egg

fomation missing yolk all together

b. double yolk
rarely 3 yolks

c. egg within an egg

enforced return up the oviduct

Parental Care

usually female incubates eggs

12-30 days needed for incubation

incubating birds develop “incubation patches”

loss of feathers
thickening of skin
greater blood supply to area to maintain temp

in hot areas birds must shade eggs

after hatching young are fed by regurgitation

some birds (pigeons, doves, flamingos and some

penguins) produce crop milk

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 39

 secretions with a “cottage cheese”-like
consistency, very high in proteins and fats

much higher fat content than cow milk

produced by both male and female birds to

feed the young for the first few weeks

Migrations

regular extensive seasonal movements of birds

between summer and winter regions

origins: more northerly birds were forced south as

winters got colder

enables bird to live in optimal climate all the time

generally breed in northern latitudes
overwinter in southern latitudes

as north and south ranges moved further apart

due to continental drift the migration routes
got longer and longer

just less than half of all birds migrate

(for 1000’s of years thought most hibernated)

Why Migrate?

migration increases the amount of space available

for breeding
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 40
  reduces aggressive territorial behavior

avoids climate extremes favors homeostasis

 less energy needed to maintain internal

stability

north in summer

 long summers, abundance of insects to rear

young, not many predators

much variation in methods of migration

 most birds migrate at night,

esp smaller birds

mainly before midnight and immediately before dawn

 protection from predators

 can spend day feeding

some larger birds migrate in day

eg. hawks, shore birds, black birds, pelicans, bluejays

 most fly at <3000’

very rare to find one >5000’

 most travel at 20-50 mph

some 80-100mph
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 41
 ancients thought they migrated in 1 night
 would have required speeds of 180-240 mph

 main timing factor is changes in day length

Arctic tern has longest migration route:

nests from Mass to Greenland and Alaska

spends winters off shores of Anarctic near Falkland Islands

 experiences more hours of daylight than any other bird

only crossing the equator does it esperience long nights

navigation cues:

birds use a variety of navigation clues including

sun and stars and olfactory cues. Some can
even use the earth’s magnetic fields

1. visual
follow familiar migratory routes

most migratory birds have well established routes

2. earth’s gravitational field

some birds have crystals of magnetite around their eyes
and in their nasal cavities of upper beak

3. celestial cues

4. sun’s position in sky & ‘time sense’

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 42

Classification of Birds

birds today are divided into 2 major groups with 27-28

orders:

1. flying birds
largest group of these are perching birds
 comprise >1/2 of all bird species

2. flightless birds (ratites)

originally from flying forms

flightlessness almost always evolved on islands with few

predators

lost use of wings; keel and flight muscles degenerate

 lived with few predators in isolated areas

tendency to become quite large

eg. moas of New Zealand to 500 lbs

eg. elephant bird of madagascar is largest bird that ever

lived

2 M tall, 450kg(~1000 lbs)

developed very powerful legs

 can achieve very fast running speeds

eg. emu ~ 30 mph,

eg. ostrich ~ 42 mph and up to 60 mph

swimming birds:
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 43
 most have webbed feet

birds with webbed feet for paddling have legs

far back on body and tend to be clumsy
walkers

those that dive skillfully are usually not very good

fliers

some chase prey underwater by paddling with

wings

some very effective swimmers

eg. gentoo penquin  22mph

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 44

 Bird Ecology
1. pollination
eg. hummingbirds
do not have a highly developed sense of smell
but do have excellent sense of vision
frequently bright red or yellow flowers
little if any odor
fused petals with nectary
produce copius quantities of nectar
long floral tubes prevent most insects from reaching the
nectar
eg. fuschias, petunias, morning glories, salvias, cardinal
flowers, trumpet creepers, columbines, penstemons

2. disperse seeds
eg. edible fruits
attracts birds or mammals
may eat whole fruit or spit out pits
if swallowed seeds resistant to digestive juices
squirrels and birds bury fruits and seeds
nuts stored underground are forgotten

eg. passively carried by birds

hooks or spines to catch in feathers
in mud on feet of birds, etc.
burs, beggars ticks, devils claw, etc.

3. pest control
eg. Birds eat many things: beetles, flies, spiders, earthworms,
rotting fish, offal, poison oak berries, weed seeds, etc

eg. raptors & owls - eat mice, rats, snakes

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 45

 Human Impacts
1. meat and eggs

20 B birds are used for food (world/yr):

chickens 20 B/yr
turkeys 304 M/yr
ducks 773 M/yr
geese 209 M/yr

US exports:
120,000 tons of wings to china
2 M tons of leg quarters to Russia

US also processes:

1.6 M tons of feathers/yr

 ground u into feather meal as animal feed or plastic
fortifiers

intestines go to rendering plants where they

are ground up and used in pet foods and
fertilizers
US exports ~50,000 tons to South Africa

~330,000 tons of feet/yr exported to China

scientists have recently bred a “featherless

chicken

 grows faster
 don’t need to pluck it
Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 46
 ~ 91 Billion eggs produced US each year

2. extinct or endangered species due primarily

to human activities
2/3rds of bird species are declining in numbers

eg. about 20% of world’s bird species have

gone extinct in historic past

eg. Passenger Pigeon (Ectopistes migratorius)

inhabited eastern N America
200 yrs ago was the world’s most
abundant bird
 3-5 Billion
once accounted for ~1/4th - 1/3rd of all N
Am birds
 1830’s Audubon saw a single flock
estimated at 10 miles wide and 100’s
miles long (~1 Bill birds)

were easily slaughtered for meat (pigeon pie)

 they wouldn’t fly away if threatened

over 20 yrs of hunting and habitat loss at end of

1800’s the population was decimated

last wild bird was shot in 1900

last individual (Martha) died at the Cincinnati Zoo in

1914

eg. Ivory Billed Woodpecker

3. Other direct effects of humans on birds

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 47

 100 M – 1 B birds are killed by glass collisions/yr
in US

140,000 – 328,000 birds die from wind turbines/yr

in US

500,000 - 700,000 birds are killed by getting

tangled in fishing nets and hooks/yr world

4. Introduced pests
eg. starling

eg. house sparrow

eg. brown tree snake  Guam 1950 caused extinction

of 9 of 18 native bird species; 3 sp of bats and several
lizards

5. Domesticated Birds and Bird as Pets

some birds have been truly domesticated:

eg. chickens, turkeys, geese, ducks, pigeons

a few are semidomesticated

eg. hawks and falcons

earliest domestication ~1700 BC in Persia

Europe ~300 BC

12% of pet sales are birds (19% dogs; 5% cats)

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 48

 ??5 M live birds are sold worldwide
4.2-13 M US/yr ??world??

in US (1980’s): 500,000 birds sold as pets

European Countries buy 3/4th ‘s of live birds

illegal trade:
bird collectors will pay $10,000 for a rare hyacinth macaw
from Brazil

$12,000 for a pair of golden-shouldered parakeets

from Australia

mortality rate of live animal trade is enormous:

~50 animals caught or killed for every live animal
that gets to “market”

6. bird watching
more lucrative than bird hunting

7. hunting

91 M birds are hunted each year worldwide

21 M waterfowl
 2injured/ 1 taken

500,000-700,000 seabirds killed as bycatch

8. research
5 M birds are used for research each yr

9. wildlife photography, art

Animals: Phylum Chordata-Vertebrata-Birds; Ziser Lecture Notes, 2015.11 49

 Phylum Chordata – Vertebrates
Reptilia
8000 species worldwide; 340 sp in US & Canada

reptiles include lizards, snakes, turtles, and are

generally looked on as cold, creepy or scary
creatures

over earth’s history they have been an extremely

diverse and successful group

even today they are a very successful group

they occupy a great variety of terrestrial habitats

in some habitats such as deserts they are the

dominant vertebrate group

some have even returned to an aquatic lifestyle in

oceans and freshwaters
______________________________________

reptiles were the 1st vertebrates no longer tied to

water, even for reproduction

 1st truly terrestrial vertebrates

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 1

First Reptiles

310 MY ago, 50 MY after the appearance of the first

amphibians, some amphibians developed the
ability to lay eggs on land

 the first reptiles

while amphibian adults can live on land

they must have water to reproduce

complete independence from water didn’t occur

until the evolution of a self contained egg
capable of storing water (=cleidoic egg or
amniotic egg)

the appearance of this new type of egg

allowed the evolution of reptiles, birds &
mammals

the earliest example of a true reptile was a

lizard-like, partly aquatic animal

~1.5’ long

probably ate mostly insects

 shows characteristics of both groups

reptilian skeleton
still had lateral line system
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 2
 probably didn’t yet lay shelled eggs

by 225 (end of Permian) most amphibians of the time

had become extinct

the amphibians were “quickly” replaced by a great

diversification of reptiles

shortly after reptiles arose, they began diversifying

onto land at time of:
variable climate; tending to get drier

diversification of plants including flowering plants created

numerous habitats

increase in the number of insects and amphibians provided

an ample food supply
_______________________________________________

largest living reptile:

eg. Komodo dragon (Varanus komodensis)

10 feet long; 300 lbs

eg. Australian saltwater crocodile

up to 28 feet long

eg. leatherback sea turtle

8 feet long; 1500 lbs

largest reptile ever:

eg. largest of all reptiles: Diplodocus hallorum, giant sauropod

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 3

 120+ feet long (37 m); 30-80 tons

 largest animal ever to have walked on land

smallest living reptile

Carribean lizard <3/4ths of an inch nose - tip of tail

smallest snake – 4” thread snake from Barbados

reptiles were much more abundant and diverse

~160-100MY ago

= age of reptiles (Mesozoic)

lasted >165 M Y

Skin

a second major innovation of reptiles is a thick, tough,

dry, waterproof skin

their skin consists of the same 2 basic layers of

vertebrate skin: epidermis & dermis

 but epidermis much thicker than amphibians

 and contains lipids and waxy keratin

 very effective water proofing

 the skin of reptiles also contains scales but

unlike fish scales:
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 4
 reptile scales are in the epidermis,
not the dermis

**epidermal (not dermal) scales**

reptile scales ≠ fish scales
(epidermal) (dermal)

[scutes of turtles are modified scales]

also reptile scales are made of keratin, a

waxy protein, not enamel and dentin
as in some fish
(structures made completely of keratin are
often referred to as “horny”)

reptiles scales are homologous to feathers

and fur of birds and mammals

some extinct forms (dinosaurs & relatives) had

primitive feathers or hair in addition to
scales

reptile skin provide excellent protection from drying,

abrasion, predators

outer layer of epidermis is shed periodically

some reptiles retained the ability to grow bony

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 5

 dermal plates (equivalent to fish scales) under
their scales producing large rigid scales
eg. turtles , alligators

thick dermis contains chromatophores

in some the chromatophores can rapidly

change color for camoflage
eg. green anoles

eg. inland taipans, fierce snakes up to 10 ft long,

can get darker in winter and lighter in
summer

eg. australian copperhead can also change color

in some reptiles the thick keratinized (horny) skin

has been modified into claws, scutes (large
scales of turtles), horns and rattles

Skeleton & Support

new features appear in reptile skeleton:

fewer skull bones

teeth still simple and peg like but in more efficient

and stronger jaws

 more biting force

nasal cavity separated from mouth by shelf of

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 6
 bone = palate

 easier to breath while eating

limbs stronger, more flexible & closer to body

 better designed for walking

toes with claws

Movement

more powerful muscles than amphibians

limbs are stronger and more flexible for walking

chameleons have opposable toes for arboreal life

millions of adhesive fibers on the feet of geckos are so

effective for climbing that they allow the animals
to hang from a ceiling by a single toe

one group of extinct reptiles were the second group of

animals (and the first vertebrates) to be able to
fly

some reptiles can glide:

eg. Draco = flying dragon

uses extended ribs

can glide up to 50’

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 7
 eg. gliding gecko Ptychizoon sp.

eg. some gliding snakes

some can fly up to 60 ft horizontally as they drop from

trees

they can flatten their bodies and undulate to “catch a

breeze”

most land reptiles can burrow into mud

eg. turtles and small lizards

most reptiles swim with ease

some groups of reptiles have lost their limbs and and

use their scales to grip the ground while rib
muscles move them foreward

Feeding and Digestion

most reptiles are carnivores

jaws are more efficient for crushing and gripping prey

(fish use “suction” for feeding; their jaws are not
very muscular; amphibians can’t “chew”)

tongue is muscular and mobile

used to help catch prey

tongue never attached to front of mouth

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 8
 in some tongue serves as touch receptor

most reptiles have teeth

crocodilian teeth are constantly replaced

in some reptiles (snakes & a few lizards) salivary

glands are modified into poison glands to take
down prey
eg. king cobras are one of the deadliest species of
venomous snakes

one snake (the tiger keelback snake in Asia) is the

only known vertebrate that sequesters toxins from
its prey (frogs and toads)
it stores these toxins in glands behind its head

it uses these stored toxins to ward off prey

while having its own anticoagulant toxin to capture its own prey

some large snakes are ambush predators and kill their

prey by suffocation

doesn’t directly squeeze it to death

 once wrapped around their prey, each time the prey the snake
tightens its grip, until prey can no longer inhale

since meals tend to be scarce they gorge

eg. some adult pythons can swallow an adult pig whole

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 9

 within days of a meal their organs enlarge by at
least 1/3rd, some to double their size – greatly
increasing their metabolism
the change in metabolic activity is greater than that of
a racehorse going from rest full speed in a quarter mile
race

after the meal is digested the organs shrink back

to their fasting size.

again, teeth are not used for chewing

stomach often has pebbles to help grind food

(=gastroliths) common find at dinosaur sites

liver and pancreas produce wide variety of digestive

enzymes

Respiration

Reptiles never have gills

since skin is thick and dry most reptiles depend

completely on lungs for gas exchange

lungs are more developed, more folding, more

surface area

more efficient

air is sucked into lungs, not gulped as in fish and

amphibians
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 10
 can expand and contract the rib cage to inhale &
exhale

some have fully developed ribcage and sternum to

facilitate breathing through lungs

still relatively low breathing rate since still cold

blooded

(1/10th that of birds and mammals of same size)

but most can’t breath while running since any of

same muscles are used for both purposes

a few (crocodilians) with muscular diaphragm

also a few reptiles can exchange gasses through skin

eg. sea snakes, soft shelled turtles

also, aquatic turtles can extract oxygen from water

they “breath” through their mouth and cloaca
(can pump water in and out, highly vascularized)

no vocal cords like amphibians

 reptiles can only hiss

Circulation

like amphibians, most with three chambered hearts

with 2 atria & 1 ventricle

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 11
 but partial septum separates the ventricle

 slows mixing of oxygenated &

unoxygenated blood

two separate circuits of bloodflow:

pulmonary and systemic

the respiratory & circulatory systems of reptiles

provides more oxygen to tissues than that of
amphibians

more efficient heart; higher blood pressure

but still less efficient than that of birds and

mammals

crocodilians have 4 chambered heart which

completely separates the pulmonary and systemic
circuits
similar to that of birds and mammals

overall, still low metabolic rate = cold blooded

(~1/10th rate of birds or mammals of same size)

some reptiles in past were warmblooded

Nervous System

more advanced than amphibians

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 12
 cerebrum increased in size but no ‘cortex’
 allows more complex behaviors

cerebellum less important than in birds and

mammals
eg. motor functions in many reptiles is at least
partly controlled by ganglia along spinal cord

but still reptiles are not as dependent on their brain as

mammals are
eg a turtle ‘lived’ 18 days after brain was removed

Senses

a. vision

is most important sense organ

eyes usually with 2 moveable eyelids

some with a 3rd  nictitating membrane

most reptiles are active during the day

 have cones for good color vision

some have “third eye” pineal eye on top of head

 detects light intensity and may control

biological rhythms
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 13
 vertical pupils in nocturnal snakes (and some
mammals) are better for night time hunting, it
also gives them a deeper field of view making
it easier for them to sneak up on prey

b. smell & taste

also have well developed sense of smell

eg. crocodilians secrete a strong musk during mating
season

 olfactory epithelium in nasal cavity

Jakobson’s organ assists in sense of smell/taste

pits located on roof of mouth in lizards and

snakes

 forked tongue of snakes flicked then

touched to Jakobson’s organ to follow
chemical trails

c. heat sensors

some snakes have IR receptors

 can see body heat from warm blooded prey

d. hearing

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 14

 similar but better than in amphibians
contain inner and middle ears

external tympanum and columella (stapes)

in lizards the tympanum is recessed into a canal

some lack tympanum and use bone adjacent

to jaw to detect sound in ground, not air

Excretion & Salt/Water Balance

more efficient (metanephric) kidneys

amphibians excrete N wastes as ammonia

 requires lots of water to dilute it

most reptiles (and birds) secrete uric acid,

not urea or ammonia

 requires much less water

most excess water is reabsorbed by bladder

reptiles, birds, mammals that live near sea cannot get

rid of all salt via kidneys

most have evolved salt glands (independently)

eg. marine iguana
 salt gland empties through nostrils

eg. sea turtles

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 15
  salt gland in orbit of eyes

eg. sea snakes

 salt glands are beneath tongue

Protection/Defense

many species are well camoflaged by the color of their

skin

venomous snakes use their poisonous fangs for

protection as well as for subduing prey

rattlesnakes advertise their presence with a

threatening rattle

horned lizards can puff up their bodies causing its

spiny scales to protrude making them hard to
swallow

horned lizards can also aim and squirt a stream of

blood up to 5’ from the corners of their eyes

the blood contains foul-tasting chemicals that

deter wolves, coyotes and domestic dogs

Reproduction

dioecious

but hard to tell male from female

a few have sex chromosomes:

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 16
 in many the gene that controls gender depends on
temperature during critical period of
development

copulatory organs and internal fertilization first

became the common practice in reptiles in the
vertebrate line

all reptiles have internal fertilization

in order to enclose the embryo inside a thick

waterproof protective covering, the egg
must be fertilized before the “eggshell”
encloses it

so they don’t need water for reproduction

reptiles have 1 or 2 copulatory organs

some snakes and lizards have 1 penis

most snakes and lizards have 2 penises (=hemipenes)

 use only 1 at a time; depending in which testis has more

sperm

tauratauras lack any penis

 press cloacas together

after copulation, sperm may remain in female for

months or years before it is used to fertilize egg

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 17

a few reptiles are parthenogenetic

 egg develops without fertilization

almost all reptiles go through early development

within an amniotic egg

don’t need water for development

(only found in reptiles, birds & mammals)

with protective embryonic membranes enclosing

embryo

complete life support system

4 membranes: amnion  watery cushion
allantois  collects wastes
yolk sac  provides stored food
chorion  with allantois for respiration

embryo & membranes are enclosed within a

porous shell

can be leathery or hard shell

most reptile eggs require 4 – 6 weeks for

development before hatching

no reptiles pass through a free living larval stage

a few reptiles (some pit vipers) bear live young

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 18

 one reptile, an african skink Trachylepis ivensii ,has a functioning
placenta

a trait formerly only known for mammals

the embryos develop, attached by a placenta to the

oviducts of the mother

few living reptiles show parental care

they generally lay a large clutch of eggs and then

abandon the nest

there are some exceptions:

eg. some dinosaurs showed parental care

eg. crocodilians dig nest for 25-50 eggs & cover the eggs

hatchlings often chirp encouraging mom to uncover nest

she then picks them up and carries them to water

both mom and dad respond to distress calls

eg. a few turtle species show minimal parental care;

watching eggs for a few days and urinating on them if they

get dry

or after hatching climbing on or swimming with mom (or

any other female in the area)

many reptiles have well developed abilities to

regenerate missing body parts
eg. green anoles can lose tail
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 19
 eg. glass snake: when pursued can break of its tail with a
sharp twist

the tail twitches and writes to attract pursuer while

glass snake escapes

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 20

 Kinds of Reptiles
today only 4 groups of reptiles (3 major, 1 minor)

during mesozoic more than 12 major groups of

reptiles emerged and diversified including
dinosaurs, along with shark-like reptiles
(ichthyosaurs), plesiosaurs and flying reptiles
(pterosaurs)

 lots of niches open (no birds or mammals

yet)

Dinosaurs

~540 different species of dinosaurs have been

described

2006 study concluded that at least 70% of dinosaur genera

remain unknown

some estimate up to 900 species once existed

2007 a new kind of dinosaur that lived in excavated dens

was discovered

ichthyosaurs, plesiosaurs and pterosaurs are technically

NOT dinosaurs but other extinct groups of reptiles

the largest pterosaurs with wingspans of 30 feet, could fly

10,000 miles nonstop

dinosaurs generally lived in warmer even tropical parts

of the world but fossils are also found much closer
to poles
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 21
 Cretaceous dinosaur fossils are being found further
and further north and south.

eg. north slope,

eg. southern argentina and australia

 cold winter nights would have lasted several months

how? (1988)

dinosaurs share a group of unique features that set

them apart from all other vertebrate groups:

• their generally large size may be due to their

extremely efficient (birdlike) lungs with airsacs

• most with upright stance

 legs positioned directly beneath body

similar to mammals and birds

not like amphibians and most living reptiles

• many dinosaurs were bipedal

• considerably stronger hinge joints at knee and

ankle
same as in birds

much less flexibility than in mammals

• dinosaurs walked on their toes

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 22
 like horses

• many or most dinosaurs were warm blooded

(endothermic); body temp ~ same as
mammals
today, only birds and mammals are warmblooded

• many had feathers &/or fur in addition to

epidermal scales
feathers & fur have the same structure as reptile
scales

 all are essentially the same thing

• many showed considerable maternal care

most reptiles today have no care of young

were apparently competitively superior to mammals at

the time since mammals remained small and
inconspicuous until ALL dinosaurs disappeared

 then mammal diversity exploded

dinosaurs and most of the diversity of reptiles

disappeared ~65 MY ago

 probably meteorite impact

cretaceous extinction may have been caused by a pulse of
asteroids not just one, eg. Manson, Iowa crater 66MY old

also, extensive volcanic activity contributed to their extinction

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 23

today only 4 groups of reptiles (3 major, 1 minor)

1. Turtles (~300 species)

ancient group (only surviving anapsids)

appeared in U Triassic (200 MY ago)

very little change in body form over past 200 M

years

the most distinctive feature of turtles is their shell

no other vertebrate has developed such a

distinctive armor

dorsal carapace and ventral plastron

outer layer of keratin, inner layer of bone

formed from fused vertebrae and

covered with dermal bone=carapace &
plastron of fused scales

in some turtles the plastron is hinged to

further protect the animal from danger

the carapace & plastron are coverd by large

scales (=scutes) fused to the bone below
biologists use the number, size and color of
specific scutes to identify turtle species

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 24

 in some species the growth rings of a scute can
be used to assess an individuals general age

ribcage can’t be used for breathing

turtles must pump air into lungs like

amphibians do

aquatic forms can breath some through skin

turtles are generally slow moving grazers and

scavengers; or eat slow moving insects and
worms, etc

no teeth, instead rough horny plates

good sense of smell & color vision, poor hearing

low metabolism  live long >100 yrs

many turtles require 6-12 years to attain
maturity; some take 20 or more

internal fertilization

all turtles bury eggs in ground

nest temperature determines gender of hatchlings

no sex chromosomes

three kinds of turtles:

tortoises – terrestrial
terrapins – freshwater
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 25
 turtles – marine

eg. sea turtles

have webbed feet

up to 6’ long

migrate 1000’s of miles

live, feed and grow in sargasso sea

may take 50 years to reach sexual maturity

(low metabolism – some live to 150 yrs old)

they then return to beach where they hatched

to lay eggs

(reverse of amphibians)

as soon as they hatch sea turtle swim across 100’s of miles

of ocean

 can detect earth’s magnetic fields for navigation

eg. box turtle

shell has 2 hinges to close up from predators
omnivorous: fruits and beries
may live up to 100 years

25,000 box turtles/yr are exported to Europe

as pets
90% die in transit

eg. snapping turtle

common in ponds in eastern US
grow to 1 ft long
ferocious and short tempered
entirely carnivorous (fish, frogs, birds)
come ashore only to lay eggs

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 26

2. Lizards & Snakes (6000sp)

the largest and most diverse living group of

reptiles

appeared in fossil record in Jurassic

evolutionarily are very similar

 placed in same order

most successful group

95% of all living reptiles

very effective jaws to capture prey

very flexible
eg. snake can swallow prey several times
its own diameter

the order is about evenly divided between lizard

and snake species

2a. Lizards

terrestrial, burrowing, aquatic, arboreal or aerial

many lizards show reduction or loss of limbs

some legless forms

eg. glass lizard

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 27

 eg. worm lizard

very effective jaws to capture prey

cold blooded but can regulate temperature by

behavior to maintain a fairly constant body
temperature
 early morning basking in sun to absorb heat
 hot  turn face to sun to expose less area
 lift legs on hot substrate
 hottest part of day may retreat to burrows

the desert iguana prefers higher body

temperatures and can tolerate 117º F

 lethal to all birds, mammals, most

lizards

eg. Geckos
small, nocturnal
adhesive pads on feet  can walk upsidedown
common around houses in Austin

eg. Chameleons
catch insects with sticky tongue

eg. Iguanas
include skinks, monitors and komodo dragons
often brightly colored
some marine
have salt glands to get rid of excess salt

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 28

 2b. Snakes

while lizards are probably the most familiar and

“typical” reptiles, snakes are often considered
“strange” and even “evil” by many
for 1000’s of years, snakes were looked on as mysterious
creatures, often with magical powers and were
important in many religious practices

most are terrestrial, some are aquatic, some

marine

most lizards have moveable eyelids;

snakes eyes are permanently covered with

transparent layer; no moveable eyelids

body very similar to lizards except for legs

most snakes have lost all traces of

appendages

some retain vestiges of pelvic girdle

we often think of the loss of legs as a

disadvantage but it has actually improved
their ability to move

locomotion is radically altered, very adaptable

scales grip the ground as they make eel-like

movements
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 29
 essentially walk on their ribs:

 up to 300 ribs

each rib has separate muscles that control its

movement

snakes can climb, leap, swim, stand erect and

“run”
some species such as racers and coachwips can
outpace humans even on difficult terrain

 probably most unusual is side winding

of some desert species

sidewinders have only 2 parts of body

touching ground at any one time
 essentially walking without legs

some snakes use color for camoflage

eg. inland taipans, fierce snakes up to 10 ft long, can get darker
in winter and lighter in summer

eg. australian copperhead can also change color

no external ears or tympanic membrane

do have internal ears and can detect low

sound vibrations and soil vibrations

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 30

 most snakes depend mainly on chemical senses to
detect prey (vomeronasal or Jacobson’s organ)

use tongue as “smell” receptor

picks up chemicals with tongue

transfers tongue to Jakobson’s organ in mouth

some snakes also have pit receptors along the jaw

margins that detect IR radiation (heat)

they can detect any animal warmer than its

surroundings (esp birds and mammals, but
also other reptiles)

these receptors are the most sensitive heat

receptors known in the animal kingdom
can detect differences of 0.001oC

snakes generally have only 1 functional lung that

extends almost the whole length of the body

the other lung is greatly reduced

most rely on chemical senses to hunt

flexible jaws allow snakes to swallow prey several

times their own diameter

snakes subdues prey by suffocation(constriction)

or venom
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 31
 eg. boas and pythons  wrap and suffocate

eg. venom = saliva with a mixture of

digestive enzymes and protein fragments

 kills and starts the digestive process

some of these venoms work on the nervous

system to cause paralysis

while only 2 lizard species are venomous

(include Gila Monster)

 many snakes are venomous

a. coral snakes (ELAPHIDAE)

most dangerous
Indian cobra kills 10,000/yr
spitting cobra can blind by spitting in eyes
Tx Coral Snake
small mouth, short fangs

b. black mamba
may be world’s deadliest snake
one bite can kill a person within half hour

b. sea snakes (HYDROPHIIDAE)

are similar to cobras
most venomous of all snakes
 100 x’s more toxic than that of any
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 32
 other kind of snake

c. pit vipers
in US include rattlesnakes,
water moccasins, copperheads
fangs fold back when not in use
use pits to track warm blooded prey
 as effective in dark as daylight

Reproduction

some snakes have elaborate mating rituals

eg. red-sided garter snake

when a female garter snake emerges from

hibernation she releases a pheromone that
attracts hundreds of male snakes

they rush to her to create a large squirming

“mating ball”

mating of these snakes is a tourist attraction in

Manitoba, Canada

most snakes lay eggs

but pit vipers bear live young

3. Crocodiles & Alligators

21 species

mostly unchanged for 200 MY

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 33
 largest of the living reptiles
 up to 6M (18’)

large robust skull with massive jaws with powerful

closing muscles

the group is unlike any other living reptile group

distant cousins of dinosaurs (diapsid like dinosaurs)

more closely related to birds than other living reptiles

 have diaphragm for breathing

 bird-like breathing mechanism with air sacs
 have 4 chambered heart
 only reptile group that can make vocal sounds
 show parental care

two groups:
crocodiles  long slender snout; more aggressive

alligators  shorter, broad snouts; much less aggressive

crocodilians have scent glands and produce a

strong musk during mating season
glands in mouth and throat with slit like opening to
outside of neck

another pair within cloacal slit

the only living reptiles that can make vocal sounds

 vocal sacs on each side of throat

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 34
  bellowing mating calls

one of only a few reptile groups that show

parental care

lays 20-50 eggs per nest

tends to and protects eggs & hatchlings

gender of offspring is temperature dependent

low nest temp  females

higher nest temp  males

4. Tuataras (O. Sphenodonta)

only ~50,000 individuals survive

2 living species (endangered)

found in New Zealand; heavily protected

resemble iguanas; ~ 16” long

live in burrows

features are similar to fossils 200MY old

retain the most primitive reptilian traits

well developed parietal eye with cornea, lens & retina

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 35

 but buried beneath layer of skin

 can only detect light intensity

complete palate (nasal cavities separate from mouth

cavith

teeth fused to jaw (not in sockets)

unlike most reptiles, tuataras remain quite active

even at near freezing temperatures

live long lives; routinely to 100 yrs; some suggest

they may live up to 200 yrs

don’t reach sexual maturity until 15 or 20 yrs old

apparently can produce offspring after 100 yrs

old

females require 2 or 3 years to grow a clutch of

eggs internally; and then for another7 or 8
months more after they are fertilized

after the female finally lays the eggs they

remain in the ground for another year
before hatching

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 36

 Humans Impacts
1. Poisonous Snakebites

5 Million people are bitten by poisonous snakes

each year

causes at least 100,000 deaths and up to 400,000

amputations/yr
esp India, Pakistan & Mideast

very few have access to adequate medical care

and antivenoms are in very short supply
-2009, WHO declared snakebite a neglected disease

but in US the average American is more likely to

be killed by another person than to be bitten
by a venomous snake
8000 bitten/yr in US (~17,000 homicides)

99.8% chance of survival (~80 die/yr)

Travis county has 5 venomous snakes:

Texas Coral Snake

- small mouth short fangs; coral snakes are nocturnal;
only bite under unusual circumstances; have
up to 12 hours to get antivenom
(red touch yellow  kill a fellow
red touch black  friend of Jack)
Western Cottonmouth
- can bite underwater
Western Diamondback Rattlesnake
Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 37
 - responsible for more human deaths than any other
N. Am. snake
Blacktail Rattlesnake
- rare
Brown Banded Copperhead

most are mistrustful of snakes in general because

a few are dangerous

snakes are much more of a benefit than a threat

eg. keep rodent populations in check

humans are much more of a threat to reptiles than

they are to us

More than 20% of the world’s reptiles are now at

risk of extinction

2. Medical Research

the regenerative abilities of reptiles is under study

for possible clues to organ replacement in
humans

3. Pharmaceuticals
eg. toxins from a Brazilian viper have provided the key
ingredient in a class of drugs called “ACE inhibitors” used to
lower blood pressure

eg. protein in black mamba venom is as effective as morphine in

relieving pain without its side effects (now under
development)

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 38

 eg. protein found in Gila monster venom is used as treatment for
type II diabetes

4. Farmed Reptiles – semi-domesticated

2.6 Million crocodiles are produced each year

worldwide for food and hide

5. Reptiles as Food

eg. sea turtles

such as Kemps Ridley sea turtle

eg. Kemps Ridley sea turtle

1947: 40,000
1985: ~200
1994: 580

they like the same beaches we do

hunted for eggs and meat

in Mexico armed troops must guard

beaches during nesting season

also affected by pollution & fishing

US now requires exclusion devices on
shrimp nets

eg. Alligator meat comprises about 1/3rd of

commercial harvests in US

eg. snakes

6. World Trade in Live Reptiles/ Pet Trade

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 39
 Because reptiles are traded for such a wide variety
of reasons, there are many hundreds of
species in trade.

millions of live reptiles are sold each year for the

pet trade.

eg. In 2001 the United States imported just under

2 million live reptiles.

eg. The United States annually exports more than 8 million

red-eared slider turtles (Trachemys scripta
elegans), the world's most commonly traded live
reptile.

eg. over 500,000 were green iguanas (Iguana

iguana) from Central and South America.

eg. Other species commonly found in the pet trade include:

boa constrictor (Boa constrictor)

ball python (Python regius)
panther chameleon (Chameleo pardalis)
red-footed tortoises (Geochelone carbonaria)

reptiles are among the most inhumanely treated

animals in the pet trade.
90% of wild-caught reptiles die in their first year of
captivity because of physical trauma prior to
purchase or because their owners cannot meet
their complex dietary and habitat needs.

Because they are cheap and easily replaceable, dealers,

captive breeders, and retailers factor huge mortality
into their operating costs.

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 40

7. World Trade in Reptile Products

eg. exotic skins trade. reptile hides are used in the

“luxury fashion” trade. PETA: virtually every store
that sells exotic skins has some hand in their illicit
trade

eg. alligators are bludgeoned to death with hammers

and steel rods,

snakes and lizards are decapitated and skinned

pythons are studdend (not killed) hoses are

inserted into their mouths and they are pumped
full of water to swell up to loosen their skin, then
each snakes head is impaled on a hook and the
animal is skinned alive

eg. curios and jewelry. In many parts of the world,

"tortoise shell" curios and jewelry, which are actually
made from the shells of hawksbill sea turtles, remain
popular, as do leather items made from snakes,
lizards, and crocodilians.

eg. Tortoises and turtles fetch high prices in Asian markets

- especially in China - where their meat is eaten and
their shells are used to make traditional medicines.

eg. American Alligator; brought back from near extinction

$30M hide business now

8. Herbal Medicine
tons of turtles are harvested because chinese believe that eating
turtles will lenthen lifespan

turtle blood is available at Walmarts in China

turtle heads are consumed for labor pains

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 41

 powdered snake gall bladder is used as a cure for bronchitis

coin shakes are coiled up like a stack of coins with head on top

they are boiled into a thick black liquid that is sipped like
tea for general health

lizards are taken to treat high blood pressure

9. Industrial Products
eg. Gecko Tape

eg. special ‘hairs’ on the feet of geckos give them the power to
climb on walls and ceilings carrying up to 400 x’s their own
weight

tape that mimics the feet of geckos is now commercially

available

10. Invasive Species

eg. brown tree snakes

bioinvader of islands (eg Hawaii)

has wiped out numerous species of birds

and mammals

Animals: Phylum Chordata-Reptiles; Ziser Lecture Notes, 2015.11 42

 Phylum Chordata – Vertebrates
Amphibia
~6,000 species

one of the most significant events in vertebrate

evolution was the gradual movement from water
to land

1st vertebrate group to make transition onto land

(=tetrapods)

bacteria  arthropods  plants  amphibians  reptiles

& algae 420MY 400MY 370MY 280MY

modern amphibians still retain a unique blend of

aquatic and terrestrial characteristics

Life in Water versus Life on Land

whereas fish are adapted to an aquatic lifestyle;

all other vertebrate groups are adapted to life on
land

basic differences between water and land:

1. air contains 20x’s more oxygen than water

also with faster diffusion rate

but respiratory surfaces must be kept moist

2. air is 800x’s less dense than water

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 1
 density of water = 1g/cm3
density of air = 0.001g/cm3

water is harder to move through but does

buoy up the body

land animals need strong limbs and remodeled

skeleton to get around

appendages must be able to support body

3. air fluctuates more in temperature

ocean temperatures are constant

land has harsh seasonal cycles of freezing and

drying

4. land offers numerous new, unoccupied

habitats and untapped food resources:
eg. terrestrial arthropods and plants

5. virtually no predators on land yet

New adaptations for land life:

some other adaptations that made their appearance

after the transition onto land:

there was an increase in the number of blood

vessels supplying the respiratory organs
Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 2
 development of a pumping mechanism to get
air into and out of lungs

the ear became more important as a sense

organ

need lacrymal glands to keep eyes moist

need new method of prey capture; cant use

suction effectively

 moveable tongue now used to manipulate

food in mouth

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 3

Origin of Tetrapods (4-legged Vertebrates)

by Devonian (~400 MY ago) bony fish had developed

a significant presence in freshwater habitat

~360MY ago the earth was becoming dryer with

alternating droughts and floods

during these dry periods freshwater ponds & pools

often dried up

lungfish in Siam today spends up to 4 months per year buried in

damp soil, 2-3 ft deep

fishermen collect them with spades

some bony fish (=lungfish) living in these freshwater

habitats had lung-like sacs that allowed them to
breath air for short periods of time as well

reinforcements in their fins later enabled them to

support their weight better in shallow water and,
for short periods, on land

 lungs and limbs were originally

adaptations for fish to continue to
survive in water

amphibians are descendants of these fishes

genetic studies have recently (2007) shown that fingers first
appeared in lobe finned fish and have identified the genes that
produced “fingers” from fish fins before the origin of amphibians
(tetrapods)
Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 4
The First Amphibians

the earliest amphibians (Tiktaalik, 375 MY;

Ichthyostega, 360 MY) share many features
with these fish (Eusthenopteron):
1. both ~ 1 M long and lived during Devonian

2. skull structure was very similar

3. had “third eye” (pineal eye)

4. had middle ear that could hear sound vibrations in air

Ichthyostega actually had an ear design that allowed it to

hear better underwater than on land

 probably spent considerable time in water

5. had similar short conical teeth (=labyrinthodont); probably

predators

6. had short stocky but flexible appendages with digits

7. tail had tail fins with fin rays

8. had bony operculum (but no internal gills)

9. had lateral line system

but transition wasn’t complete

 most amphibians still need moist environment

 most must return to water for reproduction

eggs must be laid in water
Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 5
 immature stage is aquatic

once the first amhibians appeared the climate became

warmer and more humid (carboniferous)
land was covered with vast fern forests

primitive insects, some flying insects

amphibians were the dominant land animals in the

carboniferous (300MY ago)

= Age of Amphibians
-------------------------------------------------------------

most amphibians today move from pond to pond for

food during droughts

live and breed in protected moist areas:

under longs and rocks
under litter on forest floor
in flooded tree holes

some modern amphibians have adapted to a

dryer land existence:

a few don’t require water for reproduction

largest:

largest frog:

African bulfrog, Gigantorana goliath

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 6
  30 cm (~1’) long, nose to anus; 7.5 lbs;
eats prey as big as rats & ducks
[largest US bulfrog gets to 20 cm (<8”)]

largest salamander:

Chinese giant salamander

 up to 6’ long 55 lbs (lives 50 yrs iin

captivity)

Japanese giant salamander

 can get up to 4.5’ long

smallest: cuban frog

 less than 1 cm (.5”)

Body Form

three main basic body forms:

eg. salamanders: head-trunk-tail

eg. frogs: fused head-trunk, no tail

eg. caecilians: long slender snake-like body

no limbs, no post-anal tail

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 7

Skin

most with thin moist, glandular skin without scales

(~1.5-4 mm vs humans 30-80x’s thicker)

doesn’t provide much protection from abrasion,

dehydration or predators

thinness of skin and vascularization allows it to be

used for respiration if kept moist

often with many glands:

eg. mucous glands

make skin slippery  harder for predators to
get a hold

eg. poison glands

usually concentrated in areas behind eyes

when stressed poison gland secretes toxin

skin is often brightly colored

contains chromatophores in dermis

many can adjust their color for camoflage

many toxic amphibians are brightly colored as

warning coloration

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 8

 less toxic species use color for camoflage
darkening of skin color controlled by light sensitive pineal
eye which is connected to pineal gland

 triggers release of MSH from pituitary gland

Support & Movement

stronger skeleton, mostly of bone, supports body

weight on land

but legs don’t support body very well

limbs are low to the ground

body touches ground at rest

legs are not very flexible

still move in very ‘fish-like’ fashion

skeleton provides rigid framework for muscle action;

esp leg muscles
 muscle mass shifted from trunk to legs

strengthened rib cage and axial skeleton to support

internal organs
abdominal organs hang down from axial skeleton which bears
most body weight

limbs with toes for easier land locomotion

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 9
 as the first vertebrates with “legs & feet” there was apparently
some experimentation with the number of digits

early fossils are found with 5,6,7, or 8 toes

most modern amphibians have 4 toes on forelimbs and 5

toes on their hindlimbs

(almost all later tetrapods had five digits on all limbs)

 made up of the same set of bones found in all land

vertebrates

most muscles have lost the “segmentation” seen in

fish

instead muscles are modified into “opposing

pairs” to flex/extend or abduct/adduct limbs,
etc
some of the trunk muscles still retain some of the
“segmentation”

swimming:

aquatic forms have fish-like undulating swimming

motion

gliding frogs:
eg. Polypedates spp (Africa and SE Asia)

large webbed feet

can glide horizontally 30-40’ from a height of 40’

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 10

---------------------
another new structural innovation in land animals is
the presence of a “neck”

the added flexibility with an additional set of

muscles made the head much more flexible
necks are also found in some ancient fish species but most fish
lack a flexibe neck

Feeding and Digestion

most amphibians are predators (carnivores)

eat mostly insects

but some eat small mammals, birds, snakes, fish

& other frogs

some aquatic forms filter zooplankton from water

most have long flexible tongues for capturing prey

tongue of frogs is connected to front of mouth

free end produces sticky secretions to adhere

to prey

can very quickly catch insects

[some take <.5 sec to catch prey with tongue]

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 11

some amphibians have teeth to hold onto prey and
prevent its escape

food swallowed whole, not chewed

Respiration

adapations necessary for shifting from extracting

oxygen from water to extracting it from air
required major changes in both the respiratory
and the circulatory systems

amphibians can get oxygen in several ways:

a. lungs
b. through skin (cutaneous breathing)
c. mouth (buccal breathing)
d. gills

a. lungs

most amphibians have lungs

the lungs are derived from the lungs of fish

very simple lungs; essentially hollow air sacs

 amphibian lungs are not very efficient

[mammal lungs are >15 x’s more efficient]

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 12

 nostrils are now used for breathing as well as
chemosensory

nostrils open directly into mouth cavity

 cant eat and breath at the same time

and no diaphragm (breathing muscle)

 amphibians must gulp air to force it

into lungs

in most amphibians the lungs are not adequate for

getting the oxygen they need

most amphibians rely on additional structures

to supplement their lungs

b. skin

thinness of skin and blood vessels present allow it

to be used as respiratory surface
even when lungs are used for oxygen; most carbon dioxide is
lost through the skin

c. mouth

can also use mouth lining for respiration

some salamanders have dispensed with lungs and

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 13

 gills and use cutaneous or mouth respiration
only

d. gills

most amphibian larvae are aquatic and have gills

for respiration

some aquatic amphibians retain gills as adults

Circulation

air breathing also requires a restructuring of the

circulatory system

the amphibian circulatory system is improved over

that of fish

have 3 chambered heart; 2 atria, 1 ventricle

two complete circuits of blood flow

pulmonary circuit & systemic circuit

picks up O2 in lungs and returns to heart

then sends oxygenated blood to rest of

body

 much more efficient; heart is a

double pump
Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 14
 but not completely separated

there is some mixing of oxygenated and

unoxygenated blood in ventricle

Nervous System & Senses

the vertebrate brain is made up of 4 distinct functional areas:

cerebrum - higher brain functions, integration

cerebellum - coordination of movement

diencephalon - coordination of nervous and endocrine systems

brain stem - automatic internal functions

amphibian brain is about same size as fish relative to

body size

cerebrum (higher brain processing & sensory

integration), esp optic centers, are relatively
larger in amphibians than fish

cerebellum (controls movement) is relatively

smaller than fish

Senses:

a. lateral line

many aquatic species have retained the

lateral line system
in air there is not sufficient density to activate
receptors in lateral line
Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 15
 senses of smell and hearing became more
important than lateral line on land

b. use touch, pressure and temp are sensed

mainly by free nerve endings in skin

c. vision

vision is dominant sense in many amphibians

no longer a fixed open stare as in fish

eye is similar to ours with a few differences:

eye muscles to move eyeball in socket

has lacrimal gland and eyelids to protect from

drying

lower lid has a nictitating membrane

 sweeps over eye when blinking

accommodation (focus) by moving lens in and out

 not changing its shape as we do

retina has rods & cones  color vision

much visual processing occurs in the eye before

signals reach the brain

c. smell

smell has become more important

receptors still located in “nose” area

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 16
 smell due to olfactory epithelia in nasal cavities

also have “Jacobson’s Organ” in roof of

mouth

d. hearing & sound

our ear is divided into outer, middle and inner

portions
fish had only an inner ear

amphibians have both a middle and inner

ear

 eardrum is on outside of head, behind

the eyes

middle ear helps to amplify in air sound

sound waves in air are very weak

a single ear bone (=columella (stapes))

(not 3 earbones as in us)

transmits sound vibrations from eardrum

to inner ear

most amphibians have a larynx with vocal

cords

frogs pass air back and forth over vocal cords

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 17
 between lungs and vocal sac in floor of
mouth

use sound to attract a mate

better developed in males than females

 males do most of the calling

some sound is also transmitted through

forelimbs, muscles and soft tissues to
inner ear
 esp low frequency “seismic” vibrations

may warn of large predators

e. balance and equilibrium

like fish, amphibians have inner ear that

detects position and acceleration via
otolith organ and semicircular canals

Excretion & Salt/Water Balance

as in fish nitrogen wastes are eliminated as ammonia

or urea

skin and kidneys are the main way salts and water are
gained or lost
most amphibians cannot conserve water by producing a
concentrated urine

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 18

 a few desert frogs can produce uric acid

 requires much less water to eliminate

most amphibians can store urine up to 1/3rd body

wt in bladders and lymph sacs beneath skin

kidneys conserve salts by reabsorbing them from

urine to compensate

some amphibians can actually absorb water through

their skin

products from the digestive, excretory and

reproductive systems drain into a cloaca before
being released to the outside

Defense/Protection

amphibians have many enemies: snakes, birds,

turtles, raccoons, humans

 many frogs and toads in tropics are aggressive

and will fight predators

some can give a painful bite

 frogs tend to stay very still when threatened

only when they think they have been detected do

they jump in water or grasses to get away
when held, they remain motionless to catch us
Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 19
 offguard, then jump while voiding urine

 most frogs can also inflate their lungs making them

difficult to swallow

 all amphibians have poison glands in their skin

some toxins are lethal

eg. Poison Dart Frog

brightly colored (warning); one of the deadliest frogs

 poison from a single frog could kill several

humans

Choco indians of Central and South America catch

them and roast frogs over open fires then collect
the highly toxic mucus which exudes from the
frog’s skin as they die.

use the poison on the tips of their blowgun darts

eg. large toad of Panama Canal Zone can squirt a

poison that will blind

its skin is collected for fine leather

some frog toxins are hallucinogenic

(frog licking)

 a few amphibians use poisonous spines to to

protect themselves
eg. sharp ribbed newt when threatened can arch their back in
such a way that the sharp ends of their ribs actually
penetrate and poke out of the skin. As the ribs pass
Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 20
 through the layer of skin they are coated with a toxic
milky liquid to become venomous spines

eg. hairy frog does a similar thing but uses its toe bones as the
spines that it uses to slash at its attacker

 one ancient group of African frogs has poisonous

fangs that it uses to eat other animals including
other frogs

Reproduction & Development

dioecious; rarely show sexual dimorphism

mating is controlled by seasonal conditions

most amphibians breed soon after spring

emergence from hibernation

breeding season usually lasts for several weeks

no transfer organs or copulation; most amphibians

have external fertilization

eg. in salamanders male deposits spermatophore

on leaf or stick and maneuvers female over it

fertilization occurs as eggs are released

aquatic species lay eggs in clusters or stringy

masses

terrestrial species may deposit eggs in clusters

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 21
 under logs or in moist soil

in some salamanders, the adults guard eggs

eg. frog breeding is like an orgie

most larger frogs are solitary except during

breeding season

males often take possession of a perch near water

then males call to females

each species has its own unique call

amplexus: male frog holds onto female

female deposits eggs in water anchored by

sticky jelly

male deposits sperm over eggs

males will grab almost anything

often jump salamanders or other male frogs

have special release call to get males off

sometimes several males will jump on a female

many females drown from the weight holding
them under water

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 22

in most amphibians there is some parental care of the
eggs until they hatch by either the male or female

some amphibian species reproduce by

parthenogenesis

Metamorphosis

salamanders, eggs typically hatch into tadpoles in ~

1 week

with gills, suckers and spiracle

larvae resembles adult

aquatic forms retain gills

(paedomorphosus)
eg. Necturus, mud puppies

terrestrial forms lose gills and develop lungs

embryos of salamanders resemble adults

 undergo less pronounced metamorphosis

some retain gills as adults

frogs hatch as herbivorous tadpole larvae

most frogs undergo metamorphosis into adult in

a year or less
Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 23
 legs appear
tail is reabsorbed (in frogs)
lungs develop

one genus of tropical terrestrial frogs the eggs

hatch dirctly into “froglets”

no aquatic stage
__________

frogs & toads have a variety of unique reproductive

behaviors

a few tree frogs build nests: cuplike crates along

streambank

another makes waterproof depressions in tree

hollows using beeswax

some brood young in stomach

eg. Surinam toad; Pipa

completely aquatic

fertilized eggs are deposited on the back of female

the eggs sink into the spongy skin forming separate

incubation chambers

each chamber is covered by thin sheet of skin

larvae undergo metamorphosis in these chambers and

emerge as adult toads

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 24

 eg. midwife toad

female lays eggs fastened together like beads on a string

male thrusts hind legs into the egg mass and wraps them
around his body

male then takes eggs to his burrow

he comes out only at night to search for food

when larvae are about to emerge he finds a pool of water to

jump in and the larvae swim away
____________

a very few amphibians have internal fertilization

and bear live young
eg. the snakelike caecelians have internal fertilization, most bear
live young

fetuses feed on secretions and tissues they scrape from

lining of mom’s oviduct

eg. the Alpine salamander, Salamandra atra, lives in Swiss Alps

at ~4500’ and has the longest gestation period of any land
animal:

2 young are born ~ 3 years after fertilization

Hibernation

during winter most temperate frogs hibernate in

mud at bottoms of pools and streams
use energy from glycogen and fat stores

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 25

 toads tend to hibernate in humus on forest floor

some can survive freezing

eg. woodland frog is the only vertebrate able to survive being
frozen

they live north of the arctic circle

up to 65% of its body water may be frozen

heart stops completely

glucose in blood acts as antifreeze

what freezes is the water outside its cells, not water

inside cells

Migration

some amphibians have a strong homing instinct

they migrate to ponds to mate

 return to the same pond each year for mating

guided by olfactory cues

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 26

Kinds of Amphibians

3 main orders: “with tail” (O. Caudata)

“without tail” (O. Anura)
“without feet” (O. Apoda)

A. Salamanders & Newts (Order Urodela or

Caudata)
3 species of salamanders in Travis County
including the Barton Sprigs salamander, Eurycea sosorum

least specialized, resemble ancestor

limbs at right angles to trunk

 walk with “S” motion of trunk

most have aquatic larva that metamorphoses into

terrestrial adult

mainly in N America

mostly nocturnal

some up to 4’ long

live in cool damp places

including along mountain streams

 most cold tolerant of all amphibians

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 27

 a few are arboreal and avoid water

both larvae and adults are carnivores

eat worms, small arthropods & molluscs

some have teeth in roof of mouth

some with prehensile tongue up to half body

length

some without lungs or gill  breath through skin

some reproduce without metamorphosis

examples:

eg. Barton Springs salamander

eg. Hellbender (Cryptobranchus alleganiensis)

one of world’s largest salamanders (to 2.5’ (75 cm))

can live for 30 years

confined to clear stream and rivers in the Applachian

mountains

wrinkly skin through which it breaths secretes a mildly

toxic mucus to protect its thin skin from
pathogens

sometimes called a “snot otter”

feed on crayfish - swallow them whole

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 28

 once extremely common is now an endangered species
some states

has declined 77% in past 30 years

illegally caught and sold in pet stores in China and

Japan for up to $1700 each

B. Frogs and Toads (order Anura (=”no tail”))

17 species of frogs in Travis County

by far the most successful & widespread group

5283 species or 88% of all living amphibians

an ancient group
 known from Triassic (250 MY)

hind legs specialized for jumping

occupy a great variety of habitats

especially common in tropical swamps and

forests

but found in all habitats; even dry areas

frogs are more aquatic and generally live in or

near water

toads are more terrestrial and only move to water

to reproduce

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 29

 more dependent on lungs than other amphibians

those that are completely aquatic usually lack

tongue

highly specialized for jumping locomotion

some can glide like flying squirrels

eg. flying frog of tropical Asia

most have long flexible tongues attached to the

front of the mouth for capturing prey

tree frogs have large, adhesive pads on the ends

of their toes

C. Caecilians (O. Gymnophiona; Apoda)

~173 species

elongated, limbless, burrowing or aquatic animals

10 cm to >1.5 M long

in tropical forests of central and south America,

Africa, India

skin is smooth and slimy

but some with small calcified dermal scales

under skin
Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 30
 also skin has folds that make them look like
large segmented earthworms

small eyes  most ar blind as adults

sensory tentacles on snout

feed on worms and small invertebrates

skin has squirt gland that secretes irritant

causes sneezing in humans

internal fertilization, most viviparous

fetuses feed on secretions and tissues they

scrape from lining of moms oviduct

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 31

 Ecology & Human Interactions
with Amphibians

A. Beneficial Effects of amphibians

Frogs eat disease-carrying insects

Frogs are critical links between predators and

the bottom of the food chain (algae, plants,
detritus, and such)

B. As Food

not a major part of human diet

frog legs

Americans devoured more than 6.5 million pounds

of frog legs a year (1984)

led to the death of some 26 million frogs

annually.

Ninety percent came from India and Bangladesh,

which banned exports after frog declines led to
growing hordes of mosquitoes, malaria, and
increased use of pesticides.

Now Indonesia supplies most of the frogs for

restaurants

C. Education & Research

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 32
most commonly dissected laboratory animal:
in science classes and research

3-9 M US for education alone

6 M in high schools alone

3 M frogs (8% of all lab animals) are used for research

 much of our medical knowledge came from

frog dissections

 embyrological studies

 isolation of pharmaceuticals

D. Poisons

several species of tropical frogs secrete potent

neurotoxins
distasteful
induces paralysis

often brightly colored

natives in Brazil and Costa Rica use toxin to make

poison arrows

some of these toxins are hallucinogenic

leads to “frog licking”

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 33

 eg. Poison Dart Frog

one of the deadliest frogs

 poison from a single frog could kill several humans

E. As environmental Indicators

amphibians are extremely sensitive to

environmental indicators

in 80’s & 90’s noted declines

 since 80’s 120 species have become extinct

today one third of the worlds 6,000 amphibian

species are threatened

 one of largest extinction spasms in

vertebrate history

unsure of exact causes of declines:

Probable causes of decline:

1. The number one cause of amphibian decline is

habitat loss

most amphibians feed and breed in wetlands,

In the past half-century the lower 48 states

have lost more than half of their estimated
original wetlands.
Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 34
 2. pollution

deformities from animals in polluted water

3. fungal skin infection

most recently has been tied to worldwide

spread of a primitive mold pathogen

this is the first mold known to attack

vertebrates.

can kill within days

 Batrachochytrium dendrobatidis (chytridiomycota)

(including in and around central Texas)

spreads very rapidly; don’t know how it kills

frogs
Barton springs salamander has natural antibiotics in its
skin that seem to protect it from the pathogen)

scientists have mobilized to collect and save

representative species in safe haven
protected from fungus

4. deadly virus is the likely culprit in several

recent die-offs of frogs,

5. Increased exposure to ultraviolet radiation may

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 35
 damage the eggs

6. possibly caused by acid precip, deforestation

urbanization, climate change

the largest captive breeding program ever,

“Amphibian Arc” has begun to save the 500 most
endangered amphibians that cannot be protected
in the wild

Animals: Phylum Chordata-Amphibians; Ziser Lecture Notes, 2015.11 36

 Phylum Chordata: Vertebrates
The Fish Classes
Introduction
eg. lampreys, hagfish, salmon, trout, sharks, rays, tuna,
sardines, flounder, seahorses, catfish, etc etc

fish are the most diverse and successful group of

living vertebrates

 almost half of all vertebrate species

 ~28,000 living species

 ~200 new species described each year

while fish are by far the most abundant and diverse of

all vertebrate groups

they remain the least known group of vertebrates

eg. estimates are that we have collected and described only
slightly over 1/3rd of fish species in the Amazon river
basin
_________________________________________

fish are also the oldest known vertebrates

 the first true vertebrates appeared in the fossil

record 530 MY ago (early Cambrian)

 a fossil shark species from 310 MY ago was the

first vertebrate known to migrate

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 1

 it lived in rivers but swam to the sea to breed

the first fish appeared over 500 MY ago and were

small bottom-dwelling animals without jaws,
teeth, or paired fins

probably sucked up small particles of food from

the bottom sediments

for ~50 M years they were the ONLY vertebrates

on earth

they eventually developed primitive fins that

improved their swimming ability

they also developed heavy bony armor and

roamed in this form for over 20 MY

eventually most fish lost the heavy armor,

developed more streamlined bodies, jaws and
paired fins

this “modern” design produced almost all the

varieties of fish we have today
_______________________________________

all fish are aquatic & and highly adapted

for aquatic life:

fish occupy virtually every kind of freshwater and

saltwater habitat

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 2

 there are no terrestrial fish
although some can survive considerable time outside
of water and can often be found crawling on land

eg. “walking catfish”

the closest to terrestrial is the “mangrove killifish”, normally

lives in muddy mangrove swamps but when they dry
up the fish can be found stuffed into insect tracks in
rotting logs by the 100’s

size of fish:
smallest fish (also, smallest living vertebrate)

= stout infantfish, Schindleria brevipinguis, (Australia)

males 7 mm long (~1/4th “), female 8.4 mm and
weighs 1 mg

also, pygmy gobi <1/2”

largest fish = whale shark to ~50’, rumors to 70’ (40 tonnes)

most fish continue to grow throughout life

(birds & mammals stop growing at adulthood)

generally growth is temperature

dependent

annual rings are produced in scales, otoliths

and other bony parts

 the age can be accurately determined

Skin
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 3
consists of two layers:
epidermis
dermis

epidermis usually secretes slimy mucous

the slimy skin reduces friction

 up to 66%

 improve swimming efficiency

dermis sometimes tough and leathery or relatively

thin

dermis produces scales in most fish

the scales are under the slimy epidermis

the scales are made of enamel and sometimes

dentin; the same basic components as our
teeth

Skeleton

skeleton begins as cartilage framework

in most fish the cartilage is replaced by bone by

adulthood

bone is a tissue unique to the vertebrates

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 4
 originated as an external protective covering

only later evolved into an internal supporting framework

highly flexible “backbone” of cartilage or

bone is the main support and framework for
swimming muscles
# of vertebrae on a fish varies from 16 to >400

most fish have ribs but they are not used for
“breathing”

 they support swimming muscles

most fish have bone or cartilage that supports the

dorsal, ventral and caudal fins (unpaired fins)

also, most fish have paired appendages in the form

of pectoral and pelvic fins

=”appendicular skeleton”

 homologous to our arms and legs

act as rudders, for balance, feelers, weapons, sucking

discs, lures to attract prey

Movement

most of a fish’s body mass is bundles of segmented

muscle tissue = myomeres (=myotomes)

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 5

 relatively small body cavity for other organs

muscles are segmented

 zig-zag “W”-shaped bands of muscles

along sides of fish

muscles are mainly for swimming

although some can walk, crawl, burrow, and “fly”

less energy is required for swimming than most

other forms of vertebrate locomotion
 don’t need to fight gravity

Energy Consumption/kg body wt/km:

swimming: 0.39 kcal (salmon)
flying: 1.45 kcal (gull)
walking: 5.43 kcal (squirrel)

 water is 800x’s denser than air

don’t need to also fight gravity as much

each myomeres consist of short muscle fibers connected to

tough connective tissue that are also attached to the
next myomere

each myomere extends across several vertebrae

allows more power and fine control

myomeres produce “S” shaped swimming motion

fish actually “push” on the water

water is relatively dense
non compressible
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 6
the fastest fish exchange the snake-like motion for
more rigid position where most of the flexing is
toward the tail only
eg. tuna doesn’t flex body at all; all thrust is from the tail

overall, swimming speeds are not particularly fast

compared to running or flight due to the high
density of water

eg. 1 ft trout  6.5 mph

eg. 2ft salmon  14mph

the larger the fish the faster it can usually swim

barracuda is fastest fish  27 mph

usually cruising speed is much slower

most speeds reported for fish are speeds as they jump out of
water so they appear to be much faster

Maximum Speeds
sailfish 68mph
swordfish 40-60mph
marlin 50mph
bluefin tuna 30-44mph
bonito 50mph
wahoo 40-49
salmon 25mph
mackerel 20mph

flying fish can glide above the water 20-40 sec

dorsal and ventral fins improve swimming

efficiency

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 7

Feeding & Digestion

after the evolution of jaws, fish were freed from

deposit feeding and filter feeding

most modern fish are carnivores

small, numerous, sharp teeth are used to seize prey

most have very flexible jaws to engulf large prey

some can eat prey as large as they are

most fish produce new teeth continuously throughout

life

a few fish that scrape algae from hard surfaces have

teeth that can bend (rather than break) as they
scrape

most fish lack moveable tongues

if they do have tongues they are not moveable

fish swallow their food whole; they don’t “chew”

chewing would produce pieces that might clog their gills

intestine became longer to provide greater surface

area for absorbing a variety of nutrients

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 8

Respiration

most fish have gills for getting O2 from water

=thin feathery sheets with lots of blood vessels

some fish can also breath through their skin

a few fish can breath air

one fish (bearded goby) flourishes in a dead zone off the coast of
Namibia

can survive in waters where oxygen is <10% and toxic

hydrogen sulfide levels are high

 able to slow their metabolism

they return to surface waters at night

Circulation

the circulation of blood is closely tied to gas exchange

through gills

blood is pumped through arteries and veins by

simple heart

fish have a 2 chambered heart

blood flows through a single circuit:

heart arteriescapillaries veins back to heart

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 9

 blood is first pumped through gills then out to
the rest of the body

Body Temperature/Thermoregulation

fish are cold-blooded; ie poikilotherms

the body temperature of most fish is the same

as their environment

some fish maintain a higher temperature in their

swimming muscles

 as much as 20º C warmer than surrounding

water
eg. tunas, mako sharks, swordfish

other fish elevate temperature of brains and retinas

eg. marlins

these higher temperatures promote swimming and

improved nervous activities

also promote much more precise vision and faster

reflexes in predators
 such fish are some of the fastest in the world

Nervous System

fish brains are relatively small and simple compared to

other vertebrates
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 10
 but still considerably more developed than in the
invertebrates

brain is made up of several distinct functional areas:

cerebrum (higher centers) very small
cerebellum (coordination of movement) relatively large
brain stem (automatic activities) also relatively large

fish do sleep

 stay motionless for several hours

some marine species (eg. wrasses, Labridae) may bury
themselves in sand or spin “sleeping bags”  cocoons of
mucus each night to sleep

Sense Organs

in the fishes environment:

light doesn’t travel as far in water

also turbidity

sound and pressure waves travel faster and further

1. lateral line system = “distance touch”

probably the most important sense in fish

most fish depend mainly on lateral line system

for sensory information to detect food or
danger
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 11
 a set of interconnected tubes and pores along
sides of head and body

contains:
mechanoreceptors  water movements
 sound: can pick up unusually low
frequency sounds (.1-200 Hz)

electroreceptors  detect body electricity of other fish

and prey

chemoreceptors  equivalent to our sense of smell

2. eyes

paired immoveable eyes

most fish lack eyelids

in the eyes of vertebrates a circle of muscle (=ciliary muscle)

focuses the lens for near and far vison

in most fish the muscles of the lens are relaxed for near
vision

in sharks (as well as amphibians and snakes and humans)

eyes are relaxed for far vision

also, unlike our eyes that stretch the lens to

change focus; muscles in fish eyes move
lens forward or backwards to focus

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 12

 some variations in eyes of fish:

one eye of flounders migrates to same side as

other eye during embryonic development

a few freshwater fish have “bifocal” lenses to see above and

below the water at the same time

 can enhance prey perception

fish can see in color

can also detect polarized light for navigation

in dimmer waters, fish eyes get proportionately

larger

in complete darkness they are small or absent

(eg. cavefish)

3. Chemoreceptors (smell and taste)

chemicals travel well in water

for fish there is no clear distinction between smell

and taste

chemoreceptors are located in:

mouth
around head
in some over entire body
catfish and loaches on barbels around mouth
(barbels also for touch)

olfactory sacs (=nose)

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 13
 fish don’t “breath” through their nose

 nostrils are just sensory pits

most fish have a good sense of ‘smell’

prey detection

communication (with pheromones)

fish lack actual ‘taste buds’ (like we have)

but can apparently discriminate between bits

of food and trash that they take in
 will expel unwanted items that they eat

5. hearing

we are finding that sound is an important means

of communication in fish, especially deepwater
fish

fish have an inner ear with distinctive bones that

vibrate when sound waves pass through and
send a signal to the brain

4. touch

barbels in catfish, goatfish, whiting, etc

5. orientation and balance

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 14
 otolith organ and semicircular canals

for detecting orientation and changes in

motion

Excretion

kidneys like other vertebrates remove most metabolic

wastes (Nitrogen wastes)

they also play a role in salt/water balance

gills also play role in excretion

 secrete NH3 and other N wastes

Osmoregulation (salt & water balance)

the body fluids of all animals must either be in balance

with their environment or they must have
mechanisms that allow them to regulate their own
salt and water balance

almost all marine invertebrates have body fluids with

the same salt concentrations as seawater (3%)

=Isotonic to their environment

freshwater fish

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 15

 have salt content ~100 x’s greater than the
freshwater they live in

= Hypertonic
[fw salt conc=.001-.005 Moles; freshwater fish = .2-.3 M]

there is a tendency for them to gain water

and lose salts

marine bony fish

most marine fish alive today actually evolved from

freshwater fish

so they have salt content about one third that of

seawater

= Hypotonic

marine fishes has a tendency to lose water

and gain salt

scales & mucus make a fish’s body almost completely

impermeable to water

 most gain or loss of water and salt is

through gills or the lining of the mouth

to maintain salt & water balance:

freshwater fishes:
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 16
 must avoid drinking
urinate often
absorb salts ions through gill and from food

marine (bony) fishes:

drink large amounts of seawater
produce little urine
while pumping out excess salts from gills

because of these osmotic limitations, 90% of all fish

are restricted to either freshwaters or salt water

only 10% of fish are able to easily move between

freshwater and salt water = euryhaline fish

 can actively regulate to maintain salt/water

balance in their internal fluids

Reproduction & Development

only a few are hermaphrodites

only a few reproduce by parthenogenesis

most fish are dioecious

genders usually cannot be distinguished externally

most with external fertilization (oviparous)

a few have internal fertilization

some (eg. guppies) can store sperm for months

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 17
 a few with internal fertilization bear live young

most fish produce large numbers of eggs:

 less than 1/million will survive to maturity

most fish spawn at certain times of the year

 spawning is temperature dependent

 temperature is critical for survival of eggs and young

in most marine fish:

males and females come together in great schools

and release millions of egg and sperm
(probably <1/M actually survive)

eggs become part of the plankton through

embryonic and larval development

freshwater fish often have elaborate mating behaviors

many make nests

some bear live young

some show parental care

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 18

Kinds of Fish:

28,078 species

three different classes of vertebrates are categorized

as “fish”:

1. jawless fish (Agnatha)

108 species

eg. Hagfish, eg. Lampreys

2. cartilaginous fish (Chondrichthyes)

970 species

sharks, stingrays, manta rays, sawfish

3. bony fish (Osteichthyes)

27,000 species (96% of all fish)

most abundant living group

2 main types: ray finned fish & lobe finned fish

eg. bass, perch, catfish, herring, goldfish, some eels,

clownfish, sailfish, etc.

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 19

 The Agnatha
(Jawless Fish)

(108 living sp)

remnants of the earliest fish in the fossil record

oldest known vertebrates

not technically “vertebrates” since they have no vertebrae – just
a cartilage rod for support

but they are clearly relatives of the other fish groups

most ancient & primitive vertebrate group

over 500 MY fossils of jawless fish

fossil agnathans are found >500 MY
include extinct conodonts & ostracoderms

only living vertebrate group with no jaws

also lack paired fins

three main groups of agnatha:

ostracoderms - all extinct

hagfish

lampreys

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 20

Ostracoderms

circular or slit -like mouth opening without jaws

some of the earliest vertebrate fossils are in this group

(late Cambrian to Ordovician)
recently Chinese fossils pushed date back to 530 MY

flourished for 150 MY

became extinct at end of Devonian

fish armored with heavy dermal bony plates

probably for protection against eurypterids and cephalopods

no paired fins

probably stayed close to bottom

probably deposit feeders or filter feeders

may have filtered water for food

but not with cilia & mucus as in cephalochordates

used pumping action of muscular pharynx

a few may have even been predators

a later group of ostracoderms had paired

pectoral fins

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 21

  greatly improved swimming ability

also seemed to have a more sophisticated

nervous system and sense organs

 similar to todays lampreys

General Characteristics of living Agnatha:

1. all modern surviving relatives of these ancient

forms have lost their bony armor

2. can be freshwater or marine

3. long eel-like body form

4. no jaws

5. no paired fins

generally poor swimmers

6. smooth slimy skin, no scales

7. skeleton is a simple rod of cartilage = notochord,

no bone

8. gills located inside 5-16 prs of pore-like gill

openings

cartilage gill supports hold the gill-slits open

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 22
 spiracle at top of head can draw water in and over
gills

9. poor sense organs

10. parasitic or scavengers

two main groups today:

hagfish
lampreys

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 23

 Hagfish
~65 species

all are marine

about 18” long

largest known is almost 4’ long

found in deep waters

almost completely blind; eyes have degenerated

Feeding & Digestion

mainly scavengers

eat dead or dying fish, also molluscs, annelids,

etc

small eyes

although almost blind they can quickly find food

by touch and smell

enters dead or dying animal through an orifice or by

actually digging into the animal

has 2 keratinized toothed plates on its tongue

rasps bits of flesh from carcass

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 24
recently (2011) found that hagfish can absorb food
directly into their body through their skin

Circulation

has unusually low pressure circulatory system

with 1 main heart and 3 accessory hearts to boost

blood pressure through gills

Excretion

only vertebrate with body fluids the same

concentration as salt water (invertebrate trait)

Protection

secrete copious amounts of slime (500 ml/min)

 milky fluid from slime sacs along sides of body

 on contact with seawater forms a very slippery

material making them impossible to hold
can secrete enough slime to turn a bucket of water
into a gel in a few minutes

 can clog the gills of predators

 may be able to extricate themselves from jaws of

predator by “knot tying” behavior

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 25

Reproduction

breeding habits are still relatively unknown

some are apparently hermaphroditic

animal has both male and female gonads but only

one or other is functional

no larval stage as lampreys have

Conodonts

numerous fossils of small tootlike structures have

been known for years

common  used as index fossils to date sediments

weren’t sure what animal produced them

until 1980 when fossils of complete animal
were found

was an agnathan, apparently most closely

related to hagfish
had toothlike plates in throat area

notochord

swimming muscles as myomeres

paired eyes

similar inner ear structure

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 26
Human Impacts

bane to some commercial fishermen using gill or set

nets

by the time they pull catch in hagfish have

often devoured internal contents of fish

not often a pest with trawl fishing

today hagfish are collected for “leather” to make golf

bags and boots

their slime has unusual properties since it is reinforced

with strong, stretchy, spidersilk-like protein fibers
 looking at it for potential uses for stopping bloodflow in
acidents and surgeries

 also investigating its use for parachutes, packaging and

clothing

 to make synthetic fabrics similar to nylon and polyester which

are petroleum based

some species are in serious decline due to over

harvesting

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 27

 Lampreys
38 sp.; 20 sp. in North America

most 15 - 60 cm; up to 1 M long

Feeding & Digestion

most are parasitic others don’t feed as adults

parasitic adults attach to prey by sucker like mouth

rasp away flesh with teeth to suck out blood

inject anticoagulant

when finished lamprey releases its hold

host sometimes dies from wound

Reproduction

all spawn in winter or spring

 in shallow freshwater streams

male builds nest

 uses oral disc to move stones and make a

shallow depression

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 28

female joins him

female attaches to rock to hold position in current

male attaches to female and fertilizes eggs as they are

released

adults die after spawning

eggs are sticky, covered with sand

in 2 weeks eggs hatch into ammocetes larvae

 close resemblance to amphioxus (cephalochordate)

larvae drift downstream and burrow into sand

larvae are suspension feeders for 3-7 years

feeding on microscopic organisms

then rapidly metamorphose into adults:

larger eyes
oral disc with rasping teeth
nostrils shift to top of head
body shorter and rounder

all lampreys migrate up freshwater streams to breed

eggs and larvae develop in freshwater

young of marine species then migrate to ocean

until sexually mature
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 29
  others remain in freshwaters their entire lives

Human Impacts

Petromyzon first invaded the great lakes in 1913-1918

(bioinvasion)

in 1950’s destroyed great lakes fisheries

rainbow trout, whitefish, lake herring, and other

species populations were destroyed

their numbers began to decline in early 1960’s

due to depleted food

expensive control measures

 expensive larvicides placed in selected

spawning streams

today, some native species have been restocked and

are now thriving again

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 30

 Jawed Fish
Origin of Jaws & Paired Fins

oldest known jawed vertebrates appeared 420 MY

(Silurian); about 100 MY after the jawless fish first
appeared

 placoderms and acanthodians

Jaws

evolution of jaws was one of the major events in the

history of vertebrates
 freed from bottom feeding

 allowed access to a much greater variety of food sources

eg. predators

jaws evolved from gill supports (gill arches)

 in certain primitive fish the jaws resemble gill arches

 in shark embryology jaws do develop from gill arches; can

watch it happen

 cranial nerve branching and placement in jaws and gills of

cartilaginous fish resembles each other

initially, jaws just “closed the mouth”

jaws allowed feeding on larger foods:

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 31

 seaweeds, plants and animals

later jaws became armed with dermal scales that

evolved into teeth

teeth could be used to seize prey

jaws allowed predation on larger active prey

around the same time that jaws allowed eating larger

pieces of food the stomach was formed from an
expanded pouch of the gut

Paired Fins

acanthodians and placoderms were also 1st fish with

paired fins

all living jawed fish (gnathostomes) have paired

pectoral and pelvic fins
pectoral fins appeared before pelvic fins

probably originated as stabilizers for swimming

might have begun as folds in skin along sides of animal

later got muscle attachments and became moveable

In addition to moveable jaws and paired fins,

all jawed fish also have paired nostrils and

3 semicircular canals for equilibrium
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 32
4 main groups of jawed fish:

A. Acanthodians (extinct)
B. Placoderms (extinct)
C. Chondrychthyes
D. Osteichthyes

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 33

A. Acanthodians

first group of jawed fish to appear in the fossil

record (430-260 MY)

most relatively small; 4-6”

acanthodians had scales rather than bony plates

most were covered with small, diamond

shaped dermal bony scales

jaws with small sharp teeth

were probably active swimmers

live on today as bony fish

once jaws & fins evolved there was an explosion in

fish diversity

 Devonian = “age of fishes”

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 34

B. Placoderms

appeared ~20 MY after the first acanthodians

most were medium sized; 1-2’ long

some up to 30’ (10 M) long;

heavily armored fish with dermal bony plates over

front 1/3rd to 1/2 of body

the rest of the body with small bony scales or

without armor

relatively poor swimmers

were the first vertebrates to have necks

separating their heads from forelimbs

allowed them to move their heads

independently

very diverse group

some were predators

others were bottom dwellers

some had crushing mouthparts

may have eaten shellfish

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 35

 one species recently found, Materpiscis attenboroughi, 380 MY,
is the earliest known example of a vertebrate giving live
birth. The adult fish was found with bones of a developing
fetus inside

placoderms and acanthodians went extinct at end of

Devonian

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 36

 Class Chondrichthyes
(Sharks and Rays)

970 species

Last of the 4 fish groups to appear in the fossil record

yet retain many primitive fish characteristics

many have changed little from earliest fossils

all but a few are marine

most are 6-15’ long

includes the largest fish and second largest of all

living vertebrates

whale shark  up to 60’ long

 filter feeder

great white gets up to 30’ long

Body Form

either fusiform (spindle shaped) = sharks

very streamlined shape  very good swimmers

or flattened = rays
spend most time on or gliding near shallow bottoms

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 37

Skin & Scales

skin is very tough & leathery

 muscles of shark pull on skin rather than

pulling on the skeleton

bony scales reduced to small, hard, knife-like

(placoid) dermal scales embedded in skin and
stick out from skin

made of bony tissue

scales have same structure as tooth including

enamel, dentin & pulp cavity

reduced scales enhance swimming efficiency

scales are continuously shed and replaced

throughout life

Support

all members of the group have a skeleton made

mostly of cartilage

although their ancestors had bony skeletons

but sharks retained mineralization in teeth, scales

& spine

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 38

the cartilage makes the body lighter and more
bouyant

Muscles & Movement

sharks are the most graceful and streamlined of all

fish

sharks have excellent swimming ability

powerful dorsal and caudal fins for propulsion

paired appendages: pectoral and pelvic fins

but pectoral fins are rigid, not flexible

hammerhead shark uses its head for steering

since pectoral fins are not moveable

sharks are also among some of the fastest fish:

eg mako shark 60mph

eg. blue shark 43 mph

Buoyancy

all fish are slightly heavier than water

to keep from sinking, sharks must keep moving

shape of sharks tail provides lift

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 39

large liver is rich in fats and oils giving sharks near
neutral buoyancy
 don’t need to use much energy to maintain position in the
water column

in migrating sharks the liver can enlarge to 25% of the animals

weight to provide energy needed for long migrations

Feeding & Digestion

most cartilage fish are predators

yet, by nature, most tend to be timid & cautious

powerful jaws
jaws are suspended from chondrocranium by ligaments

 free movement (somewhat similar to the flexibility in the

jaws of some snakes)

teeth only grasp and tear apart prey, don’t chew

 the teeth and (dermal) scales of sharks

are essentially identical except for size
-enamel covered dentin
-same structure and composition
-both shed regularly

 teeth are enlarged scales

 form replaceable rows of teeth

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 40

 eg. easily lost, constantly replaced, usually the only
part of a shark preserved as fossils: fossil shark
teeth

sawfish, a modified ray, flails its toothed snout in

schools of fish then sucks up the injured ones

some sharks are plankton feeders:

eg. whale shark (>50’); worlds largest fish
eg. basking shark (15-40’)

a few are scavengers

skates and rays have broad, blunt, cobblestone-

like teeth for crushing clams, oyster, etc

food passes from the mouth, down the esophagus to

the large stomach
the large esophagus and stomach allow for the passage
large, unchewed pieces of food

most digestion occurs in the stomach and the first part

of the intestine

the vertebrate digestive system has two major

accessory digestive organs:

a very large multi-lobed liver

 secretes bile which is stored in the gall

bladder until needed to aid in the
digestion of fats and lipids
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 41
  liver also processes and stores nutrients
absorbed from the intestine and helps to
remove toxins that might have been taken
in

 in sharks the liver also contains huge stores

of oil to make the animal more buoyant in
the water

a glandular pancreas

 produces dozens of digestive enzymes to

help break down proteins, carbs and lipids

 the pancreas also produces important

hormones that help control sugar levels
in the body

most of the remainder of the digestive tract functions

in absorption of the food once it is digested

in sharks, part of the small intestine is modified

into a spiral valve (=valvular intestine) which
slows the movement of the food through the
system and increases the area available for
absorption of these nutrients

undigested materials pass through a short large

intestine (colon) into the cloaca and out of the
body
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 42
Respiration

gills are inside 5 pairs of gill slits similar to agnatha

a pair of spiracles behind the eyes can take in water

when mouth is occupied with food

sharks must be moving or there must be some

current to move water over the gills to extract
oxygen from the water

Circulation

all fish have a closed circulatory system with a simple,

2 chambered heart (1 atrium & 1 ventricle) and
a single circuit of blood flow

blood is pumped from the heart to the gills to pick

up oxygen, then to the rest of the body to deliver
nutrients and oxygen and transfer wastes

the blood then returns to the heart

the spleen is the main organ that produces blood

cells, filters the blood and removes “worn out”
cells

Nervous System & Senses

Senses

a. lateral line system

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 43
 a series of pores running over the surface of
the head and down the sides of the
body are the lateral line system

contain mechcanoreceptors and

electroreceptors

used to detect vibrations and current

and tiny electrical currents produced by

nearby animals

especially useful in murky waters

b. chemoreceptors esp in olfactory pits on head

open through nostrils

widely spaced nostrils allow shark to more

easily locate prey

can detect prey half mile or more distant

c. eyes not particularly good but used at close

range
the hammerhead shark has eyes widely spaced to improve
depth perception

 better for attacking prey

d. ampullary organs (of Lorenzini)

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 44

 electroreceptors in head

senses bioelectric fields around animals

used especially for final stage of attack

Excretion

sharks have relatively large kidneys

sharks retain urea to help maintain internal fluids

isosmotic to sea water

also, rectal gland assists kidney

secretes excess salts

most sharks, but only a few kinds of other fish posess

a cloaca:

a single chamber that receives products from

the intestine, kidneys and reproductive
system

opens to the outside through a single pore

Reproduction & Development

most fish and almost all vertebrates are dioecious

while most fish have external fertilization

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 45
all chondrichthyes have internal fertilization

male sharks & rays with claspers on pelvic fins

 used to transfer sperm (NOT for clasping)

usually produce only a few eggs at a time

some skates produce 2 young each time

no member of the group produces >12 at a time

many retain eggs in body till hatch

bear live young! (viviparous)

development lasts 6 months to 2 years

some sharks have primitive uterus and placenta

and provide “uterine milk” for developing
young

others get extra nutrition by eating eggs and

siblings in uterus

some sharks and skates have internal fertilization but

deposit eggs in horny (keratin) capsule

= mermaid’s purse
each “purse” may contain several eggs

often has “tendrils” to attach to objects

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 46

no parental care after eggs are laid or young are born

Other Kinds of Cartilaginous Fish:

eg. Skates and Rays

~ half of all chondrichthyes

mainly bottom dwellers

dorsoventrally flattened

enlarged pectoral fins = ‘wings’

move in wavelike fashion

gills open on underside but have large spiracle on

top to take in water

in sting rays caudal and dorsal fins have been lost

tail is slender and whiplike

armed with 1 or more spines

teeth adapted for crushing prey

(molluscs, crustaceans, small fish)

dioecious
many bear live young like sharks

eg. Electric Rays

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 47
 fish are the only animals that can directly produce
an electrical shock

the ability to produce electric shocks is confined to

only 2 groups of vertebrates:

electric rays and some bony fish

electric rays are generally slow, sluggish fish that

live in shallow waters

have some muscles modified into electric

organs to shock prey or stun predators
cells connected parallel
 high amperage, low voltage

 high power output – up to several kilowatts

electric rays were used by ancient Egyptians as

“electrotherapy” treatment for arthritis and gout

eg. torpedo ray:

contains 400 honeycomb-like prisms filled with clear jelly-

like mass

nerves connect organ to lobe of brain = electric lobe

each cell generates .02-.05 volts

 can generate combined shock of ≤35 Volts

the amount of electricity varies greatly

can only give a few shocks before it has to rest and eat

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 48

eg. Sawfish

were once common predators in coastal bays and

lagoons

they like murky water and can tolerate low

salinities

Aztecs refered it as an “earth monster”

grow to about 20’ long

use their “saw” to slash and flail about to kill or

stun prey,

then as a ladel to orient and swallow their

prey

bear live young, 6-12 at a time, each ~1’ long

already with “saw”

for milennia the fish was considered a nuisance to

fishermen

caught unintentionally as bycatch in tangled

and torn nets

they virtually disappeared by the 70’s

now a protected species

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 49

 teeth of snouts still sold for artificial spurs for
fighting cocks

fins sold for up to $3000 in asian markets for an

alternative kind of “shark fin soup”

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 50

 Ecology
1. sharks are top predators in many ocean food chains

2. Symbioses: Mutualism

eg. Shark Suckers (remoras)

an example of commensalism, although some

species may be more mutualistic by removing
parasites and pathogens from their host’s skin

shark suckers are bony fish with one of the

dorsal fins modified into a suction disc

up to 3’ long

use large “suction cup” like disc on the top of their

heads to attach to sharks

they spend most of their lives attached to the

shark and feed on debris produced from the
shark’s feeding activities

some also feed on parasites on skin of shark

some feed mainly on the hosts feces rather

than pieces of dropped food

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 51

 while common to sharks, some are also found in
rays, whales, turtles and other marine
creatures

some species are host specific

eg. Pilot Fish

about 2’ long with dark blue stripes

act as cleaner fish

keep the sharks healthy

they will eat anything; including parasites and

bodily wastes off the shark

not attached to sharks but swim along side them

for sometimes long distances

are territorial about the shark they travel with

don’t allow others to ‘horn in’

3. Symbioses: Parasitism

parasitic flatworms, annelids, roundworms and

several different kinds of Arthropods are
known to parasitize Sharks and rays.

only one species of shark is actually a parasite

itself
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 52
 = Cookiecutter Shark

they parasitize fish and sea mammals

eg. marlins, whales, larger sharks

after finding a host they fasten their highly

specialized mouth equipped with a pair of
suction-cup like lips onto their flanks

it closes its spiracles to form a tight seal

the uses its teeth to bite down and wheels in a

circle to dislodge a plug of flesh from its host

the scars left by these fish are common in large

sea animals

there are some reports reports of human “hosts”:

underwater photographers

ship wreck victims

usually night time attacks

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 53

 Examples of Cartilaginous Fish

Nektonic fish
sharks
swordfish

Benthic Fish
skates & rays
guitar fish

Hammerhead Shark
loners
slower than most sharks
use head for maneuvering
eyes far apart improve depth perception
often basks near surface of water

Guitarfish
all are tropical and warm temperate waters
sluggish bottom dwellers
eats shellfish, worms and small fish

Skates & Rays

benthic fish

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 54

 Human Impacts

1. Shark attacks

60 - 70 per year (2000-2011) ; 1-12 fatalities

especially great white (to 6 M long)

mako
tiger
bull
hammerhead

more casualities reported from Australian region

than anywhere else
in 2008 in US 4 people were killed in shark attacks; 108 were
killed by cows (blunt force trauma)

2. Shark fishing

~40 Million/yr (26-73 M 2011) are harvested worldwide

recent estimates (2012) are that shark

populations at inshore reefs worldwide have
declined by 90%

harvested for their skeletons that is dried and

powdered and used as an “herbal remedy” to
‘cure cancer’

the primary cause is China’s growing appetite for

shark fin soup
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 55
 sells for up to $100/bowl

eg. Dubai alone exports 500 tonnes of shark fins and other
shark products/ yr to Hong Kong (~ half the world
shark fin production)

its generally a legal harvest but increasingly being banned

eg. “finning” has been outlawed in US

some other countries are setting quotas

3. Medicinal/Pharmaceuticals

electric rays were used by ancient Egyptians as “electrotherapy”

treatment for arthritis and gout

chondroitin for joint treatment

extracts are being tested for anticancer drugs and

weight loss

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 56

 Class Osteichthyes
Bony Fish
most successful vertebrate class

more species than all other kinds of vertebrates

combined

27,000 sp; (96% of all fish)

~200 new species are described each year

probably 5-10,000 more undescribed species

bony fish range in size from the tiniest of all

vertebrtates to over 15’

7 mm to 17’ (oarfish) & 4.5 M blue marlin

some fossil forms may have reached up to 100’ long

have adapted to every kind of aquatic habitat

from 8000 M deep to 5200 M in Tibet

some in hot springs (44º C)

others under anarctic ice at -2º C

in totally dark caves

some make excursions onto land

Body Form

most bony fish are designed for active swimming

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 57
 but with an amazing diversity of body form:

fusiform shape (eg. tuna)

powerful tail

fastest fish, often live in open ocean

streamlined bodies to reduce friction

rod shaped (eg. barracuda)

elongated, arrow-like fish with powerful tails, pelagic

predaceous fish

flattened/depressed (eg. flounder)

flattened bodies in bottom forms

spherical shape (eg. puffer fish)

when threatened, can inflate body so they can’t be
swallowed

ribbon shape (eg. wolffish, eels)

slow swimmers, secretive, move easily wriggle into
cracks and crevices for protection or to ambush
prey

laterally compressed (eg angelfish)

camoflage; viewed head on are almost invisible; also
allows quick, sharp turns

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 58

 grotesque forms (eg. anglerfish)
many deepwater forms; cryptic or mimic for protection

unusual shape of seahorse head is perfectly suited for

attacking its tiny crustacean prey

Skin & Scales

most bony fish have thin, overlapping dermal scales

in dermis

replaced heavy bony dermal scales of extinct

groups

light and flexible to enhance swimming ability

several types of scales:

eg ganoid  bony, don’t overlap (more primitive)

others do overlap

some have completely lost scales

unlike sharks, bony fish do not shed scales

they grow throughout life

 can be used to age fish

skin color

skin of fish shows a variety of colors and texture

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 59

 most bony fish can control their color to some
degree due to special skin cells in dermis

=chromatophores

controlled by nervous system

can be: silver, yellow, orange, black

blended for other colors

allows fish to change color to blend with

substrate

color changing is most highly developed in

flounder (flatfish) species

skin color serves a variety of functions:

eg. countershading

most open ocean fish have dark backs and light bellies
making it more difficult for predators to spot
them in open water

fw fish shades of green, brown, blue above

and silver or yellow white below

 from below blends with sky,

from above blends with substrate

eg. concealment:

eg. coral reef fish are highly colored

but on reef cant see them
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 60
 eg. often have blotches, spots and bars
 ~army camoflage

eg. mimicry:

another form of camoflage

eg. pipefish, anglerfish, sargassum fish

take coloration, texture and form of seaweed

eg. distraction:

eg. false eyespots

draw a predator to the back of the animal

allowing fish to escape in other direction

eg. butterfly fish

eg. advertising:

attract attention for a special service

eg. cleaner fish help remove skin parasite

their distinctive color is recognized by their

“customers” and they are not harmed by them

eg. warning:

many highly colored fish stand out from their

surroundings

 warn potential predators that they are

poisonous

eg. lionfish

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 61

bioluminescence

the skin of some fish also contains light emitting

organs

= photophores

may be on head; lateral line, sides of belly, on

barbels, etc

found in many unrelated groups

Support

cartilaginous skeleton is laid down in embryo

it is then replaced by well ossified bony

skeleton in fetus and adults of most species

a few species retain cartilage skeleton

all fins contain bony or cartilage rods = rays

connected by thin membranes

 makes them stronger

 they can also be folded down for more

maneuverability

pectoral and pelvic fins are both moveable and

controlled by muscles for more precise control
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 62
dorsal fin is moveable and sometimes becomes
highly specialized for:
camoflage

venomous spines (eg. scorpion fish)

lures (eg. anglerfish)

sucker (shark suckers)

Movement

most of body mass is segmented myotomes

even more so than in cartilaginous fish

(2/5ths of body volume in most; 3/4ths in tuna)

movement is mainly swimming,

but some can walk, crawl, burrow, or “fly”

very flexible and moveable pectoral and pelvic fins

pectoral fins used to steer and swim

Buoyancy

most bony fish today have swim bladder to control

buoyancy

swim bladder arose from lungs of some primitive

air breathing bony fish
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 63
 lungs evolved 1st in fish  then swim bladders!

most early bony fish evolved and diversified in

freshwaters

they lived in swamp waters with low O2 levels

needed accessory oxygen source

alternating wet and dry temps made lungs

essential for survival

called a lung if used mainly for respiration;

called swimbladder if used to control buoyancy

by adjusting the volume of gas (O2)in the swim

bladder a fish can achieve neutral buoyancy and
remain suspended indefinitely with no muscular
exertion
control of buoyancy probably coevolved with fin modifications to
improve and refine locomotion

most pelagic fish have swim bladders (except tuna)

bottom fish generally lack swim bladders

(eg. flounder)

in fish that migrate to & from great depths have the

additional problem of the gasses being forced into
blood
eg. humans suffer the bends when returning to surface as
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 64
 gas comes out of solution and forms bubbles in the
blood

fish are able to prevent gasses from entering body

fluids at great depths and keep body fluids at
equilibrium with oxygen in surface waters
their swim bladders can resist pressures >240 atm (up to
8000’ depth) (=pressure inside compressed gas
cylinders) and still keep oxygen in blood at 0.2 atm
(~sea level pressure)

Feeding & Digestion

bony fish diversified into a variety of feeding types:

a. plankton feeders

most common feeding type

most pelagic species and commerical fish are

plankton feeders (zooplankton)
eg. herring, anchovies, menhaden

travel in large schools

plankton are strained with sieve like gill rakers

b. predators

teeth used to seize prey

gill arches of some fish are modified into

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 65
 “pharyngeal jaws” for chewing, grinding &
crushing

a few have molar-like teeth behind gills to grind

food

eg. Piranhas are known as viscious predators - mainly through

movies and TV

several dozen species of carnivores; 6-10” long

found in South American Rivers of the Amazon Basin

jaws bristle with unique, sharp, densly packed teeth

bad rap; can be very aggressive but are only rarely known
to bite and injure humans

but are considered a nuisance to commercial and sport

fishers

sold for aquaria but illegal in most states in US

occasionally found in US rivers but generally can’t survive

cold winters

c. herbivores

many freshwater fish eat plants, grasses, algae,

etc
eg mollys, some cichlids, head standers, etc

d. omnivores
eg. angelfish, goldfish, guppies

e. scavengers & detritivores

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 66
 eg. catfish, suckers & minnows

f. parasites
eg. Toothpick fish (Candiru)

parasitic freshwater catfish in Amazon river

eel shaped, translucent  impossible to see in water

up to 6” (15 cm) long

most feared fish in these waters, more than piranha

they can detect respiratory currents and swim toward gill

openings of other fish where they feed on prey’s blood

some species lie in wait in murkey bottom mud

sample water for nitrogen wastes from gills of fish

eg. ammonia, urea

once detected they dart towards the gill cavity with a burst
of speed

once inside gills they lodge themselves in place with its

spines

gnaws a hole toward a major blood vessel and gorges itself

for a few minutes only

it then dislodges itself and sinks back to bottom of river to

digest its food

victim usually bleeds to death

is known to attack people and swims into an orifice;

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 67

 vagina, anus, penis

locates its target when people urinate near the fish

has been known (and videotaped) swimming up a

urine stream into penis of victim

almost impossible to remove without surgery

Respiration

most bony fish get oxygen through gills

the gills of bony fish are much more efficient at extracting
oxygen from water

gills are located inside pharyngeal cavity

gills are highly vascularized filaments of soft

feathery tissues covered with a thin epidermis

often have “gill rakers” that filter water before

passing over gills

to remove food and bits of debris that

might clog the gills

gills are covered by a bony flap = operculum

offers protection and reduces friction when

swimming

 operculum can also actively pump water

across gills
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 68
 fish can still “breath” even if not moving

Nervous System & Senses

bony fish have a simple nervous system and the sense

organs as described for fish in general

however, bony fish make much more use of sound

than the other two fish groups
at least 1000 fish species make and use sounds

clicks, grunts, thumps

we have an outer ear that collects sound waves, a

middle ear with an ear drum and ear ossicles that
vibrate and transmit the sound waves to the inner
ear for processing

fish have no “outer ear” or ear drum

sound travels further and faster in water than it does

in air

there is a great variation in fishes ability to produce

and detect sound in water

The Hearing Ranges of Fish (humans ~20-20,000 Hz)

Common Name Hearing Range in Hz

Atlantic Salmon 40 to 350
Bonito/Tuna 100 to 900
Red Piranha 80 to 1,500
Goldfish 40 to 3,200
Brown Bullhead 100, to 4,000
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 69
 Stone Moroko 100 to 8,000
Atlantic Cod 20 to 38,000
American Shad 200 to 180,000
Gulf Menhaden 200 to 180,000

used mainly to attract mates or ward off predators

in fish, the swim bladder has secondary function in

hearing

swim bladder is connected to inner ear by a set

of bones = Weberian ossicles

sound vibrations are picked up by swim

bladder and transferred to ear through
ossicles

acts kind of like an eardrum and ear ossicles in

humans

helps to amplify even very faint sounds in

water

Defenses

1. color and shape can be used for camoflage

2. puffer, blowfish and porcupine fish

fill their elastic stomach with water or air until it’s

large and spherical in shape

predators may die from choking

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 70
3. venomous fish

some fish (including puffers) are highly venomous

eg.Puffer

contain high concentrations of potent toxin concentrated in

internal organs

if a puffer is swallowed they can sicken or kill its

predator

In Japan some eat puffer (fugu) as a delicacy specially

prepared by trained chefs

eg scorpionfish lionfish:

dorsal spine can inject venom

produce painful would, rarely fatal

4. Electric Fish

water conducts electricity

the ability to produce electric shocks is confined to

only 2 groups of vertebrates: electric rays and
some bony fish

bony fish: electric eel & electric catfish

eg. electric eel

in rivers in South America

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 71

 grows to 3 - 7 ft long

electric organ is modified muscle tissue

 up to 40% of body weight

most powerful electric organ of all fish

can produce up to 600 volts @ 1 ampere to stun or

kill prey

the eel is well insulated and does not get shocked

 can give several 100 shocks up to 300 V

each/second

doesn’t need to actually touch victim

electric field extends several feet around fish

eg. electric catfish

found in the Nile river

organ seems to have evolved from the skin rather than

from muscle tissue (?)

consists of a uniform layer of gelatinous tissue

Reproduction & Development

most bony fish are dioecious

only a few are hermaphrodites

genders cannot be distinguished externally

most with external fertilization (oviparous)

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 72
 a few bear live young (eg. guppies)

unlike sharks & rays, most bony fish produce large

numbers of eggs:
eg. herring  25,000
eg. lunpfish  155,000 - 1M
eg. halibut  3.5 M
eg. sturgeon  635,500
eg. cod  4 - 6 M

 less than 1/million will survive to maturity

most fish spawn at certain times of the year

 spawning is temperature dependent

 temperature is critical for survival of eggs and young

in most marine fish: eggs are released and become

part of the plankton through embryonic and larval
development

A few fish make nests and show fairly elaborate

mating behaviors and parental care

and its most often the male who puts in the

“extra effort”

eg. Stickleback
male constructs very elaborate nest of grass and
weeds bound by mucous threads
(only example of “case building” in a vertebrate
animal)
then looks for a mate to entice inside
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 73
 if gentle persuasion doesn’t work, he may drive 1 or 2
females into nest until enough eggs are laid
then he jealously guards them for many days until
they hatch

eg. Tilapia
starting in late spring they begin to spawn
males build circular nests in streambed
females choose mate based on nest
both spawn over nest
after eggs hatch mother will carry the young in her
mouth for a few weeks then they are on their own

eg. Seahorses
seahorses are only vertebrates in which the male actually
becomes pregnant
male contains a brood pouch, completely sealed except for
a tiny hole
female lays eggs in males pouch
male squirts sperm directly into pouch to fertilze eggs
males nurture their young, provide food and oxygen and
get rid of waste products
young remain there for ~ 10 days till hatching
male convulses (as if in labor pains) and muscular
contractions eventually force all the seahorses out of
the pouch
almost immediately, the female shows up, an new courtship
ritual and the male may again become pregnant by the
next day

eg. Some marine catfish

eggs are incubated in males mouth
young continue to be carried an protected in males mouth
after they hatch
male doesn’t eat for ≥ 1 month

eg. Some other Catfish species

females attach eggs to underside and she carries them
around till they hatch

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 74

some fish show absolutely no parental care or interest
eg. Gambusia
in an aquarium will eat all their young as soon as they are
born

fish continue to grow throughout life

annual rings are produced in scales, otoliths and

other bony parts

 the age can be accurately determined

Migrations

A. Catadromous (“down running”) fish

some fish spend most of their lives in freshwater

but return to sea to spawn

= catadromous
eg. some eels

each fall large #’s of female eels are seen swimming down
rivers toward the sea

when adults leave rivers in Europe and N America they

reach ocean and swim at great depths to Sargasso Sea

takes several months to reach this area; here they spawn

and die

they tiny larvae begin their return trips to the coastal rivers
 takes up to 3 yrs in Europe

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 75

 each spring large #’s of young eels appear in coastal rivers
swimming upstream

males remain in brackish waters near mouth

 female continue 100’s of mile upriver

by 8-15 years females >1 M long; return to sea to rejoin

males and spawn

B. anadromous (“up running”) fish

other spend most of adult life in the sea, and

return to freshwater to spawn
eg. Atlantic species (eg. salmon & steelhead trout)

make spawning runs year after year

eg. pacific species ( sockeye, sliver, humpback & chum salmon)

make one spawning run then die

adults spend 4 yrs at sea yet can unerringly return to

parent stream
 only a few stray go to wrong stream

when salmon return to site where they were hatched, they

spawn and die

the following spring the newly hatched fry “imprint” on the

stream as they drift downstream to the sea

How do they find the mouth of the river when they are
returning to spawn?

apparently can navigate by orienting to sun’s

position
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 76
 but they can also navigate on cloudy days

may also be able to use earth’s magnetic field

probably also use ocean currents, temperature

gradients, food availability to reach general
area

annual run of wild salmon today is ~3% of the 10-16

Million fish that ran 150 yrs ago

Salmon runs in Pacific NW have been devastated by

stream degradation:
eg. logging, dam construction (50 dams)

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 77

Preadaptations to Terrestrial Environment

all other vertebrate classes are primarily terrestrial

and evolved from fish ancestor

(NOT ‘in preparation’ for moving onto land, but adaptations that
made the transition possible)

a. Air Breathers

many fish can survive out of water for a short time

by breathing air in a variety of ways:

lungs, skin, gulping air, rectum, air chambers in

head, etc
eg. some fish (eg. Anabantoids) can breath air via
labyrinthine chamber in head behind gills

eg. lungs of lungfish & gars

eg. bowfin (Amia) at low temp use mostly gills, at

higher temperature use mostly lungs

eg. some Corydoras Catfishes can process air in the hind part of
the gut

eg. freshwater eels commonly make excursions onto land in

rainy weather

do gas exchange through moist skin

eg. electric eel has degenerate gills and must get

most of its oxygen by gulping air

eg. Indian climbing perch (Anabas) spends most of its time on

land near water’s edge

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 78

 has special chamber above much reduced gills for
respiration

can also absorb oxygen from air if skin is moist

eg. mudskipper can be out of water for long periods but prefers
to keep tail in water to absorb oxygen from water through
its skin

eg. a species living in African streams and ponds that often dry
up during dry season. fish burrow down into mud and
secrete copius amounts of slime and mix it with mud to
form a hard “coccoon” until next rainy season

b. some fish can walk on land with their strong

fins
eg. Indian climbing perch (Anabas) spends most of its time on
land near water’s edge

only climb in wet weather

eg. freshwater eels commonly make excursions onto land in

rainy weather

eg. walking catfish

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 79

Fish Life Histories

eels, Anguilla sp
hatch in spring
begin life in larval form, ribbonlike
larvae enter estuaries and bays turn into “glass eels”
transparent but resembling adult
move to freshwaters and live 8-15 years
stay hidden in weed beds ledges ormud during daylight
eat whatever is available
can come ashore searching for food: worms and grubs
can breath through skin
slime layer protects from salinity changes
highly developed sense of smell
light sensitive skin also responds to low frequency sounds
after 10 years in freshwaters, gonads enlarge, fish change shape
and become a bronze or silver eel
eyes enlarge and become more sensitive to light
body fat increases
begin migration to sea to Sargasso sea
need fairly high water temperatures to reach sexual maturity
large eel can produce 2.5 Million eggs

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 80

 Fish Ecology and Symbioses
1. Fish occupy almost every conceivable aquatic
habitat

in most they play a significant role in several

levels of the food chain; predators, herbivores,
scavengers, etc

especially abundant in coastal areas

2. Symbioses: Mutualism
eg. cleaner fish

~50 species of cleaner fish

esp along coasts and coral reefs

generally remove dead skin and parasites

cleaner fish advertise their services with conspicuous

coloration

eg. Hippo cleaner fish

several species of fish clean Hippos

some feed on vegetable matter in feces of hippo

some clean cracks in the bottom of feet

some clean hippo tail bristles

Hippos visit cleaning sites where these fish gather and

invite fish to clean them

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 81

 eg. remora

attaches to shark

eats excess foods, removes skin parasites, etc

eg. clown fish

sea anemones

eg. Pistol shrimp and goby fish

pistol shrimp dig holes in sediment looking for food

they are almost blind and when they bring a load of sand
out of their burrow they are highly susceptible to
predation

gobys live in the tunnels made by the shrimp for protection

goby actively protects the shrimp from harm

they communicate through antennae of shrimp and tail of

fish

when predator comes both rush into the tunnel

at night they both go into the burrow and shrimp closes the
opening

3. Symbioses: Parasitism
eg. some fish mimic the bright colors of cleaner fish to actually
feed on the skin and scales of their host

eg. candiru (toothpick fish)

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 82

 Examples of Bony Fish
Fish of the Nekton
squirrelfish
jacks
butterfish Benthic Fish
triggerfish founder
cowfish moray eels
filefish snake eels
tuna batfish
sailfish seahorses
anchovies pipefish
menhaden sea robins
lookdowns soles
sheepshead tongue fish
drum stargazers
redfish

Sharksucker (Remora)
flat oval sucker on head
attach to hosts by suction
up to 2 feet long
hitchhikers: free ride and eat food scraps
also remove parasites and clean damaged skin

Lookdown
prefer to stay near bottom
scrounge around for small shrimp, worms, etc
thin sharp turns, lots of maneuverability

Sheepshead
especially in brackish coastal waters, around mangrove roots and
pilings
crush shells of barnacles, stone crabs and fiddler crabs, mole
crabs, coquinas in surf

Stargazer
live on sand and mud bottoms
eyes protrude above
have electric organs
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 83
Sea Robin
bottom dwellers
to one foot long
use free pectoral rays as tiny legs

Pipefish
males brood young
live in submerged grass beds nearshore
hidden predatory

Sea Horse

in ancient religious writings it was believed that everything seen

on land was also represented in the ocean; eg grape fish,
sword fish, sawfish, sea horse, also sea bulls, sea goats,
sea lions; only some were true beings of the sea
very slow swimmers; use mainly small pair of pectoral fins for
locomotion
male broods young
well camoflaged
prefer grassy areas of estuaries and nearshore

Moray Eel
lives in burrows and crevices
predator
elusive, sly, powerful, viscious

Toadfish
sedentary; poor swimmers
well camoflaged
feed on fish and shellfish, some scavenging
prefer oyster reefs and shelly habitats around pilings; quiet areas
large gills can thrive in low oxygen waters

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 84

 Human Impacts of Fish
1. As Food: Commercial Fisheries

we have harvested fish throughout all of human

history
piles of fish remains have been found in archaeological sites
90,000 years old

A. Marine Fisheries

marine food sources (including fish) provide about

25% (directly and indirectly) of animal protein

commercial fishing employs 200 Million people

worldwide

2.6 billion people worldwide depend on fish for protein

(2002)

60% of all fish consumption is by the developing

world (2008)

main fish taken commercially:

anchovies
pollock
mackerel
herring

location of largest fishing grounds:

China
Peru
Chile
Japan
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 85
 United States

main uses of commercial fish are as food for humans

and livestock

estimates suggest that seafood production from wild

fish stocks will not be sufficient to meet growing
world demand in the next century

total marine fish catch has peaked at about 100

Million tons and remained stable, in spite of
increased efforts to catch fish

most commercial fisheries are near shore where most

pollution and damage occurs

open ocean catch has been increasing

~ 11% of total (2002)

per capita (per person) fish catch is decreasing as

population expands

11 of worlds 17 major fisheries are overfished and in

decline

estimates that ~ half of all commerical fisheries stocks

are “fully exploited” and another quarter are
overexploited, depleted or slowly recovering

most conclude that global marine fish

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 86

 production is not sustainable at the
current levels

additionally, about 20% ($10 – 20 Billion) of fish catch

is illegally caught

an evolutionary consequence of overfishing: fish are

adapting by growing slower and staying smaller in
size

a few of the problems:

a. subsidies have encouraged overfishing which

makes it a nonsustainable resource
the world spent $124 billion to catch $70 billion of fish

the difference was paid for by taxpayers

b. most commercial fisheries are near shore

where most pollution and damage occurs

c. of the world fish catch only two thirds are used

directly for human consumption,

the rest is converted to fish meal and oil,

and pet and livestock food

d. a recent study estimates that about 20% of

commercial fish are illegally caught at a cost
of $10-20 Billion in lost profits

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 87

 e. much of what is collected is wasted as
“bycatch”, especially in industrial countries,

examples:
eg. shrimpers typically discard 5 to 8 times as many
creatures than they keep

eg. gulf of mexico shrimpers killed 34 million red

snapper and over 3000 sharks in one year

eg. open ocean fishermen use large drift nets

(25’deep & 50 miles long), set out 30,000
miles of net a night worldwide

18 miles of net is lost per night

1000 miles per year become ‘ghost nets’ and

trap and entangle fish for decades as they
float in the ocean

these nets killed 42 million seabirds, marine

mammals and other nontarget animals

e. there has been an increase in biomass

fishing:

collecting all life in an area and grinding it

up for meal, to use as animal feed & for
fish farming

decimates communities in an area

estimates are that at least half of the

world’s continental shelves are

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 88

 scoured by trawlers at least once
every year

B. Freshwater Fisheries

fish from inland waters accounted for 10% of total

catch (2002)

many river basins, especially in developing countries

support intensive fisheries

inland fish are considered to be the most threatened

group among all vertebrates used by humans

C. Aquaculture (fish farms)

global production from aquaculture (2012)

= 69 million tons

in 2013, for the first time ever, the world is producing

more farmed fish than farmed beef
if the current trend continues humans will consume more farmed
fish than wild-caught fish by 2015

fastest growing animal protein sector

especially in developing countries

aquaculture produces more than 220 species

carp are the largest group

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 89

China and other Asian countries are the largest
producers least environmentally damaging since they farm
scavenger species

salmon and other carnivorous fish favored by the west require

much heavier food and energy inputs and create huge
amounts of pollution

2. Pets

15-30 Million fish of up to 1000 species are sold

globally each year

20 million fish are sold each year as pets in US alone

marine animals only $200 M - 300 M sales annually

Outdoor fish ponds have been around for at least 2000 years

The Romans were the first to bring fish indoors - for fresh food

10th century Chinese kept bowls of goldfish

in Victorian England marine aquariums became the rage

3. Commercial Uses

eg. fish scales used for decorative beads

eg. fish meal: made from fish heads, fish offal and
trimmings

also converted into liquid form and used as

additions to animal feed.

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 90

eg. unsold or surplus fish are composted to produce
high nitrogen fertilizer

4. Pharmaceuticals

eg. oil extracted from fish livers are good source of

Vitamin D

eg. fish oil especially rich in omega-3 fatty acids

eg. puffer fish  tetrodotoxin as pain releiver

1 fish can produce 600 doses

5. Research

3.5 - 7 M fish used for research in us each yr

6. “Herbal” uses of fish

eg. ancient greek writings and herbals from China and

other countries have touted the healing properties
of seahorses for 1000’s of years
pulverized and made into a tea used to calm bladder, treat
asthma, soothe boils, pustules and ulcers, and as an
aphrodisiac

today seahorse powders and tablets are taken to treat throat

infections, high cholesterol, kidney and liver disease

today, at least 70 tons (25 Million) seahorses are harvested

worldwide, each year, to be roasted, crushed and dissolved
to make traditional medicines; whole ones are used as
talismen to improve luck in fishing
Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 91
  none of the uses have been shown to have any
scientifically valid value

Animals: Chordates - Fishes; Ziser Lecture Notes, 2015.11 92

 Phylum Chordata
48,000 species

very diverse phylum but considerably less diverse

than eg. arthropods or molluscs

also many fewer species than most other major phyla

most advanced phylum of animal kingdom; more

complex anatomy
eg nervous system

eg. circulatory system

one to which we belong along with fish, amphibians

reptiles, birds and other mammals

some of the largest or most massive animals

major identifying characteristics:

1. Notochord
flexible rodlike structure

enclosed by a fibrous sheath

extends the length of the body

in larva and/or adult

provides basic support and serves as main axis for

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 1

 muscle attachments to permit “fishlike”
undulatory movements

first part of skeleton to form in embryo

in primitive chordates the notochord persists through life

in most chordates the notochord is replaced by a vertebral

column of bone

remnants of the notochord remain as “intervertebral

discs”

2. Dorsal tubular nerve cord

in most invert groups; nerve cord is ventral & paired with paired
ganglia along it’s length

in chordates the nerve cord is a single dorsal hollow nerve cord

front end usually enlarged to form brain

3. Pharyngeal (gill) slits

slit-like opening sleading from throat to outside

first evolved as a filter feeding apparatus

still used by some to filter water for food

in others as gills

in some groups they are only found in embryo

and lost as adults

4. Endostyle or Thyroid Gland

specific kind of tissue found only in chordates

was originally part of the feeding apparatus

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 2
 endostyle secretes mucus and traps food inside the
pharyngeal cavity

eg. lamprey larva

in most chordates the same tissue has become an endocrine

gland in the neck region that helps control metabolism

5. Post-anal tail
in aquatic chordates it provides motility

 especially designed for propulsion inwater

especially for larval forms and fish

fish later added fins to increase efficiency

in terrestrial chordates it became a tail for leverage and balance,

not for movement

in humans “tailbones” is its remnant

Origin of Phylum

oldest known chordate fossil is from the Ediacaran

(600-543 MY) in Australia

6 cm long

Three major Subphyla:

Subphylum: Urochordata (tunicates)

Subphylum: Cephalochordata (lancelets)

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 3

 Subphylum: Vertebrata (vertebrates)

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 4

 Subphylum Urochordata
(tunicates, sea squirts)

1600 sp

all marine

widely distributed in all marine waters

at all depths

size ranges from microscopic to 1 ft in diameter

most are bag-like sessile suspension feeders as adults

they often live in large colonies attached to rocks and

other hard surfaces on the ocean floor

adult tunicates seem to lack the major characteristics

of the phylum

adults resemble sponges more than they do other

chordates
 most are sessile
 lack a coelom
 pump water through siphons

but tadpole-like larva has typical chordata features

adults have tough, nonliving, tunic covering body

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 5

 secreted by mantle

forms 2 siphons

one group, salps, are barrel shaped pelagic animals

with transparent gelatinous bodies

live singly or in colonial chains up to several

meters long

Feeding & Digestion

filter feeders

incurrent siphon  pharynx (branchial sac) slits atrium

 excurrent siphon

water is drawn in through the incurrent siphon

into a basket-like pharynx (branchial sac)

mucus is secreted by glandular endostyle in groove

along base of pharyngeal basket

uses mucus and cilia to move food toward mouth

complete digestive tract

food moves from the pharynx into the stomach for

processing

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 6

the filtered water passes into the atrium and out the
excurrent siphon

Respiration

pharynx also serves as a respiratory organ

Circulation

simple open circulatory system with small ventral

heart and 2 major blood vessels

blood is pumped through sinuses within the

animal’s tissues

 blood alternately flows in each directions

Nervous System

nerve with ganglia and plexus of nerve fibers

Excretion

no specialized excretory organs

nitrogens wastes are in the form of crystals deposited

in the tunic

Reproduction & Development

all tunicates are hermaphrodites with single ovary and

testis
Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 7
fertilization produces an elongated free-swimming
larva

= “tadpole larva”

larva reveals true chordate nature

has all 5 major chordate characteristics

has digestive system but doesn’t feed

swims about for hours to days, until it finds a suitable

substrate to settle on

attaches to substrate by adhesive discs

once firmly attached the larva undergoes an

exceptionally rapid metamorphosis into the adult:

in just 10-15 seconds the tail, notochord,

nerve cord, brain sensory structures and
muscles are crushed into a mass of tissue

over the next few hours, the internal organs c

hange their positions, the siphons adult
nervous system develop

loses its tail and most chordate features and becomes

an adult

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 8

some believe vertebrates arose from a neotenous
tunicate

Human Impacts of Tunicates

antiviral, antitumor

including possible treatment for malignant

melanoma
 the most dangerous form of skin cancer

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 9

 Suphylum Cephalochordata
(lancelets)

only 29 species (5 species in US)

closest living relatives to vertebrates

they display several intermediate characteristics

between invertebrates and vertebrates

the adult animal has all major chordate

characteristics

yet lacks the internal skeleton of vertebrae and

bone that most vertebrates have

burrowers and swimmer

slender, translucent, laterally compressed, fishlike or

eel-like body

ventral side of body is flattened

bears 2 folds o skin = metapleural folds

while they can swim they spend most of their time

partially buried in sandy bottoms of coastal waters
with their heads protruding

3-7 cm long

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 10

 often found with tails buried in sand

instead of tunic, outer body is covered by soft

epithelium

have fish-like fins with reinforcing fin rays

internal structure is very simple with with basic

chordate characteristics:

springy notochord for support supports body

while swimming or burrowing

Movement

with well developed “V”-shaped bundles of swimming

muscles

= myotomes (=myomeres)
provide fish-like movement by contracting
against notochord

also have dorsal hollow nerve cord

Feeding & Digestion

are filter feeders

mouth surrounded by oral hood with tentacles

(=oral cirri) an a wheel organ
Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 11
 cilia on these structures draw in water and food

pharynx strains food from water

food is trapped in mucous and enters the mout for

processing

digestive system is similar to tunicates but slightly

more developed

 has hepatic caecum (or liver) as accessory

digestive organ

food is drawn into the intestine by mucus and cilia

water passes through pharynx into atrium and out

the atriopore

Circulation

closed circulatory system

circulatory system similar to fish but no heart

Respiration

some respiration occurs through the pharynx

but most gas exchange occurs by diffusion

through the ventral body wall
Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 12
Nervous System

hollow nerve cord above notochord

anterior end of nerve cord = very small brain

 almost no cephalization

pairs of spinal nerves innervate each myotome

single ocellus at front of head

Excretion

simple excretory system of nephridia similar to those

in annelids

Reproduction & Development

all are dioecious with males and females

larvae resemble adults but are covered by cilia

cilia are used for swimming and drawing food

toward the mouth

retain chordate features in adult

resemble small fish

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 13

 Subphylum Vertebrata
General

most complex (?advanced) group of animals

larger, more active  need more food

one explanation for their diversity and dominance is

that, in general, vertebrate species have a lot
more genes
 amphioxus has as many genes as an ant or fly

eg flies  10,000 genes

eg. annelids  13,000 genes

but mice and humans  ~20,000 genes

fish became the 1st true vertebrates

oldest known fossils of a vertebrate:

560 MY old 2.5” long found in Australia

a similar 530 MY old fossil was found in China

while only 1 invertebrate group (the insects)

developed the powers of flight all vertebrate
groups have developed some ability to fly or at
least glide through the air

Major Characteristics:

1. internal jointed skeleton of bone or cartilage

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 14
 an endoskeleton permits unlimited growth

much more efficient design

is a living skeleton

grows with animals (not a case)

doesn’t need to shed regularly

probably began as cartilage then later became

calcified into bone

cartilage grows fast to form initial skeleton

a hardened skeleton is also ideal for muscle

attachments

especially in areas of high mechanical stress

real bone emerged as external protective dermal

plates

protected the head and brain and anterior part

of the body
eg. ostracoderms, placoderms

these bony plates later became modified into

scales of some fish

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 15

 since bone is living tissue it also becomes
important as a mineral reservoir (esp.
phosphorus & calcium)
eg humans: calcium needed for:
muscle contractions
nerve impulses
clotting
secretions
heart beat
etc

in the most primitive vertebrates its not much

more than a cartilage rod

= notocord

skeletons of lampreys and sharks and rays and

some bony fish (eg. sturgeon) remains mainly
cartilage in adults

in most vertebrates it is divided into:

axial skeleton
“braincase” - surrounds brain
vertebral column
ribcage

appendicular skeleton (limbs)

jointed appendages: pectoral & pelvic
eg. fins, legs, wings,

2. Segmented skeletal muscles (myotomes)

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 16
 became “W” shaped instead of “V” shaped as in
amphioxus

provided more control over body movements

3. complex skin

multilayered: epidermis, dermis

contains

numerous of sensory receptors

glands (oil, sweat, wax, scent, poison, etc)

keratin structures: scales, hair, feathers

4. more efficient digestive system

digestion shifts from moving food by cilia and

mucus to using muscular contractions (=
peristalsis) to move food through GI tract

vertebrates were the first to add an acid

producing stomach that more effectively
digests proteins

additional digestive glands:

pancreas & liver improve digestive efficiency

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 17
5. efficient respiratory systems closely tied
to circulation of blood

the original function of the pharyngeal slits to

filter water for food becomes functional
gills

6. increasingly efficient circulatory system

with pumping heart (2,3, or 4
chambered)

ventral heart
most inverts have dorsal heart

closed circuits of arteries and veins

RBC’s (erythrocytes) containing hemoglobin

for efficient distribution of oxygen to
tissues

7. most complex and best developed

nervous system of all animals

usually well developed head with sense organs

and brain (cephalization

lifestyle shift from filter feeding to predation

increased emphasis on brain and senses

better sensory and motor integration

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 18

 CNS = brain & spinal cord
 central processing and
coordination

PNS = nerves (eg. cranial nerves,

spinal nerves)
 conduct impulses to brain from
sense organs and from brain
to muscles and glands

senses:
complex eyes

inner ears for sound and balance

improved taste and smell

lateral line for water vibrations

electroreceptors to detect prey

8. Improved efficiency of excretory system

paired true kidneys (most cephalochordates had

none)

collect and get rid of metabolic wastes & toxins

greater role in salt and water balance

9. almost all are dioecious and reproduce

only sexually

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 19

Origin and Evolution of Chordates

similarities with echinoderms & hemichordates:

radial cleavage
deuterostomes
same coelom formation

oldest known fossils:

a. Pikaia (Burgess Shale; middle Cambrian, ~500 MY)

5 cm long; “V” shaped myotomes

probably a cephalochordate

b. Haikonella (early Cambrian, 530 MY)

chordate features: notochord, phayrnx, dorsal nerve cord

also some vertebrate features but not a vertebrate:

pharyngeal muscles
paired eyes
enalrged brain

urochordates are probably the most primitive

surviving group

but adults are too specialized

Neoteny ?

 larval form achieves sexual

reproduction; accelerated
development of reproductive organs

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 20

cephalochordates are clearly similar to ancestral
vertebrate form

most primitive vertebrates were jawless

 fossil agnathans predate all gnathostomes in

fossil record (>500 MY)

 oldest group = conodonts

conodonts are clearly related to group represented

by living lampreys and hagfish

lampreys may have evolved from ostracoderms:

small bottom dwellers
covered with bony plates
jawless
extinct
probably endoskeleton of cartilage not bone

Classification of Vertebrates

the classes of vertebrates are often grouped into

clades according to criteria of major evolutionary
significance:

A. jaws present or absent

agnatha = jawless
gnathostomes = mouth with jaws

B. fins versus walking legs

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 21
 pisces = paired fins for swimming
tetrapods = paired limbs for terrestrial locomotion

C. offspring develop within a fluid-filled sac

(=amnion)
anamniotes = do not develop within fluid filled
sac; eg fishes and amphibians
amniotes = do develop within a fluid filled sac;
eg. reptiles, birds, mammals

from most primitive to most advanced:

jawless fish 35sp; >500MY

fishes sharks and rays 850sp;
bony fish 21,000sp; 420MY
amphibians 4300sp; 360MY
reptiles 7000sp; 280MY
birds 9700sp; 150MY
mammals 4600sp; 200MY

Animals: Chordates & Introduction to Vertebrates; Ziser Lecture Notes, 2015.11 22

 Phylum Chaetognatha
(arrow worms)

means “hairy jaws”

>100 sp

marine

warm shallow waters

most are planktonic

swim and drift on surface at night; sink deeper

during the day

very common

voraceous predators in planktonic community

a few species are benthic forms

Body Form

small straight bodies

torpedo or dart like

1 - 12 cm long

Body Form
1
head - unsegmented trunk - postanal tail

Body Wall

think cuticle covers body

single layer of epidermis

on sides of animal becomes multilayered

well developed coelom

used as hydrostatic skeleton

Movement

highly mobile and nearly transparent

 effective predators

use caudal fins and longitudinal muscles to dart at prey

horizontal fins act as stabilizers

Feeding & Digestion

predators of copepods, fish and each other

use water vibrations to detect prey

2
teeth and chitinous spines on head

use curved spines around mouth to capture prey

complete digestive tract

Circulation

simple hemal system

Nervous System

well developed nervous system

consists of nerve ring around esophagus

with cerebral ganglia and other ganglia

sense organs:

eyes

sensory bristles

may also be able to detect water vibrations, water

currents and chemicals

no respiratory or excretory systems

Reproduction

3
all are protandrous hermaphrodites

function first as male then as female

cross or self fertilization

eggs are planktonic

Evolutionary Relationships

some think they’re related to nematodes or annelids??

others believe they are related to deuterostomes

molecular phylogeny suggests they are not closely

related to either protostomes or deuterostomes

 they apparently split some time before the

protostome/deuterostome split

4
 Phylum Hemichordata
(acorn worms & graptolites)

85 species

worm-like deuterostomes

hemichordates share characteristics with both

echinoderms and chordates

share 2 significant features with Chordates:

pharyngeal gill slits

dorsal nerve cord

Once thought they were chordates but defining
characteristic, notochord, not present as was originally
thought

widely distributed but secretive

most are sedentary or sessile bottom dwellers

usually in shallow waters

some are colonial in tubes

2 main classes, similar embryos but very different

adult forms:

1
 Enteropneusts (acorn worms)
Pterobranchia (graptolites)
Enteropneusts (acorn worms)

mucus covered wormlike body divided into three

regions:

tonguelike proboscis

most active part of the animal

short collar

long trunk

head and collar resemble acorn

trunk can be over 2 M long

very soft and easily broken

live in U – shaped burrows

some live under rocks and seaweeds

Movements

burrowers use hydrostatic proboscis to excavate a

burrow

Feeding and Digestion

2
all are ciliary mucus feeders

proboscis probes sediment and collects food

particles in mucous strands

cilia carry particles to groove at edge of collar

food is directed to mouth

some feed like earthworms

by ingesting mud or sand then gut extracts

organic debris

water swallowed while feeding exits through

pharyngeal slits (=gill slits, but not really for
respiration)

Respiration

paired row of pharyngeal slits (=gill slits) just behind

collar

cilia keep a current of water flowing through slits

no gills but gas exchcange ocurs across skin as water is

passed through pharyngeal slits

Circulation

3
open circulatory system

middorsal vessel

leads to networks of blood sinuses

colorless blood doesn’t carry oxygen well

mainly seems to have excretory function

Nervous System

dorsal and ventral nerve cord

dorsal cord is partly hollow as in chordates

the skin contains a network of nerve fibers

Reproduction & Development

most species reproduce only sexually

only 1 species is known to reproduce asexually

separate sexed = dioecious

external fertilization

direct development in most

some produce tornaria larva

4
very similar to bipinaria larva of echinoderms

5
 Pterobranchia (Graptolites)

~20 living species

sedentary, colonial

small; individual zooids 1 - 7 mm

zooids live in colonies inside collagenous tubes

zooids not interconnected

external appearance they look more like

ectoprocts

tentacles with cilia for gathering = lophophore

graptolites are important index fossils for geologists

Body Form

body of proboscis, collar and trunk

Reproduction

some are dioecious

many are hermaphrodites

6
 The Story So Far

have now covered all major phyla but one

and most minor phyla

have moved from very simple body plans without

true tissues or organs

to complex body plans with well developed

digestive, respiratory, circulatory, nervous,
endocrine, excretory and reproductive system

from sessile animals with radial symmetry to highly

motile animals with bilateral symmetry and a
distinct head

from acoelomate to pseudocoelomate to

eucoelomate animals

the complex animals we have discussed so far were

protostomes
ie. mouth develops first during embryonic
stagess

all the remaining phyla are deuterostomes

ie. anus is first to develop

echinoderms, arrow worms, acorn worms &

chordates

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 1

 Phylum: Echinodermata
means “prickly skin”

6225 living species; >20,000 fossil species

“If there are animals from another planet already here,

they’re probably starfish.”

“Echinoderms are the Bohemians of the animal kingdom”

-Burnet & Matsen

include: starfish, sea cucumbers, basket stars, brittle

stars, sea lilies, etc

has an extremely abundant and diverse fossil record

15 classes of extinct species

 more extinct classes than any other animal group

much more diverse fossil record than species existing

today

all marine; found in all oceans at all depths

some of the most abundant of all marine

invertebrates
 unable to osmoregulate; even rare in brackish waters

almost all are bottom dwellers

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 2

 a few are pelagic swimmers

a few are commensal

in general they are not often prey to other species

mostly drab colors

but a few are red, orange, purple, blue, etc

Distinctive Characteristics of Phylum:

1. most with pentamerous (=pentaradial) radial

symmetry

2. no distinct head or brain (no cephalization)

3. most have endoskeleton of calcium plates

4. unique water vascular system for feeding and

movement

5. dermal branchiae for gas exchange

6. no real circulatory system

7. no excretory system

8. sense organs poorly developed

9. pedicellariae for protection

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 3
Origin of Echinoderms

the origin of the phylum is obscure

earliest known echinoderm was Arkarua from Vendian

(560 M Y ago)

echinoderms counter the direction of the story of

evolution so far

in fast paced world they live in the slow lane

echinoderms current body plan probably evolved from

a bilateral ancestor which became sessile:

fossil record shows that attached forms were

once plentiful

radial symmetry is an adaptation toward a

sessile existence
eg. sponges, cnidaria

conditions seem to have favored survival of

motile descendents

only major invertebrate phylum with affinities for

vertebrates
 deuterostome, bilateral, coelomate

Body Form

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 4

most evident feature: radial symmetry

no distinct head

oral vs aboral surface

radial symmetry is a secondary trait

 larvae are bilateral then after
metamorphosis they become radial

in most its pentamerous radial symmetry

Body Wall

epidermis

outer surface covered by epidermis

made up of: epithelial cells
ciliated mucous cells
ciliated sensory cells

nerve plexus in basal part of epidermis

dermis

below epidermis is thick dermis

made of connective tissues

lots of collagen fibers

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 5

 secretes skeletal pieces = ossicles

= endoskeleton

ossicles are bony plates made of calcium

crystals
each ossicle represents a single crystal of magnesium
rich calcite (6(Ca,Mg)C03 )

formed within cells of dermis

in many classes ossicles have bony projections

for defense

unlike any other phylum, echinoderms can vary

rigidity of dermis

pliability of collagen fibers is under nervous

control

= “catch collagen”
soft and pliable  rigid

allows animal to hold various postures for

long periods without muscular effort

beneath dermis is layer of outer circular and

inner longitudinal muscle

true coelom lined with peritoneum

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 6

Movement

movement & food gathering done predominantly by

water vascular system

a second, separate coelomic compartment unique

to echinoderms

derived from coelom and lined with ciliated

epithelium

the whole system operates hydraulically

filled with fluid (mainly sea water and some

proteins and cells

internal canals connect ot the outside through the

madreporite

leads to stone canal (contains calcareous

deposits)

joins ring canal just inside and around the mouth

long radial canals extend into each arm

in arm, lateral canals branch off radial canals

have valves to prevent backflow

lead to small muscular sacs that serve as fluid

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 7
 reservoirs

= ampullae

connected to muscular tube feet

tube feet are concentrated in ambulacral

groove

the tip of the tube feet are flattened, forming

suckers

suctionlike cups can produce strong

force

tube feet used to cling to substrates, move and to

feed

most echinoderms don’t have large muscles

muscles mainly used to move tube feet

but some also attached to ossicles to allow them to

bend and flex

water vascular system also compensates for the

absence of a blood circulatory system

Feeding & Digestion

echinoderms are particle feeders, scavengers or

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 8
 predators

no parasitic species

simple, usually complete digestive tract

but functional anus is often reduced

stomach has 2 chambers: cardiac & pyloric

digestive enzymes are secreted into stomach by

pyloric caecae

Respiration

tiny saclike projections extend through epidermis

= dermal branchae (or papulae)

 exchange respiratory gasses

 get rid of ammonia (N-wastes)

the same functions are also shared by tube feet in

most groups

Circulation

echinoderms rely mainly on coelomic circulation for

transport of gasses and nutrients

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 9

 ciliated lining circulates fluids around body cavity
and into dermal branchiae

coelomic fluid contains amoeboid cells

they do have a blood vascular system (= hemal

system )with heart but its usually rudimentary is
rudimentary

and its function unclear

 may play some role in distributing nutrients

Nervous System

no brain or centralized processing area

circumoral ring and radial nerves branching from it

helps coordinate movement of arms and movement of

the starfish in general

tube feet are innervated by nervous sysem

 enables all feet to move in single direction

if circumoral ring is cut, podia in all arms become
uncoordinated; no movement is possible

few specialized sense organs

have some simple tactile, chemical and

photoreceptors and statocysts
Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 10
Protection

in many starfish the body surface bears small jaw-like

pedicellariae

some are stalked, some sessile (unstalked)

 protect against animals and debris that settle on

the animals surface

Excretion

removal of nitrogen wastes (mainly ammonia) is

through the body surface, dermal branchiae
and tube feet
some amoeboid cells can also engulf nitrogen wastes and move them
to the outside through the dermal branchiae or tube feet

Reproduction & Development

sexes typically separate  dioecious

external fertilization

produce characteristic ciliated, free-swimming,

planktonic larva

= bipinnaria

bilateral symmetry
Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 11
 undergoes metamorphosis to become radially
symmetrical adult

early developmental stages are similar in all classes

some can also reproduce asexually by

fragmentation

many also have excellent powers of regeneration

 can regenerate from 1/5th of oral disc & a single

arm
but may require up to a year

some deliberately cast of an arm as a means of

asexual reproduction

don’t seem to age  can liver forever?

Ecology

a wide variety of other animals make their homes in or

on echinoderms, including:
algae, protozoa, ctenophores, turbellaria, barnacles, copepods,
decapods, snails, clams, polychaetes, fish and other echinoderms

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 12

Echinoderm Classification

Class: Asteroidea
(starfish, sea stars, sea daisies),
1500 living species

Class: Ophiuroidea
(brittle stars, basket stars, serpent stars)
>2,000 living species;

Class: Echinoidea
(sea urchins, heart urchins, sand dollars & sea
biscuits)
950 living species

Class: Holothuroidea
(Sea Cucumbers)
1150 living species

Class: Crinoidea
(sea lilies, feather stars)
625 living species

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 13

 Class Asteroidea
(sea stars, starfish)

~1500 species

free moving

inhabit all seas except low salinity areas

bottom dwellers

mostly found on hard rocky surfaces

many live in deep ocean

also common along littoral zone in coastal waters

where they may congregate in very large numbers

1 cm to 1 M diameter
eg. giant Pycnopodia has over 20 arms and is the size
of a manhole cover

often brightly colored: red, orange, blue, purple, green

etc

best representatives of the basic features of the

phylum

body composed of rays (arms) projecting from a

central disc
Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 14
 arms not sharply set off from central disc

in some arms are very short

eg. Culcita  a pentagon with no arms

mouth and 100’s of tube feet underneath

typically pentamerous symmetry

most with 5 arms

sunstar up to 40 arms

some have up to 50 arms

Oral Surface

mouth in center of oral surface

wide furrows project from mouth into each arm

= ambulacral grooves

each groove contains 2-4 rows of podia

(=tube feet)

margins of each groove are guarded by moveable

spines

tip of each arm has 1 or more tentacle-like sensory

tube feet and a red pigment spot (=eye spot)

Aboral Surface
Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 15
 inconspicuous anus in center of disc

large sievelike madreporite toward one side

aboral surface bears numerous pedicellariae

keeps integument free of sponges, corals

also used in feeding and defense

Movement

mainly by tube feet

can adhere to any solid surface by the suction

created and slowly creep along

~few cm/minute

Feeding and Digestion

many sea stars are scavengers

a few are suspension feeders

feed on small plankton and organic debris

mucous strands carry food to the mouth

most asteroids are carnivores

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 16
 feed on molluscs, crustaceans, polychaetes and
other echinoderms

use chemoreceoptors to detect and locate prey

some can locate buried prey and dig down to

get them
eg. some swallow prey whole and regurgitate
undigested ossicles & spines, etc

eg. some attack larger seastars and begin eating the

end of an arm and work their way up

eg. many are able to evert their stomachs through the

mouth to engulf and eat prey

eg. some feed exclusively on bivalves

 some, such as asterias, are notorious

predators of oysters

wraps itself around its prey

exert steady pull on valves

[force of 12.75 newtons (equivalent to human

lifting 1000lbs wit 1 hand)]

~ a half hour the adductor muscles of bivalve fatigue

and relax slightly

only need 0.1mm gap to insert stomach and

digest oyster

takes 2.5 - 8 hrs to digest a bivalve

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 17

digestive system is arranged radially

mouth at the center of the disc

leads to short esophagus

opens to large stomach that fills most of the

inside of the oral disc

stomach divided into large cardiac region and

small, aboral pyloric region

pyloric ceca (digestive glands) , 2 per arm, drain

into pyloric region

products of digestion in stomach are carried to

pyloric caecae to complete digestion and
absorption

short tubular intestine opens through the anus on

aboral side

Respiration

dermal branchiae (papulae) extend through ossicles

to surface of the skin

these plus tube feet provide most of the gas

exchange for sea stars

in burrowing species, dermal branchiae are protected

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 18
 in channels below umbrella-like spines

Reproduction & Development

Asexual reproduction

many starfish regularly reproduce asexually

 central disc divides in half and animal breaks into two
parts;

each regrows missing part

starfish can also regenerate from an arm

or others an arm and a small piece of the central

disc

Sexual Reproduction

most are dioecious

gonads in small area at base of each arm

when filled with eggs sor sperm they almost

completely fill arm

some lay egg masses

others brood eggs

a few are viviparous

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 19

 but most produce free swimming larvae

gametes released through pores near base of

each arm

1 breeding season per year

1 female may shed 2.5 M eggs

larvae are planktonic, free swimming bipinnaria

larva

metamorphosis converts bilateral larva to radial

juvenile

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 20

Examples of Sea Stars:

eg. sea Daisies

2 known species

discovered in 1986

 inside a piece of wood collected a half mile below surface

at first given their own CLASS; genetic analysis shows them to

be asteroids

disc shaped animals with fringe of short spines

most are <1cm in diameter

contain tube feet around the periphery of the disc

lack arms, mouth, gut and anus

ventral surface may actually be the lining of the stomach which

digests food externally

covered by membranous velum which absorbs

nutrients

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 21

 Class: Ophiuroidea
(brittle stars, basket stars, serpent stars)

~2000 sp

not as diverse in structure as asteroids

but probably the most advanced class of echinoderms

also, the most active of the phylum

found in all types of marine benthic habitats

mainly benthic

tend to be secretive

in cracks and crevices on hard substrates

some can burrow

a few can swim

up to 12 cm diameter

most are fairly drab, a few are highly colored

leathery skin and few cilia

have arms with central disc but:

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 22

 long thin arms sharply set off from disc

no ambulacral groove

 tube feet (podia) play little role in

locomotion

visceral organs are confined to central disc

typically 5 arms

but in basket stars they repeatedly branch to

produce tentacle like mass

Movement

water vascular system:

madreporite opens on oral surface

tube feet have no ampullae

tube feet mainly used in feeding;

not used much for locomotion

muscles are much more important in this group

locomotion by snake-like arm movements

ossicles of arms are arranged into flexible columns

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 23
 (called “vertebrae”) connected by muscle strands

can rapidly clamor over ricks and seaweed

no arm preferences

 can move easily in any direction

Feeding & Digestion

brittle stars are carnivores, scavengers, deposit

feeders or filter feeders

deposit and suspension feeders collect small organic

particles from the water or sediment and use
mucous strands to send food toward mouth

basket star extends it arms into the water to catch

plankton

some carnivores use their arms to capture prey and

“hand” it to mouth

others”ambush” their prey

use arms to hold central disc off grand to form a

kind of shelter

when fish takes an interest in the area under

the starfish it quickly wraps its arms
around it
Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 24
mouth on oral side has 5 jawlike plates

digestive system does not extend into arms

digestive system has sack like stomach,

no intestine or anus

 ie. incomplete digestive tract

Respiration

no dermal branchiae

brittle stars also have internal sacs called bursae

5 pairs of invaginations open toward oral surface at

genital slits

bursae are connected to outside by slits on

margins of each arm

water circulates in and out of the bursae for gas

exchange

many species can actively pump water in and out

of bursae

can also use tube feet for respiration

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 25

Circulation

hemal system is reduced as in starfish

Nervous System

same as asteroids

Excretion

bursae may also be main excretory organ

Reproduction & Regeneration

sexual reproduction

mostly dioecious, a few species are monoecious

gametes discharged into bursae then through

thegenital slits to the outside

some brood their young in the bursae

most species produce a free swimming larva =

ophiopleuteus

it metamorphoses into the juvenile

asexual reproduction & autotomy

brittle stars can spontaneously cast off arms

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 26
 the cast of pieces can regenerate into whole
brittle stars

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 27

 Class Echinoidea
(sea urchins, heart urchins,
sand dollars & sea biscuits)

~950 sp.

widely distributed in all seas

all are benthic

remain close to the substrate

typical urchins seem to prefer hard substrates

some, eg sand dollars and heart urchins like to burrow in

softer sandy substrates

compact body enclosed within a test (or shell) of

closely fitting ossicles or plates

plates are sutured firmly together

most are more or less hemispherical in shape

no arms, but 5 ambulacral areas on test through

which very long tube feet extend

many have developed a secondary bilateral symmetry

with numerous long moveable spines

most 6-12 cm dia; a few to 36 cm

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 28
many colors: brown, black, purple, green, white, red

Body Form

oral side:

mouth contains a comlex chewing mechanism of 5

converging teeth attached to muscle bands

= Aristotle’s lantern

used to scrape and chew algae from rocks

around mouth are circle of heavy modified tube

feet

also a ring of gills

aboral side:

anal region

genital opening

madreporite

Movement

use very long tube feet and prehensile spines

in most urchins, moveable spines cover most of the

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 29
 body

have ball & socket joints with tubercles on

test

Feeding & Digestion

most sea urchins are grazers

scrape algae from substrates with teeth

a few boring sea urchins feed on encrusting algae on

walls of their burrows

Respiration

mainly by tube feet

in sand dollars the respiratory podia are arranged in flower petal
pattern on aboral surface

some have modified tube feet that form branched gills

Hemal System and Nervous System are similar to

basic plan

Protection

spines:

collagen fibers can make spines stiff and erect for

protection
Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 30
 in some urchins spines are hollow and can inject a
painful poison

pedicellaria:

all echinoids have pedicellariae

some types contain poison glands

pedicellaria are also used to keep surface clean

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 31

 Class: Holothuroidea
(Sea Cucumbers)

~ 1150 sp

rule the deep ocean benthos

 make up 90% of biomass on deep ocean floor

often on sandy or muddy bottoms

some crawl on sea floor

others hide beneath rocks

some are burrowers

range from 3 cm to 1 M long

most are black, brown, or olive green

sea cucumbers are among the strangest of the

echinoderms:

like sea urchins have no arms

have ambulacral areas instead

tend toward bilateral symmetry:

polar axis is elongarted so some become

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 32
 long and wormlike or

“cucumber shaped”

“U-shaped”

with mouth and anus are on opposite ends

bottom side = “sole” on which they crawl

body has a leathery appearance

in most the ossicles are greatly reduced to

microscopic plates embedded in body wall

a few are covered in armor of calcareous

plates

mouth is always surrounded by 10-30 tentacles

(modified tube feet) which are part of the
water vascular system

tentacles are highly retractile

 can be completely retracted into mouth

tube feet can also be modified into sensory

papillae, fins, sails, etc

Body Wall
Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 33
nonciliated epidermis with thick dermis below

microscopic ossicles embedded in dermis

beneath dermis is layer of circular then longitudinal

muscle

Movement

large fluid filled coelomic cavity serves as a

hydrostatic skeleton

relatively sluggish

lie on bottom or burrow into mud

many hard bodied forms live beneath stones or in coral

crevices

there are a few deep water pelagic (swimming) forms

move by wavelike contractions of body wall

have tube feet modified into fins, sails, or

medusa-like bells

Feeding & Digestion

mainly deposit feeders and suspension feeders

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 34

use tentacles to collect food and deliver it to mouth

mouth opens into a muscular pharynx

then to esophagus and stomach

some holothurians lack stomach

most have a long, looping intestine

leads to anus which opens into cloaca

cloaca = chamber in which digestive system, excretory system
(if present) and reproductive system all open into

Respiration

most have a respiratory tree consisting of two highly

branched tubes

its attached to the cloaca

pumping action of cloaca circulates water through

it
a tropical pearlfish makes its home in the respiratory tree of
some sea cucumbers

leaves at night to search for food

Circulation

water vascular system is similar to other classes

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 35

but madreporite is not on the surface of the body but
in the coelom

its filled with coelomic fluid rather than

sea water

coelom is connected to sea water via ducts in the

wall of the cloaca

holothuroideans have the best developed hemal

system of all echinoderms

lack beating heart

peristalsis of dorsal vessel circulates blood

blood is like coelomic fluid

involved in some transport of gasses and nutrients

Nervous System

as in other classes

Excretion

respiratory tree also serves as main excretory organ

Protection

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 36

many sea cucumber are capable of evisceration
the front or back end ruptures and the internal organs are
expelled

seems to be a seasonal phenomenon

possibly when food is scarce or in order to eliminate wastes

stored inside the internaltissue

the organs are later regenerated

may be a protective mechanism

a few sea cucumbers posess a large mass of white,

pink, or red tubules (= tubules of Cuvier)
attached to the base of their respiratory tree
when irritated or attacked, the anus is directed toward
the intruder and the tubules are shot out of the anus

in some the tubules are sticky

in others they release a toxin

small crabs and lobsters may be rendered completely

harmless and helpless

the sea cucumber later regenerates the tubules for the

next attack

Reproduction

most are dioecious; a few are hermaphrodites

with single gonad

only group of echinoderms with single gonads
Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 37
some brood their young inside coelom

most have external fertilization

free swimming larva = auricularia

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 38

 Class: Crinoidea
(sea lilies, feather stars)

~625 species

an ancient group; many fossil species

some are stalked sessile animals

others are free living and motile

 can swim or crawl short distances

many are deep water forms

most live at depths of 100 M or more

but a few are found in the intertidal zone

Body Form

flower shaped body

sometimes attached to the end of a stalk

body disc, = calyx, is covered in leathery skin

covering calcareous ossicles

upper surface of calyx bears mouth and anus

arms have pinules giving feather-like appearance

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 39
 stalk if present, sometimes has cirri

no madreporite, spines or pedicellariae

the water vascular system uses coelomic fluid

no madreporite to take in seawater directly

Feeding and Digestion

crinoids are suspension feeders

very slow metabolism

can live for 1000’s of years

Reproduction

dioecious

either brood eggs or release them to produce

doliolaria larva

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 40

 Ecology

sea stars are often the top predators in some benthic

communities

though unpalatable to most organisms to some they

are the preferred meal:

eg. some fish with strong teeth

eg. sea otters

Economic/Human Impacts
1. echinoderms never attack humans

don’t transmit any diseases

although handling poisonous forms can kill

2. “crown of thorns” starfish destroys Pacific coral

reefs

feed on coral polyps

sometimes attack in “herds”

the number of reef attacks is increasing

sometimes results in extensive damage

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 41
 very expensive to control outbreaks

3. sea urchins destroy kelp forest

but are preyed on by sea otters

4. predatory starfish can devastate commercial clam

or oyster beds
eg. a single starfish can eat a dozen clams or oysters in a day

sometimes an infestation is treated with quicklime

 destroys dermal branchiae and kills animal

 but also kills many other soft bodied invertebrates;

but not the oysters who temporarily close their

shells

5. as food:

eg. in China and Pacific Islands sea cucumbers are

boiled and dried and eaten as a delicacy
or used as a food flavoring
in some areas collecting has severely depleated their
populations

eg. roe (gonads & eggs) are sold, raw or roasted,

as a delicacy in Japan and in sushi restaurants
>30M pounds of urchins were harvested in 1986

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 42

6. echinoderms have been widely used in
developmental research
“we know more about the embryology of echinoderms
than probably any other embryo”

Animals: Phylum Echinodermata; Ziser Lecture Notes 2008 43

 Synopsis of Phylum Arthropoda (Arthropods)

Identifying Characteristics of Phylum:

-includes: crabs, crawfish, shrimp, spiders, scorpions, mites, ticks, millipedes, centipedes,
insects (dragonflies, butterflies, ants, wasps, beetles, etc)
-more species in this phylum than in ANY phylum of ANY kingdom of life
-more widely distributed over the earth than any other animal phylum
-paired jointed appendages  the only invertebrate with this trait
-hard (sclerotized) exoskeleton of chitin completely covers body; must molt to grow
-segmented body; allows infinite possibilities for adaptive modifications
-well developed head (cephalization)
-numerous sense organs; antennae & compound eyes are characteristic sense organs of arthropods
-several pairs of jointed feeding appendages
-very active and energetic animals  most active invertebrate group; virtually every form of animal
movement is found in arthropods: walking, running, crawling, burrowing, swimming, flying,
-were the first animals to move onto land; allowed wide and rapid distribution and dissemination
-arthropods have a very complex muscular system; some with 1000’s of muscles
-virtually every mode of feeding: carnivores, herbivores, omnivores, parasites
-well developed complete digestive tract: mouth - esophagus - stomach - intestine – anus
-variety of respiratory systems; gills, book gills, lungs book lungs, trachea
-arthropods have a simple open circulatory system  coelom a haemocoel filled with blood;
has dorsal heart and blood vessels; blood with pigments to carry oxygen:
-ladderlike nervous system similar to annelids
-well developed sense organssimple eyes = ocelli, compound eyes , antennae, chemoreceptors, tactile
hairs & spines, statocysts
-arthropods have a variety of efficient excretory systems; antennal glands, malpighian tubules, coxal
glands, some aquatic forms excrete through skin or gills
-mostly dioecious with lots of variation in developmental stages, a few reproduce parthenogenetically

Subphylum: Trilobita
-completely extinct subphylum; 4,000 fossil species; earliest arthropod group
-named for the division of the body into 3 longitudinal lobes
-highly specialized marine bottom dwellers (benthos) from shallow flats and reefs to deeper waters
-dominated marine benthos for 300 Million years

Subphylum: Myriopoda (centipedes and millipedes)

-long worm-like segmented body divided into a head and trunk
-head with 1 pair of antennae and poorly developed clusters of ocelli
-with paired appendages on almost all segments
-mostly terrestrial; generally found in moist humus or in damp areas under rocks and logs
-mandibles for feeding
-insect-like tracheae for respiration
-insect-like malpighian tubules for excretion
centipedes: means “100 feet”; body usually flat in cross-section; one pair of walking legs on most
segments; predators with pair of poison fangs
-millipedes: means “1000 feet”; body usually round in cross section; 2 pairs of legs on most “segments”;
most are scavengers

Subphylum: Chelicerata
-include horse shoe crabs, sea spiders, scorpions, spiders, ticks, mites, sea scorpions
This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions
 -most members of the group are terrestrial1st group of animals to successfully make transition to land
-head is fused to thorax = cephalothorax, abdomen behind this
-main feeding appendages are chelicerae (pincer-like or fang like) used to grab or pierce or tear prey
-most also have second feeding appendage = pedipalp
-only arthropod group without antennae
-most have 4 pairs of walking legs
-aquatic species have book gills; terrestrial species use book lungs or tracheae
-aquatic species have simple and compound eyes; terrestrial species have several pairs of simple eyes
-excretory organ of most Chelicerates are malpighian tubules and coxal glands at base of some legs
-dioecious; some with elaborate mating rituals; some with considerable parental care

Human Impacts of Chelicerates:

-spiders are directly beneficial as predators each kill 1000’s of insect crop pests
-large infestations of some mites can damage food and ornamental plants by sucking their juices
-venomous species  a few are deadly; eg. black widow, eg. brown recluse, eg. scorpions
-arachnid Diseases and Parasites: ticks, mites, chiggers, etc.
-more serious impact on humans is as disease vectors: eg mites and ticks
-scientists are experimenting with venom genes to use as biological control against insect pests
-spider silk is being investigated for a variety of possible uses.

Subphyulum Crustacea
shelled creatures; “the insects of the sea” eg: lobsters, crayfish, shrimp, crabs, water fleas,
copepods, barnacles, pill bugs, etc
-crustaceans are mostly aquatic, the great majority are marine
-in most crustaceans today, the body is usually divided into a cephalothorax, abdomen and tail
-cephalothorax usually with 2 pairs of antennae & compound eyes; often has carapace extending over
the sides of the animal
-abdomen usually with pairs of jointed appendages on most segments
-generally have many pairs of appendages; most appendages are biramous
-use jaw-like mandibles as main feeding structures; also maxillae and maxillipeds
-great variation in feeding types: predators, suspension feeders, scavengers, etc
-respiration in small crustacea: no special organs ; in larger crustacea usually with feathery gills
-in some crustacea development is direct but most crustaceans produce a variety of distinctive larval forms
as the animal develops eg. nauplius

Ecological & Economic Impacts of Crustaceans

-crustaceans feed a vast number of other animals in the oceans and in freshwaters
-many symbiioses; mutualistic, commensal and parasitic forms
-more than 10 million tons of crustaceans are harvested for food each year (2007)
-crayfish are commonly sold and used as bait either live or only the tail meat
-crayfish & land crabs are kept as pets
-many crustaceans are serious pests; eg rice crabs, crayfish, barnacles, etc
-many Crustaceans are endo- and ectoparasites on other organisms
-some act as intermediate hosts for human parasites; eg Guinea worm, tapeworms

Subphylum: Hexapoda (insects)

-most successful & widespread group of all life; today insects have spread into all major habitats
-were the first animals to fly
-body in three parts: head, thorax and abdomen

Animals: Arthropods-General Ziser Lecture Notes, 2012.10 2

 -head with large compound eyes, several (usually 3) simple eyes (=ocelli), 1 pair of antennae,
mandibles and other mouthparts for feeding
-thorax divided into three segments 6 legs; most also have 2 pairs of wings
-abdomen contains reproductive organs; females have ovipositor to lay eggs
-great diversity of leg types; walking, running, jumping, swimming, digging, climbing, grasping
-insects feed upon almost every kind of organic substance; the same basic mouthparts are modified in
many ways to facilitate different methods of feeding:
-respiraton by tracheal system with spiracles  adaptation to air
-they have a rich supply of sense organs located all over the body; these contribute to a rich diversity of
insect behaviors
-many insect species exist as colonies eg. ants, bees, wasps, termintes, some beetles
-insects defend themselves in a variety of ways; hard exoskeleton, quick reflexes, defensive postures,
spines & bristles, use of sound, warning and camoflage colors, chemical defenses, stings
-excretions using malpighian tubules to absorb metabolic wastes from blood and drain into intestine
-wide range in life spans for adult insects: hours to years
-insects are dioecious; most have internal fertilization; mating is an important part of an insects behavior
set; insects usually lay many eggs; some lay eggs on specific plant or animal
-most insects also go through several distinct developmental stages as they grow from egg to adult
-insects often have complex development including metamorphosis; most of the rest have incomplete
metamorphosis
-some insects go dormant in adverse conditions; or diapause for extended dormancy

Ecological & Economic Impacts of Insects

-insects are the most important organisms in most terrestrial ecosystems; without insects, most of the
terrstrial life on earth would disappear
-Important in Recycling of nutrients eg. 90% of all dead animals are eaten by ants
-insect pollinators are keystone species in some terrestrial ecosystems
-insects have formed a wide variety of symbioses with virtually every major kind of living organisms
-in the US a 2006 study estimates that insects directly or indirectly contribute more than $57 Billion to
our economy
-some insects have been semi-domesticated: eg honeybees, silkworms, mealworms, crickets
-commercial products; eg. chitin, shellac, dyes
-venomous Insects: ants, bees, wasps, hornets, blister beetles, etc
- parasites & Diseases & Vectors; mosquitoes, bed bugs, lice, fleas, flies
-insect and spider silk is being investigated for a variety of purposes
-insects as food; in many parts of the world, insects are considered delicacies
-blood sucking insect, Dipetalogaster maximus, is used as a high tech syringe
-insects as chemical detection devices; “wasp hound” = a portable hand held odor detector
-scientists hope to harness the activities of termite bacteria to break down cellulose to produce ethanol and
biofuels

Animals: Arthropods-General Ziser Lecture Notes, 2012.10 3

 Synopsis of Phylum Annelida
(segmented worms, bristle worms)

Identifying Characteristics of the phylum

-large successful phylum in water & on land; include earthworms, sand worms, bristle worms, clam
worms, fan worms, leeches
-worldwide distribution: marine, brackish, freshwater and terrestrial; some live in tubes they secrete or
make with sand or shell
-elongated wormlike body with head-body-pygidium; true coelom present
-most show some degree of cephalization with a distinct head (=prostomium) tentacles, palps and
sensory structures; mouth with pharynx and chitinous jaws
-body with well developed metamerism (=segmentation); seen in just a few other major phyla;
segments are separated by tissue = septae; allows more efficient hydrostatic skeleton
offers a way to achieve greater size:
-most annelids have paired appendages on most segments = parapodia; used for locomotion,
respiration, in some, parapodia modified into fans and mucous bags for feeding terminal -
-body wall a single layer of epidermis; epidermis secretes a thin flexible cuticle and setae
-beneath epidermis is two layers of muscle tissue
-coelom is filled with fluid (except leeches) & serves as hydrostatic skeleton for burrowing, crawling
and swimming
-complete digestive tract “tube within a tube” design; with pharynx, esophagus, crop, gizzard,
intestine (with typhlosole on dorsal surface), anus; chloragogue cells line surface of intestine
-respiration by gills, parapodia, or through the body wall
-closed circulatory system with dorsal and ventral vessel and several pairs of pumping hearts; blood
contains pigments to carry oxygen
-pair of cerebral ganglia; paired ventral nerve cords; ladderlike connections in each segment
-senses; simple photoreceptors, some with complex eyes, statocysts, chemoreceptors, tentacles, palps
-one pair of nephrida (=metanephridia) in each segment
-both asexual and sexual reproduction; monoecious or dioecious; larva, if present = trochophore

Class: Polychaeta (Sand Worms)

-largest, most diverse and most primitive class of Annelids
-sand worms, bristle worms, fan worms, clam worms, etc
-mostly marine; a few found in freshwater
-deposit feeders, filter feeders, predators, scavengers; some have elaborate filtering structures
-distinct head with mouth and sense organs
-most body segments have appendages = parapodia with setae
-free swimming polychaetes are mostly predators; sedentary polychaetes are filter or deposit feeders
-most are dioecious; gonads appear as temporary swelling of peritoneum at certain seasons
-some polychaetes live most of the year as sexually immature individuals = atokes; become sexually
mature and swollen with gametes = epitokes

Class Oligochaeta (Earthworms)

-mostly terrestrial; most abundant ‘worms’ on land; also many live in freshwaters
-relatives of sand worms but no parapodia and very few setae
-no distinct head
-most are scavengers on decaying organic matter; mainly burrowers; eat as they burrow then let
digestive system extract nutrients
-typhlosole in intestine improves absorption of nutrients
-no respiratiory organs or parapodia like polychaetes; breath through skin, no lungs or gills
This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions 1
 -earthworms are hermaphrodites; cross fertilize each other; use clitellum to form coccoon

Class Hirudinea (Leeches)

-mainly freshwater; a few marine and terrestrial
-many are carnivores; some are parasites
-body is dorsoventrally flattened with anterior and posterior suckers
-coelom is filled with connective tissue and muscle
-no parapodia ; no setae; leeches have poor hydrostatic skeleton
-most are predators of snails, worms and insect larvae; some are scavengers; some are blood sucking
parasites
-very slow digestion; can live for almost a year on one meal
-most exchange gasses through skin; a few aquatic forms have gills
-hermaphroditic; cross fertilize during copulation
-do have clitellum to produce coccoon that receives eggs and sperm

Ecological and Economic Impacts of Annelids

Polychaetes
-detritus food chains; prominent in marine food webs
-beardworms entire ecosystem not based on photosynthesis; common in hydrothermal vent
communities
-Major decomposers of deep sea whale carcasses
-human food (samoa)
Oligochaetes
-detritus food chains
-important in keeping soil fertile since they are constantly turning over earth and mixing
organic matter into it
-Food for birds and other animals
-Food for Humans
-Fishing bait
Hirudinea
-medicinal uses; in past centuries medicinal leech, Hirudo, was used to suck out “bad blood”
today leeches used in medicine to speed healing of reattached fingers and limbs
-commonly used in biology labs
-leeches have become leading research models for understanding how the nervous system works
-some chemicals used by the leech in obtaining and digesting blood are being studied for treating
circulatory diseases
-leeches have also affected history: eg. land leeches of India

This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions 2
 Synopsis of Phylum Mollusca (Molluscs)

Identifying Characteristics of Phylum:

-second largest phylum of animals in terms of number of known species
-most versatile body plan of all animals
-triploblastic with true coelom (eucoelomate); protostome
-bilateral symmetry; some with secondary assymetry
-soft, usually unsegmented body consisting of head, foot and visceral mass
-body usually enclosed by thin fleshy mantle
-mantle usually secretes hard external shell
-complete digestive tract, many with a radula, a rasping or scraping feeding organ, stomach,
digestive glands, crystalline style, intestine
-respiratory system of gills in aquatic forms or "lung"-like chamber in terrestrial forms
-most with open circulatory system; body cavity (coelom) a haemocoel while cephalopods have a
closed circulatory system
-CNS is a ring of ganglia in head area with paired nerves and ganglia extending to other parts of
the body
-usually 1 pair of nephridia (=metanephridia) often called kidneys (not really true kidneys)
-marine forms with characteristic trochophore larva; freshwater bivalves with glochidia larva

Class: Polyplacophora (Chitons)

-fairly sedentary; may move short distances to feed
-head and cephalic sensory organs reduced
-shell contains 8 overlapping plates on dorsal surface; can roll up like pill bugs/armadillo
-most feed using radula to scrape algae from surface
-mantle forms a girdle around margins of plates
-broad ventral foot attaches firmly to substrate
-gills suspended in mantle cavity along sides of thick
-flat muscular foot

Class: Scaphopoda (Tusk Shells or Tooth Shells)

-single tubular shell open at both ends
-mantle wraps around viscera and fuses to form tube
-feeds mainly on detritus and protozoa caught by cilia on foot or using captacula
-radula carries food to gizzard for crushing

Class: Bivalvia (Clams)

-shell is laterally compressed; 2 valves (right & left); consists of 3 layers; periostracum, prismatic
layer, nacreous layer
-“bulging” part of shell on dorsal side near hinge = umbo
-shell held together dorsally by hinge and adductor muscles extending between shells
-shell is secreted by mantle; their mantle can also produce “pearls”
-posterior portions of mantle come together to form incurrent and excurrent siphons
-most bivalves are filter feeders: gills are used to filter food out of water
-in stomach, food is sorted; a gelatinous rod (= crystalline style) spins slowly (by cilia) & dissolves
to release digestive enzymes
-3 chambered heart wraps around intestine in pericardial cavity on dorsal side of body
-freshwater bivalves have internal fertilization; gills become brood chambers; produce bivalved
glochidia larvae

This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions
Class: Gastropoda (Snails)
-means “belly foot”
-largest and most successful class of molluscs
-unlike clams, snails and slugs have a distinct head with brain, sense organs (ocelli, tentacles,
chemoreceptors) and mouth
-mouth with radula
-elongated body with foot below for gliding
-mantle secretes a single shell, often with operculum, and forms dorsal surface of animal
-most shells show some degree of coiling; in addition to coiling, some animals also show tortion
- most gastropods are herbivores; use radula to scrap algae off of hard surfaces
-simple gills are variously modified in aquatic forms
-terrestrial snails have mantle cavity that serves as a “lung” with pneumostome
-many gastropods perform elaborate courtship ceremonies

Class: Cephalopoda (Octopi and Squid)

-means “head foot”
-most fossil forms had very large heavy shells kept buoyant by gas filled inner chambers; only a
few today with large external shell; some have internal shell completely enclosed by mantle;
some have completely lost shell and mantle encloses and protects animal
-in most cephalopods the mantle serves as the animals outer covering
-the surface of the mantle is covered by pigment cells called chromatophores
-the “head-foot” is elongated into 8 or 10 tentacles (up to 90 in nautilus) and 2 longer arms
-mouth at center of arms; contains chitinous beak or jaws
-all cephalopods are predators; use tentacles and arms to capture and handle prey
-closed circulatory system more efficient for gas exchange and transport
-largest brain of any invertebrate, generally considered the cleverest of all invertebrates and rival
mammals in some ways
-most cephalopods have an ink sac for protection
-during mating, before copulation, males often make color displays to compete against rival males
-sperm encased in packets = spermatophores

Ecological Impacts of Molluscs

1. important in food webs in aquatic ecosystems and even in terrestrial ecosystems
2. snails are major source of calcium for birds
3. oysters are keystone species since they tend to form reefs nearshore
4. freshwater bivalves are now the most threatened group of invertebrates in the US
5. Bioinvasives

Human Impacts of Molluscs

1. tusk shells used as money (=wampum) by native Americans
2. as food: oysters, scallops, mussels, octopus, squid
3. precious “stones”
4. ink  sepia dye
5. cuttlebone from cuttle fish
6. Pharmaceuticals
7. shell collecting
8. pollution control
9. destructive species

This is NOT “what’s on the test”; this is a summary of the major points from lab and lectures; the lecture & lab notes are the sources of exam questions
 Phylum Tardigrada
(Water Bears) bodies are short, plump and cylindrical

~900 sp.; 2 fossil species 5 apparent body segments

tiny, highly specialized animals 4 pairs of stubby unjointed legs

<1.2 mm long; most .3-.5 mm each leg with 4 claws

seldom encountered unless looked for Body Wall

most live in temporary water films on mosses and lichens on body covered by cuticle secreted by epidermis
trees, stones or forest floor (limnoterrestrial)
smooth or ornamented
some live in freshwaters

some on bottom of detritus or aquatic algae or mosses in some cuticle is divided into segment-like plates
also a few marine forms
cuticle contains chitin and is periodically shed

short, barrel-shaped body with distinct head and 4 musculature of separate muscle bands is
pairs of short knob-like legs arthropod- like

most species are cosmopolitan contain both smooth and striated muscles

easily dispersed by wind the number of cells that comprise the epidermis is
constant within a species
many are cosmopolitan
à used to help identify species
à easily dispersed by wind
Movement
2 known fossil tardigrades have been described from
Cretaceous amber move slowly crawling on 4 pairs of telescopic legs
Animals: Phylum Tardigrada, Ziser Lecture Notes, 2011.11 1 Animals: Phylum Tardigrada, Ziser Lecture Notes, 2011.11 2

using hooks to grab substrate Nervous System

st
1 3 prs à to move foreward
nervous system is arthropod-like
4th pair à to move backwards
relatively large brain
has both a pseudocoelom and a small coelom
two ventral nerve cords
Feeding & Digestion
segmental ganglia
most tardigrades feed on plant or algae cells
have pair of eyespots
à use a pair of stylets to pierce cell
(nematode-like) most have sensory bristles and spines especially on
head
then suck out fluids with muscular pharynx
Reproduction & Development
others feed on detritus in soil
reproduction is typically from late fall to early spring
some are predators and consume body fluids of
nematodes and other soil animals most are dioecious with single gonad

no circulatory or respiratory system in some species males are unknown and reproduction
is probably by parthenogenesis
most of body is a hemocoel
1-30 eggs are laid at a time
respiration mainly through body wall
eggs are sometimes left behind in the shed cuticle as
Excretion the females emerge

some tardigrades have malpighian tubules branching direct development less than 2 weeks
from intestines (arthropod trait)
Animals: Phylum Tardigrada, Ziser Lecture Notes, 2011.11 3 Animals: Phylum Tardigrada, Ziser Lecture Notes, 2011.11 4
tardigrades apparently can live up to 60 years takes 4 minutes to 7 hours to revive from cryptobiotic
state
Cryptobiosis (suspended animaltion)

tardigrades are typically active only a few months out

of the year

similar to some nematodes and rotifers

can enter state of cryptobiosis

water content goes from 85% to 3%

body becomes barrel-shaped

can remain dormant 4 to 7 years

tardigrades may rank as the most resilient animals

active or in cryptobiosis tardigrades have

survived:
without water for 120 years
temperatures as high as 304ºF (151ºC)
temperatures near “absolute zero” -458ºF (-272ºC)
doses of X-rays and gamma rays that are lethal to almost
all other life forms
lack of oxygen
preservatives (eg. absolute alcohol, ether)
exposure to vacuum of space for 10 days
pressures 6 times that at the bottom of the ocean (6000 x’s
pressure at sea level)

Animals: Phylum Tardigrada, Ziser Lecture Notes, 2011.11 5 Animals: Phylum Tardigrada, Ziser Lecture Notes, 2011.11 6
 Phylum Onycophora à can conform to fit in small places
(Velvet Worms)
mostly nocturnal
70 species
àtend to avoid light
an ancient phylum
à has changed little since the Cambrian (560MY) defend themselves by squirting sticky liquid up to 30”

believed to be a phylum closely related to arthropods Body Form

have chitin cuticle on outside of body long cylindrical segmented, wormlike body
antenna
mandibles
tracheae entire body is covered by bumps (=tubercles) which
similar nervous sytem are covered by tiny scales
haemocoel à giving the body a velvet appearance
annelid like simple eye at base of antennae

have 14-23 pairs of stubby legs

wormlike segmented body; 1.4-15 cm (to 6”) long
each leg has a pair of claws
àlooks like slugs with legs
anterior end with pair of antennae and ventral
thought to be a mollusc when first discovered in 1826 mouth

blue, green, orange or black Body Wall

originally a marine animal (Burgess Shale) a. body covered by thin and flexible cuticle

today all are terrestrial same layers as those found in arthropods

found in rainforests and semitropical habitats b. epidermis secretes cuticle

live in crevaces and under logs c. beneath epidermis are 3 layers of smooth, annelid-
like muscle
Animals: Phylum Onycophora; Ziser Lecture Notes, 2009 1 Animals: Phylum Onycophora; Ziser Lecture Notes, 2009 2

arthropod like tracheal system of tubes opening to

muscle unstriated à contract and move slowly the outside by spiracles

d. body cavity a true fluid-filled coelom Circulation

Movement have arthropod like haemocoel with open circulatory

system and dorsal tubular heart
fluid filled coelom is used as hydrostatic skeleton
Excretion
onycophorans crawls slowly on peglike legs
each segment contains a pair of nephridia similar to
Feeding & Digestion those in annelids

well developed digestive system Nervous System

mouth with a pair of long claw-like mandibles similar to that in annelids and arthropods

most species are predaceous small annelid-like eye at base of each antenna

àfeed on snails insects and worms poor vision

have slime glands that shoot milky fluid that quickly Reproduction & Development
congeals into slime
all are dioecious
can shoot slime up to 30”
sperm packet is deposited on female
once captured they secrete enzymes (salivary
secretions) into prey to partially digest it before lay eggs or bear live young (ovoviviparous)
eating

Respiration
Animals: Phylum Onycophora; Ziser Lecture Notes, 2009 3 Animals: Phylum Onycophora; Ziser Lecture Notes, 2009 4
 Subphylum: Hexapoda
eg. roaches were the main insects of the time
(insects)
eg. some dragonflies had 2’ wingspans
~1.1 M species; probably millions more
today insects have spread into all major habitats
there are more species of insects than all other animal
species dominant fauna of all freshwater and soil habitats

à entire field of study = Entomology some live in deep underground caves

most successful & widespread group of all life on top of world’s highest mountains

adapted to land before most other terrestrial animals some insects live in unusual habitats:
except for a few Chelicerates
some flies occur by the millions in brine lakes eg. Great Salt
(Devonian 390 MY) Lake, where hardly any other life forms are able to
survive
adaptations to land (even deserts):
some insects live in hot springs up to 120º F (49º C)
waxy cuticle
varnish layer can close spiracles
many species are found inside ice in anarctica
extract & retain fluids from food and metabolism
(some don’t need any liquid water at all)
larvae of “petroleum flies” live in pools of petroleum around
diapause & resistant eggs
oil wells

had 40 MY to evolve and diversify before serious a few insect species have been found breeding in brine vats
competition for space and resources from other holding human cadavers at medical schools
animal phyla
the “short-circuit” beetle bores into lead cables

à still no birds around yet but only a very few are truly marine; (Why?)

à still not a lot of parasites of insects yet most are <2.5 cm (1/4”)
by carboniferous (~300MY ago ) there many different àsmall size helps them escape enemies
kinds of insects
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 1 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 2

àneed very little food to sustain themselves A few kinds of insects have been semi-domesticated:
range from < 1 mm to 25 cm honeybees

eg. Atlas moth of India has a wingspan of almost 1 foot
eg. Walkingstick of India is up to 15” long
insects invented agriculture & animal husbandry
àmany ants and termites cultivate fungi within
their burrows
à some ants guard aphids that let the ants “milk”
them for nectar-like secretion
3 trillion bees are kept and managed for pollenation and
honey production
silkworms (moth larvae spin coccoons of silk)
each coccoon is made from over a half mile of silk
mealworms & crickets are used to feed pets (or humans) or
for fishing bait
Body Form

may have been 1st to invent slavery body in three parts: head, thorax and abdomen
à there are ~35 species of “slave making ants” that regularly
raid the nests of other ant species and take young back to Head
their own colony
large compound eyes
there they work as they would in their home colony
searching for food, raising young, etc
several (usually 3) simple eyes (=ocelli)
always very closely related species

ants will always do the work of whatever colony they are in

1 pair of antennae
when they emerge as adults
many kinds of antennae
were the first animals to fly
eg. grasshoppers, crickets and cockroaches have long
antennae
à 130 MY before pterosaurs
eg. butterflies have knob on end
insects: 400 MY; devonian
reptiles: 200 MY pterosaurs; late jurassic , eg. moths antennae are featherlike
birds: 150 MY; coexisted with pterosaurs for~90MY
bats: 54 MY (Eocene)

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 3 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 4
 antennae are very complex sense organs in
which different segments control different wings are extensions of cuticle formed by
aspects of an insects life: epidermis
eg. in ants:
Abdomen
one segment detects nest odor and helps prevent
an ant from entering the wrong colony segmented (10-11 segments)
another segment identifies offspring of a specific
queen reproductive organs

another segment detects the ants own feeding females have pincher like or syringe like
trail
ovipositor to lay eggs
another segment helps detect what is needed by
the immature ants it is tending in some social insects it is modified into a
stinger
mandibles and other mouthparts for feeding
Insect Movement
Thorax

divided into three segments most kinds of movement are created by muscular
system (striated muscles like us)
(pro- meso- & metathorax)
insects have a more elaborate muscular system than
à each with sclerites: any other invertebrate group

tergum, pleura, sternum insects have more muscles than most animals
including us
each thoracic segment bears 1 pair of legs
eg. humans have ~700 individual muscles; some insects
have 900 or more muscle organs; some caterpillars
à total 6 legs; thus hexapods have 4,000

most insects also have 2 pairs of wings on insects are remarkably strong, given their small size
thorax
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 5 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 6

eg. a bee can pick up 50 times its weight swimming (diving beetles, many insect larvae)

eg. beetles are the strongest insects; up to 60 x’s their B. Wings

weight

[but their small size relative to weight makes them only appear strong most insects have two pairs of wings
à if insects were as large as humans they would be little if
any stronger than us]
some use both pairs to fly (eg. butterflies)
A. Legs
some the 1st pair cover and protect second pair
(eg. beetles)
great diversity of leg types:
a few have only 1 pair (flies, mosquitoes)
insect legs are adapted for the same kinds of
movements as vertebrates: a few are wingless (lice, fleas)

walking, running, jumping, swimming, Insect Flight

digging, climbing, grasping
insects were the first animals to fly
legs in 5 segments à 130 MY before any other animal

sometimes modified for unsure of evolutionary origin of wings:

jumping (grasshoppers, crickets, fleas) 1. originated as small flaps that first allowed gliding
= “flying squirrel theory”
eg. grasshoppers can jump 20 times length of
body 2. were 1st used as solar collectors to raise body
temperature
à equivalent jump for human would be
1/3rd length of football field later for gliding, then for flight

eg. fleas are probably the best jumpers 3. originated from gills of aquatic forms many have
can jump 8 “ high and 13 inches in length hinged gills on thorax

à equivalent feat in human flying in most insects is like swimming for us

à leap tall bldg in a single bound
storage of pollen (bees)
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
due to size and relative density of air
wings are separate from legs and other
7 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 8
 appendages eg. the common housefly is one of most talented
aerodynamicist on the planet

don’t just move up and down à more maneuverable in flight than birds, bats or bees

à common pattern resembles “butterfly eg. hoverflies are even better

stroke” of human swimmers
can make six turns a second
direct and indirect flight muscles
hover, fly strait up and down, backwards, do somersaults ,
land upsidedown etc
the rate of wing beating varies considerably:
yet: has brain smaller than a sesame seed
fastest
eg. midge >1000bps has only a dozen muscles for flying
eg. housefly beat ~200bps
eg. mosquito ~300bps but are loaded with sensors
eg. honey bee ~ 190bps compound eyes
wind sensitive hairs
more typical beat: antennae
eg. locust ~20bps three light sensors = ocelli
eg. dragonfly ~25bps

slowly beating:
flight greatly improved dispersal ability
eg. white butterfly ~12bps
eg. swallowtail ~5bps some insects are able to migrate 1000’s of
miles or fly at high altitudes:
speed of insect flight also varies greatly:
eg. monarch butterfly flies slowly (à 6 mph)
from slowest insect flight speeds: but can fly 100’s of miles at a time

eg. mosquitoes ~2mph eg. painted lady migrates 4000 miles

eg. houseflies ~5 mph
eg. butterflies ~6mph eg. some butterflies have been seen at 20,000’
eg. honey bee ~7 mph
some wings are only temporary structures
to quite fast
eg. males and queen ants use wings only for mating flight,
eg. hawkmoth ~35mph
then they drop off
eg. horse fly ~30mph
eg. dragonflies à 25 mph
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 9 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 10

Feeding & Nutrition

d. some are predaceous
insects feed upon almost every kind of organic
substance e. some are parasitic

most feed on plant juices and tissues the same basic mouthparts are modified in various
= phytophagous ways to facilitate different methods of feeding:

many feed on dead or decaying organic matter a. chewing mouthparts

= saprophagous
in many insects the mandibles are hardened into
some are predaceous and attack other insects or “jaws” that are used to snip off pieces of plant or
animal matter
smaller invertebrates
eg. many caterpillars feed on specific plants
some are parasitic and feed on blood or living
tissues from temporary hosts eg. dragonflies, some beetles have large strong
mandibles
b. siphoning mouthparts
all insects share the same basic mouthparts:
eg. butterflies & moths: the maxillae forms a long
mandibles à jaws coiled food tube for sucking nectar
hypopharynx à tongue
labrum, labium à lips
maxillae à helps to manipulate food
c. piercing mouthparts
eg. in biting flies the mouthparts resemble a beak for
insects feed on a wide variety of foods: puncturing the skin

eg. in mosquitoes, most of the mouthparts together

a. most feed on plant juices and tissues form a long hollow tube, the labium directs this
tube through the skin like a syringe to draw up blood
b. others actually eat leaves, flowers and
whole plants or animals d. sponging mouthparts

c. many feed on dead or decaying organic eg. in houseflies the labium forms a sponge-like
feeding structure to mop up liquids
matter
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 11 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 12
 larger insects use some kind of ventilation
the digestive tract is a tube divided into 3 major
regions: eg. pumping movement of abdomen

aquatic forms returned to water:

foregut à crop, salivary glands, stomach
initial food processing & temporary storage
à tracheal gills in cloaca eg dragonflies
midgut à usually has sac-like gastric caecae
enzyme secretion and absorption of nutrients à water bubbles
hindgut à intestine and rectum
rectum for water and nutrient absorption;
à return to surface to breath

Respiration Sense Organs, Communication & Behavior

insects are extremely active
an insects blood does not carry much oxygen
don’t have large gas filled sacs taking up a larger part of their they also have a rich supply of sense organs located
body for breathing all over the body
à oxygen is delivered directly to body cells
à these contribute to a rich diversity of insect
through system of tubes and passageways
behaviors
tracheal system with spiracles à adaptation to air most insect sense organs are microscopic in size and
are found on the body wall and various
many spiracles have valves that can open and close appendages:
these airways deliver oxygen to muscle cells 200,000 times small hairs, small domes
faster than our blood vessels (remove CO2 10,000 times as
fast)
keen senses but relatively simple circuits
tracheal system is lined with cuticle à must be shed eg. one signal usually results in one response for an insect

gas movement by diffusion in humans one signal can result in many different
responses

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 13 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 14

1. Vision color vision

eg. plant eating insects are especially sensitive to
vision is the most important sense for most insects green
eg. ~2/3rd’s of a fly’s nervous system is devoted to vision
eg. butterflies that feed on red and yellow flowers are
more sensitive to those colors
insects have two kinds of eyes
(but most insects can’t see red)
simple eyes that can detect light & dark
eg. some insects can see UV used during courtship

and compound eyes that are especially some insects communicate using light
effective in detecting movement
à number and duration of flashes produces unique
simple eyes are called ocelli signature for each species

2. Touch
à usually 2 or 3 on head
most sense organs that respond to touch are
à can detect only light vs dark
small hairs on epidermal setae
compound eyes
most touching involves antennae and mouthparts
with many individual lenses = facets
a single antenna can have over 5,000 sensilla
some as few as 9 facets
these touch receptors can also pick up vibrations
houseflies have 4,000
in air to respond to wind or gentle breeze
dragonflies have 28,000 facets
eg. a fly avoids a swat by ‘feeling’ the air being pushed by
your hand as you try to swat it
provide a wide field of view and particularly
good at detecting movement can also detect temperature, humidity, gravity
eg. this is another reason why many insects are hard
to swat or capture touch receptors are used in a variety of ways:

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 15 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 16
 eg. mole cricket uses antennae to recognize nymphs in
dark burrow
eg. social insects can detect unwelcomed visitor to their
nest and attack them
eg. social insects often stroke and groom each
other with antennae and mouthparts
eg. waggle dance of honeybees involves touching to
communicate location of pollen
3. Hearing & Sound
insects have many different kinds of organs for
hearing
the insects with the best developed hearing are
those that communicate using sound
the simplest are hairs that respond to touch
many insects have hearing organs inside
their legs that respond to vibrations
passing through the ground or a plant
eg. ants come out of nest if you stomp on the
ground
cicadas and crickets detect sound with an
ear-like tympanum
moths, crickets, mantids etc can hear between
25,000 - 45,000 Hz
[humans hear sound waves from 20 - 20,000 Hz]
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
4. Taste, Smell & Chemical Communication
both taste and smell are types of chemoreceptors
they detect specific chemicals
smell à if the chemical is volatile and travels some
distance
taste à if the insect is in direct contact with the source
chemoreceptors are found on all parts of the
insects body
àon setae, antennae, mouthparts, legs
àbut are mainly on antennae and feet
à wasps and crickets know where to lay eggs
because they have chemoreceptors on their ovipositors
receptors for taste and smell have a pore that
receives the chemical to be detected
eg. a single sense organ on the antenna of a polyphemus
moth has 18,000 pores for chemicals to enter
most insects have 4 of the same taste
sensations as do humans: sweet, sour,
bitter and salty
organs of smell do not detect as many different
odors as does a human nose
but it is tuned more finely
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
àcan detect ultrasonic signals of bats to avoid
being eaten
sound may be used to individuals of the same
species
eg. males and female mosquitoes are attracted to
each other by sound of their buzzing wings
eg. tapping sound of death watch beetles (eat wood
on old house -heard by those keeping watch over
dead person before burial) used to attract a mate
many insects that can hear have structures for
making various sounds to communicate with
each other or to other species
some rub body parts together to make sounds
= stridulation
eg. chirping sound of crickets, katydids, beetles and
ants can be used to communicate with group or
seek a mate
some have drum like membrane that can
vibrate to make a sound
eg. cicadas are loudest of all insects
à can be heard half mile away
à songs differ for each species
hissing cockroach blows air out its spiracles to
make sound
many insects can make supersonic sounds
17 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 18
à can detect differences between very
similar chemicals and in much lesser
amounts
some can detect airborne chemicals 5 miles
away
eg. male of lesser emperor moth can smell the
pheromone of the female up to 6 miles away
smell is used to find suitable plant for laying eggs
used by flies to find dead animals
some parasitic wasps can track host species by
smell
smell is also used to communicate within a species
eg. sex pheromones attract mates
eg pheromones that cause grouping behavior
eg. ants release alarm pheromones if disturbed or
threatened
eg. ants, caterpillars and other insects produce trail
pheromones to map route to food
5. in addition to external senses insects also have a
variety of proprioceptors
can monitor body positions, eg legs and wings,
internal pressures
19 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 20
 could not walk or fly without this information
eg. ants, bees, wasps, termites, some beetles

Behavior & Communication they can work much more efficiently as a group than
as solitary insects
insects have keenly developed senses
they have a very structured division of labor
à can be used for communication
and behavioral responses highly dependent on each other

insects communicate in a number of ways using their generally unable to survive long on their own
external senses
à they work almost as a single individual
insect behavior is mostly instinctive (innate)
in social insects workers change tasks depending on
eg. caterpillar is programmed to eat a certain plant
their age
eg. courtship, egg laying, migrations are preprogrammed
younger workers engage in safer chores like
but they do have the ability to learn “home maintenance”;
eg. bee dance
the older workers do the riskier foraging
eg. notice landmarks near nest or food outside the nest
eg. when Colorado potato beetles first attempt to mate,
they are not very good at identifying their own species
they change jobs based on their own
or even distinguishing head from tail; with repeated assessment of their remaining longevity
attempts they get better at it
eg. when harmed in some way they switch to the more
research has recently shown that insects have a similar “fight or risky jobs
flight” response to stress as vertebrates have (same genes)
eg. honey bees
Colonial Insects
Each colony consists of a single queen, about 100 male
drones, and 60,000 workers
only about 10 of the 20 or so orders of insects have
developed some type of social organization
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 21 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 22

Organization of the colony is maintained by pheromones &

secretions produced by the queen
Workers live 4-5 weeks. Young workers feed brood, drones and
queen. Older workers construuct comb using wax from
abdominal wax glands. They also clean the hive and guard
the entrance. Older workers forage.
Bees feed on pollen and nectar; pollen for protein, nectar to
make honey
Foragers that return with nectar perform dances that provide
information on the distance, direction and quality of the
food source; eg. round dances indicate food within about
300 ft of the hive, waggle dances indicate the food is
further away. Nectar is shared with other workers to verify
the quality of the food source.
Honey is made by evaporating water from nectar and adding
glandular secretions. Each colony can produce up to 2 lbs of
honey per day.
The queen deposits an egg in each brood cell. unfertilized eggs
produce drones, fertilized eggs produce workers, fertilized
eggs fed royal jelly produce queens
eg. ants
similar division of labor with one or more queens, drones,
workers, soldiers, etc
in relative size an ant has one of the largest brains among
insects
à each ants brain consists of ~ 1 M cells with the
processing power equivalent to a Mac II computer
(1987-1 meg)
à as a colony of 40,000 individuals, the colony has a
processing power equivalent to the brain of a
mammal

a large bee colony can store up to 100 lbs of honey and 5 lbs of
Insect Defenses & Weaponry
pollen per colony for the winter months
1. hard exoskeleton
Large colonies in late spring produce new queens. Queens are
selectively bred in a special queen cell. They are fed “royal the hard exoskeleton of insects serves as an
jelly” by workers. Only one queen will ultimately survive.
effective defense against many dangers
The ‘old queen’ with about half of the workers will swarm to
eg. large beetles, including weevils, have a particularly thick
establish a new colony
exoskeleton
The new queen makes a single mating flight in the spring with
about a dozen drones. The drone that succeeds in some insects have a “fracture line” in each
impregnating her plugs her genital opening with his broken appendage that allows the leg to break off
off penis after insemination to insure that no other males easily if caught by a predator
can copulate with her. Most drones die shortly after the
mating flight; surviving drones are kicked out of the hive eg. many crane flies, walkingsticks, grasshoppers
The queen acquires enough sperm to last her lifetime of 4-5
years; she can deposit up to 3000 eggs/day.
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 immature insects that sacrifice a leg can often
regrow it after several molts
2. quick reflexes
as the most active invertebrates, most insects can
easily avoid threats by quick escape; flying or
running
most insets have sensitive hairs that can detect
the change in air pressure of a fast moving
hand
eg. cockroaches can respond & escape from an approaching
shoe in less than 50 milliseconds
eg. resting houseflies can move to avoid a swat in 30-50
milliseconds
3. “hold your ground” or “play dead”
some insects, when threatened, remain in place
and adopt a defensive posture
eg. tortoise beetles have strong adhesive pads on their feet;
they lie flat against a leaf and hold on when threatened
eg. cuckoo wasps curl up into a hard rigid ball like pill bugs
for protection
some insects simply “play dead”
if on trees and shrubs; they release their grip
and disappear into the undergrowth
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
mimicry eg. viceroy butterfly
one group of beetles can change color bypumping
fluids into grooves and channels in its clear
exoskeleton
7. chemical defenses
many insects wage chemical warefare using
noxious chemicals to ward off predators
in some cases they manufacture their own toxins
others use chemicals extracted from host
plants
eg. stink bugs secrete a foul smelling chemical from glands
in their exoskeleton when threatened
eg. some species of cockroaches and termites secrete a
sticky substance from anal glands to immobilize ants,
spiders, centipedes or other predators
some of these chemicals can be painful
or cause temporary blindness
eg. Bombadier beetle à shoots hot explosive
secretions toward predator from glands in their
abdomen
each gland has two chambers, one contains
hydroquinones and hydrogen peroxide;
the other contains various enzymes;
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
4. spines, bristles and hairs
eg. many predators will avoid a mouthful of spines and bristles
found on many caterpillars
eg. parasitic flies or wasps have more difficulty getting close
enough to lay eggs in or on the potential host bearing
spines and bristles
5. sound
eg. hissing cockroaches emit a threatening hiss
eg. some moths can mimic the sound of “bad tasting” moths to
avoid being attacked by bats
eg. tiger moths can detect the ultrasonic sounds made by bats
if at low intensities (bat is relatively far away), they just
change course and fly away
at higher intensities (bat is much closer), they quickly drop
from the air in an evasive looping dive
6. Color as warning or camoflage
some insects have bright red or orange colors
exposed only when they are threatened by
predator that may scare predator away
others with large eyespots that may
intimidate
in other color and shape are used as camoflage
camoflage eg. walking stick
25 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 26
when the two are mixed a series of explosive
reactions occur
peroxide breaks down to oxygen and water
and the concotion heats to boiling and blasts
out as a toxic cloud
some caterpillars click their jaws loudly when
predators approach as a warning they will
vomit a distasteful fluid if they don’t back off
in some species of termites, when “soldiers”
become too old to defend the colony they blow
themselves up spraying a toxic brew which
kills them and their attackers
stinging insects use pain as an effective defense
or a way to subdue prey
some insect toxins that cause pain are
expelled by hollow body hairs
simply brushing against them causes them
to break and release their painful
chemicals
eg. saddleback caterpillars
eg. pus caterpillars
some social insects, usually infertile workers,
have their ovipositors modified into
stingers attached to poison glands
eg. bees, wasps, hornets, ants, etc
27 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 28
 sting-pain index based on human perception the goal of this initiation rite is to keep the sleeve
on for a full ten minutes without showing any
0. sting doesn’t penetrate skin signs of pain

1. sharp, sudden, effect similar to a stray spark when finished, the boys’ (now men) arms are
temporarily paralyzed because of the venom, and
eg. fire ant sting they may shake uncontrollably for days

2. a piercing pain Excretion

eg. honeybees, common wasps & hornets, acacia ant
malpighian tubules (2-100’s) absorb metabolic
3. bold, caustic, burning and unrelenting wastes from blood and drain into intestine
eg. paper wasps, harvester ants
Reproduction & Life Cycles
4.blinding, fierce, excruciating and debilitating
wide range in life spans for adult insects:
eg. tarantual hawk, warrior wasps
hours to years
4+. worst sting: pure intense, brillian pain
eg. adult mayflies live for less than a day; do not eat
eg. bullet ant (Paraponera clavata) sting produces
sole purpose is to reproduce
wave after wave of throbbing, all-consuming pain
that continues unabated for up to 24 hrs
eg. species with longest lifespan may be a tropical
termite whose queen may live for 50 years
Fortunately, bullet ants are not aggressive except
when defending themselves or their colony
Mating
Bullet ants are used by some indigenous people in
their initiation rites to manhood: insects are dioecious
the ants are first knocked out by drowning them
in a natural chloroform most show sexual dimorphism with male trying to
attract the females:
then hundreds of them are woven into sleeves
made out of leaves, with stingers facing inward. eg. male rhinoceras beetles with long “horns”

when the ants awaken, boys slip the sleeve onto eg. male stag beetles with large mandibles
their arm.

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 29 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 30

eg. male butterflies often more colorful eg. praying mantis courtship

most have internal fertilization females have predisposition to eat their mate before, during
or after mating

mating is an important part of an insects behavior set eat head first à remove “off switch” of male copulating

to find mates (during mating, the subesophageal ganglia (head)

inhibit ventral ganglia from triggering intensive
copulatory activity)
most use pheromones
eg. Aphids
some use light; eg. lightning bugs
female aphids can reproduce without sex they produce
others use sound or color
clones of unfertilized eggs that develop into females

some mark and protect territories from other males yet they do have sex annually; but not for the males sperm.
They are after “sexually transmitted diseases”
eg. some dragonfly species
beneficial bacteria from the male that can help them
often have courtship rituals using pheromones &/or digest plant food, resist extreme temperatures, kill
sound to attract mate internal parasites

eg. cicadas, crickets, grasshoppers the female can also pass these infections on to her future
clones
eg. deathwatch beetles bang their heads against the sides of
their tunnels to signal a mate many female insects mate only once/lifetime and store
the sperm
eg. mosquitoes and midges use high frequency sound of their
wingbeats to attract a mate
insects usually lay many eggs
eg. “lightning bugs” use distinctive patterns of light to attract
mates eg. queen honeybee >1 M/lifetime

internal fertilization eg. termites lay 10,000 or more/day

male must insert sperm into females seminal receptacles eg. cockroaches can lay >1 million eggs in a year

there is a great diversity in insect reproduction: but some only 1 egg per mating

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 eggs are sometimes enclosed in protective case eg. butterflies: larva=caterpillar
eg. flies: larva=maggot
eg. roaches; praying mantis eg. beetles: larva=grub

some lay eggs on specific plant or animal once adult emerges it no longer molts

many insects “mark” plant with chemical to b. most of the rest have incomplete
dissuade other females from depositing eggs metamorphosis (hemimetabolous):
there eggànymphà adult
à ID of the chemical they use may be
useful as a natural insect deterrent nymph resemble adult without wings

some may care for young after hatching wings develop externally as budlike
growths on thorax
Development
grow by successive molts
most insects also go through several distinct
eg. grasshoppers, cicadas, mayflies,
developmental stages as they grow from egg to stoneflies, dragonflies
adult
c. a few have direct development
insects often have complex development including
metamorphosis egg à juvenile à adult

eg. silverfish, springtails

a. 88% have complete metamorphosis
(holometabolous): metamorphosis is regulated by hormones
eggàlarvaàpupaà adult
3 major endocrine glands in insects:
larva usually has chewing mouthparts brain produce
ecdysial gland juvenile hormone
pupa is a nonfeeding stage before corpora allata & molting hormone
metamorphosis; often in a cocoon
Some insects overwinter as immature stage
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 33 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 34

à don’t complete development until they are Monarch butterflies:

literally frozen in winter months
millions migrate over 2500 miles in autumn from
Diapause breeding grounds in the eastern US and
Canada to a winter haven in Mexico
some insects are able to enter a state of dormancy to
survive adverse conditions; eg temp, humidity the fall migrants are great great grandchildren
of the monarchs that left Mexico the
if this dormancy is for an extended period and previous spring
triggered by daylength = diapause
yet they return to the same dozen or so
Migration roosting areas

many insects migrate they can orient by the sun and can detect
earth’s magnetic field
eg. locusts, butterflies, moths, dragonflies, grasshoppers, some
beetles 2013 saw a 59% drop in migrating monarchs since 2012;
partly due to use of a herbicide that kills monarch’s
generally if an insect is seen: food plant

more than 8 or 10 ft above the ground

flying in a relatively straight direction
ignoring prominent food sources on its journey

it is probably migrating

most fly at high altitudes and catch seasonal winds to

migrate

a few, like monarch butterflies fly at much lower levels

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 35 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 36
 So, Why are insects the most
abundant and most diverse group of
living organisms on Earth?
insects have been the dominant life form for over
400 MY.
-they have witnessed the rise and fall of the
dinosaurs
-they have survived at least 4 of the earth’s mass
extinctions
-they continue to thrive as humans have spread
over the planet and are in the process of
creating our current mass extinction of life
Their success is largely due to a unique combination
of just a few major features:
1. Exoskeleton
their exoskeleton is made of chitin bound with
several proteins making hard and rigid for
protection, yet with flexible and elastic
“hinges” for freedom of motion. This “shell” is
coated with a waterproof waxy layer
their exoskeleton gives insects support needed
for any kind of habitat; protection from
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
eg. A leaf miner spends its entire life tunneling
in a single leaf
eg. in some ant species, an entire colony may
live inside a single acorn or plant gall
eg. parasitic wasps in at least 7 families
complete their entire development within
the egg of other insects.
small size is also an advantage in avoiding
predation
they can easily hide in a crack in a rock,
beneath the bark of a tree or the petal of a
flower
their exoskeleton is hard enough for them to
burrow between individual grains of sand
and to squeeze through the tiniest of
cracks
3. Flight
insects are the only invertebrates that can fly
they flew 100MY before the first flying vertebrate
and they still have the most effective flight
mechanics:
-an insects flight muscles generate twice the
power per muscle mass than vertebrate
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
predators and environmental hazards and
prevents desiccation in dry habitats
muscles are attached directly to the body wall to
combine maximum strength with optimal
leverage and strength
many insects can lift many times their own
weight
2. Small Size
most species are relatively small (<1”)
for an animal with an exoskeleton, small size is a
distinct advantage. The larger the insect the
thicker and more cumbersome it would have
to be
in the real world, “Mothra” and “the Fly” could
never walk much less get off the ground to
fly
another advantage of small size is there is only a
need for minimal resources to survive and
thrive
food requirements are so modest that an
insect may spend its entire life on a single
plant or animal and never exhaust its food
supply
37 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 38
flying muscles, mainly due to
characteristics of their exoskeleton
flight provided a new escape from predators and
an extremely effective means of dispersal
efficient energy use allows some insects to travel
great distances or remain airborne for
extended periods
eg. more than 200 species of moths, dragonflies, locusts,
flies and beetles migrate over long distances
4. Reproductive Potential
insects produce large numbers of viable eggs in
very short time spans
à they produce a large #’s successful
offspring
5. Metamorphosis
in more advanced insects the freeliving larval
stage that lives in microhabitats different from
the adult of the same species
only 9 of the 28 different kinds of insects have
“complete metamorphosis” yet they represent
about 86% of all living insect species
39 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 40
 Ecological Impacts of Insects
eg. 90% of all dead animals (mainly insects) end up as food
1. insects are the most important organisms in most in ant nests
terrestrial ecosystems eg. 99% of human and animal wastes is decomposed at
least partly by insects & other invertebrates
insects are at the heart of healthy ecosystems:
eg. herbivores
insects are an integral part of every landbased
a large number of insects feed on plants but only a small
foodchain percentage of them are considered pests

because of their number, variety and influence many have proven to be effective controls against
bioinvasive plants
on larger organisms and whole ecosystems
eg. a moth that eats prickly pear cactus in Australia
eg. soil insects
eg. a European beetle was introduced to California to
many types of insects spend part or all of their lives control an invasive “goatweed”
underground
eg. pollinators are keystone species in some
many feed on humus or decaying plant matter terrestrial ecosystems (more under
they aerate the soil, recycle organic wastes, fertilize the soil symbioses)

eg. springtails – many millions/acre sexual reproduction in most plants is made possible by the
process of pollination
eg. ants – nest in soil, feed above ground
the majority of pollinators are insects
eg. termites, burrowing bees, wasps, beetles and flies
without a certain insect species to pollinate the dependent
plant species will decline or go extinct
eg. scavengers
eg. carnivores
eg. termites, ants, wood boring beetles, dung beetles, flies

feed on decomposing plants and animals and recycle their keep populations of their prey in check.
nutrients
eg. parasitoids
remove rotting materials
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 41 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 42

insects with parasitic larval stages Insect Symbioses

eggs are laid inside body of host and larval stage hatches insects have formed a wide variety of symbioses with
and grows feeding on host eventually killing it
virtually every major kind of living organisms
at metamorphosis becomes freeliving adult
Bacteria
work with carnivores to control population sizes

eg. many fly and wasp species; tachinids, almost all animals, including insects have bacterial
ichneumonids symbionts

without insects, most of the terrstrial life on earth some that live in digestive system are mutualistic
would disappear
àhelp break down hard to digest fibers and
starches
most of the rest of the world is literally helpless
without them àmake essential vitamins

vertebrates wouldn’t last more than a few months àprotect from pathogens

if all insects suddenly disappeared àprovide additional nutrients from food eaten

invertebrates, especially insects, are “the little

eg. cellulose digesting bacteria
things that run the world”
-EO Wilson
some species of bacteria, some protists, and
most fungi are able to produce the proper
2. Insect Symbioses
enzymes to digest cellulose in plants
insects have formed a wide variety of symbioses
virtually no animal species can produce
with virtually every major kind of living
cellulose digesting enzymes
organisms
animals that are strict herbivores rely on
cellulose digesting microorganisms to
extract nutrients from plant materials
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 43 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 44

eg. termites
many species of cellulose digesting bacteria and
protist that digest the bits of wood eaten by
the termites
a diet restricted to wood only (not softer green
parts of plants) is very low in nitrogen and
termites also have nitrogen fixing bacteria
that converte N2 into useable nutrients for
the termites
Protozoa
eg. cellulose digesting flagellates in the gut of termites
rd
1/3 to 1/2 of a termites weight are these symbiotic
protozoa
work with bacteria to break down cellulose fibers
Fungi
Animal “Fungus Farms”
eg. Atta ants (leafcutter ants)
mounds up to 18 ft underground
the ants maintain active fungal gardens to feed
larvae and adult in the colony
àthey cultivate fungi within their mounds
àthey feed the fungi bits of leaves, bark and other
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
fungus never forms mushrooms in mound
but when rain is imminent
termites carry bits of mycelia out of mound
spread them around the ground
several days after rain mushrooms appear and
produce spores
the termites reappear and collect the genetically mixed
hyphae and take them back into the nest
abandoned mounds produce crops of mushrooms
for several years after termites leave
à African people consider them a rare
delicacy
Plants
eg. Pollination
a large part of the success of flowering plants is due to
the variety of pollenation
à provides much better mixing of genes
flowering plants have coevolved with many kinds of
animals through most of their history
this close relationship between plants and
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
plant material
àthey weed out debris that might spoil their garden
àhave wheelbarrow-like depression on their backs
for carrying spores
fungus is transferred from colony to colony by ants
freeliving cultures of fungus have never been
found
fungus needs ants to provide them with food
in Central and South America leaf cutter ants are
the dominant herbivores in food chains
à consume 17% of total leaf production
eg. African termites (Microtermitinae)
build chimneys up to 30 ft tall of feces and mycelia
eat mostly plant material but don’t have cellulose
digesting microorganisms living in their gut
bring it home and eat it before entering
mound
then go in and defecate
use their feces to grow fungi (Termitomyces)
all stages of termite readily eat the fungus
45 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 46
pollinators has resulted in coevolution
à mutual adaptations for mutual benefits
plant: petals, scent, nectaries
animal: special body parts and behaviors
We like to think of animals as the most ‘evolved’ lifeforms
à but plants lack brains, muscles, speech yet they don’t need them
- they borrow animals to do their bidding
In some instances the relationships have become very
specialized such that only a single species of
animal can pollinate a particular species of plant.
insect pollinators
petals colorful and large
often with nectaries
much less pollen produced
insects can be attracted by showy flowers, smell &/or nectar
eg. in irises the insect cannot get to the nectar without collecting
pollen
eg. milkweed produces packest of pollen that entangle the legs
of any insect that lands on its flower
eg. some plants depend on a single species of insect for
pollenation:
eg. some orchids
eg. long tongued hawk moth
eg. fig wasp and fits
eg. yucca plants are pollenated solely by the yucca
47 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 48
 moth
usually white or dull in color
eg. orchard fruits are almost entirely insect pollenated (beetles cant see as well as bees)
mainly by bees
--------------------- flowers tend to have a strong yeasty, spicy or
fruity odor, sometimes unpleasant to us
a. bee & wasp pollinated
secrete no nectar but may supply food as
pollenate more flowers than any other group pollen or in special storage cells in petals

20,000 different species of bees are important pollinators most eat parts of the flower
for many plants
eg. magnolias, lotus
bees pollenate mainly spring flowers
c. carrion flies
bees have very good vision
tend to be dull red or brown or white
à flowers are generally brightly colored
often have foul odors resembling rotting meat
predominately blue or violet markings
eg. skunk cabbage
rarely pure red (pure red appears black to them)
eg. carrion flower = Rafflesia
flowers generally shallow (short mouthparts)
a parasitic flowering plant of SE Asia
honeybees are attracted to nectar
they also gather pollen early botanists thought it might be a fungus

flowers often delicately sweet and fragrant parasitic plant with no true roots or leaves
parasitic on vines in genus Tetrastigma
flowers often have lines or distinctive (Vitaceae)
markings that function as “honey guides”
à lead bees to nectar produces a huge speckled flower on little stem
with no leaves
some of these markings are only seen in
UV light largest known flower:
à invisible to us, not bees flower bud size of basketball
flower up to 1 m across; weighs up to 10 kg
b. beetle pollinated
smells like rotting meat
flowers generally large and open
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 49 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 50

fruit is eaten by three shrews and other mammals wasps have very good vision

d. butterflies flower preferences similar to bees

but wasps pollenate mainly in summer
prefer sweet fragrance and provide nectar
wasps are much less “hairy” so not very good pollenators
fused petals force insect to crawl into flower for nectar
flowers are generally brightly colored
nectaries are usually at bases of deep spur that
only butterflies and moths can reach with their predominately blue or violet markings
mouthparts
flowers generally shallow (short mouthparts)
some butterflies can detect red flowers
flowers often delicately sweet and fragrant with nectar
eg. daisy family: butterfly bush= Buddelias, or fruit juices
goldenrods, blazing star
eg. fig wasps exclusively pollenate almost 1000
eg. monarchs à pollenate milkweeds species of tropical fig trees

e. moth pollinated eg. some species of orchids resemble certain wasp

females that attract only one species of wasps
white or yellow large tubular flowers
heavy fragrance
eg. Plant Galls
open at night, closed in daylight
galls are any deviation from the normal pattern of
eg. yuccas, night blooming jasmine, night blooming plant growth on a particular organ or part due to
cereus
the presence of another organism
f. ants
all plant organs are attacked: roots, stems, leaves,
dense clusters of small flowers near ground
flowers and fruits
sticky pollen
à leaves are most common organ attacked
with nectar
many galls superficially resemble plant parts such as berries,
eg. some composites, poppies, nuts and other fruit types

h. wasps some galls look like fungi and for years some were misidentified
as a fungal species
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 51 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 52
 much more nutritious than eating typical leaves,
some galls resemble strange sea urchins and coral polyps stems or roots.

galls on trees and shrubs often appear in the spring the gall insect typically becomes trapped inside the
gall and becomes a victim for a diversity of
when the leaves first appear parasites itself

à when they are highest in food quality à creates a complex insect community
within the gall.
plant galls have been known since antiquity (350 BC)
eg. some oak galls support as many as
eg. plant galls were known and used extensively in medicine, 22 species
industry and as human food for over 1000 years in China,
India and parts of Europe eg. another kind of gall had 75 different
species present
plant galls can be produced by a variety of organisms
but mainly various species of insects a gall appears to be a parasitic relationship

à~13,000 gall forming insects are known àthe gall is formed as the insect attacks and
(~2% of all known insects)
wounds the plant
à 70% of galls are produced by flies & wasps
eg. Ant/Plant Symbioses
most insects are highly host and organ specific
eg. ants and acacia trees
insects especially form leaf galls one species of acacia tree is always found with a
colony of a particular ant species associated with it
gall forming insects are highly specialized
plant feeders: the tree is small (5’-6’) and its branches are covered
with pairs of sharp thorns up to 3” long.
tend to be sedentary and feed only on certain cells
No acacia of species Acacia drepanolobium has been
found that did not contain colonies of the ant species,
they induce the plant tissue to grow rapidly inside the
Crematogaster mimosae and C. nigriceps.
gall to provide adequate food
these ants live only on these trees; up to 20,000/tree
the stimulated growth created by the insect
produces a high protein, nutritious meal
ants get room and board:

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 53 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 54

at every 6th pair of thorns is a swelling (=pseudogall) stings

which is a hollow space inhabited by the ants
sting can penetrate elephant hide and can numb small
at base of each leaf are nectaries that exude a sugar mammals (and humans) for several days
solution similar to flower nectar

acacia trees get protection from herbivores

àgrab intruder with strong mandibles and sting it

eg. antelope gets a face full of stings and leaves

eg. ants and tropical epiphytes

some epiphytes have swollen knobby tuber with with hollow

chambers inside inhabited by ants

ants accumulate humus and their excretions nourish the plants

in some species the ants collect seeds of various epiphytes and

plant them in their nests to create “hanging gardens”

they collect animal feces to nourish seeds

some epiphytes reward the ants by producing starch grains or

sugary secretions

eg. ants and rattan vines

certain rattan (Korthalsia) vines have swollen areas that house

ants

if a mammal accidentally brushes against plant the ants make a

rattling noise by beating their mandibles on the hollow
chambers and scare the animal off

eg. ants and Barteria tree

in West Africa

ants (Pachysima) defend the tree against herbivores by potent

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 Economic Impacts of Insects
in the US a 2006 study estimates that insects directly
or indirectly contribute more than $57 Billion to
our economy
1. as nutrition for wildlife, insects contribute ~$50
Billion/yr to US economy from fishing, hunting,
hiking, and enjoying wildlife
2. pest control
insects often prey on agricultural pests saving up
to $4.5 billion/yr in crops and pesticides
$50 Million worth of beneficial insects are sold worldwide
per yr; esp in greenhouses
eg. ladybugs
3. Pollination services
without pollinators many plants cannot be
fertilized to produce seeds
90% of worlds flowering plants are animal
pollinated
àincluding 80% of world’s 1330 cultivated
crop species
à 1/3rd of US agricultural crops are insect
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 57
1999 $126 Million worth of honey was produced in US
eg. in 2000, beekeepers worldwide harvested 125 million
pounds of beeswax (US goes for $1-2.10/lb)
used in foods, candles, sealing wax, polishes & other
industrial uses
in US ~1/2 of honeybee colonies have been
lost in last 50 years à 25% in last 5 yrs
alone
since 2007, thousands of bee hives have been
decimated
threats to pollenators:
habitat loss & fragmentation
loss of nesting and overwintering sites
intense exposure to pesticides and
herbicide
introduction of exotic species
newest threat is a parasitic fly that kills
entire hives
eg. silkworms (moth larvae spin coccoons of silk)
eg. silk
silk has been collected and spun for over 4000 years
primarily an oriental industry
in 2000 silk producers harvested 225 Million lbs of silk
from domesticated silkmoths
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 59
pollinated (~100 commercial crops)
in US à responsible for pollinating over $20 Billion
worth of agricultural products in 1998
economic value of insect pollinators worldwide is
$217 Billion/yr (08)
in US, excluding honey bees, other insect
pollinators contribute ~$3 Billion/yr to
agriculture
4. dung beetles burying animal dung help to recycle
nutrients, fertilize the soil, and reduce flies and
disease carrying insect on grazing land at a value
in US of $380 million/yr
5. A few kinds of insects have been semi-
domesticated:
eg. honeybees
3 trillion bees are kept and managed for pollenation and
honey production
honeybee pollination services are 60-100x’s more
valuable than the honey they produce
eg. Honey Production:
honey production dates back to ancient egypt
honey bees are not native to the US, introduced in
1600’s
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 58
today silk is being replaced by synthetic fibers but its
still a major industry in the southeast
eg.China, worlds largest silk producers (150
Million lbs/yr) had silk exports worth $2.8
Billion in 1999
before making a coccoon, silkworms are fattened up on a
diet of mashed mulberry leaves.
the worms then anchor a thread onto a branch and reel the
silk from a tiny hole in their mouth
each coccoon is made from over a half mile of silk
the coccoons are collected and plunged in boiling water to
kill the silkworm and remove the sticky glue that holds
the coccoon together
workers find the end of the filament and thread it onto a
bobbin and collected as long filaments
silk can also be reconstituted into its original jelly-like form
and respun using a variety of techniques to produce
the thype of thread required or poured into molds, cast
into transparent films or freeze-dried to make sponge
or foam
eg. mealworms & crickets
used as animal foods and fish bait
6. commercial products
eg. chitin
second most abundant organic polymer
can be used to coat fruits to slow ripening
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 60
 eg. shellac
made from lac à a resin produced by certain species of
scale insects in India and SE Asia
the “lac insect” form encrustations up to .5” thick on twigs
that are scraped off melted and dried into flakes
eg. dyes
cochinile bugs (scale insect) on prickley pear cactus à red
dyes; the color is from the dried bodies of the insects
dyes have also been made for other species of scale insects
and from the galls produced by gall-wasps
eg. tannic acid
some galls from gall wasps are used to extract tannic acid
used to make ink and other products
7. Venomous Insects
ants, bees, wasps, hornets, blister beetles, etc
some caterpillars have poisonous hairs or spines
that are attached to poison gland
with any contact spine penetrates skin and
injects poison
eg. asp or pus caterpillar
spines remain poisonous even after they are
shed
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
eg. Lice
>3000 species of lice; parasitize almost all species of birds
and mammals; most are scavengers, a few are
parasites
they spend their entire lives on the body of their hosts
many feed on keratin of skin, feathers, scales
others feed on secretion; scabs, mucus, blood, oils
some suck on blood intermittently for hours at a time
looked on today with disgust and loathing but:
high proportion of some populations (50%) esp
children have them
common in jails, camps, etc
in some countries lice are believed to be an indication
of robust health and fertility
human lice suck blood and usually feed at night
up to 1 mg of blood per meal
eggs = nits, deposited on hairs or clothing
eg. head lice
prefer fine hair of head
eg. body lice
generally live on clothing when not feeding
a female can lay 80-100 eggs at a time
head and body lice can spread:
typhus cholera
trench fever trench fever
relapsing fever impetigo
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
8. Parasites & Diseases & Vectors
eg. mosquitoes
besides being blood parasites are also vectors for
malaria, yellow fever, dengue fever, elephantiasis, etc
mosquitoes indirectly kill more people each year than any
other cause (infect up to 1 Billion each year and kill up
to 3 million)
eg. bed bug
associated with birds, bats, and other mammals including
us
worldwide distribution
after mating, the female must feed to be able to lay fertile
eggs
4-5mm nocturnal insects that feed on blood
in day, hide in bedding and cracks and crevices anywhere in
room
can live up to a year even without food
in large infestations produces distinctive “bedbuggy smell”
bed bug infestations spread so quickly in apt buildings
because there is extensive interbreeding
à the whole infestation is likely to be a “family affair”
bedbugs also quickly adapt; pesticide resistance and can
quickly spread it to the ‘family’
61 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 62
eg. crab louse
mainly in coarse hairs of body:
pubic area, armpits, beard, eyebrow, eyelashes
almost exclusively confined to caucasians
almost always venereally transmitted
each female can lay 25 eggs at a time
eg. Fleas
over 1000 species of fleas
ectoparasites of birds and mammals (warm blooded)
larvae live in nests
have compressed bodies and backward spines and
bristles to help them move through fur
piercing, sucking mouthparts
long powerful legs à enormous jumping power
eg. human flea (Pulex irritans; really a pig flea)
can jump 13” horizontally
7.75” vertically
equivalent jump for human:
450’ broad jump
275’ high jump
most breed and lay eggs in nests of hosts
cat and dog fleas lay eggs in fur of host
cat, dog, rat fleas readily attack humans
most fleas suck blood wherever they can find it
some species can survive up to 2 years without food
fleas are fairly indiscriminant in host choice
à since they change hosts easily they easily
63 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 64
 transmit diseases: typhus, plague, etc
David Harum: “A reasonable amount of fleas is good for a
dog, they keep him from broodin’ on bein’ a dog”
eg. Rabbit Flea (Spilopsyllus)
parasite’s ovaries develop in response to
corticoid hormones of pregnant host’s
reproductive cycle
àshortly after rabbits are born
levels of pituitary hormones in young stimulate
fleas to copulate and lay eggs
afterwords, most fleas return to mom and
comlete regression of their gonads occurs
eg. Flies
eg. blood sucking flies à blackflies, sandflies, no-see-ums,
horseflies, stable flies
eg. myiasis
some flies lay eggs in flesh and maggots feed on
tissue until they molt into adults
blow fly larvae are used as surgical aids
only eat dead tissues à clean wound
they destroy bacteria
wound heals much quicker
9. Medical applications
eg. maggots used to clean wounds by eating dead tissue from
wounds
secrete a fluid containing enzymes that speed healing
certain species of blowfly larvae; 5-10/cm2 for 2-3 days
will eat and remove ONLY the dead and damaged tissues
and leave the healthy tissues alone
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 65
eg. silk is already used to make underwear for British and
American troops in Afganistan
much more resistant to shrapnel injuries, doesn’t tear
– even if the shrapnel enters the body it is
enclosed in the silk and therefore much
easier to extracrt.
especially important for a part of the body that is often
of most concern to the soldiers when injured
eg. films of silk have been used to produce artificial corneas
– “its amazingly transparent”
eg. may one day be used as scaffolding to get stem cells to
mend bones and muscles
eg. controlled releases of antibiotics and other medicines
inside the body
2. Insects as food
in industrialized world we often react in disgust at
the thought yet we eat insects all the time
eg. FDA allows up to 60 insect fragments/100 grams of
chocolate; up to 30 pieces/100g of peanut butter; 1
insect fragment per gram of macaroni
in many parts of the world, insects are considered
delicacies
eg. central African children eat ants and grubs as they play
eg in SE Asia street vendors are thronged for their fried
crickets
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 67
heal some kinds of wounds much more effectively and
much more quickly than other treatments
have produced a prototype gel that healed wounds more quickly
hope to produce wound dressings impregnated with the enzymes
10. Chinese Herbal Medicine
weaver ants are pulverized into a powder used to treat asthma
powdered cockroach are used for stroke
silkworm feces is used to treat typhus
bee venom, honey and other bee products are used to treat
miscellaneous ailments
dried cicadas are boiled in a soup to improve eyesight
Future Applications
1. Insect and spider silk is being investigated for a
variety of purposes
20x’s stronger than steel
it does not trigger an immune response
can be easily produced at low temperatures and
pressures compared to other similar polymers
is biodegradable
organisms such as bacteria, potato and tobacco
plants and goats have been genetically
engineerred to produce silk
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 66
eg. in africa a certain species is sought after so much that
biologists are concerned about its extinction
eg Aborigines in Australia will travel several hours to find a
cache of honey ants to eat
113 nations eat bugs of over 1000 species
eg. in Mexico, >200 insect species are consumed. Demand
is so high that 40 species are now threatened with
extinction
we happily eat crabs, shrimp, and lobster; the
“insects of the ocean” but are repulsed by the
thought of grilled cicadas
what do they taste like?
ants have a lemon tang
giant water bugs tase of mint
fire ant pupae taste like watermelon
gram for gram many insects are more nutritious
than beef or pork
eg. 3 crickets could provide an individuals daily
iron requirement
eg. many insects have a fairly high concentration of
essential amino acids
eg. and the fats in bugs are considerably healthier than the
fats in meat
eating insects would be considerably “greener”
than our current meat diet
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 68
 and it’s a sustainable industry that causes much 5. insects as chemical detection devices
less destruction of the environment
insects have extremely sensitive olfactory senses

3. If eating insects directly is too distasteful, how military and security services need ultra sensitive,
about feeding them to livestock: flexible and portable odor detectors

manure à maggots à our meal sniffer dogs

cost ~$15,000 each,
protein content of maggot cakes equals that of soy & takes 6 months to train
fishmeal require dedicated handler

in addition fats and oils can be extracted for fuel “wasp hound” = a portable hand held odor detector

chitin used as binder and adhesive with a team of black wasps as its sensor

manure becomes a commodity: so far the wasps have learned to respond to

chickens $660 M/yr just about any odor tried
pigs $5 B/yr
wasps show coiling behavior when they detect
if it works with manure, how about sewage? food molecules

other insects (eg. bees) when they detect food

4. blood sucking insect, Dipetalogaster maximus, they stick out their tongues
is used as a high tech syringe
they can detect an odor at concentrations of a
can take up to 4 ml in one meal few ppt

donor doesn’t feel a thing à equivalent to finding a grain of salt

in a swimming pool
used as a way to get blood samples from wild animals that
are difficult to sample in other ways after a couple day stint as sniffers they are
returned to hive and a new batch is trained
eg. can measure stress hormone levels in nesting
terns without having to capture them can also be used to
assure food quality,
eg. used to survey rabies infections in bats check for contaminants
removes blood sample using a needle; they maybe even help diagnose diseases such as
recover quickly cancer or TB

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 69 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 70

6. Termites as geological prospectors

desert termites in Africa and Australia need damp soil for their
mounds. They can go down 100’ to find it.

rather than drilling wells to find mineral deposits deep under the
sand, geologists can sample a termite mound to find gold

7. Scientists hope to harness the activities of termite

bacteria to break down cellulose to produce
ethanol and biofuels

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 71

 Subphylum: Hexapoda
eg. roaches were the main insects of the time
(insects)
eg. some dragonflies had 2’ wingspans
~1.1 M species; probably millions more
today insects have spread into all major habitats
there are more species of insects than all other animal
species dominant fauna of all freshwater and soil habitats

à entire field of study = Entomology some live in deep underground caves

most successful & widespread group of all life on top of world’s highest mountains

adapted to land before most other terrestrial animals some insects live in unusual habitats:
except for a few Chelicerates
some flies occur by the millions in brine lakes eg. Great Salt
(Devonian 390 MY) Lake, where hardly any other life forms are able to
survive
adaptations to land (even deserts):
some insects live in hot springs up to 120º F (49º C)
waxy cuticle
varnish layer can close spiracles
many species are found inside ice in anarctica
extract & retain fluids from food and metabolism
(some don’t need any liquid water at all)
larvae of “petroleum flies” live in pools of petroleum around
diapause & resistant eggs
oil wells

had 40 MY to evolve and diversify before serious a few insect species have been found breeding in brine vats
competition for space and resources from other holding human cadavers at medical schools
animal phyla
the “short-circuit” beetle bores into lead cables

à still no birds around yet but only a very few are truly marine; (Why?)

à still not a lot of parasites of insects yet most are <2.5 cm (1/4”)
by carboniferous (~300MY ago ) there many different àsmall size helps them escape enemies
kinds of insects
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 1 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 2

àneed very little food to sustain themselves A few kinds of insects have been semi-domesticated:
range from < 1 mm to 25 cm honeybees

eg. Atlas moth of India has a wingspan of almost 1 foot
eg. Walkingstick of India is up to 15” long
insects invented agriculture & animal husbandry
àmany ants and termites cultivate fungi within
their burrows
à some ants guard aphids that let the ants “milk”
them for nectar-like secretion
3 trillion bees are kept and managed for pollenation and
honey production
silkworms (moth larvae spin coccoons of silk)
each coccoon is made from over a half mile of silk
mealworms & crickets are used to feed pets (or humans) or
for fishing bait
Body Form

may have been 1st to invent slavery body in three parts: head, thorax and abdomen
à there are ~35 species of “slave making ants” that regularly
raid the nests of other ant species and take young back to Head
their own colony
large compound eyes
there they work as they would in their home colony
searching for food, raising young, etc
several (usually 3) simple eyes (=ocelli)
always very closely related species

ants will always do the work of whatever colony they are in

1 pair of antennae
when they emerge as adults
many kinds of antennae
were the first animals to fly
eg. grasshoppers, crickets and cockroaches have long
antennae
à 130 MY before pterosaurs
eg. butterflies have knob on end
insects: 400 MY; devonian
reptiles: 200 MY pterosaurs; late jurassic , eg. moths antennae are featherlike
birds: 150 MY; coexisted with pterosaurs for~90MY
bats: 54 MY (Eocene)

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 3 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 4
 antennae are very complex sense organs in
which different segments control different wings are extensions of cuticle formed by
aspects of an insects life: epidermis
eg. in ants:
Abdomen
one segment detects nest odor and helps prevent
an ant from entering the wrong colony segmented (10-11 segments)
another segment identifies offspring of a specific
queen reproductive organs

another segment detects the ants own feeding females have pincher like or syringe like
trail
ovipositor to lay eggs
another segment helps detect what is needed by
the immature ants it is tending in some social insects it is modified into a
stinger
mandibles and other mouthparts for feeding
Insect Movement
Thorax

divided into three segments most kinds of movement are created by muscular
system (striated muscles like us)
(pro- meso- & metathorax)
insects have a more elaborate muscular system than
à each with sclerites: any other invertebrate group

tergum, pleura, sternum insects have more muscles than most animals
including us
each thoracic segment bears 1 pair of legs
eg. humans have ~700 individual muscles; some insects
have 900 or more muscle organs; some caterpillars
à total 6 legs; thus hexapods have 4,000

most insects also have 2 pairs of wings on insects are remarkably strong, given their small size
thorax
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 5 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 6

eg. a bee can pick up 50 times its weight swimming (diving beetles, many insect larvae)

eg. beetles are the strongest insects; up to 60 x’s their B. Wings

weight

[but their small size relative to weight makes them only appear strong most insects have two pairs of wings
à if insects were as large as humans they would be little if
any stronger than us]
some use both pairs to fly (eg. butterflies)
A. Legs
some the 1st pair cover and protect second pair
(eg. beetles)
great diversity of leg types:
a few have only 1 pair (flies, mosquitoes)
insect legs are adapted for the same kinds of
movements as vertebrates: a few are wingless (lice, fleas)

walking, running, jumping, swimming, Insect Flight

digging, climbing, grasping
insects were the first animals to fly
legs in 5 segments à 130 MY before any other animal

sometimes modified for unsure of evolutionary origin of wings:

jumping (grasshoppers, crickets, fleas) 1. originated as small flaps that first allowed gliding
= “flying squirrel theory”
eg. grasshoppers can jump 20 times length of
body 2. were 1st used as solar collectors to raise body
temperature
à equivalent jump for human would be
1/3rd length of football field later for gliding, then for flight

eg. fleas are probably the best jumpers 3. originated from gills of aquatic forms many have
can jump 8 “ high and 13 inches in length hinged gills on thorax

à equivalent feat in human flying in most insects is like swimming for us

à leap tall bldg in a single bound
storage of pollen (bees)
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
due to size and relative density of air
wings are separate from legs and other
7 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 8
 appendages eg. the common housefly is one of most talented
aerodynamicist on the planet

don’t just move up and down à more maneuverable in flight than birds, bats or bees

à common pattern resembles “butterfly eg. hoverflies are even better

stroke” of human swimmers
can make six turns a second
direct and indirect flight muscles
hover, fly strait up and down, backwards, do somersaults ,
land upsidedown etc
the rate of wing beating varies considerably:
yet: has brain smaller than a sesame seed
fastest
eg. midge >1000bps has only a dozen muscles for flying
eg. housefly beat ~200bps
eg. mosquito ~300bps but are loaded with sensors
eg. honey bee ~ 190bps compound eyes
wind sensitive hairs
more typical beat: antennae
eg. locust ~20bps three light sensors = ocelli
eg. dragonfly ~25bps

slowly beating:
flight greatly improved dispersal ability
eg. white butterfly ~12bps
eg. swallowtail ~5bps some insects are able to migrate 1000’s of
miles or fly at high altitudes:
speed of insect flight also varies greatly:
eg. monarch butterfly flies slowly (à 6 mph)
from slowest insect flight speeds: but can fly 100’s of miles at a time

eg. mosquitoes ~2mph eg. painted lady migrates 4000 miles

eg. houseflies ~5 mph
eg. butterflies ~6mph eg. some butterflies have been seen at 20,000’
eg. honey bee ~7 mph
some wings are only temporary structures
to quite fast
eg. males and queen ants use wings only for mating flight,
eg. hawkmoth ~35mph
then they drop off
eg. horse fly ~30mph
eg. dragonflies à 25 mph
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Feeding & Nutrition

d. some are predaceous
insects feed upon almost every kind of organic
substance e. some are parasitic

most feed on plant juices and tissues the same basic mouthparts are modified in various
= phytophagous ways to facilitate different methods of feeding:

many feed on dead or decaying organic matter a. chewing mouthparts

= saprophagous
in many insects the mandibles are hardened into
some are predaceous and attack other insects or “jaws” that are used to snip off pieces of plant or
animal matter
smaller invertebrates
eg. many caterpillars feed on specific plants
some are parasitic and feed on blood or living
tissues from temporary hosts eg. dragonflies, some beetles have large strong
mandibles
b. siphoning mouthparts
all insects share the same basic mouthparts:
eg. butterflies & moths: the maxillae forms a long
mandibles à jaws coiled food tube for sucking nectar
hypopharynx à tongue
labrum, labium à lips
maxillae à helps to manipulate food
c. piercing mouthparts
eg. in biting flies the mouthparts resemble a beak for
insects feed on a wide variety of foods: puncturing the skin

eg. in mosquitoes, most of the mouthparts together

a. most feed on plant juices and tissues form a long hollow tube, the labium directs this
tube through the skin like a syringe to draw up blood
b. others actually eat leaves, flowers and
whole plants or animals d. sponging mouthparts

c. many feed on dead or decaying organic eg. in houseflies the labium forms a sponge-like
feeding structure to mop up liquids
matter
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 larger insects use some kind of ventilation
the digestive tract is a tube divided into 3 major
regions: eg. pumping movement of abdomen

aquatic forms returned to water:

foregut à crop, salivary glands, stomach
initial food processing & temporary storage
à tracheal gills in cloaca eg dragonflies
midgut à usually has sac-like gastric caecae
enzyme secretion and absorption of nutrients à water bubbles
hindgut à intestine and rectum
rectum for water and nutrient absorption;
à return to surface to breath

Respiration Sense Organs, Communication & Behavior

insects are extremely active
an insects blood does not carry much oxygen
don’t have large gas filled sacs taking up a larger part of their they also have a rich supply of sense organs located
body for breathing all over the body
à oxygen is delivered directly to body cells
à these contribute to a rich diversity of insect
through system of tubes and passageways
behaviors
tracheal system with spiracles à adaptation to air most insect sense organs are microscopic in size and
are found on the body wall and various
many spiracles have valves that can open and close appendages:
these airways deliver oxygen to muscle cells 200,000 times small hairs, small domes
faster than our blood vessels (remove CO2 10,000 times as
fast)
keen senses but relatively simple circuits
tracheal system is lined with cuticle à must be shed eg. one signal usually results in one response for an insect

gas movement by diffusion in humans one signal can result in many different
responses

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1. Vision color vision

eg. plant eating insects are especially sensitive to
vision is the most important sense for most insects green
eg. ~2/3rd’s of a fly’s nervous system is devoted to vision
eg. butterflies that feed on red and yellow flowers are
more sensitive to those colors
insects have two kinds of eyes
(but most insects can’t see red)
simple eyes that can detect light & dark
eg. some insects can see UV used during courtship

and compound eyes that are especially some insects communicate using light
effective in detecting movement
à number and duration of flashes produces unique
simple eyes are called ocelli signature for each species

2. Touch
à usually 2 or 3 on head
most sense organs that respond to touch are
à can detect only light vs dark
small hairs on epidermal setae
compound eyes
most touching involves antennae and mouthparts
with many individual lenses = facets
a single antenna can have over 5,000 sensilla
some as few as 9 facets
these touch receptors can also pick up vibrations
houseflies have 4,000
in air to respond to wind or gentle breeze
dragonflies have 28,000 facets
eg. a fly avoids a swat by ‘feeling’ the air being pushed by
your hand as you try to swat it
provide a wide field of view and particularly
good at detecting movement can also detect temperature, humidity, gravity
eg. this is another reason why many insects are hard
to swat or capture touch receptors are used in a variety of ways:

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 15 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 16
 eg. mole cricket uses antennae to recognize nymphs in
dark burrow
eg. social insects can detect unwelcomed visitor to their
nest and attack them
eg. social insects often stroke and groom each
other with antennae and mouthparts
eg. waggle dance of honeybees involves touching to
communicate location of pollen
3. Hearing & Sound
insects have many different kinds of organs for
hearing
the insects with the best developed hearing are
those that communicate using sound
the simplest are hairs that respond to touch
many insects have hearing organs inside
their legs that respond to vibrations
passing through the ground or a plant
eg. ants come out of nest if you stomp on the
ground
cicadas and crickets detect sound with an
ear-like tympanum
moths, crickets, mantids etc can hear between
25,000 - 45,000 Hz
[humans hear sound waves from 20 - 20,000 Hz]
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
4. Taste, Smell & Chemical Communication
both taste and smell are types of chemoreceptors
they detect specific chemicals
smell à if the chemical is volatile and travels some
distance
taste à if the insect is in direct contact with the source
chemoreceptors are found on all parts of the
insects body
àon setae, antennae, mouthparts, legs
àbut are mainly on antennae and feet
à wasps and crickets know where to lay eggs
because they have chemoreceptors on their ovipositors
receptors for taste and smell have a pore that
receives the chemical to be detected
eg. a single sense organ on the antenna of a polyphemus
moth has 18,000 pores for chemicals to enter
most insects have 4 of the same taste
sensations as do humans: sweet, sour,
bitter and salty
organs of smell do not detect as many different
odors as does a human nose
but it is tuned more finely
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
àcan detect ultrasonic signals of bats to avoid
being eaten
sound may be used to individuals of the same
species
eg. males and female mosquitoes are attracted to
each other by sound of their buzzing wings
eg. tapping sound of death watch beetles (eat wood
on old house -heard by those keeping watch over
dead person before burial) used to attract a mate
many insects that can hear have structures for
making various sounds to communicate with
each other or to other species
some rub body parts together to make sounds
= stridulation
eg. chirping sound of crickets, katydids, beetles and
ants can be used to communicate with group or
seek a mate
some have drum like membrane that can
vibrate to make a sound
eg. cicadas are loudest of all insects
à can be heard half mile away
à songs differ for each species
hissing cockroach blows air out its spiracles to
make sound
many insects can make supersonic sounds
17 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 18
à can detect differences between very
similar chemicals and in much lesser
amounts
some can detect airborne chemicals 5 miles
away
eg. male of lesser emperor moth can smell the
pheromone of the female up to 6 miles away
smell is used to find suitable plant for laying eggs
used by flies to find dead animals
some parasitic wasps can track host species by
smell
smell is also used to communicate within a species
eg. sex pheromones attract mates
eg pheromones that cause grouping behavior
eg. ants release alarm pheromones if disturbed or
threatened
eg. ants, caterpillars and other insects produce trail
pheromones to map route to food
5. in addition to external senses insects also have a
variety of proprioceptors
can monitor body positions, eg legs and wings,
internal pressures
19 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 20
 could not walk or fly without this information
eg. ants, bees, wasps, termites, some beetles

Behavior & Communication they can work much more efficiently as a group than
as solitary insects
insects have keenly developed senses
they have a very structured division of labor
à can be used for communication
and behavioral responses highly dependent on each other

insects communicate in a number of ways using their generally unable to survive long on their own
external senses
à they work almost as a single individual
insect behavior is mostly instinctive (innate)
in social insects workers change tasks depending on
eg. caterpillar is programmed to eat a certain plant
their age
eg. courtship, egg laying, migrations are preprogrammed
younger workers engage in safer chores like
but they do have the ability to learn “home maintenance”;
eg. bee dance
the older workers do the riskier foraging
eg. notice landmarks near nest or food outside the nest
eg. when Colorado potato beetles first attempt to mate,
they are not very good at identifying their own species
they change jobs based on their own
or even distinguishing head from tail; with repeated assessment of their remaining longevity
attempts they get better at it
eg. when harmed in some way they switch to the more
research has recently shown that insects have a similar “fight or risky jobs
flight” response to stress as vertebrates have (same genes)
eg. honey bees
Colonial Insects
Each colony consists of a single queen, about 100 male
drones, and 60,000 workers
only about 10 of the 20 or so orders of insects have
developed some type of social organization
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 21 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 22

Organization of the colony is maintained by pheromones &

secretions produced by the queen
Workers live 4-5 weeks. Young workers feed brood, drones and
queen. Older workers construuct comb using wax from
abdominal wax glands. They also clean the hive and guard
the entrance. Older workers forage.
Bees feed on pollen and nectar; pollen for protein, nectar to
make honey
Foragers that return with nectar perform dances that provide
information on the distance, direction and quality of the
food source; eg. round dances indicate food within about
300 ft of the hive, waggle dances indicate the food is
further away. Nectar is shared with other workers to verify
the quality of the food source.
Honey is made by evaporating water from nectar and adding
glandular secretions. Each colony can produce up to 2 lbs of
honey per day.
The queen deposits an egg in each brood cell. unfertilized eggs
produce drones, fertilized eggs produce workers, fertilized
eggs fed royal jelly produce queens
eg. ants
similar division of labor with one or more queens, drones,
workers, soldiers, etc
in relative size an ant has one of the largest brains among
insects
à each ants brain consists of ~ 1 M cells with the
processing power equivalent to a Mac II computer
(1987-1 meg)
à as a colony of 40,000 individuals, the colony has a
processing power equivalent to the brain of a
mammal

a large bee colony can store up to 100 lbs of honey and 5 lbs of
Insect Defenses & Weaponry
pollen per colony for the winter months
1. hard exoskeleton
Large colonies in late spring produce new queens. Queens are
selectively bred in a special queen cell. They are fed “royal the hard exoskeleton of insects serves as an
jelly” by workers. Only one queen will ultimately survive.
effective defense against many dangers
The ‘old queen’ with about half of the workers will swarm to
eg. large beetles, including weevils, have a particularly thick
establish a new colony
exoskeleton
The new queen makes a single mating flight in the spring with
about a dozen drones. The drone that succeeds in some insects have a “fracture line” in each
impregnating her plugs her genital opening with his broken appendage that allows the leg to break off
off penis after insemination to insure that no other males easily if caught by a predator
can copulate with her. Most drones die shortly after the
mating flight; surviving drones are kicked out of the hive eg. many crane flies, walkingsticks, grasshoppers
The queen acquires enough sperm to last her lifetime of 4-5
years; she can deposit up to 3000 eggs/day.
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 immature insects that sacrifice a leg can often
regrow it after several molts
2. quick reflexes
as the most active invertebrates, most insects can
easily avoid threats by quick escape; flying or
running
most insets have sensitive hairs that can detect
the change in air pressure of a fast moving
hand
eg. cockroaches can respond & escape from an approaching
shoe in less than 50 milliseconds
eg. resting houseflies can move to avoid a swat in 30-50
milliseconds
3. “hold your ground” or “play dead”
some insects, when threatened, remain in place
and adopt a defensive posture
eg. tortoise beetles have strong adhesive pads on their feet;
they lie flat against a leaf and hold on when threatened
eg. cuckoo wasps curl up into a hard rigid ball like pill bugs
for protection
some insects simply “play dead”
if on trees and shrubs; they release their grip
and disappear into the undergrowth
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
mimicry eg. viceroy butterfly
one group of beetles can change color bypumping
fluids into grooves and channels in its clear
exoskeleton
7. chemical defenses
many insects wage chemical warefare using
noxious chemicals to ward off predators
in some cases they manufacture their own toxins
others use chemicals extracted from host
plants
eg. stink bugs secrete a foul smelling chemical from glands
in their exoskeleton when threatened
eg. some species of cockroaches and termites secrete a
sticky substance from anal glands to immobilize ants,
spiders, centipedes or other predators
some of these chemicals can be painful
or cause temporary blindness
eg. Bombadier beetle à shoots hot explosive
secretions toward predator from glands in their
abdomen
each gland has two chambers, one contains
hydroquinones and hydrogen peroxide;
the other contains various enzymes;
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
4. spines, bristles and hairs
eg. many predators will avoid a mouthful of spines and bristles
found on many caterpillars
eg. parasitic flies or wasps have more difficulty getting close
enough to lay eggs in or on the potential host bearing
spines and bristles
5. sound
eg. hissing cockroaches emit a threatening hiss
eg. some moths can mimic the sound of “bad tasting” moths to
avoid being attacked by bats
eg. tiger moths can detect the ultrasonic sounds made by bats
if at low intensities (bat is relatively far away), they just
change course and fly away
at higher intensities (bat is much closer), they quickly drop
from the air in an evasive looping dive
6. Color as warning or camoflage
some insects have bright red or orange colors
exposed only when they are threatened by
predator that may scare predator away
others with large eyespots that may
intimidate
in other color and shape are used as camoflage
camoflage eg. walking stick
25 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 26
when the two are mixed a series of explosive
reactions occur
peroxide breaks down to oxygen and water
and the concotion heats to boiling and blasts
out as a toxic cloud
some caterpillars click their jaws loudly when
predators approach as a warning they will
vomit a distasteful fluid if they don’t back off
in some species of termites, when “soldiers”
become too old to defend the colony they blow
themselves up spraying a toxic brew which
kills them and their attackers
stinging insects use pain as an effective defense
or a way to subdue prey
some insect toxins that cause pain are
expelled by hollow body hairs
simply brushing against them causes them
to break and release their painful
chemicals
eg. saddleback caterpillars
eg. pus caterpillars
some social insects, usually infertile workers,
have their ovipositors modified into
stingers attached to poison glands
eg. bees, wasps, hornets, ants, etc
27 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 28
 sting-pain index based on human perception the goal of this initiation rite is to keep the sleeve
on for a full ten minutes without showing any
0. sting doesn’t penetrate skin signs of pain

1. sharp, sudden, effect similar to a stray spark when finished, the boys’ (now men) arms are
temporarily paralyzed because of the venom, and
eg. fire ant sting they may shake uncontrollably for days

2. a piercing pain Excretion

eg. honeybees, common wasps & hornets, acacia ant
malpighian tubules (2-100’s) absorb metabolic
3. bold, caustic, burning and unrelenting wastes from blood and drain into intestine
eg. paper wasps, harvester ants
Reproduction & Life Cycles
4.blinding, fierce, excruciating and debilitating
wide range in life spans for adult insects:
eg. tarantual hawk, warrior wasps
hours to years
4+. worst sting: pure intense, brillian pain
eg. adult mayflies live for less than a day; do not eat
eg. bullet ant (Paraponera clavata) sting produces
sole purpose is to reproduce
wave after wave of throbbing, all-consuming pain
that continues unabated for up to 24 hrs
eg. species with longest lifespan may be a tropical
termite whose queen may live for 50 years
Fortunately, bullet ants are not aggressive except
when defending themselves or their colony
Mating
Bullet ants are used by some indigenous people in
their initiation rites to manhood: insects are dioecious
the ants are first knocked out by drowning them
in a natural chloroform most show sexual dimorphism with male trying to
attract the females:
then hundreds of them are woven into sleeves
made out of leaves, with stingers facing inward. eg. male rhinoceras beetles with long “horns”

when the ants awaken, boys slip the sleeve onto eg. male stag beetles with large mandibles
their arm.

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 29 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 30

eg. male butterflies often more colorful eg. praying mantis courtship

most have internal fertilization females have predisposition to eat their mate before, during
or after mating

mating is an important part of an insects behavior set eat head first à remove “off switch” of male copulating

to find mates (during mating, the subesophageal ganglia (head)

inhibit ventral ganglia from triggering intensive
copulatory activity)
most use pheromones
eg. Aphids
some use light; eg. lightning bugs
female aphids can reproduce without sex they produce
others use sound or color
clones of unfertilized eggs that develop into females

some mark and protect territories from other males yet they do have sex annually; but not for the males sperm.
They are after “sexually transmitted diseases”
eg. some dragonfly species
beneficial bacteria from the male that can help them
often have courtship rituals using pheromones &/or digest plant food, resist extreme temperatures, kill
sound to attract mate internal parasites

eg. cicadas, crickets, grasshoppers the female can also pass these infections on to her future
clones
eg. deathwatch beetles bang their heads against the sides of
their tunnels to signal a mate many female insects mate only once/lifetime and store
the sperm
eg. mosquitoes and midges use high frequency sound of their
wingbeats to attract a mate
insects usually lay many eggs
eg. “lightning bugs” use distinctive patterns of light to attract
mates eg. queen honeybee >1 M/lifetime

internal fertilization eg. termites lay 10,000 or more/day

male must insert sperm into females seminal receptacles eg. cockroaches can lay >1 million eggs in a year

there is a great diversity in insect reproduction: but some only 1 egg per mating

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 31 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 32
 eggs are sometimes enclosed in protective case eg. butterflies: larva=caterpillar
eg. flies: larva=maggot
eg. roaches; praying mantis eg. beetles: larva=grub

some lay eggs on specific plant or animal once adult emerges it no longer molts

many insects “mark” plant with chemical to b. most of the rest have incomplete
dissuade other females from depositing eggs metamorphosis (hemimetabolous):
there eggànymphà adult
à ID of the chemical they use may be
useful as a natural insect deterrent nymph resemble adult without wings

some may care for young after hatching wings develop externally as budlike
growths on thorax
Development
grow by successive molts
most insects also go through several distinct
eg. grasshoppers, cicadas, mayflies,
developmental stages as they grow from egg to stoneflies, dragonflies
adult
c. a few have direct development
insects often have complex development including
metamorphosis egg à juvenile à adult

eg. silverfish, springtails

a. 88% have complete metamorphosis
(holometabolous): metamorphosis is regulated by hormones
eggàlarvaàpupaà adult
3 major endocrine glands in insects:
larva usually has chewing mouthparts brain produce
ecdysial gland juvenile hormone
pupa is a nonfeeding stage before corpora allata & molting hormone
metamorphosis; often in a cocoon
Some insects overwinter as immature stage
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 33 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 34

à don’t complete development until they are Monarch butterflies:

literally frozen in winter months
millions migrate over 2500 miles in autumn from
Diapause breeding grounds in the eastern US and
Canada to a winter haven in Mexico
some insects are able to enter a state of dormancy to
survive adverse conditions; eg temp, humidity the fall migrants are great great grandchildren
of the monarchs that left Mexico the
if this dormancy is for an extended period and previous spring
triggered by daylength = diapause
yet they return to the same dozen or so
Migration roosting areas

many insects migrate they can orient by the sun and can detect
earth’s magnetic field
eg. locusts, butterflies, moths, dragonflies, grasshoppers, some
beetles 2013 saw a 59% drop in migrating monarchs since 2012;
partly due to use of a herbicide that kills monarch’s
generally if an insect is seen: food plant

more than 8 or 10 ft above the ground

flying in a relatively straight direction
ignoring prominent food sources on its journey

it is probably migrating

most fly at high altitudes and catch seasonal winds to

migrate

a few, like monarch butterflies fly at much lower levels

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 35 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 36
 So, Why are insects the most
abundant and most diverse group of
living organisms on Earth?
insects have been the dominant life form for over
400 MY.
-they have witnessed the rise and fall of the
dinosaurs
-they have survived at least 4 of the earth’s mass
extinctions
-they continue to thrive as humans have spread
over the planet and are in the process of
creating our current mass extinction of life
Their success is largely due to a unique combination
of just a few major features:
1. Exoskeleton
their exoskeleton is made of chitin bound with
several proteins making hard and rigid for
protection, yet with flexible and elastic
“hinges” for freedom of motion. This “shell” is
coated with a waterproof waxy layer
their exoskeleton gives insects support needed
for any kind of habitat; protection from
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
eg. A leaf miner spends its entire life tunneling
in a single leaf
eg. in some ant species, an entire colony may
live inside a single acorn or plant gall
eg. parasitic wasps in at least 7 families
complete their entire development within
the egg of other insects.
small size is also an advantage in avoiding
predation
they can easily hide in a crack in a rock,
beneath the bark of a tree or the petal of a
flower
their exoskeleton is hard enough for them to
burrow between individual grains of sand
and to squeeze through the tiniest of
cracks
3. Flight
insects are the only invertebrates that can fly
they flew 100MY before the first flying vertebrate
and they still have the most effective flight
mechanics:
-an insects flight muscles generate twice the
power per muscle mass than vertebrate
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
predators and environmental hazards and
prevents desiccation in dry habitats
muscles are attached directly to the body wall to
combine maximum strength with optimal
leverage and strength
many insects can lift many times their own
weight
2. Small Size
most species are relatively small (<1”)
for an animal with an exoskeleton, small size is a
distinct advantage. The larger the insect the
thicker and more cumbersome it would have
to be
in the real world, “Mothra” and “the Fly” could
never walk much less get off the ground to
fly
another advantage of small size is there is only a
need for minimal resources to survive and
thrive
food requirements are so modest that an
insect may spend its entire life on a single
plant or animal and never exhaust its food
supply
37 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 38
flying muscles, mainly due to
characteristics of their exoskeleton
flight provided a new escape from predators and
an extremely effective means of dispersal
efficient energy use allows some insects to travel
great distances or remain airborne for
extended periods
eg. more than 200 species of moths, dragonflies, locusts,
flies and beetles migrate over long distances
4. Reproductive Potential
insects produce large numbers of viable eggs in
very short time spans
à they produce a large #’s successful
offspring
5. Metamorphosis
in more advanced insects the freeliving larval
stage that lives in microhabitats different from
the adult of the same species
only 9 of the 28 different kinds of insects have
“complete metamorphosis” yet they represent
about 86% of all living insect species
39 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 40
 Ecological Impacts of Insects
eg. 90% of all dead animals (mainly insects) end up as food
1. insects are the most important organisms in most in ant nests
terrestrial ecosystems eg. 99% of human and animal wastes is decomposed at
least partly by insects & other invertebrates
insects are at the heart of healthy ecosystems:
eg. herbivores
insects are an integral part of every landbased
a large number of insects feed on plants but only a small
foodchain percentage of them are considered pests

because of their number, variety and influence many have proven to be effective controls against
bioinvasive plants
on larger organisms and whole ecosystems
eg. a moth that eats prickly pear cactus in Australia
eg. soil insects
eg. a European beetle was introduced to California to
many types of insects spend part or all of their lives control an invasive “goatweed”
underground
eg. pollinators are keystone species in some
many feed on humus or decaying plant matter terrestrial ecosystems (more under
they aerate the soil, recycle organic wastes, fertilize the soil symbioses)

eg. springtails – many millions/acre sexual reproduction in most plants is made possible by the
process of pollination
eg. ants – nest in soil, feed above ground
the majority of pollinators are insects
eg. termites, burrowing bees, wasps, beetles and flies
without a certain insect species to pollinate the dependent
plant species will decline or go extinct
eg. scavengers
eg. carnivores
eg. termites, ants, wood boring beetles, dung beetles, flies

feed on decomposing plants and animals and recycle their keep populations of their prey in check.
nutrients
eg. parasitoids
remove rotting materials
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 41 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 42

insects with parasitic larval stages Insect Symbioses

eggs are laid inside body of host and larval stage hatches insects have formed a wide variety of symbioses with
and grows feeding on host eventually killing it
virtually every major kind of living organisms
at metamorphosis becomes freeliving adult
Bacteria
work with carnivores to control population sizes

eg. many fly and wasp species; tachinids, almost all animals, including insects have bacterial
ichneumonids symbionts

without insects, most of the terrstrial life on earth some that live in digestive system are mutualistic
would disappear
àhelp break down hard to digest fibers and
starches
most of the rest of the world is literally helpless
without them àmake essential vitamins

vertebrates wouldn’t last more than a few months àprotect from pathogens

if all insects suddenly disappeared àprovide additional nutrients from food eaten

invertebrates, especially insects, are “the little

eg. cellulose digesting bacteria
things that run the world”
-EO Wilson
some species of bacteria, some protists, and
most fungi are able to produce the proper
2. Insect Symbioses
enzymes to digest cellulose in plants
insects have formed a wide variety of symbioses
virtually no animal species can produce
with virtually every major kind of living
cellulose digesting enzymes
organisms
animals that are strict herbivores rely on
cellulose digesting microorganisms to
extract nutrients from plant materials
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 43 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 44

eg. termites
many species of cellulose digesting bacteria and
protist that digest the bits of wood eaten by
the termites
a diet restricted to wood only (not softer green
parts of plants) is very low in nitrogen and
termites also have nitrogen fixing bacteria
that converte N2 into useable nutrients for
the termites
Protozoa
eg. cellulose digesting flagellates in the gut of termites
rd
1/3 to 1/2 of a termites weight are these symbiotic
protozoa
work with bacteria to break down cellulose fibers
Fungi
Animal “Fungus Farms”
eg. Atta ants (leafcutter ants)
mounds up to 18 ft underground
the ants maintain active fungal gardens to feed
larvae and adult in the colony
àthey cultivate fungi within their mounds
àthey feed the fungi bits of leaves, bark and other
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
fungus never forms mushrooms in mound
but when rain is imminent
termites carry bits of mycelia out of mound
spread them around the ground
several days after rain mushrooms appear and
produce spores
the termites reappear and collect the genetically mixed
hyphae and take them back into the nest
abandoned mounds produce crops of mushrooms
for several years after termites leave
à African people consider them a rare
delicacy
Plants
eg. Pollination
a large part of the success of flowering plants is due to
the variety of pollenation
à provides much better mixing of genes
flowering plants have coevolved with many kinds of
animals through most of their history
this close relationship between plants and
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
plant material
àthey weed out debris that might spoil their garden
àhave wheelbarrow-like depression on their backs
for carrying spores
fungus is transferred from colony to colony by ants
freeliving cultures of fungus have never been
found
fungus needs ants to provide them with food
in Central and South America leaf cutter ants are
the dominant herbivores in food chains
à consume 17% of total leaf production
eg. African termites (Microtermitinae)
build chimneys up to 30 ft tall of feces and mycelia
eat mostly plant material but don’t have cellulose
digesting microorganisms living in their gut
bring it home and eat it before entering
mound
then go in and defecate
use their feces to grow fungi (Termitomyces)
all stages of termite readily eat the fungus
45 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 46
pollinators has resulted in coevolution
à mutual adaptations for mutual benefits
plant: petals, scent, nectaries
animal: special body parts and behaviors
We like to think of animals as the most ‘evolved’ lifeforms
à but plants lack brains, muscles, speech yet they don’t need them
- they borrow animals to do their bidding
In some instances the relationships have become very
specialized such that only a single species of
animal can pollinate a particular species of plant.
insect pollinators
petals colorful and large
often with nectaries
much less pollen produced
insects can be attracted by showy flowers, smell &/or nectar
eg. in irises the insect cannot get to the nectar without collecting
pollen
eg. milkweed produces packest of pollen that entangle the legs
of any insect that lands on its flower
eg. some plants depend on a single species of insect for
pollenation:
eg. some orchids
eg. long tongued hawk moth
eg. fig wasp and fits
eg. yucca plants are pollenated solely by the yucca
47 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 48
 moth
usually white or dull in color
eg. orchard fruits are almost entirely insect pollenated (beetles cant see as well as bees)
mainly by bees
--------------------- flowers tend to have a strong yeasty, spicy or
fruity odor, sometimes unpleasant to us
a. bee & wasp pollinated
secrete no nectar but may supply food as
pollenate more flowers than any other group pollen or in special storage cells in petals

20,000 different species of bees are important pollinators most eat parts of the flower
for many plants
eg. magnolias, lotus
bees pollenate mainly spring flowers
c. carrion flies
bees have very good vision
tend to be dull red or brown or white
à flowers are generally brightly colored
often have foul odors resembling rotting meat
predominately blue or violet markings
eg. skunk cabbage
rarely pure red (pure red appears black to them)
eg. carrion flower = Rafflesia
flowers generally shallow (short mouthparts)
a parasitic flowering plant of SE Asia
honeybees are attracted to nectar
they also gather pollen early botanists thought it might be a fungus

flowers often delicately sweet and fragrant parasitic plant with no true roots or leaves
parasitic on vines in genus Tetrastigma
flowers often have lines or distinctive (Vitaceae)
markings that function as “honey guides”
à lead bees to nectar produces a huge speckled flower on little stem
with no leaves
some of these markings are only seen in
UV light largest known flower:
à invisible to us, not bees flower bud size of basketball
flower up to 1 m across; weighs up to 10 kg
b. beetle pollinated
smells like rotting meat
flowers generally large and open
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 49 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 50

fruit is eaten by three shrews and other mammals wasps have very good vision

d. butterflies flower preferences similar to bees

but wasps pollenate mainly in summer
prefer sweet fragrance and provide nectar
wasps are much less “hairy” so not very good pollenators
fused petals force insect to crawl into flower for nectar
flowers are generally brightly colored
nectaries are usually at bases of deep spur that
only butterflies and moths can reach with their predominately blue or violet markings
mouthparts
flowers generally shallow (short mouthparts)
some butterflies can detect red flowers
flowers often delicately sweet and fragrant with nectar
eg. daisy family: butterfly bush= Buddelias, or fruit juices
goldenrods, blazing star
eg. fig wasps exclusively pollenate almost 1000
eg. monarchs à pollenate milkweeds species of tropical fig trees

e. moth pollinated eg. some species of orchids resemble certain wasp

females that attract only one species of wasps
white or yellow large tubular flowers
heavy fragrance
eg. Plant Galls
open at night, closed in daylight
galls are any deviation from the normal pattern of
eg. yuccas, night blooming jasmine, night blooming plant growth on a particular organ or part due to
cereus
the presence of another organism
f. ants
all plant organs are attacked: roots, stems, leaves,
dense clusters of small flowers near ground
flowers and fruits
sticky pollen
à leaves are most common organ attacked
with nectar
many galls superficially resemble plant parts such as berries,
eg. some composites, poppies, nuts and other fruit types

h. wasps some galls look like fungi and for years some were misidentified
as a fungal species
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 much more nutritious than eating typical leaves,
some galls resemble strange sea urchins and coral polyps stems or roots.

galls on trees and shrubs often appear in the spring the gall insect typically becomes trapped inside the
gall and becomes a victim for a diversity of
when the leaves first appear parasites itself

à when they are highest in food quality à creates a complex insect community
within the gall.
plant galls have been known since antiquity (350 BC)
eg. some oak galls support as many as
eg. plant galls were known and used extensively in medicine, 22 species
industry and as human food for over 1000 years in China,
India and parts of Europe eg. another kind of gall had 75 different
species present
plant galls can be produced by a variety of organisms
but mainly various species of insects a gall appears to be a parasitic relationship

à~13,000 gall forming insects are known àthe gall is formed as the insect attacks and
(~2% of all known insects)
wounds the plant
à 70% of galls are produced by flies & wasps
eg. Ant/Plant Symbioses
most insects are highly host and organ specific
eg. ants and acacia trees
insects especially form leaf galls one species of acacia tree is always found with a
colony of a particular ant species associated with it
gall forming insects are highly specialized
plant feeders: the tree is small (5’-6’) and its branches are covered
with pairs of sharp thorns up to 3” long.
tend to be sedentary and feed only on certain cells
No acacia of species Acacia drepanolobium has been
found that did not contain colonies of the ant species,
they induce the plant tissue to grow rapidly inside the
Crematogaster mimosae and C. nigriceps.
gall to provide adequate food
these ants live only on these trees; up to 20,000/tree
the stimulated growth created by the insect
produces a high protein, nutritious meal
ants get room and board:

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 53 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 54

at every 6th pair of thorns is a swelling (=pseudogall) stings

which is a hollow space inhabited by the ants
sting can penetrate elephant hide and can numb small
at base of each leaf are nectaries that exude a sugar mammals (and humans) for several days
solution similar to flower nectar

acacia trees get protection from herbivores

àgrab intruder with strong mandibles and sting it

eg. antelope gets a face full of stings and leaves

eg. ants and tropical epiphytes

some epiphytes have swollen knobby tuber with with hollow

chambers inside inhabited by ants

ants accumulate humus and their excretions nourish the plants

in some species the ants collect seeds of various epiphytes and

plant them in their nests to create “hanging gardens”

they collect animal feces to nourish seeds

some epiphytes reward the ants by producing starch grains or

sugary secretions

eg. ants and rattan vines

certain rattan (Korthalsia) vines have swollen areas that house

ants

if a mammal accidentally brushes against plant the ants make a

rattling noise by beating their mandibles on the hollow
chambers and scare the animal off

eg. ants and Barteria tree

in West Africa

ants (Pachysima) defend the tree against herbivores by potent

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 55 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 56
 Economic Impacts of Insects
in the US a 2006 study estimates that insects directly
or indirectly contribute more than $57 Billion to
our economy
1. as nutrition for wildlife, insects contribute ~$50
Billion/yr to US economy from fishing, hunting,
hiking, and enjoying wildlife
2. pest control
insects often prey on agricultural pests saving up
to $4.5 billion/yr in crops and pesticides
$50 Million worth of beneficial insects are sold worldwide
per yr; esp in greenhouses
eg. ladybugs
3. Pollination services
without pollinators many plants cannot be
fertilized to produce seeds
90% of worlds flowering plants are animal
pollinated
àincluding 80% of world’s 1330 cultivated
crop species
à 1/3rd of US agricultural crops are insect
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 57
1999 $126 Million worth of honey was produced in US
eg. in 2000, beekeepers worldwide harvested 125 million
pounds of beeswax (US goes for $1-2.10/lb)
used in foods, candles, sealing wax, polishes & other
industrial uses
in US ~1/2 of honeybee colonies have been
lost in last 50 years à 25% in last 5 yrs
alone
since 2007, thousands of bee hives have been
decimated
threats to pollenators:
habitat loss & fragmentation
loss of nesting and overwintering sites
intense exposure to pesticides and
herbicide
introduction of exotic species
newest threat is a parasitic fly that kills
entire hives
eg. silkworms (moth larvae spin coccoons of silk)
eg. silk
silk has been collected and spun for over 4000 years
primarily an oriental industry
in 2000 silk producers harvested 225 Million lbs of silk
from domesticated silkmoths
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 59
pollinated (~100 commercial crops)
in US à responsible for pollinating over $20 Billion
worth of agricultural products in 1998
economic value of insect pollinators worldwide is
$217 Billion/yr (08)
in US, excluding honey bees, other insect
pollinators contribute ~$3 Billion/yr to
agriculture
4. dung beetles burying animal dung help to recycle
nutrients, fertilize the soil, and reduce flies and
disease carrying insect on grazing land at a value
in US of $380 million/yr
5. A few kinds of insects have been semi-
domesticated:
eg. honeybees
3 trillion bees are kept and managed for pollenation and
honey production
honeybee pollination services are 60-100x’s more
valuable than the honey they produce
eg. Honey Production:
honey production dates back to ancient egypt
honey bees are not native to the US, introduced in
1600’s
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 58
today silk is being replaced by synthetic fibers but its
still a major industry in the southeast
eg.China, worlds largest silk producers (150
Million lbs/yr) had silk exports worth $2.8
Billion in 1999
before making a coccoon, silkworms are fattened up on a
diet of mashed mulberry leaves.
the worms then anchor a thread onto a branch and reel the
silk from a tiny hole in their mouth
each coccoon is made from over a half mile of silk
the coccoons are collected and plunged in boiling water to
kill the silkworm and remove the sticky glue that holds
the coccoon together
workers find the end of the filament and thread it onto a
bobbin and collected as long filaments
silk can also be reconstituted into its original jelly-like form
and respun using a variety of techniques to produce
the thype of thread required or poured into molds, cast
into transparent films or freeze-dried to make sponge
or foam
eg. mealworms & crickets
used as animal foods and fish bait
6. commercial products
eg. chitin
second most abundant organic polymer
can be used to coat fruits to slow ripening
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 60
 eg. shellac
made from lac à a resin produced by certain species of
scale insects in India and SE Asia
the “lac insect” form encrustations up to .5” thick on twigs
that are scraped off melted and dried into flakes
eg. dyes
cochinile bugs (scale insect) on prickley pear cactus à red
dyes; the color is from the dried bodies of the insects
dyes have also been made for other species of scale insects
and from the galls produced by gall-wasps
eg. tannic acid
some galls from gall wasps are used to extract tannic acid
used to make ink and other products
7. Venomous Insects
ants, bees, wasps, hornets, blister beetles, etc
some caterpillars have poisonous hairs or spines
that are attached to poison gland
with any contact spine penetrates skin and
injects poison
eg. asp or pus caterpillar
spines remain poisonous even after they are
shed
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
eg. Lice
>3000 species of lice; parasitize almost all species of birds
and mammals; most are scavengers, a few are
parasites
they spend their entire lives on the body of their hosts
many feed on keratin of skin, feathers, scales
others feed on secretion; scabs, mucus, blood, oils
some suck on blood intermittently for hours at a time
looked on today with disgust and loathing but:
high proportion of some populations (50%) esp
children have them
common in jails, camps, etc
in some countries lice are believed to be an indication
of robust health and fertility
human lice suck blood and usually feed at night
up to 1 mg of blood per meal
eggs = nits, deposited on hairs or clothing
eg. head lice
prefer fine hair of head
eg. body lice
generally live on clothing when not feeding
a female can lay 80-100 eggs at a time
head and body lice can spread:
typhus cholera
trench fever trench fever
relapsing fever impetigo
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12
8. Parasites & Diseases & Vectors
eg. mosquitoes
besides being blood parasites are also vectors for
malaria, yellow fever, dengue fever, elephantiasis, etc
mosquitoes indirectly kill more people each year than any
other cause (infect up to 1 Billion each year and kill up
to 3 million)
eg. bed bug
associated with birds, bats, and other mammals including
us
worldwide distribution
after mating, the female must feed to be able to lay fertile
eggs
4-5mm nocturnal insects that feed on blood
in day, hide in bedding and cracks and crevices anywhere in
room
can live up to a year even without food
in large infestations produces distinctive “bedbuggy smell”
bed bug infestations spread so quickly in apt buildings
because there is extensive interbreeding
à the whole infestation is likely to be a “family affair”
bedbugs also quickly adapt; pesticide resistance and can
quickly spread it to the ‘family’
61 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 62
eg. crab louse
mainly in coarse hairs of body:
pubic area, armpits, beard, eyebrow, eyelashes
almost exclusively confined to caucasians
almost always venereally transmitted
each female can lay 25 eggs at a time
eg. Fleas
over 1000 species of fleas
ectoparasites of birds and mammals (warm blooded)
larvae live in nests
have compressed bodies and backward spines and
bristles to help them move through fur
piercing, sucking mouthparts
long powerful legs à enormous jumping power
eg. human flea (Pulex irritans; really a pig flea)
can jump 13” horizontally
7.75” vertically
equivalent jump for human:
450’ broad jump
275’ high jump
most breed and lay eggs in nests of hosts
cat and dog fleas lay eggs in fur of host
cat, dog, rat fleas readily attack humans
most fleas suck blood wherever they can find it
some species can survive up to 2 years without food
fleas are fairly indiscriminant in host choice
à since they change hosts easily they easily
63 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 64
 transmit diseases: typhus, plague, etc
David Harum: “A reasonable amount of fleas is good for a
dog, they keep him from broodin’ on bein’ a dog”
eg. Rabbit Flea (Spilopsyllus)
parasite’s ovaries develop in response to
corticoid hormones of pregnant host’s
reproductive cycle
àshortly after rabbits are born
levels of pituitary hormones in young stimulate
fleas to copulate and lay eggs
afterwords, most fleas return to mom and
comlete regression of their gonads occurs
eg. Flies
eg. blood sucking flies à blackflies, sandflies, no-see-ums,
horseflies, stable flies
eg. myiasis
some flies lay eggs in flesh and maggots feed on
tissue until they molt into adults
blow fly larvae are used as surgical aids
only eat dead tissues à clean wound
they destroy bacteria
wound heals much quicker
9. Medical applications
eg. maggots used to clean wounds by eating dead tissue from
wounds
secrete a fluid containing enzymes that speed healing
certain species of blowfly larvae; 5-10/cm2 for 2-3 days
will eat and remove ONLY the dead and damaged tissues
and leave the healthy tissues alone
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 65
eg. silk is already used to make underwear for British and
American troops in Afganistan
much more resistant to shrapnel injuries, doesn’t tear
– even if the shrapnel enters the body it is
enclosed in the silk and therefore much
easier to extracrt.
especially important for a part of the body that is often
of most concern to the soldiers when injured
eg. films of silk have been used to produce artificial corneas
– “its amazingly transparent”
eg. may one day be used as scaffolding to get stem cells to
mend bones and muscles
eg. controlled releases of antibiotics and other medicines
inside the body
2. Insects as food
in industrialized world we often react in disgust at
the thought yet we eat insects all the time
eg. FDA allows up to 60 insect fragments/100 grams of
chocolate; up to 30 pieces/100g of peanut butter; 1
insect fragment per gram of macaroni
in many parts of the world, insects are considered
delicacies
eg. central African children eat ants and grubs as they play
eg in SE Asia street vendors are thronged for their fried
crickets
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 67
heal some kinds of wounds much more effectively and
much more quickly than other treatments
have produced a prototype gel that healed wounds more quickly
hope to produce wound dressings impregnated with the enzymes
10. Chinese Herbal Medicine
weaver ants are pulverized into a powder used to treat asthma
powdered cockroach are used for stroke
silkworm feces is used to treat typhus
bee venom, honey and other bee products are used to treat
miscellaneous ailments
dried cicadas are boiled in a soup to improve eyesight
Future Applications
1. Insect and spider silk is being investigated for a
variety of purposes
20x’s stronger than steel
it does not trigger an immune response
can be easily produced at low temperatures and
pressures compared to other similar polymers
is biodegradable
organisms such as bacteria, potato and tobacco
plants and goats have been genetically
engineerred to produce silk
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 66
eg. in africa a certain species is sought after so much that
biologists are concerned about its extinction
eg Aborigines in Australia will travel several hours to find a
cache of honey ants to eat
113 nations eat bugs of over 1000 species
eg. in Mexico, >200 insect species are consumed. Demand
is so high that 40 species are now threatened with
extinction
we happily eat crabs, shrimp, and lobster; the
“insects of the ocean” but are repulsed by the
thought of grilled cicadas
what do they taste like?
ants have a lemon tang
giant water bugs tase of mint
fire ant pupae taste like watermelon
gram for gram many insects are more nutritious
than beef or pork
eg. 3 crickets could provide an individuals daily
iron requirement
eg. many insects have a fairly high concentration of
essential amino acids
eg. and the fats in bugs are considerably healthier than the
fats in meat
eating insects would be considerably “greener”
than our current meat diet
Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 68
 and it’s a sustainable industry that causes much 5. insects as chemical detection devices
less destruction of the environment
insects have extremely sensitive olfactory senses

3. If eating insects directly is too distasteful, how military and security services need ultra sensitive,
about feeding them to livestock: flexible and portable odor detectors

manure à maggots à our meal sniffer dogs

cost ~$15,000 each,
protein content of maggot cakes equals that of soy & takes 6 months to train
fishmeal require dedicated handler

in addition fats and oils can be extracted for fuel “wasp hound” = a portable hand held odor detector

chitin used as binder and adhesive with a team of black wasps as its sensor

manure becomes a commodity: so far the wasps have learned to respond to

chickens $660 M/yr just about any odor tried
pigs $5 B/yr
wasps show coiling behavior when they detect
if it works with manure, how about sewage? food molecules

other insects (eg. bees) when they detect food

4. blood sucking insect, Dipetalogaster maximus, they stick out their tongues
is used as a high tech syringe
they can detect an odor at concentrations of a
can take up to 4 ml in one meal few ppt

donor doesn’t feel a thing à equivalent to finding a grain of salt

in a swimming pool
used as a way to get blood samples from wild animals that
are difficult to sample in other ways after a couple day stint as sniffers they are
returned to hive and a new batch is trained
eg. can measure stress hormone levels in nesting
terns without having to capture them can also be used to
assure food quality,
eg. used to survey rabies infections in bats check for contaminants
removes blood sample using a needle; they maybe even help diagnose diseases such as
recover quickly cancer or TB

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 69 Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 70

6. Termites as geological prospectors

desert termites in Africa and Australia need damp soil for their
mounds. They can go down 100’ to find it.

rather than drilling wells to find mineral deposits deep under the
sand, geologists can sample a termite mound to find gold

7. Scientists hope to harness the activities of termite

bacteria to break down cellulose to produce
ethanol and biofuels

Animals: Arthropods-Hexapoda Ziser Lecture Notes, 2015.12 71

 Crustaceans a river shrimp, Macrobrachium jamaicense, was
collected from Devils River, Tx: 3’ long including
antennae, body was 10.5” long, 3 lbs
=shelled creatures; “the insects of the sea”
some crustaceans live for several decades; some molt
~67,000 species throughout life

eg: lobsters, crayfish, shrimp, crabs, water fleas, so continuous increase in size
copepods, barnacles, pill bugs, etc
eg. Lobsters may be the longest lived
crustaceans are mostly aquatic, mostly marine Crustaceans
one was collected that weighed 35 lbs
inhabit most waters of the earth: ocean, arctic,
freshwaters, high mountain creeks and lakes was estimated to be 50 yrs old;
thermal springs, brine waters
à ~ 4’ long total

many are at the base of aquatic food chains body only was ~2’ long,
part of zooplankton
claw was an additional 20”

only a few are terrestrial; eg. sow bugs or pill bugs
crustaceans vary in size from microscopic (<0.1 mm)
to >12’
eg largest (longest) is giant Japanese crab
à up to 12’ from end of claws to tail and a weight of
40 lbs (20 kg)
eg. crayfish & freshwater shrimp
some crustaceans are quite colorful; blue, red,
orange, yellow
many are bioluminescent
Distinguishing Characteristics of Crustacea:
1. the crustacean body is divided into a
cephalothorax, abdomen and tail (telson)

largest crustaceans in freshwaters often have carapace extending over abdomen

some up to 2’ and weigh 9 lbs and gills

Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 1 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 2

in some groups carapace forms clamshell like valves that lots of variation in appendages between groups
encloses body
eg in decapods (crayfish, crabs, lobsters, etc):
2. they are the only arthropods with 2 pairs of
antennae 1st 2 pair à antennae with chemoreceptors

next 5 pr à feeding appendages; including

3. most with compound eyes, smaller ones only with cephalothorax mandible, maxilla and maxillipeds
simple eyes
next 5 à walking legs including cheliped and
4. crustaceans use jaw-like mandibles as main gills

feeding structures; abdomen next 5 à called swimmerets; used to carry

eggs and as copulatory organ
also maxillae and maxillipeds
telson last à uropod = swim fin

5. abdomen usually with pairs of jointed appendages

on most segments
most crustaceans can cast off legs or pinchers to
escape predators and later regrow them
they generally have many pairs of appendages
range from 3 to 50 pairs of limbs 6. unique larval stage in marine forms:

many are biramous appendages nauplius larva

à 2 branches from base segment most have single larval eye throughout several larval
stages

appendages are modified for a variety of uses some retain the larval eye into adulthood and never
sensory develop compount eyes
feeding
defense Body Form
walking
swimming
reproduction the most ancient crustaceans resembled some kind of
respiration aquatic centipede; long segmented body
with similar segments and similar appendages
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 3 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 4
 over time, great variety of body types arose abdomen usually with pairs of jointed
appendages on most segments
only one small group of these primitive
crustaceans remain today telson (tail)
only 10 species in their own class ‘Remipedia’
has its own paired appendages
in most crustaceans today, the body is usually divided
into a cephalothorax, abdomen and tail = uropods that with the telson form a finlike
(telson) tail

some have distinct head and thorax Movement

often have carapace extending over the sides of Crustaceans generally have many pairs of appendages
the animal larger species 5 or more pairs of walking legs

in some groups carapace forms clamshell like ‘valves’ that

encloses the whole body
most appendages are biramous

in others the carapace covers cephalothorax but not they branch like a “wishbone”; one of the
abdomen branches usually has a gill attached at its
base
cephalothorax
most crustaceans can cast off legs or pinchers and
feeding and sensory appendages
regrow them
5 or more pairs of walking legs
voluntary (striated) muscle tissue arranged in
st antagonistic groups
including chelipeds (pinchers) on 1 pair
eg. flexors & extensors
abdomen
similar to vertebrates

segmentation is most apparent in the abdomen

Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 5 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 6

Feeding & Digestion legs to strain water for food

4. scavengers
use jaw-like mandibles as main feeding
structures eg. isopods, amphipods

5. parasites
also maxillae and maxillipeds
eg. fish lice, tongue worms
chelipeds (1st walking legs) are also used to catch,
tear off and put pieces of food to the mouth crustaceans have a well developed digestive system:

great variation in feeding types: cardiac stomach with gastric mill for grinding

1. predators gastric mill has hardened “teeth”

eg. crabs use large claws used to break open shells to feed some have gastroliths in stomach to help
eg. mantis shrimp grinde food

is an ambush predator, extremely carnivorous and pyloric stomach for sorting

aggressive

called “split thumb” in Bermuda and West Indies digestive gland secretes digestive enzymes

front end looks like praying mantis Respiration

has “jackknife claws”
in small crustacea: no special organs
live in solitary burrows

eyes are stalked and constantly watch for prey

à exchange across body surface

2. herbivores in larger crustacea: respiration usually by feathery

gills
3. suspension feeders
on bases of walking legs
eg. barnacles sit upside down in shell and use
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 7 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 8
 crustacean blood has the ability to clot
in some sides of carapace form gill chambers
that enclose gills Nervous System:

have an appendage called a “bailer” that as in other Arthropods, similar to Annelids

creates a water current across gills
2 pairs of ganglia around the esophagus that supply
Circulation sense organs of the cephalothorax

open circulatory system double ventral nerve cord with a pair of ganglia in
each segment along the rest of the body
no veins or capillaries

fluid filled coelom = hemocoel Sense Organs

dorsal heart in hemocoel sense organs are well developed in crustaceans

heart has ostia (holes) to draw in hemolymph 1. most have compound eyes and simple eyes
(blood) (ocelli)

hemolymph is pumped through 2 or more arteries for compound eye consists of many separate units called
ommatidia
distribution
each with cornea divided into many small squares or hexagons
most crustaceans have some kind of blood pigment to called facets
better distribute oxygen to tissues each ommatidium behaves like a separate eye

hemolymph may be bluish, reddish or colorless: each ommatidium has a moveable pigment component that
allows it to adjust to differing light intensities
most: hemocyanin à bluish pigment with Copper
in daylight only light rays that strike the cornea directly
others: hemoglobin à red pigment with Iron reach the photoreceptor cells

some: no pigments in dim light rays can activate the photoreceptors in more
than one ommatidium

Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 9 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 10

zooplanktonic crustaceans show particular

sensitivity to light as they undergo daily some crabs have striae or ridges on inner side of
migrations up and down the water column to chelae that they rub against tubercles on
stay in the best light regime for feeding yet carapace
being able to avoid visually hunting predators
some crabs have tympanic membranes on their
2. chemoreceptors (taste) on mouthparts, 1st leg segments to pick up these sound
vibrations
3. crustaceans uniquely have 2 pairs of antennae
sound is used for warning, to frighten enemies,
4. tactile hairs and spines spread over body mating rituals, etc

5. statocysts for orientation 7. Light emitting organs & communication by

light
at base of antennae
many crustaceans have light emitting organs
saclike; opens to surface by pore (=photophores) that use luciferase to
produce light
take in sand grains which trigger hair
cells to provide info on orientation
many crustaceans give off rich blue sparks of light
6. hearing: communication by sound when disturbed
eg. some ostracods and a few copepods, even some
many crustaceans make underwater noises to freshwater decapods
communicate
krill have light organs with lens and reflector to
eg. pistol crabs snap claws together producing sound focus and intensify the light beam
like pulling a cork from a bottle

eg. one species of mantis shrimp makes a vigorous many pelagic crustacea flash brilliantly during
rasping noise by rubbing uropods against mating swarms
underside of telson

eg. Florida spiny crab produces sound like moist

Endocrine System
fingers rubbing against a window pane
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 11 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 12
hormones help to control:
located in ventral part of the head anterior to
molting the esophagus
although molting (ecdysis) is hormonally controlled the
cycle is triggered by environmental stimuli to nervous generally a coiled tube leading to a sac
system which opens at the base of the
antennae or maxillae
eg. temperature, day length, humidity (land crustacea)
or some combination
extracts N-wastes from hemolymph
molt-inhibiting hormone levels decrease surrounding it and secretes it to the
molting hormone increases
outside

once molting is initiated it proceeds automatically also used to regulate water & salts

body coloration (K & Ca+ conserved; SO4 & Mg excreted)

chromatophore (pigment cells) behavior is controlled by [no malpighian tubules]

hormones from glandcells in the base of the eyestalks

Reproduction
heart rate
most are separate sexed (dioecious)
blood sugar levels
but a few are hermaphrodites including barnacles
sexual development
some have courtship rituals involving chemical, visual
Excretion
and auditory cues:
nitrogen wastes are excreted through skin (if no gills) eg. fiddler & ghost crabs

or through gills & antennal glands male attract females during the day by waving its claws and
both day & night by producing distinctive drumming
sounds

antennal glands (or maxillary/green gland)

Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 13 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 14

when a female touches a male the frequency increases 2 pr antennae

dramatically 3 prs appendages
1 pr mandibles
female can only mate after final molt
all function as swimming appendages at this stage

some develop large “apron” for carrying eggs from the nauplius stage often develop other distinctive
larval forms for different groups of crustaceans
copulation: male delivers sperm packet to
eg. zoea larva – some crabs
receptacle using modified swimmerets
eg. protozoea larva – shrimp
a few groups, mainly those living in temporary waters
eg. copepodid larva - copepods
reproduce by parthenogenesis
later there is a more dramatic change =metamorphosis
eg. brachiopods, ostracods, isopods and a few crayfish
into the adult
males are rare or unknown in some species

eggs are generally released into the water

some retain their eggs until they hatch in brood

pouches
eg. Most crabs and shrimp

in some crustacea such as crayfish, development is

direct with no larval stage

but most crustaceans produce a variety of distinctive

larval forms as the animal develops

many marine crustaeans begin with a characteristic

larval form

= nauplius larva
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 15 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 16
 Classification of Crustacea eg. water fleas, fairy shrimp, tadpole shrimp, brine shrimp)

Class Ostracoda (13,000 sp. (70,000 fossil sp.))

The subphylum Crustacea is divided into 4 major
small, body flattened side to side and protected by bivalve-
classes (with 2 additional classes for a few rare
like calcareous shell
species)
eg. seed shrimp
Class Malacostraca (40,000 sp.)
also,
largest crustacean class, extremely diverse – possibly the
most diverse class in the entire animal kingdom
Class Remipedia (17 sp.)
head-thorax-abdomen, sometimes head and thorax fused
very primitive characteristics
into a cephalothorax
à resembles a centipede in general body form but with
appendages on all major tagmata
biramous legs
with or without carapace
very poorly known à all known species are from
underwater caves
eyes on stalks

eg. isopods, pill bugs, amphipods, scuds, shrimp, crabs, Class Cephalocarida (12 sp.)
lobster, crayfish, krill, mantis shrimp, etc
occur along the coasts of the United States, in the West
Indies and Japan
Class Maxillipoda (10,000 sp.)
2-3 mm long
mostly small, reduced abdomen which generally lacks
appendages live in bottom sediment from intertidal zone to 300 m

eg. copepods, barnacles, fish lice and tongue worms also thought to be very primitive

Class Branchipoda (1000 sp.)

gills on many appendages, including some mouthparts

most have compound eyes and carapace; in some the

carapace is made of 2 valves that completely enclose
the animal

Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 17 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 18

Crustacea occupy every possible kind of habitat in the without crustaceans, animal populations in aquatic
ocean and freshwaters. ecosystems would collapse

most crustaceans live in the ocean and are one of the eg. especially the larger crustaceans; shrimp, lobsters
and crabs
“pillars of the global marine ecosystems”
eg. crabs use large claws used to break open shells to
of the 40,000 or so species of crustaceans, about feed
4000 of these are found in freshwaters.
eg. fiddler crab uses largest claw for social interactions

Most Crustacean, both marine and freshwater inhabit only uses small claw for scavenging food from
benthic and planktonic habitats sand

the females brood eggs or brood young in a ventral

Benthic Crustaceans brood chamber (=marsupium)

most crustaceans are benthic organisms; they live on, on freshwater sediments the larger
in, or near the bottom sediment of both marine crustaceans are mainly crayfish and
and freshwater habitats shrimp
there are even a few freshwater crabs
benthic marine crustaceans include mainly the
larger species many eat plant life on the floors of lakes, ponds and
streams
benthic crustaceans play an important role in eg. isopods generally much smaller, <.5”
decomposing dead or decaying animals, algae and
plants (except for one group of deep water isopods that grow more
than a foot long)

crustaceans feed a vast number of other animals isopods are mainly scavengers
in oceans and in freshwaters
mainly found crawling on the substrate or under rocks
and submerged plants
their numerous symbioses help to control populations
of other animals seldom found in open water

Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 19 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 20
 common in both marine and small freshwater lakes and
streams
some crustaceans construct tubes in sediment, wood and
rock
eg. boring isopods can destroy wooden pilings in less
than 2 yrs
some isopods can tunnel through limestone rock
some have been known to burrow through the
insulation of undersea cables shorting them out
a few cave adapted forms occur in subterranean
waters
eg. amphipods (side-swimmers or scuds)
much more common and diverse in marine benthos than in
freshwaters
shrimp-like appearance: strongly compressed laterally
but no carapace
generally much more active at night than during
daytime
amphipods are voracious feeders
omnivorous scavengers; feed on all kinds of plant and
animal matter
a few are parasites
like decapods, the females brood eggs and young in a
ventral brood chamber (=marsupium)
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11
viable eggs have been collected from dried ponds and
revived after 20 years
some marine crustaceans are sessile (=attached)
eg. Barnacles (=Cirripedia)
sessile: secrete shell of several calcium plates in
which they live
common in intertidal areas
when tide goes out they can close their shells
every time a wave comes in they extend their legs to filter
the water
considered a kind of mollusk until 1830
but once they were discovered to produce a nauplius
larva it was clear they were a kind of crustacean
there are 2 main kinds of barnacles:
some with stalk = goose barnacles
some without = acorn barnacles
eggs hatch into motile, nauplius larvae then a cypris larva
after swimming a short time the larva secretes a
strong polysaccharide cement from its antennae
and attaches to the substrate
à the strongest adhesive known
adults secrete chemicals that attract the larvae to settle
near them to facilitate reproduction
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11
eg. Ostracoda (=seed shrimp)
much more common in freshwater habitats
much smaller
mainly benthic animals that inhabit all types of
substrates in standing and running water
a few actively swim just above the substrate
generally use their antennae to move
enclosed in bivalve carapace that completely covers
the entire animal
their shells are so strong that they fossilize well
ostracods are the oldest known crustacean group in
the fossil record
marine species are especially important to
paleontolotists in dating sediment
(65,000 fossil species vs 13,000 living species)
generally feed on bacteria, fungi, algae and detritus
ostracods have the largest sperm in the animal kingdom in
both relative and absolute terms.
Ostracod sperm can be up to ten times the length of
the male's body!
Some male ostracodes need a special organ (Zenker's
organ) to aid in sperm transport.
ostracods lay their eggs on sediment or aquatic vegetation
the eggs can hatch immediately or years later
21 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 22
the carapace develops into a mantle that secretes
calcareous plates
legs develop in feathery cirri for filtering water
animal sits up-side-down in shell and extends legs to filter
feed
almost all are hermaphrodites yet they cross fertilize with
internal fertilization
a few species are dioecious with the dwarf males
attached to the female
they don’t feed and die after inseminating the
female
barnacles are preyed on especially by starfish and snails
Planktonic Crustaceans
many smaller crustacea are part of the plankton in
both marine and freshwaters = zooplankton
larger planktonic crustaceans are the primary
herbivores feeding on the phytoplankton (algae
and bacteria)
small planktonic crustaceans such as copepods,
ostracods and krill are essential links between
producers and larger consumers in aquatic food
webs
krill and copepods are extremely abundant in the
worlds oceans
23 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 24
 these planktonic crustacea may be the animals with the
greatest biomass on the planet
depending on these zooplankton are a diverse
community of predators
including other crustaceans, squid, fish, and
whales
eg. Copepods
copepods are one of the most abundant and diverse
groups of crustacean
over 12,000 species (some believe this to be about
15% of total species out there)
may be the most abundant aquatic animals on the
planet
copepods are abundant in both marine and freshwater
environments
but much more abundant and diverse in the ocean
most common as plankton but many species also occur on
or in the sediment as benthic organisms
small, slender, clearly segmented body
large pair of antennae used for movement
feathery legs to filter food
feed in a variety of ways: scraping food from hard
surfaces, filtering articles from the water, seizing
and biting prey
extremely important food source for marine fish
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11
many cladocerans show a dramatic change in body shape
over the year = cyclomorphosis
large eyes – looks like a single eye but is actually 2
compound eyes that are fused together
very large antennae that are used for locomotion
inside the carapace are 5 or 6 pairs of feet used to
filter the water for food
most cladocera are filter feeders and eat algae, protozoa
and bacteria
they also eat aquatic plants
a few are predators on microscopic animals
most cladocera reproduce sexually by fertilization of egg
with sperm, and parthenogenetically
use parthenogenesis for most of the year
switch to ’regular’ sexual reproduction under adverse
conditions
female carries her eggs around in a brood pouch
enclosed in carapace
eggs hatch and young swim free – direct development
eggs play a key role in dispersal to new habitats
resistant to freezing and drying
the eggs of some have remained viable for up to 300
years
Temporary or Highly Saline Waters
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11
majority of the diet of commercial fish is
copepods
some freshwater forms are important vectors for diseases
such as guinea worm
eg. Krill (Euphasids)
small shrimp-like animals extremely abundant in
marine plankton
often occur in swarms up to 30,000 individuals/m3
a major part of the diet of whales, seals, penguins and
cephalopods among others
eg. whales eat 2-3 tons of krill per meal
large numbers of crustacean larvae along with
fish larvae making up a significant portion of
the biomass of the open ocean
(=microzooplankton)
are a significant source of fish food in marine ecosystems
eg. Water Fleas (=Cladocerans)
cladocera are especially abundant in permanent
freshwater ponds & lakes, among marginal
vegetation
à important part of freshwater zooplankton
much less common in the ocean
body is enclosed within a bivalve shell called a
carapace that covers the thorax and the
abdomen but not the head
25 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 26
certain specialized crustaceans are the dominant
animals in highly saline or alkaline environments
or in temporary waters such as playas
most crustaceans inhabiting these temporary habitats are from
the Class Branchiopoda
“breath through their feet”
à feathery gills at base of walking legs
the branchipoda generally inhabit temporary pools, ponds and
playas
and are generally completely absent from permanent
bodies of water
feed mainly on algae, bacteria, protists and microscopic animals
they typically appear in the spring and disappear in late summer
or autumn as habitat dries
to survive most produce very drought resistant eggs that can
survive dried or frozen for years in lake beds
the eggs of most hatch into nauplius larvae
eg. fairy shrimp, tadpole shrimp, clam shrimp
eg Fairy Shrimp
live in vernal pools and hypersaline lakes worldwide
including pools in deserts, ice covered mountain lakes
and Antarctica
common, especially during the cool months of year, but
seldom seen unless pursued
27 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 28
 fairy shrimp are most easily distinguished from all other
similar crustaceans in that they swim upside down.
not usually found in ponds that contain fish
one of few crustacean groups with distinct head
stalked compound eyes
no carapace
some grow up to an inch long
11 pairs of swimming legs
use legs to swim upside down
graceful movements, often transparent
feed by filtering algae, bacteria, protozoans etc from the
water or by scraping algae from surfaces
Important food source for many birds and fish
Many species of fairy shrimp compete intensely for mates.
Since mating usually occurs just after the female
moults, males often grasp her and are towed
around while waiting for her to moult.
Some females have a chain of these attached males,
including some which have died while waiting
includes: brine shrimp (Artemia)
only animals that flourish in the Great Salt Lake of
Utah and other hypersaline environments
today they are cultured extensively as fish food
eggs sold as novelties; “sea monkeys”
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11
enclosed within a carapace of 2 valves to resemble a small
clam
they extend their second antennae out of the carapace to
swim
they feed on detritus or plankton by drawing water into the
carapace
most reproduce sexually
some reproduce by parthenogenesis
Terrestrial Crustaceans
while the vast majority of crustaceans are aquatic,
some groups are semiaquatic or terrestrial
eg. land crabs burrow above tide line into the water table
can survive days out of water
eg. pill bugs & sow bugs (isopods)
isopods are the only group of crustaceans with truly
terrestrial representatives
have very delicate gill-like respiratory organs that
must be kept moist
found in damp places under stones and logs
able to roll up for protection (=rolly pollys)
young develop in brood pouch
some salt water relatives are found along coasts and
live in seaweed, along rocks and algae
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11
eg. Tadpole Shrimp (Triops sp.)
resembles a baby horseshoe crab
considered a “living fossil” this genus has remained
essentially unchanged for the last 70 M years (time of
the dinosaurs)
large shield-like carapace covering most of the body
look somewhat like tiny horseshoe crabs
the name Triops refers to it’s 3 eyes
at end of abdomen are two long filamentous extensions
omnivorous: they eat anything organic includng their
siblings (they are cannibalistic)
usually found in vernal pools, worldwide, scratching at the
sediment or straining the water for food
several of their activities have human benefits:
they eat culex mosquito larvae which transmits west
Nile virus
used in Japan to eat weeds in rice paddies
their presence in the Western US usually indicates the
presence of endangered spadefoot toads
the eggs are also sold to raise as “pets” called
‘aquasaurs’ or ‘trigons’ or just ‘triops’
eg. Clam Shrimp (Conchostraca)
laterally compressed
29 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 30
some bore into wood causing destruction of pilings
and warves
eg. beach fleas or sand hoppers (amphipods)
some are almost terrestrial; found crawling around on
piers and jetties
Symbioses
many examples have already been cited
a. numerous commensal relationships with other
invertebrates
eg. many bivalves harbor commensal crabs within their
shells
eg. crabs and shrimp also live inside sponges, worm
tubes, etc
eg. some barnacles in symbiosis with humpbacks and other
whales
stick on skin; esp head, flippers and flukes
appear to cause little damage except for some species
that seem to burrow into the skin but don’t seem
to cause serious inflammation
feed on scraps produced by whale feeding
b. mutualistic interactions
eg. decorator crabs attach sponges, anemones, etc to their
shells for camoflage
eg. cleaner shrimp remove skin parasites from fish
31 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 32
 then they develop into a nonparasitic stage until they
set up “cleaning stations” along the reefs that fish know to are ready to feed again
visit
they repeat this cycle several times during their lives
remove parasites, dead skin and other debris
eg. tongue biter: enter host as males through gills and
some cleaner shrimp even enter the mouths of fish to clean change gender as they mature
gills
she attaches herself to fish’s tongue and sucks blood
c. a very diverse variety of crustaceans have become
fishes tongue disappears and the parasite becomes the
parasitic functioning ‘tongue’ of the fish

eg. Fish Lice (Branchiura, eg. Argulus) eg. copepods

one of the simplest and most common forms of crustacean

almost half of all copepod species are either parasitic or
parasites are the fish lice
commensal with other organisms
parasites on marine and freshwater fish
there is a hugh diversity of shapes and lifestyles
cling tightly to fish with a set of antennae modified into larg
Parasitic copepods often cling to the outside of their hosts
barbed suckers
others invade the gills, nostrils or mouth of their host
have flattened bodies, compound eyes and maxillae
modified into suckers to attach to the sides of fish
eg. Dryodora: attaches to the rectum of fish and
mouth borne on a long proboscis used to suck blood and develop into pustule looking sacs
mucus from their host
eg. Anchorworms (eg. Lernaeocera): some species
after feeding on host the parasite detaches and drifts embed themselves on the eyes of sharks and feed
downstream on the vitreuous jelly inside.

many species can tolerate both fresh and salt waters eg. Trebius: ectoparasite on the embryo of
angelsharks (inside mom)
eg. some isopod parasites
eg. Acrothoracica
mainly ectoparasites
related to copepods and barnacles
eg. Gnathiidae: are similar to ticks in that they attach and
gorge on blood parasitize corals and echinoderms

Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 33 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 34

female cypris larva attaches to a crab and injects

eg. Tongue Worms (Pentastomida) a mass of eggs

parasites that live in the respiratory tract of vertebrates, these cells migrate to intestine of host and
usually reptiles in the tropics develop rootlike growths that permeate the
hosts body
also some fish, bird and mammals (including humans)
develops an extensive system of branches extending
so unlike other crustaceans that until recently they were into every appendage
classified in their own phylum, pentastomida
a saclike growth appears under the crabs abdomen
wormlike; 2-13 cm long; >70 sp, 4 fossil genera where eggs and sperm form (Sacculina is a
hermaphrodite)
4 clawlike appendages at anterior end
the crabs metabolism is completely altered:
mouth with protuberance
the cells of the parasite multiply and differentiate
no resp, circ or excretory organs into a reproductive form which produce an
egg mass in the female hosts apron
their life cycle involves 2 hosts
the host protects, ventilates and grooms the egg
the intermediate host is vertebrate prey of final host mass as if it were her own

the larvae live in it’s blood if crab is a male:

eg. Tantulocarida body assumes shape of a female

reduced length of some segments
crustaceans that parasitize other crustaceans broadening of abdomen
testes reduced or converted to ovaries
among the smallest arthropods
à both male and female resemble mature female
can be less than 0.1 mm long bearing eggs: physically and behaviorally

eg. Sacculina

probably the most bizzare of all parasitic animals

Sacculina is a highly modified barnacle that has

become a parasite of crabs

Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 35 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 36
 Economic Importance of
Crustaceans
many are at the base of aquatic food chains
part of zooplankton
1. as food
eg. crab, lobster, crayfish, shrimp
more than 10 million tons of crustaceans are
harvested for food each year (2007)
à mostly shrimp, crab, lobsters and
prawns
the heyday of lobster fishing was in the
1890’s:
1892 yield was 24 M lbs of lobster;
25 pounders were common
80% of all crustaceans are harvested in Asia,
mainly China
some crab are harvested by breaking off claws
and throwing rest back
blue crab are held captive until they molt, the
viscera are removed and they are sold as
“soft-shelled” crab
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11
Aquaculture of freshwater crustaceans now
accounts for 2.4 million tons of seafood eaten/
year (2011)
mainly shrimp and crabs
a similar amount was cultured in brackish
water aquaculture
some are allergic to seafood
mainly an allergy to tropomyosin – a muscle
protein
but a few other proteins may also cause
allergies
2. bait
crayfish are commonly sold and used as bait either
live or only the tail meat
sometimes causes problems with bioinvasions
3. pets
crayfish are kept as pets in freshwater aquaria
land crabs are often sold in pet stores
also brine shrimp and tadpole shrimp
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11
crayfish (crawfish) are commonly eaten in the
southern US and in other countries
Louisiana produces 70-90% of all commercial
crayfish, most of it from aquaculture
recent (2007) annual harvest of ~55,000 tons
Krill are now being harvested for human
consumption around the Antarctic
can harvest 12 tons/hour
but they are difficult to process
goose barnacles are popular food on the coasts of
Spain, Portugal and parts of N. Africa
France is developing a goose barnacle fishery
to export them
292 tons/yr worldwide (2011)
In Libya are many brine lakes from which large
quantities of brine shrimp (Artemia) are
harvested
collected in nets and sun dried until they form
a “foul-smelling black paste” then eaten
native Americans used to harvest them from
the Great Salt Lake
37 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 38
4. many crustaceans are serious pests
a. cause crop destruction
eg. rice crabs in China and India eat rice;
burrows may drain rice fields destroying crops
eg. crayfish destroy young cotton plants
b. boring & fouling organisms
borers destroy warves & docks and wooden
hulled boats
undermine sea walls and bore into stone
destroy underwater cables
adhere to ships reduce efficiency and increase
hull decay
eg. barnacles
5. some act as intermediate hosts for human
parasites
eg. Guinea worm
larva is in copepods; swallowed in contaminated water
grow in lymphatic system
up to 3’ long
female produces blister like lesions on lower extremeties to
lay eggs in water
eg. fish tapeworm
39 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 40
 larva in Cyclops and Diaptomus
eaten by fish eg. Cladocera are easily cultured and used to study behavior,
evolution, speciation and animal ecology
humans eat uncooked fish

6. Some Crustaceans are used for dissections in

biology classes
eg. crayfish

7. Real and imagined medicinal uses:

many crustaceans have been used as remedies

since antiquity

cures based on crustaceans were described in

medical texts from the 1st century AD
eg. ashes from freshwater crabs used to treat rabid dogs

eg. decapod gastroliths rich in calcium used to treat e

xcessive stomach acid

eg. the hard parts, flesh, and juices had diuretic properties
and were used to treat urinary and gynecological
problems

eg. during the 16th century crabs were thought to cure TB

and cancers

eg. even today up to 20 species of crustaceans are used

medicinally in Latin America

used as remedies for asthma, bronchitis, epilepsy, and

hemorrhages

8. Several Crustacea are used as experimental

models in research:
Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 41 Animals: Arthropoda - Crustacea; Ziser Lecture Notes, , 2016.11 42
 Phylum Arthropoda
General Characteristics of phylum shared by members of all subphyla:
-chitinous, hardened exoskeleton that must be shed to grow
-obvious segmentation (metamerism)
-paired, jointed appendages on many segments

Subphylum: Trilobita
body plan: head, thorax, pygidium
compound eyes
antennae
mandibles for feeding?
branched (biramous) lappendages
respiration by gills?
able to roll up like pill bugs
once most common arthropod, now completely extinct

Subphylum: Myriopoda (centipedes, millipedes)

body plan: head, long trunk
lack compound eyes
single pair of antennae
mandibles for feeding Major Groups:
unbranched legs on most segments Chilopoda (centipedes)
respiration by tracheae Diplopoda (millipedes)

Subphylum Chelicerata: (spiders, horshoe crab, scorpions, mites, ticks)

body plan: cephalothorax, abdomen
most lack compound eyes
no antennae Major C l a s s e s:
chelicerae for feeding (no mandibles) Merostomata (horshoe crabs)
four pairs of unbranched legs Arachnida (spiders, scorpions, mites & ticks)
respiration by gills, book lungs, book gills or tracheae Pycnogonida (sea spiders)

Subphylum Crustacea: (crabs, shrimp, crayfish, barnacles, pill bugs, water fleas)
body plan: cephalothorax, abdomen, tail
compound eyes
two pairs of antennae Major C l a s s e s:
mandibles for feeding Malacostraca (shrimp, crab, pill bugs, amphipods)
branched (biramous) appendages Branchiopoda (water fleas, brine shrimp, fairy shrimp)
respiration by gills Maxillipoda (copepods, seed shrimp, barnacles)
only subphylum that is mostly aquatic

Subphylum: Hexapoda (beetles, flies, bugs, crickets, mayflies, dragonflies, moths, wasps, etc.)
body plan: head, thorax, abdomen
compound eyes
single pair of antennae
mandibles for feeding
three pairs of unbranched legs Major Groups:
two pairs of wings Apterygota (wingless insects; springtails, silverfish)
respiration by tracheae Pterygota (flying insects; dragonflies, butterflies, etc)
includes only invertebrates that can fly
 Phylum Arthropoda
Subphylum: Trilobita
Subphylum: Myriopoda
Class: Chilopoda (centipedes)
Class: Diplopoda (millipedes)
Class: Pauropoda
Class: Symphyla
Subphylum: Chelicerata
Class: Merostomata (horshoe crabs)
Class: Pycnogonida (sea spiders)
Class: Arachnida (spiders, scorpions, mites, ticks)
Subphylum: Crustacea
Class: Branchiopoda (tadpole shrimp, clam shrimp, water fleas, brine shrimp, fairy shrimp)
Class: Maxillipoda (seed shrimp, copepods, fish lice, barnacles)
Class: Malacostraca (shrimp, crabs, lobsters, crayfish, pill bugs, isopods, krill, amphipods,)
Class: Remipedia
Class: Cephalocarida
Class: Pentastomida (tongue worms)
Subphylum: Hexapoda
Class: Insecta
Order: Protura
Order: Diplura
Order: Collembola (springtails, snowfleas)
Order: Thysanura (silverfish, bristletails)
Order: Ephemeroptera (mayflies)
Order: Odonata (dragonflies, damselflies)
Order: Orthoptera (locusts, grasshoppers, walking sticks, praying mantis)
Order: Dermaptera (earwigs)
Order: Plecoptera (stoneflies)
Order: Isoptera (termites)
Order: Embioptera (webspinners)
Order: Psocoptera (book lice, bark lice)
Order: Zoraptera
Order: Mallophata (biting lice
Order: Anoplura (sucking lice)
Order: Thysanoptera (thrips)
Order: Hemiptera (true bugs; squash bugs, assassin bugs, chinch bugs, stink bugs)
Order: Homoptera (cicadas, aphids, scale insects, leafhoppers, treehoppers)
Order: Neuroptera (dobsonflies, at lios, lacewings)
Order: Coleoptera (beetles, fireflies, weevils)
Order: Strepsiptera
Order: Mecoptera (scorpionflies)
Order: Lepidoptera (butterflies, skippers, moths)
Order: Diptera (true flies; fruit flies, house flies, mosquitoes, blow files, crane flies)
Order: Trichoptera (caddisflies)
Order: Siphonaptera (fleas)
Order: Hymenoptera (ants, bees, wasps, hornets)
 Arthropods - Chelicerates c. respiratory system
to extract oxygen directly from air instead of water
an ancient group with good fossil record
need moist thin surface protected inside body
include horse shoe crabs, sea spiders, scorpions, ! several experiments:
spiders, ticks, mites, sea scorpions book lungs
tracheae
most living species are fairly small
some can open and close spiracles

but some fossil sea scorpions (eurypterids) were d. modification of appendages

5-6’long; a few up to 9’
for life on land (gravity)
most members of the group are terrestrial strong muscles and strong support
!1st group of animals to successfully make transition to land
e. change in sensory systems
! oldest known fossils of terrestrial animals are chelicerate
arthropods from 420 M years ago (Silurian) change from chemical senses (~taste; more reliable in
water) to vision, vibrations of air & soil not water
(=sound), and airborne chemical senses (~smell)
Transition to land
f. Internal fertilization
required major adaptations:
gametes must remain moist
a. Ability to resist drying
leads to anatomical changes for copulation
already had waxy cuticle
more elaborate courtship rituals
! impermeable to water

b. improved excretory system g. eggs must be protected from drying

egg must contain enough nutrients for early development
for salt and water balance
or animal must bear live young
conserve water by getting rid of N wastes as urea or uric
acid rather than as ammonia (most aquatic animals)
h. ability to withstand extreme
Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 1 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 2

temperatures cephalothorax contains:

or become inactive during temperature extremes
feeding appendages; including chelicerae
eg. diapause and pedipalps

Distinctive Characteristics of Chelicerates: usually 4 pairs of walking legs

1. head is fused to thorax = cephalothorax chelicerates lack antennae

2. named after their main feeding appendage Feeding & Digestion

! chelicerae (pincer-like or fang like)
used to grab or pierce or tear prey
most also have second feeding appendage
= pedipalp
4. only arthropod group without antennae
chelicerae are either pincher like or fang like
pedipalps are usually larger and may also have
“pinchers”
in some the pedipalps resemble the walking
legs

5. most have 4 pairs of walking legs Respiratory System

Body Plan aquatic species have book gills

body consists of a cephalothorax and abdomen terrestrial species use book lungs or tracheae

cephalothorax book lungs consist of parallel air pockets

extending into haemocoel
no distinct head
! book lungs resemble book gills of Limulus
head is fused with thorax to form a but are internal
cephalothorax

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 3 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 4
Senses
= modified nephridia open at coxa of 1st and 3rd
1. Vision walking leg

aquatic species have simple and compound Reproduction

eyes
dioecious
terrestrial species generally have several pairs of
simple eyes or ocelli some with elaborate mating rituals

2. mechanoreceptors some with considerable parental care

! sensory hairs and bristles used for “touch”

some also can detect sounds; used for

“hearing”

Circulation

open circulatory system as in all arthropods

body cavity a haemocoel

in some, blood contains the blood pigment

hemocyanin to transport oxygen

Excretion

excretory organ of most Chelicerates are

malpighian tubules

also have coxal glands at base of some legs

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 5 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 6

Classification of Chelicerates Class Merostomata

Class Merostomata (4 species) Eurypterids (Water Scorpions)
ancient. formerly diverse. group of aquatic chelicerates
extinct group
today consist of 3 genera and 4 species of horse shoe crabs
! lived from Cambrian to the Permian
Class Pycnogonida (~1000 species) (especially Silurian & Devonian)

sea spiders
largest of all fossil arthropods
Class Arachnida (>100,000 species)
!some up to 6’ long
spiders & scorpions, mites, ticks, etc
body plan is a blend between horseshoe crabs and
scorpions

cephalothorax with simple and compound

eyes, chelicerae & pedipalps

largest chelicerae of the subphylum

pedipalps resemble the 4 pr of walking legs

4 pr of walking legs

all but the last pair of walking legs had

“pinchers”

last pair of legs have paddlelike structures for

swimming

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 7 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 8
 segmented abdomen Horseshoe Crabs
[not “true” crabs which are crustaceans]
spikelike telson
also an ancient group; but still survives today
sluggish bottom dwellers
! almost unchanged for 300 million years
marine, brackish and probably freshwaters
can grow up to 2 feet in total length and weigh up to
feeding: 6 lbs

they were the “killing machines” of the oceans females usually larger than males

probably mostly ate trilobites generally found on sandy bottoms in deeper water

one paleontologist: “ I’d much rather be in a pool with a can swim upside down above sediment
6’ shark than a 6’ eurypterid”

Body Form
Eurypterids may have been the first animals to move
onto land
large cephalothorax, abdomen & long telson

cephalothorax with compound and simple eyes

Feeding & Digestion

eats clams, snails and sandworms

mouth is in center of legs

! chews food with bases of legs = gnathobases

have stiff spines

uses chelicerae to help get and break up food

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 9 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 10

Ecological and Human Impacts

Respiration
used by humans as cheap fertilizer for past 100 yrs,
breaths by flat, leaflike book gills on underside of until 1970 when the last processing plant closed
abdomen
pharmaceuticals are also extracted from them
the book gills can also be used for swimming
today millions are taken to be used as bait for eels,
Reproduction & Development conch and whelk

spawn in early summer (May & June) are the most important food source for loggerhead
turtles
nights of full and new moons they arrive by 1000’s
on Atlantic beaches (100 years ago they arrived eggs are also a valuable food source for migrating
by the millions) shore birds

female always dragging 1 & sometimes 2 or 3 !overharvesting of horseshoe crabs has

correlated with a decline in shorebird numbers
males up onto the beaches with her
the dramatic decline in shorebirds caused New Jersey to
female burrows into sand to lay several 1000 eggs ban the collecing of horseshoe crabs recently, now
at a time on beach subject to $10,000 fine

a horseshoe crab reserve has been recently created in

may lay 120,000 eggs per season
Delaware Bay where they are now permanently
protected
one or more males deposit sperm on the eggs
then female covers them with sand today, the horseshoe crab population has stopped declining

a similar species is now almost extinct in Japan

in 8-10 days, eggs hatch into larvae = “trilobite
larva”

horseshoe crabs live up to 20 years

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 11 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 12
 Class Pycnogonida Class Arachnida
(sea spiders) >100,000 species
1000 species most chelicerates belong in this class
shallow coastal waters and deep oceans include spiders, scorpions, mites, ticks, & “daddy
long-legs”
common in all oceans, especially polar waters
body with cephalothorax & abdomen
also, often found on soft tissues of sponges,
hydroids and soft corals, anemones, and clams cephalothorax with eyes, feeding appendages
and 4 pairs of walking legs
spider-like in appearance but with very small body and
very long legs abdomen may or may not be segmented
most with 8 legs; but some with 10 or 12 contains digestive, respiratory and
reproductive systems
many species have chelicerae and pedipalps
Feeding & Digestion
small head with 2 prs of eyes
most are predators with fangs, claws, venom glands
mouth at tip of long proboscis or stingers
are fluid feeders fangs = modified chelicerae
! feed on cnidaria and soft bodied animals claws = modified pedipalps
gut branches and gonads extend throughout entire stinger = last abdominal segments
length of each leg
generally predigest soft tissues then suck in
no resp or excretory systems
Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 13 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 14

predigested liquid with strong sucking pharynx Senses

Respiration most are predators

trachea, book lungs or both ! relatively simple eyes but very good vision

book lungs consist of parallel air pockets extending Excretion

into haemocoel
main excretory organ in arachnids are malpighian
oxygen diffuses from air into blood tubules

Circulation =closed end tubules that branch off the hindgut or

open circulatory system as in all arthropods
dorsal tubular heart that moves fluid around internal
organs
rectum and float in haemolymph
unique organs of arachnids and insects
also have coxal glands at the base of each leg

hemocyanin is used to transport oxygen =modified nephridia

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 15 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 16
 Classification of Arachnida A. Scorpions
Order: Scorpionida (scorpions) (~2,000 sp)
~2000 living species
Order: Thelyphonida (vinegarroons or whip scorpions) (~100 sp)
-book lungs
-coxal glands & malphiggian tubules for excretion most ancient of all arachnids
Order: Schizomida (short-tailed whip scorpions) (~230 sp)
mainly in tropics and subtropics
Order: Palpigradi (microwhip scorpions) (3,000 sp)
Order: Amblypygida (whip spiders) (~150 sp) but some temperate
-no tail
-short, wider, flat
-whiplike pedipalps especially deserts
-common in South Africa
Order: Pseudoscorpioida (pseudoscorpions) (3,300 sp) secretive
-very small, no tail
-poisonous, active at night = nocturnal
-widely distributed
-tracheal system
-coxal glands usually stay in underground burrows in day
Order: Solifugida (wind scorpions, sun spiders) (~1000 sp)
-resemble spiders Texas has 18 species of scorpions
-not poisonous but bite may become infected from bacteria Austin has 2 species
-in dry areas, most nocturnal
-SW US
Body Form
Order: Ricinulei (hooded tickspiders) (60 sp)
Order: Acarina (mites & ticks) (30,000 sp) cephalothorax & abdomen
-large order
-single prosomal carapace, united epistosomal segments
-pedipalps may be chelate or leg like cephalothorax (=prosoma)
-smaller ones have no respiratory organs, larger ones have
trachea short
Order: Opilionida (daddy-long-legs, harvestmen) (6,300 sp)
Order: Aranea (true spiders) (40,000 sp) contains large median eyes

2-5 pairs of smaller lateral eyes

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 17 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 18

even death
chelicerae and pedipalps
a few are deadly to humans
4 pair of walking legs
~25 of the 1500 species (2%)
abdomen (=opisthosoma)
none in Texas
subdivided into a wider mesosoma of 7 segments eg. Centruroides

and a thinner tail-like metasoma of 5 segments found in Mexico

kills 100’s – 1000’s/yr in Mexico

at tip of abdomen is recurved stinger
effect of poison is very rapid
Feeding & Digestion immediate drowsiness
excessive salivation
sluggish tongue
come out at night to prey on spiders and insects severe contractions of jaw muscles
fever to 104 or 105
detect prey by sensing vibrations in sand with hairs on may be hemorrhaging in stomach, lungs and
intestines
legs and possibly pectines waves of convulsions
cardiac irregularities
large prey are subdued by injecting paralyzing venom breathing difficulties
from stinger at tip of tail
death usually occurs in <3hrs
rarely sting humans ! only if provoked
if patient lives this long will probably
survive
venom is a neurotoxin that causes paralysis
Reproduction & Development
very painful but rarely fatal
scorpions have an elaborate courtship ritual and show
in children the more poisonous species may cause
convulsions parental care like spiders
vomiting
Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 19 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 20
male and female
at birth 1-100 babies are are born
hold each other by pinchers (chelae)
touch chelicerae after birth, babies climb onto mom’s back
and dance back and forth
! she carries them around for a week or so
male then stings female before they set out on their own

! to subdue her aggression

does not permanently harm her

male deposits spermatophore on ground

(a solid packet of sperm)

male maneuvers female until her genital pore is over

the spermatophore

she flips open lid of spermatophore releasing sperm

into her genital opening

in some species male then flees; in others he remains

to be cannibalized
! provides food to nurture young

development can take up to a year

scorpions are viviparous

! embryos develop inside mom

development may take several months to a year

absorb nutrients from mom

born alive
Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 21 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 22

B. Spiders then retracts like pocket knife

large group; 40,000 species some spiders also use chelicerae to:
carry prey
Body Form grasp objects
dig burrows
two part body
pedipalps help in manipulating food
cephalothorax and unsegmented abdomen
also sensory
connected by small pedicel
and to transfer sperm to female
cephalothorax
4 pairs of walking legs
simple eyes, no compound eyes, can detect
movement with sensory setae
in some may actually form image
! tactile; cover legs and detect vibrations
yet some with very good vision in web and in air

eg. jumping spider, Portia, has eyes with spatial acuity abdomen
better than most mammals and birds (better than a
cat or pigeon)
unsegmented
(but retinas have only a few 1000 receptors; humans
have ~200 M/eye) with spinnerets and up to six silk glands for
webmaking
most have chelicerae modified into a
“switch-blade” like fang
Feeding and Digestion
! extends and injects venom from poison
all spiders are predators
gland into prey
fang-like chelicerae inject venom into prey
Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 23 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 24
 eg. one African spider can liquefy a 2” fish or small
snake in <3 hrs
venom can be:
spider then uses pharynx and “pumping stomach” to
1. neurotoxin suck up liquefied prey

! affects nervous system and muscles spiders have very low metabolism:
to cause paralysis or death most can live for long periods without eating

2. digestive enzymes eg. tarantulas ! several months

eg. Black Widow ! 200 days

! digest proteins to kill and liquify prey
Spider Webs
3. pain inducing amines
virtually every aspect of spider biology depends on its
! make prey unable to fight back ability to produce silk:

after biting prey, spider backs off while toxin kills or used to wrap prey
paralyzes it silk is also used in web as “trip lines”

many spiders wrap prey with silk to further males produce sperm webs
immobilize it
females weave coccoons and build nursery webs

after prey has been subdued: juveniles use it to “balloon” to new habitats

spider may mash it with chelicerae and spiders have six different kinds of silk glands
regurgitate digestive enzymes over it
!each produces a different kind of silk:
or sperm web
drag line
if venom consisted of digestive enzymes injected coccoon
into prey the spider sucks up liquified prey various parts of web

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 25 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 26

silk is made of liquid proteins (keratins) that harden spider plucks web to determine where prey is located in
web
as it is secreted from glands
weight for weight spider silk is 20 x’s stronger than steel some other variations of web and silk use:
eg. Bola Spiders
silk is extruded from spinnerets
produce single sticky strand
!tension of pulling out silk changes its structure
from a liquid to a solid string they throw at flying insects (eg moths) to catch them

eg. many spiders live in webs of other spiders

after use it is eaten and amino acids are recycled into
new silk eat host’s prey or host

3 basic types of webs: Reproduction

1. sheet webs when ready to reproduce, male stops feeding

most common constructs “sperm web”

leads to funnel shaped retreat in which spider awaits prey
deposits drop of semen on it
conspicuous on lawns after heavy dew
picks up and stores semen in tip of pedipalp
2. cobwebs
loosely woven goes in search of a mate (pheromones)

depend on sticky threads to snare prey copulation is usually preceded by elaborate courtship
rituals
3. orb webs
most well known ! to ensure the female doesn’t treat suitor as
prey
most intricate geometry

general pattern varies by species if successful, male inserts pedipalp with sperm into
female genital opening
Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 27 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 28
female stores sperm in seminal receptacle eg. Black Widow

after mating male goes out in search of another has been found in all 50 states; common in central
female Texas

female remains in web and deposits 100’s or 1000’s of female is the most venomous
eggs in egg web
venom is a neurotoxin
often lots of maternal care
! venom is 15x’s more toxic than that of a
sexual cannibalism is also common among spiders prairie rattlesnake

after mating female will pounce on male but so little that only ~1% of bites are fatal

fatherly meal improves hatching success for eggs native Americans of California rubbed arrows with
mashed spiders for more effective hunting
fishing spiders are so aggressive that some
females pounce and eat the male suitor before causes synapses to release Ach causing:
muscular spasms
the nuptuals then lay unfertilized eggs that abdominal rigidity
don’t develop cramps
sweating
Examples of Spiders salivation
high blood pressure
sometimes convulsions
the two most dangerous spiders in US are
Black Widow and eg. . Brown Recluse
Brown Recluse.
South Central US, including Texas
both are common in Texas
venom contains digestive enzymes
both have a tendency to live in homes, outhouses and
outbuildings
Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 29 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 30

!its venom contains enzymes that destroy wander the jungle floor at night
blood cells
in day they hide inside termite mounds, under logs
this induces WBC’s to attack surrounding and rocks or in banana plants and bromeliads
tissues
in some bites very little venom is released;
creates large crater-like wounds that may in others a large amount is injected
require months to heal
bite can cause:
its bite can be fatal to children
intense pain

eg. Phoneurtria sp. (Brazilian wandering spider) sweating

also called banana spiders but NOT the same as the relatively acute allergic reaction
harmless Nephila, also called banana spiders
uncomfortable penile enlargement - can lead to impotence
most dangerous of all spiders (being studied as erectile dysfunction meds)

death (esp in children injected with large dose)

! most toxic venom
eg. camel ‘spiders’ (=wind scorpions)
species responsible for most spider bite deaths
mideastern species
Costa Rica and throughout South America east of have reputation as one of the nastiest arachnids but are
the Andes relatively harmless

large spiders: 10-12 cm (4-5”) leg span it is said: they are bigger than a human hand
extremely aggressive
will climb onto the belly of a camel and eat the camel
very aggressive alive

truth: they are as large as a 5 yr old child’s hand

! will attack anything (including humans) who they can run up to 10 mph (fastest moving nonflying
appear aggressive arthropod)
they don’t eat camels or people

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 31 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 32
C. Harvestmen (Daddy Longlegs; Harvestmen)
feed as scavengers on inverts and plants
~6,300 sp.
not predators
most conspicuous in late summer (harvest time)
cephalothorax has stink glands for protection
easily distinguished from spiders
Reproduction
Body Form
dioecious
cephalothorax & abdomen
internal fertilization
broadly joined abdomen and cephalothorax
rather than pedicel of spiders male has penis
cephalothorax lay eggs (oviparous)
no antennae Evolutionary Relationships
simple eyes new research shows they are more closely related to
scorpions than to spiders
no poison fangs

long thin legs

can be cast off to escape (=autotomy)

abdomen

unlike true spiders, abdomen is segmented

Feeding & Digestion

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 33 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 34

D. Mites and Ticks after mating, the female broods the offspring inside her
body

30,000 species described there may be up to 30 young, typically all are female except
for one male
! most authorities believe unknown species outnumber known
these offspring reach sexual maturity before they are even
(est: 500,000 - 1 M undescribed species) born

most economically important group of Chelicerates the lone male fertilizes all the females in the brood and dies
before being “born”

Body Form the females are born pregnant with the next generation

cephalothorax and abdomen completely fused the advantage to this bizzare life cycle is a very short
generation time; 3-5 days

no signs of segmentation eg. mites

have projecting mouth region = capitulum mites usually < 1mm

contains mouthparts: ! some mites are so small they can only be

identified with electron microscope
chelicerae for piercing or tearing food
many species are freeliving
Feeding & Digestion
feed on decaying vegetation;
some are blood sucking parasites
some are predators
Reproduction
some are blood sucking parasites during all or
some mites have a highly unusual reproductive part of their life cycles
strategy; eg. Acarophenax
these mites are parasites of beetles some mites have become adapted to live as
internal parasites in the lungs and air sacs of
Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 35 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 36
 snakes, birds and mammals
feed on dermal tissue of many vertebrates
a. follicle mites (Demodex) including humans

mutualists with humans the irritation is largely due to sensitization to

in the saliva that it injects to liquify skin
found in hairs of face especially around nose, cells (not due to burrowing)
and in ear wax
12-24 hrs after infection itching is at its worst
~ 1 in 5 people harbor them, proportion increases in
older folks
eg ticks
usually symptomless
ticks are much larger than mites
in a few may cause redness or irritation
all ticks are parasites during some part of
in dogs different species of same genus their life cycle
cause mange
feed on blood and lymph
b. dust mites
most infest mammals
feces of dust mites sometimes causes
allergies many attack birds

eg. chiggers (redbugs) a few attack cold blooded vertebrates

“there is probably no creature on earth that some show host preference; others are
can cause more torment for its size than a nonselective
redbug”
attracted by animal smells from a distance
is actually a larvae: minute, reddish; of up to 50’
0.2x0.15”; barely visible to eye
! tend to collect on game trails
Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 37 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 38

prodded by chemicals or heat

some lay up to 12,000 eggs at a time
best removed by gentle pulling
ticks surpass all other arthropods in the numbers
and variety of diseases that they can carry

they are vectors for some of the world’s main

human diseases

second only to the mosquito in importance

transmit viral, fungal, & bacterial diseases

eg. deer tick ! lyme disease

eg. wood tick !rocky mtn spotted fever

eg. tularemia (rabbit fever)

their bite can be serious

anticoagulants are sometimes toxic

may cause fever and inflammation

wounds made by ticks are very likely to

become infected

especially if “head” is torn off

may even result in blood poisoning

most ticks will not let go even if touched or

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 39 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 40
 Human Impacts of Chelicerates: excavate tortuous burrows in skin
esp fingers, wrists, elbows, groin, back of knees
mating occurs in burrows
most arachnids, though feared, are actually harmless eggs hatch and larval mite excavates 2-3 mm/day
to humans eg. chiggers
“there is probably no creature on earth that can cause
1. spiders are directly beneficial as predators more torment for its size than a redbug”
! each kill 1000’s of insect crop pests red bugs are mite larvae that are very small and can
easily go through clothing and burrow in skin
minute, reddish; 0.2x0.15”; barely visible to eye
2. large infestations of some mites can damage food adults are not parasites
and ornamental plants by sucking their juices chiggers don’t actually burrow into skin but secrete
saliva to dissolve skin cells for food
the irritation is largely due to sensitization to saliva
2. Venomous species ! a few are deadly and its enzymes
12-24 hrs after infection itching is at its worst
eg. black widow
eg. follicle mites
eg. brown recluse
cause mange in dogs
in humans related species are commensals not
eg. scorpions, esp Centruoides
parasites
live in hair follicles esp eyebrows
3. Arachnid Diseases and Parasites:
4. more serious impact on humans is as disease
eg. ticks vectors:
all ticks are vertebrate parasites
esp of birds and mammals eg. mites and ticks:
feed on blood and lymph Rocky Mtn Spotted Fever
some lay up to 12,000 eggs Lyme Disease
the bite can be serious relapsing fever
anticoagulants are sometimes toxic tularemia
may cause fever and inflammation Cattle Diseases

eg. mites 5. Chinese herbal Medicine

eg. dust mite Black scorpions are sold for $12/lb
feces in housedust causes dust allergies
Future Applications
eg. itch mite (scabies)
Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 41 Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 42

6. scientists are experimenting with venom genes to

use as biological control against insect pests

venom gene in virus ! infect & kill insect pest

7. spider silk is being investigated for a variety of

possible uses.
(very similar to insect silk, eg silkworms)

20x’s stronger than steel

it does not trigger an immune response

can be easily produced at low temperatures and

pressures compared to other similar polymers

is biodegradable

Animals: Arthropods-Chelicerates Ziser Lecture Notes, 2012.10 43

 Subphylum: Myriopoda
myriopods = many feet

mainly centipedes and millipedes

may be most primitive group of living arthropods

long worm-like segmented body

almost all segments are similar

mostly terrestrial, many freshwater, very few marine

species

general features:

body plan: head & long segmented trunk

1 pair of antennae

mandibles for feeding

uniramous appendages on each segment

tracheae for respiration

typical annelid-like nervous system

excretion by malpighian tubules attached to

digestive tract (similar to insects)
1
4 main classes (only 2 are major):

1. Chilopoda (centipedes)

2. Diplopoda (millipedes)

3. Pauropoda

4. Symphyla

2
 Class: Chilopoda (centipedes)

“100 feet”

3000 species

some tropical species grow to 25 cm (10”) long

usually found in cool, moist habitats

under longs and rocks

Body Form

head and long segmented trunk

usually somewhat flattened

head

with pr of antennae

pair of eyes

each eye consists of a group of simple ocelli

 cant form image

main mouthparts are mandibles

also 1-2 prs of maxillae

3
 usually with pair of poison fangs

trunk

long wormlike segmented trunk

almost all segments are similar

on each of the body segments is a pair of

walking legs
from 15 to 191 pairs

but none have 50 pairs (its always an odd #)

Feeding and Digestion

most are very active nocturnal predators

mainly use antennae to detect prey;

eyes not well developed

smaller ones eat worms and insects

larger ones, such as Scolopendra of Texas, can eat

lizards and small mice

mandibles are primary feeding appendages

1st pair of appendages is a pair of poison fangs to

subdue prey
4
  apparently not harmful to humans
sting causes slight pain
disappears in a few hours

Reproduction

prolonged courtship – several hours

male deposits a spermatophore

then transfers it to female with mouth

female often cares for young in underground burrow

5
 Class: Diplopoda (millipedes)

>10,000 species

“1000 feet”

usually in moist dark places

under logs and stones

can burrow through rotting logs

Body Form

body usually round in cross section

but locally, flattened millipedes are more common

head & long segmented trunk

head

head with 2 clusters of simple eyes (=ocelli)

1 pr antennae

feeding appendages are mandibles and maxillae

trunk

most “segments” after 1st 4 are fused in pairs (=diplosegments)

millipedes are “double footed”

6
 2prs legs/segment)

none have 1000 feet

most have fewer than 50 legs (25 prs)

maximum  752 legs (376 prs)

=Illacme plenipes; endemic to small area in California; thought

extinct until rediscovered in 2006

some can curl up like pill bugs for defense

like crustaceans, cuticle is often reinforced with

calcium salts

Feeding & Digestion

most are scavengers:

feed mainly on decaying vegetation

a few are herbivores

feed on living plants

much less active than centipedes

eyes not well developed  essentially blind

mainly use antennae as primary sense organ

7
some are able to spray defensive chemicals from
“stink” glands along sides of body for protection

eg. toxins, irritants, HCN, prussic acid

eg. a European species secretes a tranquilizer similar to
quaaludes

predator (eg spider) eating it

becomes totally relaxed for several days
Reproduction

dioecious

in most appendages on the 7th segment are modified

as copulatory organs in males

after copulation female lays eggs in nest and guards

them

8
 Adaptive Radiation in Molluscs
Class: Monoplacophora
shell: forms a single dorsal conical shell
head: reduced
mantle: covers undersurface of shell
gills: 5 or 6 pairs
foot: broad flat ventral, for creeping
radula: present
larva:

Classes: Caudofoveata & Solanogastres (formerly Cl. Aplacophora)

shell: none, but with calcareous scales and spicules in mantle
head: reduced
mantle: encloses animal
gills: absent, or present in cloaca
foot: reduced to small ridge within ventral groove
radula: present in some, for piercing
larva: trochophore

Class: Polyplacophora (Chitons)

shell: modified into eight overlappingdorsal plates
head: present
mantle: greatly enlarged, modified into "girdle" around base of shells
gills: present
foot: broad flat ventral, for gliding movement
radula: present
larva: trochophore

Class: Scaphapoda (Tusk Shells or Tooth Shells)

shell: anteroposteriorly elongated into tapering
tusk-like tube open at both ends
head: reduced to short proboscis
mantle: lines inside of shell, used for respiration instead of gills
gills: none; oxygen diffuses across mantle
foot: conical, elongated ventrally and used for burrowing
radula: present
larva: trochophore & veliger
Class: Bivalvia (Clams)
shell: two lateral, usually symmetrical, hinged valves
head: absent
mantle: lines inside of both shells; forms siphons for water flow
gills: most with pair of large gills; also used for feeding and as
marsupium
foot: ventral, wedge-shaped, very muscular, used for burrowing
radula: absent
larva: marine forms with trochophore & veliger; fw - glochidia

Class: Gastropoda (Snails)

shell: most with single shell often coiled spirally
head: present and well developed
mantle: lines inside of shell; in pulmonates forms lunglike sac
gills: some with single gill; terrestrial forms without
foot: ventral, very large, used for gliding movement
radula: highly developed as grater, rasp, brush or comb
larva: marine forms with trochophore & veliger

Class: Cephalopoda
shell: most reduced to internal pen or absent
mantle: forms external protective covering over visceral mass
gills: 2 to 4 present within mantle cavity
foot: modified to form siphon and tentacles
radula: present, takes in food cut with beak
larva: none
 Subphylum: Trilobita
distinct head with pair of antennae and
completely extinct subphylum; 4,000 fossil species compound eyes

earliest arthropod group heads of many were armed with long sharp
spines
appeared before the “Cambrian Explosion”
not sure of any specialized mouthparts
named for the division of the body into 3 longitudinal
lobes thorax

most 2-7 cm; largest 70 cm (~2.5’) 2-29 segments

all marine; no freshwater fossils are known segments added as animal grew

highly specialized marine bottom dwellers (benthos) each segment had pair of biramous
from shallow flats and reefs to deeper waters appendages

a few may have been free-swimming or planktonic ! one portion a walking leg

dominated marine benthos for 300 Million years ! the other part with a fringe of
filaments; probably gills
flourished during Cambrian and declined by Ordovician became
extinct at end of Permian ( base segment of each leg had bristles or
teeth
Body consists of a head (cephalon), thorax, &
pygidium
! probably used to grind food and move
food toward mouth
all three divided into longitudinal lobes in cross
section ! “trilobite”
pygidium
head
posterior end of several fused segments
Animals: Arthropoda - Trilobites; Ziser Lecture Notes, 2012.10 1 Animals: Arthropoda - Trilobites; Ziser Lecture Notes, 2012.10 2

Ecology & Feeding

trilobites were among the most prominent ancient

marine animals

trilobites were able to dig into bottom sediment to find

food or to conceal themselves from predators

could also roll up like pill bugs for protection

some may have been herbivores on seaweeds or beds

of algae

others may have fed on small invertebrates; sponges,

bryozoa

still others may have been filter feeders

Animals: Arthropoda - Trilobites; Ziser Lecture Notes, 2012.10 3

 Arthropods - General àexcellent for protection
includes: crabs, crawfish, shrimp, spiders,
àalso waterproof à good for life on land
scorpions, mites, ticks, millipedes, centipedes,
insects (dragonflies, butterflies, ants, wasps,
3. segmented body
beetles, etc)
allows infinite possibilities for adaptive
1,100,000 known species;
modifications
at least 2-3 M more species
more species in this phylum than in ANY phylum of ANY 4. well developed head (cephalization)
kingdom of life
with numerous sense organs
half of all know species of every kingdom of life

includes 2/3rds of all known animals antennae & compound eyes are characteristic
sense organs of arthropods
more widely distributed over the earth than any other
animal phylum brain (ganglia)

àlive in virtually every habitat on earth 5. several pairs of jointed feeding appendages
common in all terrestrial, freshwater and marine
habitats
6. very active and energetic animals

Distinctive Characteristics of Arthropods à most active invertebrate group

can walk, jump, burrow, fly
1. “jointed legs”
some can fly over 30 mph
à the only invertebrate with this trait
some can run up to 10 mph
2. hard (sclerotized) exoskeleton of chitin
completely covers body
Animals: Phylum Arthropoda-General; Ziser Lecture Notes, 2012.4 Animals: Arthropods-General Ziser Lecture Notes, 2015.11 2

___________________________________ food
Arthropods are one of the most ancient phyla with pollination
many fossils drugs, dyes, silk, honey, wax
crop pests
à polychaetes (annelids) and arthropods probably arose vectors of disease
from a common ancestor over 600 M years ago

à one of the few animal phyla that existed before the were the first animals to move onto land
Cambrian explosion à Silurian 420 MY ago

shortly after the Cambrian explosion arthropods spider-like and centipede-like animals appeared shortly after
plants moved onto land
quickly became the dominant lifeforms and have
dominated the fossil record since (predators; must have been prey around also)

one of the oldest animal species on earth (has remained were the 1st animals to fly
unchanged) is Triops cancriformis
150 MY before flying reptiles, birds, bats
insects à 330 MY; Carboniferous
à 180 M yrs à requires no males
pterosaurs à 170 MY;late Jurassic
birds à 150 MY; (coexisted with pterosaurs
many unusual forms now long extinct for ~90 MY)
bats à ~40 MY; late Eocene
in terms of numbers of individuals:
à opened up a whole new set of ecosystems and
200 M individual arthropods for every habitats
person on earth before anything else began to compete for the same
resources
most <6 mm (1/4”) long
allowed wide and rapid distribution and dissemination
largest: Japanese crab 12’’; largest ever found was 19’ (5.79M), across the globe
40lbs (18kg)

smallest: mite <0.1 mm

tremendous economic importance to humans

Animals: Arthropods-General Ziser Lecture Notes, 2015.11 3 Animals: Arthropods-General Ziser Lecture Notes, 2015.11 4
Arthropod Body Plan Body Wall

segmented body body is completely covered with hard exoskeleton

also folds into mouth and anus to form lining of foregut and
allows infinite possibilities for adaptive hindgut
modifications
cuticle also lines tracheae
lots of fusion of segments into a variety of body
plans: main component is chitin (a starch) but much thicker
than the thin flexible chitin of previous animal
head & trunk phyla such as segmented worms and roundworms

cephalothorax & abdomen in some chitin is further hardened with proteins

head - thorax - abdomen and calcium deposits (eg. crustacea)

paired jointed appendages exoskeleton is secreted by epidermis (hypodermis)

arthropods are the only invertebrates with jointed structure:

appendages
two major layers each further subdivided into
appendages are also highly adaptable to suit finer layers
almost infinite functions:
outer thin epicuticle: hardened (= sclerotized) protein
sensoryà antennae, palps with waxy surface for waterproofing

feeding à mandibles, chelicerae, etc much thicker, inner procuticle (includes exocuticle and
endocuticle): thick outer layer of chitin above a
locomotion à walking, climbing, swimming, thinner inner layer that remains thin and flexible
flying, walking, swimming,
some crustaceans (eg. lobsters & crabs) have a much
reproduction thicker and stronger procuticle

often impregnated with Calcium salts

Animals: Arthropods-General Ziser Lecture Notes, 2015.11 5 Animals: Arthropods-General Ziser Lecture Notes, 2015.11 6

also structures for respiration, swimming &

greatly increases its strength
mating
exoskeleton is often highly colored:
camoflage many spines act as tactile organs (touch)
recognition
warning with the advantages of this exoskeleton it has one
major drawback:
various microscopic canals run through cuticle and
open to outside: àanimals can’t grow without shedding and
regrowing a larger exoskeleton
pore canals à calcium salts for sclerotization in crustacea
Molting
wax canals à secrete waxy covering for water proofing

dermal gland ducts à unknown function the problem is solved by molting

exoskeleton consists of many separate hardened a complex process requiring environmental

plates with flexible hinges between factors and the interaction of various
à areas where cuticle hasn’t been hardened
hormones

the exoskeleton also contains various folds
(apodemes), flaps and spines:
muscles are attached to fingerlike inner
extensions of skeleton (=apodemes)
à when muscle pulls it moves part
eg. lobster closes claws
some parts modified for feeding
Animals: Arthropods-General Ziser Lecture Notes, 2015.11
includes actual shedding of old cuticle
= ecdysis
eg. insects go through a fixed # of molts till
adulthood, then they don’t molt anymore
eg. spiders & some crustaceans molt indefinite #
of times throughout their lives
a. molting is usually initiated by environmental
cues or a buildup of pressure in the body
à causes the release of molting hormone (=ecdysone)
7 Animals: Arthropods-General Ziser Lecture Notes, 2015.11 8
 b. triggers epidermis to secrete enzymes similar to muscle bundles that move our
(proteases and chitinases) that digest and bones
dissolve the inner layers of old cuticle
(procuticle) and it separates from body wall insects have more muscles than most animals
including us
c. epidermis secretes new procuticle
eg. humans have ~700 individual muscles; some insects
have 900 or more muscle organs; some caterpillars
d. arthropod inflates itself with air or fluid to have 4,000
crack the old skin (at fracture lines)
also, layers of muscles surround internal organs
e. animal extricates itself from old cuticle as in segmented worms

animal is especially vulnerable at this point both striated and smooth muscle fibers

eg. soft shell crab must also shed lining of intestine

Feeding & Digestion
and tracheae at same time
virtually every mode of feeding: carnivores,
f. animal inflates itself and allows new cuticle to herbivores, omnivores, parasites
harden
arthropods typically have 4-6 pairs of feeding
Movement appendages near their mouth

virtually every form of animal movement is found in two main types of feeding appendages:
arthropods:
chelicerae à pinchers or fangs
walking, running, crawling, burrowing, swimming, flying,
etc
mandibles à jawlike
arthropods have a very complex muscular system
the jointed plates of the body and legs provide with numerous accessory feeding appendages
attachment point for muscles
Animals: Arthropods-General Ziser Lecture Notes, 2015.11 9 Animals: Arthropods-General Ziser Lecture Notes, 2015.11 10

well developed, complete, digestive tract:

eg. lungs protected internal chamber for air
mouth: salivary glands
breathing arthropods
esophagus: tube that brings food to stomach
thin walls of chamber allow exchange of gasses with
body fluids
stomach: often with specialized areas for grinding
and storing and absorbing food
eg. book lungs
eg. crop, gizzard
several hollow internal folds; reverse of book
and accessory glands that secrete enzymes
and digestive juices lungs

intestine: efficient areas for absorption of nutrients able to work in air like book lungs work in water

anus: discards unused materials eg. trachea

Respiration all terrestrial arthropods use this system for

respiration
need some kind of respiratory system since waxy
cuticle is impermeable to air is a system of branching tubules that delivers
oxygen directly to tissues
arthropods use a variety of respiratory systems
lots of different kinds depending on habitat O2 doesn’t need to travel in blood

allows for high metabolism if insects

eg. gills in most aquatic species such as
crustaeans and aquatic insect larvae and doesn’t limit body size
nymphs
insect tracheal system was an excellent
thin feathery structures or flat sheets of tissue method to get lots of oxygen to muscle
tissues
eg. book gills in some chelicerates extend from
abdomen like pages of a book à preadaptation to flight

Animals: Arthropods-General Ziser Lecture Notes, 2015.11 11 Animals: Arthropods-General Ziser Lecture Notes, 2015.11 12
Circulation
dorsal brain and double nerve cord with paired
arthropods have a simple open circulatory system ganglia in each segment

à coelom becomes hemocoel filled with blood still relatively simple, doesn’t do a lot of
as in most molluscs processing
eg. cockroach can survive 30-40 days without a head
has dorsal heart and only a few blood vessels
but much better developed sense organs
dorsal blood vessel with paired ostia in each
segment
1. Eyes
blood flows anteriorly in dorsal vessel
a. simple eyes = ocelli
out into segments and circulates around
à can detect only light vs dark
organs and back to dorsal vessel
b. compound eyes
no capillaries
with many individual lenses = facets
blood of most arthropods contains pigments to carry
oxygen: provide a wide field of view and particularly
good at detecting movement
eg. hemocyanin à bluish pigment with
Copper 2. Antennae

eg. hemoglobin à red pigment containing Iron tactile & chemical sensations

Nervous System 3. Chemoreceptors

similar to annelids:
Animals: Arthropods-General Ziser Lecture Notes, 2015.11 13 Animals: Arthropods-General Ziser Lecture Notes, 2015.11 14

in addition to being on antennae, can be found on antennal glands excretory organs at the base of
almost any body surface antennae in crustaceans used to regulate salt
balance
eg. many insects have chemoreceptors on their feet

4. Tactile Hairs & spines malpighian tubules are excretory organs unique to
Arachnids and Hexapods
equivalent to our sense of touch àbranch from hindgut or rectum

5. Statocysts collects salts and wastes and drains into the

intestine
for balance
coxal glands modified nephridia at base of legs in
the more elaborate nevous sytem with sense organs some chelicerates
allows for some of the more complex invertebrate
behaviors in some aquatic species nitrogen wastes are excreted
through skin or through gills
still mostly reflex, but with some learning
Reproduction and Development
second only to cephalopods complexity
mostly dioecious
Excretion
lots of variation in developmental stages
arthropods have a variety of efficient excretory
systems to: often quite complex

remove excretory wastes eg. larva à metamorphosis à adult

larvae = caterpillars, grubs, maggots, nauplius in
crustacea
also prevents excessive water loss on land
often with complete change in feeding and lifestyles

Animals: Arthropods-General Ziser Lecture Notes, 2015.11 15 Animals: Arthropods-General Ziser Lecture Notes, 2015.11 16
 eg. aquatic larva vs terrestrial adult
Origin & Evolution of Arthropods
eg. nymph à juvenile à adult

arthropods show many similarities to certain

a few groups reproduce parthenogenetically
segmented worms
1. metamerism with tendency for segments to become
specialized

2. similar nervous system with paired ganglia in each segment

3. some have same type of excretory system

4. spiral cleavage in primitive members

5. mesoderm derived from 4D blastomere

soft cuticle of a segmented worm was hardened by

deposits of additional proteins and calcium

the hard sections of cuticle were still separated from

each other by flexible sutures and joints

à provided protection from predators &

environmental hazards

à provided more secure site for attachment of

muscles

parts of hard exoskeleton became pivots

and levers for jointed appendages

new jointed appendages provide much more rapid

Animals: Arthropods-General Ziser Lecture Notes, 2015.11 17 Animals: Arthropods-General Ziser Lecture Notes, 2015.11 18

locomotion than hydrostatic skeleton of past Classification

as coelom became less useful for movement it because of the diversity of arthropods:
classification is complex and difficult
became more important for circulation
it is difficult to generalize about various body systems
à became a haemocoel
even taxonomists have not reached consensus on the classification
and evolutionary relationships between some group

Major Subphyla:

There are 4 main kinds of living Arthropods (plus one

extinct group we will discuss)

1. Trilobites (4,000 species)

all extinct
mostly marine

2. Myriopods (14,000 species)

“many feet”
centipedes and millipedes
mostly terrestrial
distinct head with mandibles & 1 pr antennae
many similar segments

3. Chelicerates (74,000 species)

spiders, crabs, ticks, mites, scorpions

ancient group
mostly terrestrial
chelicerae and pedipalps for feeding
no antennae
cephalothorax

Animals: Arthropods-General Ziser Lecture Notes, 2015.11 19 Animals: Arthropods-General Ziser Lecture Notes, 2015.11 20
 4. Crustacea (67,000 species)

shrimp, crab, barnacles, crayfish

mostly marine
a few freshwater and terrestrial forms
mandibles, 2 prs antennae
many appendages & many different kinds of appendages
cephalothorax

5. Hexapoda (>1,100,000 species)

most successful animal group

87% of all arthropods
62% of all animals
50% of all life on earth
mostly terrestrial
a few freshwater, hardly any marine
distinct head with mandibles & 1 pr antennae
body consist of head, thorax and abdomen
3 prs of legs, most with 2 prs of wings

Animals: Arthropods-General Ziser Lecture Notes, 2015.11 21

 Phylum Echiura anterior flattened proboscis (=prostomium)
(Spoon Worms)
can be extended and retracted
140 species
posterior cylindrical trunk
sausage shaped worms
Proboscis
1 cm to 50 cm
has ciliated groove giving it a spoon-like
all marine appearance

most live in shallow waters; a few deep water forms proboscis is very mobile

!many burrow in sand or mud sweeps on mud to find organic debris

!other live in rock and coral crevaces can extend up to 10 times its retracted length
eg. Bonellia is 7 cm (~3.5”) long and can extend its
!a few live inside dead sand dollars, mollusc proboscis 1.5 meters (4.5’)
shells, or annelid tubes
no tentacles
they enter shells when young and get too large to
leave
Trunk
generally are deposit feeders
trunk is gray, reddish brown, or rose
Body Form
body has several sets of setae
cylindrical and somewhat sausage shaped
!hooked, anterior setae used for digging
resemble sipunculans in size and general habits burrows

body in two parts: !setae at posterior end for anchorage

Anials: Phylum Echiura; Ziser Lecture Notes, 2012.10 1 Anials: Phylum Echiura; Ziser Lecture Notes, 2012.10 2

Reproduction & Development

! circles of setae around posterior end for
anchorage and burrow maintenance dioecious

Feeding and Digestion show sexual dimorphism;

most are deposit feeders males often much smaller

collect small particles of detritus gametes shed into water

digestive system is extremely long and coiled external fertilization

mouth is at base of prostomium produces trochophore larva

anus is at posterior end of trunk metamorphosis to wormlike adult

Circulation Human Impacts

simple closed circulatory system in arctic spoon worms were once eaten by eskimos

Excretion

excretion by nephridia

Nervous System

simple nervous system

circumenteric nerve ring

ventral nerve cord

Anials: Phylum Echiura; Ziser Lecture Notes, 2012.10 3 Anials: Phylum Echiura; Ziser Lecture Notes, 2012.10 4
 Phylum Sipuncula generally drab colors
(Peanut Worms)
no setae
~320 species; a few fossil forms
Body
unsegmented benthic worms with tentacles
wormlike body is divided into two parts
look like small sausages or peanuts
anterior section = proboscis (=introvert)
most < 10cm long; range 2mm to >72 mm
posterior end = swollen trunk
all are marine
introvert
most species have worldwide distribution
bears mouth surrounded by scalloped fringe, lobes
or tentacles
mainly in shallow waters
anterior retractile tentacles
widespread on mud and sandy shores
are ciliated and grooved
a few to 5000 M
trunk
sedentary; most construct burrows in mud or sand
lined with mucus
muscular walls
a few live in coral crevices
Body Wall
one species bores into wood
has thick soft cuticle secreted by epidermis
extend tentacles to feed
beneath epidermis are layers of circular then
when some species contract their tentacles into the body they longitudinal muscles
resemble a peanut
Animals: Phylum Sipuncula, Ziser Lecture Notes, 2012.10 1 Animals: Phylum Sipuncula, Ziser Lecture Notes, 2012.10 2

eucoelomate = true coelom hemerythrin to carry oxygen

large, fluid filled coelom Nervous System

traversed by muscles and connective tissue brain with circumoral ring

strands
ventral unsegmented nerve cord
Feeding & Digestion
some simple sense organs
are nonselective, suspension or deposit feeders
Excretion
food is collected by cilia and mucous on tentacles
1 pair of sac-like nephridia
food is drawn into mouth when tentacles are retracted
Reproduction & Development
“J” Shaped digestive tract with anus
asexual reproduction occurs by transverse fission
bands of muscles control tentacles and anchor
digestive tract and help to stir its contents à posterior 1/5th of body pinches off to produce a
new individual
Respiration
sexual reproduction
gas exchange through body wall
almost all are dioecious
Circulation
ovaries and testes develop seasonally
no circulatory system
gametes develop in coelom and exit through
have two chambered coelom with circulating coelomic nephridiopore
fluid to transport gasses, nutrients and wastes
external fertilization
coelomic fluid contains red blood cells with
Animals: Phylum Sipuncula, Ziser Lecture Notes, 2012.10 3 Animals: Phylum Sipuncula, Ziser Lecture Notes, 2012.10 4
 larvae usually a trochophore larva

à may indicate molluscan affinities

Sipunculus most comon genus

Evolutionary Affinities

sipunculids are schizocoelus protostomes

may be distantly related to annelids

Animals: Phylum Sipuncula, Ziser Lecture Notes, 2012.10 5

 Phylum Annelida true coelom present
(segmented worms, bristle worms)
mesoderm on inside of body wall and outside
15,000 species of digestive system
layers of muscles inside body wall and on outside of
large successful phylum in water & on land digestive tract

include earthworms, sand worms, bristle worms, clam

worms, fan worms, leeches with head-body-pygidium
some with bizzare forms
worldwide distribution:
head (prostomium & peristomium)
marine, brackish, freshwater and terrestrial
most annelids show some degree of
Body Form cephalization with a distinct head
(=prostomium)
elongated wormlike body
tentacles, palps and sensory structures
<1mm to 3 meters
peristomium behind prostomium contains the
hollow tube-within-a-tube design mouth

one of the most successful animal designs with pharynx and chitinous jaws

à room for development of complex organs body with well developed metamerism
with muscle layers (=segmentation)

à allows for circulation of body fluids most prominent distinguishing feature

à provides hydrostatic skeleton seen in just a few other phyla: eg arthropods, chordates

segments are separated by tissue = septae

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 1 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 2

epidermis a single layer of cells (columnar epithelium)

each segment has it’s own set of muscles and
other organs epidermis secretes a thin flexible protective cuticle

allows more efficient hydrostatic skeleton for most annelids have setae à small chitinous bristles
burrowing and movement
secreted by epidermis
offers a way to achieve greater size:
repeated on each segment (ie. “bristle worms”)
rather than increasing size of each organ
used as anchors while burrowing
à each organ is repeated in each segment
to prevent capture
allows organs of each segment to become more
specialized for various functions such as some used for swimming
digestion, respiration, reproduction,
locomotion, etc or as protection or camoflage

the segmentation is both external and internal beneath epidermis is two layers of muscle tissue
essential features of segmentation:
thin layer of circular muscle
several systems (eg. nervous, excretory) show serial
repetition thick layer of longitudinal muscle (obliquely
striated)
segmentation is produced during embryonic
development
enhances use of hydrostatic skeleton
NOT the same as asexual budding as in tapeworms
allows for peristaltic movement for digging
terminal pygidium with anus
through sediment
Body Wall
body cavity a true coelom
Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 3 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 4
 lined with peritoneum (squamous epithelium) across surface

lines inside of body wall & outside of digestive 3. swimming:

tract
mainly polychaetes and leeches
also layers of muscle along digestive tract
undulating body movements
peritoneum also form mesenteries that hold
blood vessels and the septae between parapodia help in polychaetes
segments
Feeding & Digestion
Movement
complete digestive tract “tube within a tube” design
coelom is filled with fluid (except leeches) which serves
as hydrostatic skeleton muscle layers allow modification of tract into
various structures:
annelids have 3 general types of movements:
muscular pharynx- to take in food, often with
1. burrowing: eversible pharynx with jaws

waves of peristaltic contractions sweep crop – food storage

down body
gizzard – food grinding
1st animal elongates àcontraction of circular muscle

2nd animal sortens à contraction of longitudinal muscle intestine – digestion and absorption of
nutrients
setae anchor hind end of body while front end
pushes foreward anus – elimination of undigested wastes

2. crawling: Respiration

polychaetes use parapodia alternately to move through body wall in most species
Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 5 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 6

oxygen:
body wall is richly supplied with capillaries to
hemoglobin (Fe) red - most annelids
absorb and transport oxygen hemerythrin (Fe) red
chlorocruorin (Fe) green
some marine forms respire through parapodia
(only 4 blood pigments known in animal kingdom & annelids
have 3 of them)
a few species have gills
blood also contains amoeboid cells which engulf
Circulation foreign particles (like our WBC’s)

body cavity is filled with coelomic fluid which helps annelids therefore have a double transport system for
move food and wastes around foods, gasses, wastes

most annelids also have a closed circulatory system fluid filled coelom
that more efficiently carries nutrients and wastes circulatory system with heart & vessels

several pairs of “pumping hearts” keep blood flowing à foods, wastes and respiratory gasses are
carried both in blood and in coelomic fluid
dorsal and ventral vessels connected by capillary
network Nervous System
dorsal vessel sends blood anteriorly
have both CNS and PNS
ventral vessel sends blood posteriorly
CNS: a pair of dorsal cerebral ganglia above the
dorsal vessel is main pump pharynx and ventral nerve cord

several pairs of aortic arches (=”hearts”) help to with paired fused ganglia in each segment
keep pressure up in ventral vessel
PNS: nerves branch off fused ganglia to supply
blood: body wall and body organs

most with dissolved blood pigments to carry

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 7 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 8
Senses: secondary sex characteristics

simple single celled photoreceptors or clusters of regeneration

cells ocelli (= eyespots)
Excretion
a few polychaete eyes have cornea, lens, retina
one pair of nephrida (=metanephridia) in each
à can form images segment similar to that in molluscs
(a few polychaetes have protonephridia or both)
statocysts in some for balance
nephric tubule:
nuchal organ à ciliated pit in head area
nephrostome = funnel like opening into previous segment
also found in some molluscs and a few other
invertebrates coiled ciliated tubule surrounded by capillaries

may function in chemoreception bladder like structure

tentacles & palps à well developed sense of nephridipore = opening to outside

touch
function:
other simple chemoreceptors
wastes from coelom are drawn in
free nerve endings à tactile??
salts and organic wastes from blood are
discharged into duct
Endocrine System
useful stuff is selectively reabsorbed
neurosecretory cells in brain and ganglia
in earthworms and leeches chlroagogue cells
secrete hormones that regulate:
collect NH4 or urea and deposit in blood or
take directly to nephrostome
reproduction

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 9 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 10

some nitrogen wastes are also excreted through body distinct head with eyes and tentacles
wall segments with parapodia and lots of setae

excretory organs also help in salt and water balance no clitellum

Reproduction and Development Class: Clitellata (Earthworms & Leeches)

Annelids have both asexual and sexual reproduction subclass: Oligochaeta (Earthworms)
mainly terrestrial and freshwater
quite variable within the phylum
head absent
Asexual fewer setae, no parapodia

most can bud to some degree subclass: Branchiobdellida

other spontaneously fragment commensal on crayfish

no setae
Sexual
posterior sucker only
monoecious or dioecious
subclass: Hirudinea (Leeches)
most annelids are hermaphrodites terrestrial, freshwater or marine

larva, if present = trochophore no parapodia or setae

fixed # of segments with “false segments” (=annuli)

Classification of Annelida anterior and posterior suckers

Class: Polychaeta (Bristle Worms) Class: Echiura (Spoon Worms)

mostly marine shallow marine burrowing forms

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 11 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 12
 once considered a separate phylum

Class: Polychaeta
(Sand Worms)

means “many setae”; also called bristle worms

10,000 species; 2/3rds of all Annelid species

sand worms, bristle worms, fan worms, clam worms,

etc

largest, most diverse and most primitive class of

Annelids

all are aquatic; mostly marine; worldwide distribution

a few found in freshwater

most 2-4” long (5-10 cm) ; some up to 10’ (3 M)

often brightly colored

deposit feeders, filter feeders, predators, scavengers,

live in crevasses, old shells, burrows or construct

tubes

some have elaborate filtering structures

eg feather duster worms

a few are pelagic à part of the plankton

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 13 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 14

both parapodia and setae are moved by internal

muscle bands
important in marine food chains
these parapodia have a variety of functions and
Body Plan create many bizzare shapes:

distinct head with mouth and sense organs & à crawling or digging in the sediment; use parapodia
wormlike body or trunk with repeating segments as as legs

body segments with flaplike parapodia à swimming, use parapodia as paddles

Head à as gills for respiration

à used as anchors while burrowing or to prevent

have distinct head capture

head has retractable pharynx with chitinous à to create feeding currents inside tubes

jaws used to capture prey à converted into feathery appendages to filter water

lots of different kinds of sense organs à as protection or camoflage

1. chemoreceptors (nuchal glands) on palps and à in some, parapodia modified into fans and mucous
tentacles bags for feeding or to create water currents

2. touch receptors also on tentacles for locating food and most polychaetes are active swimmers, crawlers or
shelter burrowers in the sediment
3. eyes (simple eyes = ocelli; and more complex eyes)
Feeding & Digestion
some can focus an image = esp predators
1. predators
very similar to cephalopod and vertebrate eyes

Body or Trunk the most active polychaetes are predators

eg. clam worm or sand worm (Nereis)
on each segment are a pair of flaplike parapodia,
each with many bristles (=setae) up to 10” long

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 15 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 16
 live in mucus tubes in or near low tide; but can also swim
most are small; some up to 19 cm
males - irridescent bluish-greenish color
carnivores
females - light green with yellow, orange-red mottling
many are commensals with other marine inverts
most active at night
2. filter feeders
move out onto sand to search for food

use their jaws to capture small animals many polychaetes burrow or live in tubes rather
than crawling around on the sediment
jaws open as pharynx is everted

jaws close as pharynx is retracted many sedentary polychaetes are filter feeders

eg. Blood Worms (Glycera) eg. Fanworms, tubeworms, featherduster worms)

red worms, all marine, several species secrete many kinds of tubes:
firm calcareous tubes
found in shallow waters glue sand grains together
bits of shell cemented together
poor swimmers but good burrowers some burrow

carnivores most have long feathery tentacles that they extend to

filter feed
on their proboscis are 4 hollow jaws that can inject poison
into prey resemble colorful flowers when feeding

eat other worms and organisms in the sediment cilia on tentacles move collected particles toward
mouth
painful to humans
tentacles can be quickly retracted when threatened
harvested extensively in NE US for bait
often develop specialized food gathering structures
eg. Scale Worms
for filter feeding
very abundant
leads to tagmosis à fusion and reduction of
metamerism
flattened and coverd with scales formed by the modified
parapodia
Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 17 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 18

eg. Chaetopterus (parchment worm)

secretes parchment like tube

Excretion

creates a continuous current through its tube to feed protonephridia and in some metanephridia or both
tubeworm must maintain a flow of water to get oxygen
and get rid of wastes 1 pair per segment

à uses modified parapodia as paddles opens into coelomic compartments

can emit strong bioluminescent flashes
tubule absorbs any useful materials and concentrates
burrows often shared by commensal crab wastes as fluid passes to nephridiopore

3. detritus feeders Senses:

other polychaetes eat organic detritus in or on the eyes: simple eyespots to complex organs
sediment
esp in free moving (errant) polychaetes
Respiration
in one group can form image: cornea, lens,
usually through parapodia retina

some have paired gills on some segments nuchal organs: ciliated sensory pits

eg. Tangleworms (Cirratulus grandis) chemoreceptors used in food gathering

on east and west coasts of US
statocysts in burrowers and tube building forms
yellow to green; 5-6” long
Reproduction & Development
front with great mass of long red hairlike filaments used as
gills
simple reproductive system
some have no special organ and exchange across
body surface have no permanent gonads
Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 19 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 20
 à sea is literally thick with epitokes
à gonads appear as temporary swelling of just before sun rises, epitokes burst to release gametes
peritoneum at certain seasons
anterior portion of worm returns to burrows
gametes are shed either
=synchronous mating
à through genital ducts
àensure most eggs are fertilized
à or through nephridiopore
àpredator saturation
predators have a field day; but too many prey so
à or through rupture in body wall
some are always left to reproduce

some polychaetes live most of the year as sexually àatokes safely in their burrows to repeat next year
immature individuals = atokes
a Samoan holiday to feast on epitokes
after living 1 or 2 years as benthic organisms they
become sexually mature and swollen with gametes
= epitokes
head shrinks, body enlarges, gonads develop and
produce egg or sperm

sometimes only part of the body makes the transformation,

breaks off and the rest of the worm lives to repeat
next season

eg. palolo worm

males and females gather by the millions in one spot

at night determined by phases of the moon

female releases pheromone

pheromone excites male to circle about female

swarms of epitokes appear at start of moon’s last

quarter in October or November

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 21 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 22

Ecological Roles of Polychaetes they get most of their nutrients from symbiotic bacteria
living in a large sac (=trophosome) within the trunk of
the worm
eg. detritus food chains
the worms are bright red due to hemoglobin in their blood
eg. prominent in marine food webs
the worm absorbs the hydrogen sulfide and oxygen in the
waters near the vents
eg. Beard Worms (pogonophorans)
these bind to the hemoglobin in the worms blood and are
once thought to be a separate phylum, now known to be an delivered to the symbiotic bacteria in the trunk of
unusual kind of polychaete the worm

discovered in 1900; today 150 known species the bacteria harvest energy from H2S and convert inorganic
elements into sugars for the worm
all are marine; most live in bottom ooze of deep ocean
CO2 + H2S + O2 + H2O à H2SO4 + sugars
in many the forepart bears long tentacles giving it a bearded
appearance giant tubeworms reproduce by releasing sperm and eggs
into the water
thin, transparent, segmented trunk has several pairs of setae
and is enclosed in a chitinous tube the larvae will drift through the deep water until they locate
a hydrothermal vent
the trunk ends in a small segmented opisthosoma
they will then settle to a rocky perch
the best known of the group of beardworms are the giant
tubeworms found around deep sea hydrothermal vents the young tubeworms do have a mouth and gut
and feed

some up to 6’ long, as the worm matures the mouth and gut degenerate
and the area once holding the digestive systems
with a bright red plume that extends from the tube becomes a bacteria-filled sac

giant tubeworms are part of an entire ecosystem not based tube worms seem to have few predators
on photosynthesis
although sometimes crab and shrimp will feed on
they are the only non-parasitic animals without a digestive the worm’s red plume
tract
eg. Bone eating worms (Osedax)
no mouth, digestive tract or anus

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 23 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 24
 major decomposers of deep sea whale carcasses Economic Impacts of Polychaetes
2001 found red fuzz on whale carcasses in deep ocean
eg. human food (samoa)
1000’s of polychaetes with red plumes up to 6 cm long
eg. insecticides
new genus and species of polychaete
eg. Padan – a powerful insecticide produced from a polychaete
seem to be unique to “whale fall” worm

worms have no functional mouth or gut eg. anticancer drugs

have symbiotic bacteria that digested oil in bones
eg. dolastatins from sea hare (Dolabella auricularia) has potential
anticancer properties
à they degrade hydrocarbons

the bacteria live in rootlike structures of worm that extend

in and throughout the bone

worm provides oxygen via blood vessels extending into the

roots

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 25 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 26

Class Clitellata Subclass Oligochaeta

(Earthworms)
new genetic analysis indicates that what used to be
means “few setae”
3 separate “classes” of segmented worms should
more correctly be subclasses of a “new” class:
over 3000 species
Clitellata
relatives of sand worms but:
the clitellum is part of the reproductive system of
these worms
no distinct head
it is near the head
no parapodia
the clitellum is a thick, glandular, non-segmented
and very few setae
region in these worms that secretes mucous to
hold cross-fertilizing worms together while mating most with 4 prs of short setae/segment

and it produces a sac in which eggs are placed often present in high densities:

rich soil 1 ton of common earthworms/acre

very small aquatic worms (tubificids) up to

40,000/M2 in rich muds

earthworms are extremely important in the texture

and fertility of the soil
Aristotle referred to them as “the intestines of the earth”

Darwin wondered whether there was any other animal that

has played so important part in the history of the
world as earthworms

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 27 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 28
 “Earthworms are miniature topsoil factories, they make soil.
ALL other (terrestrial) living things eventually pass
through an earthworm on the way to becoming soil.
And it is likely that nearly every atom in your body
(with very few exceptions) has been in an earthworm’s mostly terrestrial àburrow in the soil
stomach before it was part of you.”
most conspicuous ‘worms’ on land
most oligochaetes are less than a few inches long
(roundworms are much more abundant but microscopic)
some tropical earthworms get up to 3 M long
many species are common in freshwaters
eg. giant Gippsland earthworm
eg. Aquatic “earthworms”
native to Australia;
smaller, benthic, longer setae, more active
average 3’ long and 1” diameter, can reach 9’ long
better developed sense organs
dark purple head and blue-grey body
some have gills
live in deep burrow systems in clay soils along stream
banks generally eat algae and detritus

take 5 years to reach sexual maturity some with great powers of asexual budding

breed in warmer months; lay coccoons in their burrows eg. tubifex

12” worm hatches in a year red worms to 10 cm long

a protected species – being killed from tilling the land
as area converts land from grazing to farming
eg. giant Palouse earthworm
in Idaho
thought extinct but recently rediscovered
up to 3 ft long, lives in burrows 15’deep
live on bottoms of lakes, ponds and polluted streams
live in very low oxygen concentrations
have large amounts of hemoglobin
keep their heads in tubes while waving bright red tails
in heavily polluted areas banks appear bright red at low
water

spits at predators absorb dissolved nutrients (DOM) across skin

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 29 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 30

feed on decaying organic matter in the soil

one unusual group lives on glaciers
eat as they burrow then let digestive system
eg. ice worms
extract nutrients
small worms <1” long
mouth beneath prostomium
only found on surface of glaciers at temperatures below
freezing
inside the mouth is a powerful pharynx
they die at temperatures of 40º F (5ºC) or more
in some aquatic species the pharynx can be
can appear by the 100’s
everted as in sand worms to suck food in
eat algae and pollen
the digestive tract may include:
and a few oligochaetes are marine or brackish
esophagus
Body Wall
has calciferous glands that maintain calcium
protective layer of collagenous cuticle secreted by balance by secreting excess calcium from
epidermis blood into the digestive tract

surface of the body is kept moist by (lots of calcium in soil; lots gets absorbed, excess is
secreted)

pores allowing coelomic fluid to leak out

and lubricate outer surface of animal
crop – for food storage
also has numerous mucous glands
gizzard – for grinding up food into smaller pieces
Feeding & Digestion
thick and muscular
most oligochaetes are scavengers or detritus
feeders

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 31 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 32
 intestine for chemical digestion and absorption of
nutrients paired nephridia in each body segment
in aquatic forms nephridia release ammonia
in some the first part of intestine is used for
digestion in terrestrial forms nephridia release urea (conserves water)

secretes digestive enzymes in fw and terrestrial oligochaetes nephridia not only

eliminate wastes but also eliminate excess water
most of intestine is used for absorption (osmoregulation)

on dorsal surface is infolding = typhlosole also, terrestrial worms have calciferous glands

increases surface area for absorption worms eat soil; soil has lots of calcium

on outside surface of intestine are yellowish high levels of calcium in blood

chloragogue cells
calciferous glands remove excess calcium from
à equivalent to our liver: synthesizes glycogen and blood and deposit it in the intestine for
fats
removal
à they also travel through coelom to repair wounds
Sense Organs
à function in excretion: convert amino acids to urea &
ammonia
rather than concentrated in head they are distributed
Respiration all over body

no respiratiory organs or parapodia like polychaetes numerous sensory cells (chemo- and mechano-
receptors) on skin
breath through skin, no lungs or gills
chemoreceptors esp on prostomium

extensive system of capillaries in epidermis many free nerve endings à probably tactile

Excretion
Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 33 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 34

earthworms have no “eyes” but do have

numerous photoreceptors in epidermis as cocoon slips over the genital openings it
receives an egg, then sperm
Earthworm Reproduction
fertilization occurs in the cocoon
earthworms are hermaphrodites
cocoon is deposited in soil
cross fertilize each other
in 2-3 weeks a new worm emerges
copulation involves a double exchange of sperm
cells

mucous secreted from clitellum holds pair

together with genital pores aligned

can last 2-3 hours

sperm is deposited in seminal receptacle

after copulation worms return to burrows

fertilization and egg laying occur a few days later

each worm secretes a sheath of mucous around

clitellum

clitellum then secretes nourishment for egg

then envelopes mucous and food in tough

chitin-like cocoon

the worm then backs out of the cocoon

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 35 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 36
Ecological Effects of Earthworms Human & Economic Impacts of Earthworms

1. Detritus food chain 1. Food for Humans

eg. Night Crawler in some parts of Asia, Africa and Latin America people regularly
eat worms
burrow within the upper 30 cm of moist soil rich in organic
matter usually because there is not much other food available

in soft soil earthworms move by peristaltic contractions a few restaurants in the US offer them as novel food fare

setae prevent back sliding 2. earthworms improve the productivity of farm soil
this type of movement only works because
segments are separated by septa
sometimes doubling or tripling crop yields

mainly active at night 3. Fishing bait

on warm damp nights, forage for leaves and organic debris worms are commonly used for freshwater fishing

up to 54,000 earthworms /acre nightcrawlers, redworms

à turn over 18 tons of soil per year
4. Vermicomposting
prefer moist soil but if too much water they will move to surface
using worms to recycle compost
à sometimes in great numbers

à used to think they “rained” down from the sky

important in keeping soil fertile since they are

constantly turning over earth and mixing
organic matter into it
if all material ever moved through earthworm gut was piled
on surface of earth it would rise 30 miles above sea
level (5x’s height of Mt Everest)

2. Food for birds and other animals

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 37 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 38

Subclass Hirudinea
(Leeches) except for a system of spaces (=coelomic sinuses
and channels) filled with coelomic fluid
500 sp
à acts as secondary circulatory system
mainly freshwater
Movement
a few marine and terrestrial
no parapodia
most 2-6 cm long; some to 20 cm (except 1 genus)

no setae
often brightly colored
leeches have poor hydrostatic skeleton
many are carnivores; some are parasites
aquatic species use muscle layers to make undulating
body is dorsoventrally flattened
swimming movements
anterior and posterior suckers
can also use suckers to move like inchworms
fixed number of true segments
some terrestrial forms are able to “stand up” on hind
à usuall 32 plus prostomium & pygidium sucker to search for prey

each segment with 2-14 annuli (=false segments) Feeding & Digestion

Body Wall most are predators of snails, worms and insect

larvae
coelom functions as a single large chamber
protrusible pharynx with 3 jaws armed with teeth
à no septae between segments
some are scavengers
coelom is filled with connective tissue and muscle
Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 39 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 40
some are blood sucking parasites Circulation

adaptations to parasitism by leeches: many species have no blood vessels

attach to host with suckers
coelomic fluid does the work of blood in open
pierce skin with sharp teeth on end of proboscis haemocoel

while cutting, secrete local anesthetic and histamine-like may be hemoglobin in haemocoel fluid
chemical that dilates blood vessels of host

consume large blood meals Nervous System

àblood is sucked by muscular pharynx
nervous system similar to other annelids
while being swallowed, blood mixes with hirudin
(anticoagulant) to prevent clotting but leeches have two “brains”

very slow digestion à one composed of paired cerebral ganglia around pharynx as in
other annelids

gut secretes very few digestive enzymes à the other in posterior of animal consists of 7 pairs of fused
ganglia
àdepend on bacterial digestion
simple sense organs are much better developed in
can live for almost a year on one meal terrestrial species which tend to be blood suckers

may take up to 200 days to digest one meal Reproduction

can live for another 100 days afterwards
hermaphroditic
Respiration
mating process similar to earthworms
most exchange gasses through skin
à cross fertilize during copulation
a few aquatic forms have gills

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 41 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 42

do have clitellum Human Impacts of Leeches

àproduce coccoon that receives eggs and sperm 1. medicinal uses

in past centuries medicinal leech, Hirudo, was

used to suck out “bad blood”
believed many bodily disorders were the result of bad blood or
too much blood

à were collected almost to extinction in Europe

now a protected species

introduced into US but rare in nature

today leeches used in medicine to speed healing

of reattached fingers and limbs

2. commonly used in biology labs

3. leeches have become leading research models

for understanding how the nervous system works

4. some chemicals used by the leech in obtaining and

digesting blood are being studied for treating
circulatory diseases

5. leeches have also affected history:

eg. land leeches of India

live in extremely large numbers in humid forests of India

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 43 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 44
 live in trees and shrubs and fall like “drops of dew” onto any
humans passing underneath Class: Echiura
(Spoon Worms)
their mass attack caused the retreat of a British
regiment during the Sikh rebellion in India in mid
1857 (rebellion against East India Company)
140 species

sausage shaped worms

1 cm to 50 cm

all marine

most live in shallow waters; a few deep water forms

àmany burrow in sand or mud

àother live in rock and coral crevaces

àa few live inside dead sand dollars, mollusc

shells, or annelid tubes
they enter shells when young and get too large to
leave

generally are deposit feeders

Body Form

cylindrical and somewhat sausage shaped

resemble sipunculans in size and general habits

body in two parts:

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 45 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 46

anterior flattened proboscis (=prostomium)

à circles of setae around posterior end for
can be extended and retracted anchorage and burrow maintenance

posterior cylindrical trunk Feeding and Digestion

Proboscis most are deposit feeders

has ciliated groove giving it a spoon-like collect small particles of detritus

appearance
digestive system is extremely long and coiled
proboscis is very mobile
mouth is at base of prostomium
sweeps on mud to find organic debris
anus is at posterior end of trunk
can extend up to 10 times its retracted length
Circulation
eg. Bonellia is 7 cm (~3.5”) long and can extend its
proboscis 1.5 meters (4.5’)
simple closed circulatory system
no tentacles
Excretion
Trunk
excretion by nephridia
trunk is gray, reddish brown, or rose
Nervous System
body has several sets of setae
simple nervous system
àhooked, anterior setae used for digging
burrows circumenteric nerve ring

àsetae at posterior end for anchorage ventral nerve cord

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 47 Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 48
Reproduction & Development

dioecious

show sexual dimorphism;

males often much smaller

gametes shed into water

external fertilization

produces trochophore larva

metamorphosis to wormlike adult

some males are parasitic

in some species (eg green spoon worm; Bonellia viridis) the first
larvae to settle and metamorphose become females

larvae that land on top of female become males

the tiny male creeps up her body, into her mouth and migrates
down to her uterus

up to 20 males become parasitic in the females uterus giving her

an instant supply of sperm without having to search and
mate.

Human Impacts

in arctic spoon worms were once eaten by eskimos

Animals: Phuylum Annelida; Ziser Lecture Notes, 2015.10 49

 Phylum Phoronida epidermis secretes cuticle

a small group of 20 species layers of longitudinal and circular muscles

worm-like animals true coelom

2 mm up to 30cm; most <20 cm long Feeding & Digestion

all are marine benthic animals filter feeders with conspicuous set of modified
ciliated tentacles
inhabit shallow coastal waters
= lophophore
may be brightly colored: orange, pink, green, yellow
circular or crescent shaped
all secrete a chitinous or leathery tube that is either
buried in the sand or attached to rocks or shells coelom extends into tentacles

extend front end from tube to feed feed on plankton and detritus

a few species bore into mollusc shells or calcareous cilia direct food toward mouth
rock
with up to 50 ciliated tentacles in two spirals
generally solitary
à collects food
but some tend to aggregate making the seafloor à entangles it in mucous
àcilia move it to mouth
àresemble a flower bed
typical U – shaped gut leads to anus outside
Body Wall lophophore

outer flexible cuticle Respiration

Animals: Phylum Phoronida, Ziser Lecture Notes, 2009 1 Animals: Phylum Phoronida, Ziser Lecture Notes, 2009 2

gas exchange through lophophore in some tentacles brood eggs

Circulation free-swimming ciliated larva metamorphoses into

sessile adult
closed circulatory system with hemoglobin inside blood
cells at least two species reproduce asexually
àadaptation to life in anoxic or low O2 environments

no heart, some vessels constrict to pump blood

Excretion

paired metanephridia for excretion

Nervous System

simple diffuse nervous system

with nerve ring but no distinct brain

Reproduction and Development

most are hermaphrodites; some are dioecious

àbut usually cross fertilize

fertilization can be internal or external

eggs fertilized internally are released through

nephridiopore

Animals: Phylum Phoronida, Ziser Lecture Notes, 2009 3 Animals: Phylum Phoronida, Ziser Lecture Notes, 2009 4
 Phylum Brachiopoda [outcompeted by bivalves?]
(Lamp Shells)
one genus, Lingula, alive today, dates back to Ordovician
(450MY ago)
335 living species; 30,000 fossils
à may be oldest “living fossil”
filter feeders with hard protective shell
modern forms are usually 5 - 80 mm
some resemble ancient roman lamp
some fossils up to 30 cm

mostly, sessile, bottom dwelling animals (benthos) most live specimens are dull yellow or gray
a few are orange or red
not colonial like other lophophorates
resemble bivalve molluscs
most prefer shallow waters; a few deeper forms
à untill mid 1800’s was classified with them
most live attached to rocks or firm substrate
have calcareous shell and mantle
some (eg. Lingula) live in verticlal burrows in sand
and mud bottoms but resemblance is only superficial:

also an ancient group with extensive fossil record àthe two valves are dorsal/ventral
à flourished in palaeozoic seas
àventral valve is typically larger
were one of the dominant phyla after the
Cambrian explosion àpromonant lophophore as feeding organ

à had hard protective shell as the number àmost are attached to substrate by thick pedicel
of predators were increasing on ventral valve
most died out in great Permian extinction
Shell
only 1% of species alive today
Animals: Phylum Brachiopoda; Ziser Lecture Notes, 2011.1 1 Animals: Phylum Brachiopoda; Ziser Lecture Notes, 2011.1 2

shell is secreted by mantle

tentacles of lophophore capture food collected by
the smaller dorsal valve fits over larger ventral valve ciliary water currents

valves may be ornamented with growth lines, fluting, ciliated groove brings food to mouth
ridges, spines
feed on algae and organic detritus
has hole for pedicel
complete digestive tract
pedicel attaches animal to substrate
among lowest rates of metabolis of all animals
is long, fluid filled
à can survive long periods without oxygen
muscular in some; not muscular in others
à minimal food requirements
a few species have completely lost pedicel
Circulation
Mantle
circulatory system with heart
secretes shell
some cells in “blood”; function uncertain
often bears long chitinous setae
may be to move nutrients around
may be for defense
Excretion
Feeding and Digestion
system of metanephridia
brachiopods are filter feeders like other
lophophorates Nervous System and Senses

Most of the body is in the posterior part of shell while most sensory receptors are on mantle margins
lophophore fills anterior
Animals: Phylum Brachiopoda; Ziser Lecture Notes, 2011.1 3 Animals: Phylum Brachiopoda; Ziser Lecture Notes, 2011.1 4
Reproduction and Development

almost all are dioecious

produce temporary gonads

gametes discharged through nephridia

most fertilization is external

only a few species brood their eggs

direct development in some, free-swimming larvae in

other species

eg. Lingula

eg. Terebratula

Animals: Phylum Brachiopoda; Ziser Lecture Notes, 2011.1 5

 Eucoelomates a few eucoelomates (mainly echinoderms and
chordates) are deuterostomes
animals having a true coelom are referred to as being
eucoelomates à the anus is the first to develop in an embryo

eucoelomate animals have a body cavity that is there are 2 main ways that a true coelom can develop
completely lined with mesoderm in an embryo:

the mesoderm can develop into muscle layers & in most protostomes the coelom appears as
internal skeletal elements a split in the mesoderm layer of the
embryo (schizocoelous)
à a large fluid filled coelom surrounded by muscle most invertebrate coelomates are protostomes
layers makes a more effective hydrostatic
skeleton in many worms in most deuterostomes, the coelom appears
as outpocketings of the archenteron
à mesodermal layers lead to development of (enterocoelous)
arteries and veins, ie circulatory systems;
echinoderms and chordates and a few minor phyla are
better blood supply to all internal organs deuterostomes

à since mesoderm lines the digestive tract, this

allows for the development of much more
elaborate digestive organs
eg. compare the digestive tract of earthworms to Ascaris.

à mesenteries to support internal organs

most eucoelomates are protostomes

à the mouth develops first in an embryo

Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 1 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 2

Animals – Molluscs but most (80%) less than 10 cm (~4”)

110,000 living; 70,000 fossils
molluscs are mostly aquatic; found from the tropics to
second largest phylum of animals in terms of number the polar seas
of known species
occur from the bottom of the oceans to 7000 M above
some estimate there are up to 150,000 species sea level
most versatile body plan of all animals
typical fauna of all parts of the ocean, ponds,
lakes, streams and rivers, mudflats, intertidal
range from fairly simple organisms to some of the
and terrestrial habitats
most complex and specialized of invertebrates
bottom feeders, planktonic, burrowers, borers,
includes: snails, limpets, clams, mussels, chitons,
pelagic forms
octopus, squid, oysters, slugs, nautilus, tooth
shells
one group, cephalopods, are considered the most
intelligent of all invertebrates
the phylum is divided into 8 different classes but 90%
of species are in only two: bivalves & snails
the phylum originated in the sea and most of them
remain there
good fossil record; since most secrete a shell
à only bivalves and snails moved to brackish
all living classes were well established early in the
and freshwaters
fossil record
but did not become dominant until the à only snails invaded land
brachiopods mostly died out in the Permian
Extinction (~250 MY ago) mollusks are closely related to segmented worms
microscopic to 20 M (50-60’ = giant squid) and up to
à same larval form = trochophore
900 kg (1980 lbs; ~ 1 ton)
eg. Tridacna 1.5 M and 250 kg (500 lbs)
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 3 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 4
Body Plan
well developed bilateral symmetry
most are unsegmented
a few primitive forms seem to show some segmentation.
most molluscs (except bivalves) have a well
developed head
bears mouth and various sense organs
in spite of wide diversity of group most share
some basic features in their body plan:
mantle - secretes shell or becomes outer body covering
itself
visceral mass – most internal organs are embedded in
tissue rather than being surrounded by the body cavity
shell - (internal or external)
radula – specialized feeding organ in mouth
foot - usually used for locomotion
1. Mantle
the body wall of mollusks consists of an outer
layer of epidermis that extends over most of
the animal as a mantle
often contains various sense organs
eg. ocelli, sensory papillae
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10
the body cavity of molluscs is a true coelom but it
is greatly reduced to a small space around
the heart
it has become part of an open circulatory system
3. Shell
in most molluscs the mantle secretes a shell
which serves as a protective exoskeleton
great variation in shell form and structure
shell made of calcium carbonate
calcium extracted from water, soil or food
shell is composed of 3 layers:
periostracum = outer
especially thick in freshwater molluscs
à protects from acidity common in fw
in many marine forms this layer is thin or absent
prismatic layer = middle
dense prisms of calcium carbonate crystals in protein
matrix
nacreous layer = inner
glossy film of calcium carbonate sheets
secreted continuously through live
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10
glands in epidermis secretes mucous, cement and
sometimes a shell
in some mollusks the mantle hangs down to create
mantle cavity around the internal organs
houses respiratory organs; gills
sometimes the mantle itself serves as a
respiratory organ
products from the digestive, excretory &
reproductive systems empty into mantle cavity
before release
a continuous current of water is created by
cilia to bring in food and oxygen and to
remove wastes and gametes
some molluscs are able to retract head or foot into
mantle cavity for protection
2. Visceral Mass
mantle is underlain by complex layers of muscle
and connective tissues
most of the body organs are embedded in a solid
mass called the visceral mass rather than
being located in a true cavity
5 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 6
thickens
produces irridescent “mother of pearl”
shell appears in embryo
grows throughout life à often growth lines
usually used for protection
but some bivalves use shell to rasp into wood
or rock
in some such as squid and octopus the shell is
internal and greatly reduced
the mantle serves as the animals outer
covering
4. Radula
in mouth is tongue-like rasping organ = radula
found in all groups except bivalves and aplacophora
(solenogastres)
hardened file-like, up to 250,000 tiny teeth
the numbers and pattern of teeth are used to identify
certain species
new rows of teeth are continually added to the
back of the radula ad the front teeth are
worn down
7 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 8
 the foot of many bivalves contain a gland that produces
byssal threads (sea silk) for attachment
the radula is supported by a cartilage-like rod
(=odontophore) silky filaments of keratin and other proteins

used to scrape, pierce, tear or cut food attached to substrate by extremely effective cement

in some molluscs a portion of the foot is modified

radula also acts as a conveyor belt to move food
into a long tubular proboscis for feeding
toward the digestive tract

5. Foot Feeding and Digestion

all molluscs have a thick muscular foot
variously modified for locomotion:
creeping movements - produced by waves of muscular
contractions that move through foot
or by creating a slime trail from mucous glands in the
foot
eg. snails à gliding movement
detritus feeders, burrowers, borers, grazers,
carnivores, filter feeders, etc
in mouth is tonguelike rasping organ = radula
most internal organs, including the digestive system
are embedded in solid tissue called the visceral
mass, instead of being free inside a body cavity
(coelom)

or digging into sediment complete digestive tract divided into discrete,

being extended from body into sediment functionally specialized regions:
hydraulically by engorging with blood to anchor
then draw rest of body into sediment
foregut
eg. bivalves
eg. scaphopoda buccal cavity
mouth, radula, salivary glands, esophagus
receives and prepares food
or for attachment:
may secrete mucus to adhere to solid substrate midgut

eg. limpets, chitons, land snails stomach and associated digestive glands
sorting region, crushing region
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 9 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 10

crystalline style in some to grind food

some digestion takes place in digestive gland simple heart with a few vessels
hindgut
heart with 2 auricles and 1 ventricle
intestine usually long and coiled
absorption of nutrients extending from heart is aorta
formation of feces (undigested residue)

reduced coelom becomes a haemocoel surrounding

Respiration
heart
most mollusks have folded, ciliated gills (=ctenidia)
blood contains several kinds of cells:
àthin feathery sheets of tissue covered with cilia
oxygen carrying cells with pigments to improve
efficiency of oxygen transport:
also used for feeding in bivalves
haemocyanin (Cu) - most - blue
some mollusks breath through their skin some with hemoglobin (Fe) - red

many terresrial snails lack gills (=pulmonates) some white blood cells
no clotting agents or thrombocytes
àmantle is modified into a saclike “lung” for
breathing air cephalopods have a closed circulatory system

Circulation Endocrine Systems

open circulatory system in most well developed endocrine system

à blood not entirely contained within vessels à controls egg laying and growth

works good for slow moving animals Nervous System

à the most active molluscs, the cephalopods, have a closed
circulatory system
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 11 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 12
CNS is a ring of ganglia in head area with paired
nerves and ganglia extending to other parts of the
body
eg. pedal ganglia à control foot
although the molluscs have a relatively simple nervous
system
cephalopods are considered the most “intelligent”
of all invertebrates
rival some mammals in ability to reason & learn
unlike most molluscs, cephalopods are active
predators
à can recognize prey with sharp eyes
à can learn by watching
Excretion
usually 1 pair of nephridia (=metanephridia)
often called kidneys (not really true kidneys)
(metanephridium consist of a tube that opens into body cavity
at one end and drains to the outside)
in many species the nephridial tubule also acts as
gonoduct
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 13
development is direct, ie. no larval stage
Evolution of Molluscs
new molecular evidence indicates an evolutionary
relationship between Molluscs, Flatworms, Rotifers
and Annelids
Molluscs branched off the main animal line about 545
MY ago
plant and animal life on land would not arise for
another 100 MY
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 15
some molluscs have several pairs of nephridia
some excretion also occurs through body wall and gills
Reproduction & Development
most molluscs are dioecious; some, especially
gastropods, are monoecious (hermaphrodites)
many marine forms produce characteristic larva
= trochophore
same larval form is found in some flatworms,
annelids and a few other minor phyla
in some gastropods and bivalves a second larval form
develops = veliger
it is also free swimming and has a foot, shell
and mantle
in many molluscs the trochophore larvae develops in the
unhatched egg and a veliger hatches to become the only
free swimming larva in these species
some freshwater bivalves produce a parasitic larva
= glochidium
it attaches to gills of host fish and
feeds on blood
in cephalopods, many freshwater snails and some
bivalves
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 14
Classification of Phylum Mollusca
Class: Aplacophora: now Caudofoveata & Solanogastres
Class: Monoplacophora
Class: Polyplacophora (chitins)
Class: Scaphopoda (tusk shells, tooth shells)
Class: Bivalvia (mussels & clams)
Class: Gastropoda (snails & slugs)
Class: Cephalopoda (octopus and squid)
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 16
 Class: Aplacophora Class: Monoplacophora
(sometimes divided into 2 classes; Caudofoveata & Solenogastres)
means: “one plate”
means “no plate or shell’
25 species
~370 sp
small; 3mm to 3 cm
features may be closer to ancestral mollusc than any
other modern group fossils known from Cambrian

all marine; most are small; 2-140mm long àwas believed to be an extinct group until
discovered near Costa Rica in 1952

soft bodied, wormlike, no shell, no distinct head small, single, low rounded shell with ventral creeping
foot
body covered with calcareous scales or spicules
superficially resemble limpets (gastropods)
some are burrowers in muddy sediments
unlike other molluscs; show some evidence of
segmentation
àfeed on protozoa & microorganisms and detritus
some organs are serially repeated in body:
others don’t burrow and live on the sediment and feed 3-6 pairs of gills
on larger organisms including cnidaria 8 pairs of muscles
3-7 pairs of kidneys (nephridia)

radula is present in some, absent in others [it was once believed that the mollusc ancestor was a segmented
wormlike animal but current research has discredited that
some have gills for respiration hypothesis]

have radula for scraping food

no nephridia
ladderlike nervous system similar to flatworms
monoecious or dioecious
eg. Neopilina sp.
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 17 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 18

Class: Polyplacophora broad ventral foot attaches firmly to substrate

(chitons)
the grooves along sides of foot form closed
means “many plates” “tubes” open at each end

1000 species Feeding

body is flattened with convex upper surface most feed on algae using radula to scrape algae from
surface
most 2-5 cm; some to 30 cm
one predatory species (Placiphorella velata)
prefers rocky intertidal zones; captures small invertebrates by “jumping” on
them
but found to 4000 M deep
Respiration
all marine
gills suspended in mantle cavity along sides of thick
clings to rocks or hard surfaces flat muscular foot

fairly sedentary; may move short distances to feed water is pumped across gills within the “tube”

head and cephalic sensory organs reduced at low tide when exposed, can breath air to some
degree
shell contains 8 overlapping plates on dorsal surface
Nervous System
if detached, can roll up like pill bugs/armadillo
2 long nerve cords extend along body
mantle forms a girdle around margins of plates
osphradia in mantle cavity near anus
sometimes covers part or all of plates
à samples chemical in water
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 19 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 20
Reproduction Class: Scaphopoda
(tusk shells, tooth shells)
sexes separate (dioecious) in most
means “trough foot”
often with complex courting behaviors
~900 species
produce trochophore larva
slender bodies in tubular shell, open at both ends

most 2.5-5 cm; some up to 25 cm

marine molluscs

subtidal to 6000 M deep

esp Atlantic coast

sedentary (not sessile),

à burrows into mud or sand

mantle wrapped around viscera and fused to form a

tube

foot protrudes through larger end of shell

àused to burrow into sediment

as it burrows it always leaves the small end

exposed to water

Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 21 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 22

Feeding & Digestion dioecious

feeds mainly on detritus and protozoa caught by cilia produces trochophore larva
on foot

or using tentacle-like captacula

àmucous covered tentacles extending from head

captacula may also serve a sensory function

radula carries food to gizzard for crushing

Respiration

no gills

mantle wraps around viscera and fuses to form tube

movement of foot and cilia pump water through

tubular mantle cavity

àgas exchange occurs across surface of mantle

Nervous system

no eyes

captacula may serve some sensory function

Reproduction
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 23 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 24
 Class: Bivalvia shell is mainly used for protection but does have
some other uses in some species:
means: “two valves” eg. shipworms use shell to burrow into wood and
then feed on wood particles
old name for class was pelecypoda = “hatchet foot”
eg. some clams are able to bore into rock and concrete with
includes mussels, clams, scallops, oysters, shipworms spiny valves

eg. scallops use shell for propulsion by quickly closing them

1 mm to 1.5 M (Tridacna) together to force out water
up to 500 lbs
à can swim in any direction
all are aquatic
mostly marine, Mantle
many brackish,
some freshwater shell is secreted by mantle

most are filter feeders their mantle can also produce “pearls”
pearl production is a protective action toward some foreign
no head, no radula, almost no cephalization body between shell and mantle

eg. sand grain, parasite, etc

Shell
mantle secretes layers around object
shell is laterally compressed; 2 valves (right & left)
commercial pearls are produced by some oysters & some
freshwater bivalves
“bulging” part of shell on dorsal side near hinge
= umbo posterior portions of mantle come together to form
incurrent and excurrent siphons
is the oldest part of shell; grows from edges
in some bivalves the siphons can be quite long
shell held together dorsally by hinge and adductor
muscles extending between shells Foot

Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 25 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 26

thick muscular foot used for locomotion

some bivalves are predators:
changes in blood pressure as hydrostatic skeleton eg. septibranchs slurp up tiny crustaceans by creating a strong
allow foot to dig in and anchor itself in sediment suction in mantle cavity to draw in prey

then muscles constrict and pulls rest of animal in eg. razor clams thrust proboscis between shells of other
bivalves to kill and eat them

some mussels are sessile and attach to substrate by some bivalves get most of their food from symbiotic
byssal threads secreted by glands in the foot algae (dinoflagellates) that live in mantle tissue

Feeding & Digestion eg. Tridachna

food enters mouth and enters stomach

most bivalves are filter feeders:
in stomach, food is sorted
gills are used to filter food out of water
a gelatinous rod (= crystalline style) spins
food is strained out as water passes over gills slowly (by cilia) & dissolves to release
digestive enzymes
trapped in mucous and moved by cilia to mouth
particles are sorted by size:
some bivalves are deposit feeders: àlarger particles move to intestine
àsmaller particles move to digestive gland
use long proboscis like organ formed from
modified foot to collect food in sand or mud Respiration

some bivalves use their shell to bore into solid objects gills used for respiration
to extract food:
eg. shipworms “termites of the sea”
cilia on gills create incurrent and excurrent flow
have long, wormlike appearance; long siphons
use small valves as rasping organs to drill through wood oxygen is extracted from water passing over gills
and eat wood particles excavated
bacteria in gut digest the wood and fix nitrogen for host
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 27 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 28
Circulatory System in most à fertilization is external

3 chambered heart (2 atria, 1 ventricle) in eggs develop into characteristic trochophore larva
pericardial cavity
in some marine forms a second free swimming larval
wraps around intestine form is produced = veliger

beats slowly (has shell, foot and mantle)

eg. oyster:
pumps blood to gills for oxygenation and to kidney for
excretion 50 Million eggs released per season

Nervous System embryo develops into trochophore

à veliger
à spat
sense organs poorly developed
freshwater bivalves have internal fertilization
statocysts in foot
gills become brood chambers

tactile cells in mantle cavity bivalved glochidia larvae (specialized veliger) are released

eyes in scallops along mantle edge some moms can “shoot” larvae into water column

à each has cornea, lens, retina, pigment layer others produce a structure that looks like a small fish to
entice predatory fish
Reproduction
when fish bites the “bait” it gets a mouthful of
glochidia which then attach to the lungs
usually dioecious;
they parasitize gills of fish for 1 - 3 weeks generally causing little
some (eg some oyster species) are prodandrous harm to their host but dispersing far and wide

then release and sink to bottom to become filter feeders

gametes discharged into mantle cavity and out
excurrent siphon
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 29 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 30

Class: Gastropoda Body Plan

means “belly foot” unlike clams, snails and slugs have a distinct head
with brain, sense organs and mouth
70,000 living species; 15,000 fossils
sense organs: simple eyes
à 1/3rd of all molluscs tentacles
chemical receptors
largest and most successful class of molluscs
mouth with radula
extremely diverse group: snails, slugs, abalones, à rasps and scrapes algae
limpets, whelks, conchs, periwinkles, sea slugs,
sea hares elongated body with foot below for gliding

microscopic to 1 M long; most 1-8 cm mantle secretes shell and forms dorsal surface of
animal
marine, freshwater and terrestrial representatives
in slugs the mantle forms a hollow breathing
à virtually every mode of life except aerial chamber

marine: littoral to deepest ocean Shell

some pelagic in open ocean
some brackish
most have a single heavy shell for defense
freshwater: rivers, lakes, streams ponds, etc
tropics to poles some shells with operculum
terrestrial: woodlands, pastures, mosses, cliffs,
some specialized for climbing a few have no shell

usually sluggish and sedentary snails are very well protected:

some sea slugs are brightly colored àgenerally secretive habits

Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 31 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 32
 àstrong shell, some with door

àsome can produce toxic secretions

tortion
à brings gills up front for better gas exchange
àsome can deliver an active blow with sharp operculum to deter
attack
à makes more room in shell for retraction
àsome can even redeploy stinging cells from cnidarian prey
à but puts anus over head; greater chance of fouling mouth

still many are eaten by insects, fish, birds &

some shells also show spiral winding instead of
mammals
straight coil
and parasitized by a variety of helminths
makes shell more compact
slugs have lost their shell
but unbalanced à shell tipped over
(but still produce one temporarily during embryonic
development) gills, auricle, and kidney on right have been lost

most shells show some degree of coiling; a few do this assymetrical loss helped reduce fouling of
not tortion

coiling occurred early in the fossil record of à water is brought into left front side of
gastropods shell and out right side of shell
all living gastropods, whether coiled like snails or uncoiled like
limpets and slugs, descended from coiled ancestors Locomotion

in addition to coiling, some animals also show tortion unlike other molluscs, gastropods glide on their “foot”
by secreting a mucus trail
tortion was a separate evolutionary event
from coiling; occurred later in evolution a wave of muscular contractions propels them
along
animal begins with basic bilateral symmetry but
becomes assymetrical due to tortion
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 33 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 34

it takes a lot more energy to move this way than it

does to run, walk, swim or fly feed on dead and decaying organisms

the “most expensive” part is mucus production 4. some are carnivores

many snails follow the mucus trail of other feed on a variety of animals
snails to conserve their own mucus
eg. other mollusks, especially bivalves; soft corals, fish,
worms, etc
Feeding & Digestion
some carnivores can follow chemical trails to
wide variety of feeding types in the group locate their prey

1. most gastropods are herbivores often have a long tubelike proboscis that they
can thrust between the shells of bivalves and
a. use radula to scrap algae off of hard surfaces kill and digest prey
such as rocks
some use venom to subdue their prey
eg. aquarium snails
eg. cone shells
b. some are browsers
one of the most specialized of the group
eg. abalone holds seaweed with foot and breaks off pieces
with radula to eat radula consists of individual teeth; each charged with a
highly toxic venom
c. some are plankton feeders
when prey is located the snail extends wormlike proboscis
they drift in plankton as they eat algae to attract prey, eg a fish

2. some are filter feeders its proboscis is loaded with one of its toxic teeth

when close enough, the tooth is expelled like a harpoon to

eg. limpets use gill cilia to draw in water current with food paralyze the prey
particles then filters out food
this allows a slow moving predator to catch often a
3. others are scavengers much faster prey such as fish

Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 35 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 36
 the stings of some species are lethal to humans
some sea slugs are able to harvest chloroplasts from
eg. moon snail (Polinices) & oyster borer (Urosalpinx)
the algae they eat and use them temporarily for
uses radula to help bore hole through bivalves photosynthesis

also secretes digestive chemicals to help drill through shell new data indicates that not all such slugs are able to do
photosynthesis with the “stolen” chloroplasts. What are
once hole is complete, snail thrusts proboscis through hole they used for?
and spends hours or days feeding on prey
Respiration
using its radula to cut and tear off pieces of flesh

simple gills (=ctenidia) are variously modified in

some predatory gastropods are pelagic or swimming
forms that have lost their shells = Nudibranchs
eg. sea slugs
nearly all are marine
widely distributed, mostly shallow waters some pelagic
brilliant colors; some contain toxins to subdue their prey
unique coloration may be warning
some feed on sea anemones and corals
à draw color from their prey
à the undischarged nematocysts of their prey are
preserved and transported to the surface of their
bodies and used for defense
eg. sea butterflies
some secrete a mucus net to capture zooplankton
aquatic forms
terrestrial snails (=pulmonates) lack gill but have
highly vascularized mantle cavity that serves as an
air breathing “lung”
lung opens to outside through a small opening (=
pneumostome)
draw air in over moist surface of the mantle to
extract oxygen
many pulmonates have returned to water and
therefore must surface periodically to breath
Nervous System
sense organs:
simple photoreceptors that detect light and dark

then draw their web into their mouth and eat web and all a few have image forming eyes with cornea and lens
trapped food
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 37 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 38

statocysts for balance

tactile organs – one or two pairs of tentacles

after cross copulation, each partner deposits its
one pair usually contains the eyes eggs in shallow burrows

chemoreceptors – especially in incurrent areas of some sea slugs have a “disposable penis”; after
respiratory system
copulating, the penis just falls
Reproduction
it takes 24 hours to regrow a replacement
monoecious or dioecious
a few snails (eg Crepidula) are protandrous
many gastropods perform elaborate courtship
ceremonies they live stacked on top of each other

in some, courtship can last up to 20 hours each begins life as a male

some use a syringe-like stylet to inject their paretner’s brain with the original male reproductive tract
‘manipulative’ chemicals while mating to enhance their
success
degenerates and the animal regenerates
either a female or another male tract
many terrestrial snails eject a dart from dart sac
à if attached to a female males will remain males
into partners body to heighten excitement
before copulation à if too many males in the “pile” then some will
become females
some hold onto the dart and use it as a dagger,
once female, they remain female for life
stabbing their partner up to 3000 times during
mating
a few primitive gastropods eject sperm and eggs into
water
it contains a chemical that improves chances
of their sperm fertilizing the other’s eggs
in aquatic forms eggs may be attached to substrate or
float freely in plankton
as hermaphrodites, they often cross fertilize each
other
some marine forms enclose eggs in wide variety of
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 39 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 40
 tough capsules Class: Cephalopoda
eg. welk
800 living species; 10,000 fossil species
a few freshwater forms brood their eggs in oviduct or
means “head foot” the main mass of the animal
bear live young (ovoviviparus)
with a cluster of prehensile arms and tentacles
extending from one end

include squid, octopus, nautilus, cuttlefish

extensive fossil record back to Cambrian

dominant life in ancient oceans after trilobites

declined

all marine

2 cm to 60’ (giant squid, weighs almost a ton)

giant squid is most massive invertebrate

recent genetic analysis indicates that giant squid (Architeuthis

dux) anywhere in the world are all members of a single
global population.

the young apparently disperse globally the settle as adults in one

place

most active & most advanced of all molluscs

à some argue: most advanced of all invertebrates

Shell
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 41 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 42

much thicker and more muscular than in other groups

most fossil forms (eg. ammonites common in Texas)
had very large heavy shells kept buoyant by gas protects internal organs
filled inner chambers
most cephalopods are excellent swimmers
today only Nautilus has a large external shell
can forcefully expel water from the mantle cavity
eg. Nautilus
through a ventral funnel (siphon) creating
shell spiral as some gastropods “jet propulsion”

but divided into chambers allows them to maneuver quickly to flee predators
chambers not found in gastropods
the funnel can be pointed to quickly move in any
chambers connected by cord of living tissue = siphuncle direction
can adjust gas in chambers for neutral buoyancy
squid are streamlined and have lateral fins
some have internal shell completely enclosed by that greatly improve their swimming ability
mantle
octopus is better adapted to crawling than swimming
eg. cuttlefish, squid
crawls over rocks and corals & into crevices using suction discs
some have completely lost shell and mantle encloses on its tentacles to pull or anchor itself
and protects animal
some deep water octopi have webbed tentacles and
eg. octopus swim like jellyfish
shell was sacrificed for speed to avoid predation
the surface of the mantle and the rest of animal is
Mantle covered by pigment cells called chromatophores

in most cephalopods the mantle serves as the chromatophore is an elastic pigment cell
animals outer covering
tiny muscles surround each one
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 43 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 44

contraction expands chromatophore and
changes color of body
allows animal to rapidly change color
under nervous and hormonal control
can produce general darkening and lightening
flashes of pink, yellow, lavender
can form bars, stripes, spots and blotches
the ability to change color quickly offers considerable
protection (in lieu of a shell) – provides instant
camoflage
also can be used for communication
can indicate danger, protection, or used
during courtship
many deep sea forms are bioluminescent
Head & Foot
in cephalopods, the head is indistinguishable from the
foot
the “head-foot” is elongated into 8 or 10 arms (up to
90 in nautilus) and 2 longer tentacles
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10
the beak bites pieces of prey off and the radula
further tear it into pieces
from the mouth, food travels through a long thin
esophagus to a small stomach
secretions from pancreas and liver help digest
the food into a mush
food is then passed to the cecum where digestion is
completed and nutrients are absorbed into the
hemolymph (blood) for distribution
undigested wastes are passed to the rectum and out
the anus, and finally out the siphon
Excretion
Kidneys extract wastes from the hemolymph and
release wastes through the funnel
Respiration & Circulation
since the cephalopods are so active they require more
oxygen than other molluscs
most have 1 pr of large gills; no cilia
mantle cavity expands and contracts to draw water
over gills à more efficient than cilia
1-way valves allow water to enter along edges of
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10
tentacles and arms have rows of suction discs
arms have suckers along their entire length
tentacles have suckers only at distal end
as cephalopod gets closer to prey it can shoot out
tentacles to quickly capture it
a mouth is at center of arms
contains chitinous beak or jaws
the rest of the foot is modified into a funnel for
directed movement
Digestive System
almost all cephalopods are predators
only the “vampire squid” is not a predator
it is a filter feeder using tiny mucous lined filaments to fish
for organic matter
it lives in low oxygen zones where predators could not
survive
use tentacles and arms to capture and handle prey
with its suction cups
strong beaklike jaws enclose tongue-like radula
octopus and cuttlefish have poison glands to help
subdue prey
45 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 46
mantle but force water to exit through funnel for
locomotion
closed circulatory system
à more efficient for gas exchange and transport
accessory (brachial) hearts at base of each gill
improves pumping efficiency even more
Nervous System and Senses
largest brain of any invertebrate
several lobes with millions of nerve cells
more elaborate than in other classes
much of our current knowledge of nerve cell function is based
on studies of the large nerve fibers of the squid
brain is located behind mouth
octopus and cuttlefish actual “think & plan”
learn and react to their environment
level surpassed only by some vertebrates
generally considered the cleverest of all invertebrates
and rival mammals in some ways
masters of disguise
47 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 48
 most versatile use of chromatophores in the animal
kingdom, esp. cuttlefish verts: receptors point away from light source, are
behind several layers of nerve cell and have
great curiosity; can solve problems “blind spot” where optic nerve exits the eye

can easily learn behaviors by reward/punishment cephs: receptors are in front of the “wiring”, no blind
spot, more light reaches receptors
if they see another rewarded for a particular choice they will
do the same thing to hopefully get the same reward statocysts: large and more complex than in other classes
important in controlling eye position
one octopus would short circuit the light over her tank by
squirting water at it if someone forgot to turn it off at tentacles: use tentacles for tactile exploration
night
Protection: Ink Sac
can use rudimentary tools
most cephalopods have an ink sac for protection
biologists have recently (2010) reported the first
example of “tool use” by an invertebrate:
along the side of rectum
a species of octopus in Australia has been
observed carrying around coconut shells and melanin pigment (same as in our skin)
using them for cover when threatened

European Union has recently (2012) offered them the à creates smoke screen to facilitate escape
same protections as lab rats from danger

sensory organs are better developed in cephalopods Reproduction

than in other mollusk classes
cephalopods are dioecious
eyes: most have very well developed eyes
with cornea, lens retina
in the male:
eyes show remarkable convergent evolution to vertebrate
eye; cornea, lens, retina, etc sperm are produced in a single large testis and
pass to the vas deferens for storage
differences:

verts: adjust shape of lens to focus image on retina the male packages sperm in a packet called a
cephs: lens is rigid, adjust shape of eyeball to focus
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 49 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 50

male must be careful since female will often

spermatophores and stores it in mantle cavity escape so quickly that she rips the arm off the
male
each spermatophore can contain up to 10 M sperm
not too serious since male dies soon after mating
in squid a spermatophore is about the size of a rice grain
after several matings the female can select which
sperm to use to fertilize her eggs
in the female:
eggs are fertilized as female lays them
a single large ovary produces eggs
eggs attached to stones in long strings of 1000’s of
2 large nidamental glands produce a hard, foul
eggs
tasting (to deter predation) capsule that encloses
the eggs
or in some species, eggs are brooded by the female
males often fight for the attention of a female
female squid dies after laying her eggs
courtship sometimes includes numerous color changes
direct development; no free-swimming larvae
& body movements
juveniles hatch from eggs
the female typically chooses her mate
generally, cephalopods don’t show parental care but a
a specialized arm of male is used to transfer sperm to
few cases are known:
female
= hectocotylus arm eg. one group of deep water squid carries and protects and egg
sac carrying up to 3000 eggs for several months
(proximal suckers are longer and thinner than on its other
arms) eg. some octopuses spend months caring for the eggs they have
laid
male plucks spermatophore from its own mantle
protecting them from predators and keeping them clean
cavity and inserts it into females mantle cavity
by the time the eggs hatch the mothers are so exhausted
that they die outright or easily succumb to predators
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 51 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 52
Major Groups of Cephalopods
Ecological Impacts of Molluscs
1. nautiloids 1. important in food webs in aquatic ecosystems
2. ammonoids -all extinct and even in terrestrial ecosystems
3. coleoidea - all living forms except nautilus

mollusks are eaten by a large number of animals

2. snails are major source of calcium for birds

3. oysters are keystone species since they tend to

form reefs nearshore

the cracks and crevices provide homes for a large

variety of animals creating a complex
community of interactions

4. threatened & endangered mollusks

freshwater bivalves are now the most threatened

group of invertebrates in the US

some can live up to 50 years

once >300 species in rivers, lakes, etc

today: ~24 are extinct

>160 endangered or threatened

11 species in Texas about to be listed in the

endangered list (2010)

especially due to:

Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 53 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 54

now found throughout the Great Lakes

hunting and harvesting & poaching voracious eaters that disrupt food chains by competing for
same food as native species
were actively harvested by native americans for food,
tools, jewelry
extremely fertile and can disperse at any stage of life
early settlers collected massive quantities for food and
able to strongly attach to any hard surface by tough byssal
pearls
threads
even today 1000’s are collected for pearls & meat
à have found 15000 attached to a single 4” clam
often sold as pet food; some human consumption
able to completely bock water inlet pipes of water treatment
damming rivers plants and power plants

pollution, sedimentation highly costly to remove

have been found clinging to boats and are quickly spreading

since they filter the water they are throughout the US
especially sensitive to pollutants and
not yet in Texas but already have been found in Oklahoma
sediment

they are often the first species to

disappear or decline when
environmental conditions change
due to mining, industry & agriculture

5. Bioinvasives
eg. asian clam

eg. zebra mussel

named for distinctive striped shell pattern

arrived in US in 1986 via ballast water from Eiuropean

freighter into Lake Erie

Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 55 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 56
 Human Impacts of Molluscs natural pearls take many years to grow and are very
expensive
1. currency: tusk shells used as money (=wampum)
cultured pearls use “blank” of polished shell placed under
by native Americans mantle of oyster, then allows layers to deposit for 3-6
years
2. as food: oysters, scallops, mussels, octopus, squid
much quicker process, much cheaper, preferred today
eg. >2 Billion pounds of oysters are eaten each year
4. ink à sepia dye
oysters first cultivated for food for 2000 yrs

oyster middens are found from ancient roman times and are 5. cuttlebone from cuttle fish
common at coastal archaeological sites
6. Pharmaceuticals
certain species consumed cooked or raw
eg. medication derived from cone snail venom alleviates some
eg. octopus, squid & cuttlefish: 3 million tons/yr are consumed; kinds of chronic pain that even morphine can’t control
an industry valued at >$6 Billion

eg. (1990): each night 1000 SE Asian boats set out ~25,000
7. Materials Science Research
miles of nylon drift nets that extend about 30 ft down from
eg. mussels have an amazing ability to attach to rocks. their glue
the surface to catch red squid
is made from protein strands that can attach strongly to
even teflon
the bycatch from this nightly activity;
~2000 dolphins
applications to develop aquatic glues, antifouling
~300,000 tons of pomfret (a kind of fish)
paints, stronger sutures
à these are discarded
1000’s tons of tuna, swordfish, salmon
à are kept and sold on the black market 8. shell collecting

3. precious “stones” 9. pollution control

shell nacre à buttons, cameos, etc
oysters consume nitrates and ammonia helping to
some species of oysters and some fw clams produce pearls clean & purify eutrophic waters
generally produced when a parasite gets between the shell
and mantle of the animal
Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 57 Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 58

a project in Chesapeake Bay is using oyster reefs to remove

19 million pounds (8.6 M kg) of nitrogen
compounds/yr from the bay waters

10. destructive species

snails and slugs à gardens

boring snails destroy oyster beds

shipworms à damage wooden ships and warves

some are not deterred by creosote treatment

some bivalves can even bore into concrete

bioinvasions; eg zebra mussel

intermediate hosts to parasitic helminths

Animals: Phylum Mollusca; Ziser Lecture Notes, 2015.10 59

 Phylum Entoprocta a few species can move on sommer-sault fashion like fw hydra

most form “hydroid-like” colonies with individual

150 sp zooids

tiny (<5mm), sessile stalked animals; solitary or each zooid has a muscular attachment disc with
colonial adhesive glands

colonial forms superficially resembles hydroid a stalk arises from attachment disc
colonies
ends in cup shaped calyx with crown of 8-30 ciliated
all but one genus are brackish or marine tentacles

range from poles to tropics tentacles are muscular and tend to roll inward

mainly in coastal, more brackish waters Body Wall

a few are commensal on sandworms outer layer of epidermis contains sensory pits and
cilia
1 freshwater genus = Urnatella

uncertain relationships to other animal phyla epidermis secretes cuticle

superficially resemble cnidarian hydroid colonies longitudinal muscle layer in body wall
but tentacles are ciliated

also superficially resemble bryozoa (ectoprocts) body cavity a pseudocoelom

some consider them acoelomates since body cavity is filled with largely filled with gelatinous parenchyma
gelationous material

Body Form body cavity extends into tentacles and stalk

almost all species are sessile Feeding & Digestion

Animals: Phylum Entoprocta; Ziser Lecture Notes, 2009 1 Animals: Phylum Entoprocta; Ziser Lecture Notes, 2009 2

filter feeders: cilia on tentacles create water currents Reproduction

to draw in food
asexual by budding
have a complete U-shaped digestive tract lined with
cilia sexual

mouth and anus are within a ring of tentacles some are monoecious, some dioecious

Nervous System some protandric

single large ganglion between stomach and oral produce testes 1st to make sperm
surface
then ovaries to make eggs
nerves extend from ganglion to calyx, tentacles, stalk
and sensory structures fertilized eggs develop in brood pouch

sensory bristles and pits on body surface develops into ciliated larvae somewhat like
trochophore larvae of molluscs and annelids
Respiration
larva eventually settles and attaches to become
no distinct respiratory or circulatory system sessile adult

respiration primarily through epidermis, especially in Symbioses

tentacles
sometimes found living on sponges or in tubes of
Excretion sand worms

protonephridia with flame bulbs embedded in à increases filtration for food

parenchyma

drain into ducts opening to surface

Animals: Phylum Entoprocta; Ziser Lecture Notes, 2009 3 Animals: Phylum Entoprocta; Ziser Lecture Notes, 2009 4
 Phylum Gastrotricha
(Bristle Backs)
gastro=stomach; trich=hair; sometimes called “bristle
backs”
790 species
mostly meiofauna (interstitial fauna)
small group of microscopic wormlike animals
same size as rotifers (usually <1mm, to 4mm)
widely distributed and abundant in freshwater and
marine environments
benthic, part of periphyton =the layer of detritus
and small organisms found on the seabed,
submerged rocks, plants and algae
many cosmopolitan species
a few species are terrestrial and live in water films
surrounding soil particles
resemble flatworms:
thin flattened wormlike body
acoelomate; no real body cavity
Animals: Phylum Gastrotricha; Ziser Lecture Notes; 2015.10
syncytial epidermis secretes outer cuticle
cuticle is hardened in some areas to produce
scales, hooks and bristles
glide on ventral cilia
longitudinal and some circular muscles in body wall
there is no body cavity; the interior of body is filled
with mesenchyme
entire ventral surface is ciliated, or sometimes the cilia
are arranged in rows
Feeding & Digestion
feed in algae, protozoa, bacteria and detritus
mouth is at the anterior end
food is directed to mouth by cilia on head
mouth opens into an elongated muscular pharynx
has a triangular or Y-shaped lumen
the pharynx opens into a cyliindrical intestine
lined with glandular and digestive cells
Animals: Phylum Gastrotricha; Ziser Lecture Notes; 2015.10
glide on cilia like planarians
cephalic ganglia with double nerve cord
protonephridia w flame cells
but have complete digestive tract
Body Form
head & trunk
head
with brain and sensory organs
trunk
usually the body is covered with spines, bristles,
scales or plates
end of trunk usually forked
has two or more adhesive tubes or cement
glands for attachment to substrates
a few also have an elongated tail
acoelomate
some consider them pseudocoelomates but no real body cavity
Body Wall
1 Animals: Phylum Gastrotricha; Ziser Lecture Notes; 2015.10 2
the anus is on the ventral surface close to the
posterior of the body
no respiratory or circulatory systems
gas exchange by simple diffusion
a few species may be capable of anaerobic
respiration
Excretory System
protonephridia serve in removing excess water
nitrogen wastes probably diffuse through the body
wall
Nervous System
brain consists of pair of ganglia near pharynx
pair of lateral nerve cords extend the length of body
primary senses consist of bristles and cilia on the body
that function as mechanoreceptors
a few chemoreceptors
but only a few species have eyespots (ocelli)
also, have sensory bristles on head for tactile clues
3 Animals: Phylum Gastrotricha; Ziser Lecture Notes; 2015.10 4
Excretion
some are protandric
protonephridia with solenocytes rather than flame
cells a few bear live young.

solenocyte has 1 flagellum enclosed in cylinder of gastrotrichs mature rapidly and have lifespans of only
cytoplasmic rods a few days
(flame cells have a tuft of many flagella)
Classification
Reproduction and Development
Two orders:
all are hermaphrodites
O. Macrodasyida
all marine
mutual cross-fertilization interstitial
hermaphroditic
fertilized eggs are released by breaking through the no protonephridia
body wall
O. Chaetonida
mostly freshwater
can produce two kinds of eggs: only parthenogenetic females known
1 pr of protonephridia
thin-walled egg for immediate reproduction

thick-walled egg for dormancy

resistant stage àcan survive for years

direct development: no larval stage

juveniles mature in a few days after hatching

most freshwater species are parthenogenetic

Animals: Phylum Gastrotricha; Ziser Lecture Notes; 2015.10 5 Animals: Phylum Gastrotricha; Ziser Lecture Notes; 2015.10 6
 Phylum Cycliophora
feed on bacteria or bits of food dropped from host
only 3 species
can regenerate by internal budding
first discovered in 1995 on the mouthparts of the àcontinually produces new funnels by internal asexual
Norway lobster budding

as the new funnel develops the old buccal funnel

very small; less than half mm; 0.35 - 0.10 mm disingegrates

all species symbiotic on crustaceans the new funnel then emerges through the trunk and takes
the place of the old one
more recently have been found on the American Lobster and
European lobster Life cycle

Body Form complex life cycle with 3 basic life stages

saclike, bilateral body à alternation of generations

with no body cavity, acoelomate 1. asexual feeding stage

back end has short stalk with adhesive disc by which dominant stage is sessile, solitary feeding stage
they attach to their host
neither male nor female
cellular epidermis surrounded by cuticle
350 microns; saclike
Digestive System
attached to host by short stalk with an adhesive
anterior feeding organ is a ciliated buccal funnel disc at the end

sessile feeding stage is asexual

cilia around mouth moves food into esophagus
2. sexual stages
esophagus leads to à U shaped gut à anus
Animals: Phylum Cycliophora; Ziser Lecture Notes; 2015.10 1 Animals: Phylum Cycliophora; Ziser Lecture Notes; 2015.10 2

the onset of the sexual stage is believed to be female matures and breaks free
triggered by imminent molting of the lobster
host. briefly free-swimming, then attaches to lobster
mouthparts
the feeding form produces either male or female
progeny which develop internally within the female form a new dispersive larva
develops; resumably from a fertilized egg
dwarf male
the female progressively degenerates and the
a simple sac mature larva escapes

nonfeeding; no mouth, digestive system or eventually the larva settles and

anus metamorphoses into an asexual feeding
stage
two reproductive organs contain stores of
sperm Evolutionary Relationships

two penile organs taxonomy is controversial; molecular data indicates it

may be related to rotifers and acanthocephala
matures inside feeding stage before breaking
free and becoming attached to another
feeding stage that is in the process of
producing a female reproductive individual

mechanism of fertilization is unknown

female

also develops inside a feeding form

contains a single egg

Animals: Phylum Cycliophora; Ziser Lecture Notes; 2015.10 3 Animals: Phylum Cycliophora; Ziser Lecture Notes; 2015.10 4
 Phylum Loricifera
22 species described; up to 100 more collected and
yet to be described
first described in 1983 from sediment off the coast of
France
very small: .1 -.5 mm long
strictly marine and found at all depths
seem to be extremely abundant in areas of the
seafloor with methane seeps
may be used in future as indicators of
methane deposits for commercial
exploitation
meiofauna
! live in spaces between gravel & sand grains
produce protective case = lorica
body with distinct cuticle
retractable head (=introvert)
resembles front end of kinorhynchs
Animals: Loricifera; Ziser Lecture Notes 2012.9
with recurved spines for sensory and movement
(burrowing)
probably feed by puncturing cells with stylets and
sucking out the contents
complete digestive tract
may feed on bacteria
dioecious
distinct larval stage (Higgins larva)
larvae swim with propeller like “toes”
_____________
a new species of loriciferan has recently (2010) been
discovered living in salt brines on the floor of the
Mediterranean Sea
it is the first an ONLY animal that does NOT do
aerobic respiration;
ie. it lives completely without oxygen
it has no mitochondria and uses anaerobic
respiration for energy
1 Animals: Loricifera; Ziser Lecture Notes 2012.9 2
 Phylum Priapulida retractable
(Penis Worms)
used in locomotion and in feeding
(from “priapos” the greek god of reproduction)
has adhesive tubules used to capture prey
17 living & 11 fossil species (Burgess shale)
has rows of spiny sensory projections
they were probably major predators during the Cambrian (=stylets)

cylindrical cucumber shaped worms that live in sand in some species these are elongated into
or mud tentacle-like structures

.5mm to 30 cm long trunk (abdomen)

all marine benthic animals mostly in colder waters covered with tubercles and spines

àfrom shores to several 1000 M’s 30-100 superficial rings

tail or caudal appendages

à marine burrowers
hollow
some species show high tolerance for hydrogen sulfide
and low oxygen levels probably a respiratory organ

Body Form also chemoreceptor

body divided into: Body Wall

proboscis (pharynx or introvert)
appears somewhat swollen
Animals: Phylum Priapulida; Ziser Lecture Notes, 2015.9
body wall is covered with cuticle of chitin over
epidermis
cuticle is molted periodically
1 Animals: Phylum Priapulida; Ziser Lecture Notes, 2015.9 2

with rows of circular and longitudinal muscles no circulatory system

à contractions circulate fluid in body cavity but contraction of body wall circulate fluids in body
cavity
à also uses hydrostatic skeleton to move along
substrate coelomic fluid contains O2 carrying cells and
phagocytes
circular and longitudinal muscle layers in body wall
Nervous System
body cavity is pseudocoelom that acts as a
haemocoel simple nervous system

Feeding & Digestion nerve ring around pharynx

most are burrowing predators prominent mid ventral nerve cord with ganglia running
the length of the animal
feed on bacteria and soft bodied invertebrates
no specialized sense organs
eversible pharynx
nerve endings, probably for the sense of touch
in mouth, cuticle is modified into teeth around the mouth

a few species are suspension feeders Excretion

complete digestive tract excretion via protonephridia (1000’s of solenocytes)

contains microvilli to improve absorption drains through urogenital pore

terminal anus Reproduction

Circulation dioecious
Animals: Phylum Priapulida; Ziser Lecture Notes, 2015.9 3 Animals: Phylum Priapulida; Ziser Lecture Notes, 2015.9 4
reproductive system is closely associated with the
excretory system

eggs and sperm released into water

extremely slow development

produces a free swimming larva in about 3 weeks

Animals: Phylum Priapulida; Ziser Lecture Notes, 2015.9 5

 Phylum Kinorhyncha
(mud dragons) the head is completely retractable

“moveable snout” head also with retractile proboscis for burrowing

(once also called echinodera) around head is up to 7 circlet of spines (stylets)

179 species Body Wall

tiny, free living marine worms body wall covered by a cuticle, secreted by syncitial
epidermis
usually <1mm long
only a little larger than most rotifers covered with spines

found in oceans throughout the world à pole to pole epidermis with numerous mucous glands

benthic: important part of interstitial fauna no cilia on body

burrow in silt & mud
beneath the epidermis are several bands of
some have been found in algal mats longitudinal, circular and diagonal muscles

a few are commensal inside sponges or other body cavity a pseudocoelom

marine invertebrates
Feeding & Digestion
Body Form
feed on diatoms (algae) and organic matter found in
short worm-like, somewhat flattened body divided in mud
13 spiny segments
mouth on protrusible mouth cone with powerful
has head, neck and trunk pumping pharynx

no limbs complete digestive tract

Animals: Phylum Kinorhyncha; Ziser Lecture Notes; 2015.9 1 Animals: Phylum Kinorhyncha; Ziser Lecture Notes; 2015.9 2

two pairs of salivary glands and pancreatic glands paired genital organs
secrete digestive enzymes into throat area
internal fertilization with specialized penial spines
a midgut acts as a stomach and an intestine
development includes minute free-living larva
anus at posterior end of animal
progressive larval molts
No circulatory system

Excretory system of 2 protonephridia

Nervous System

circumpharyngeal ganglia and ventral nerve cord

one ganglion in each segment

senses:

simple eyespots

spines and bristles with chemoreceptors and

mechanoreceptors

Excretory system

paired protonephridia

Reproductive System

dioecious but no dimorphism

Animals: Phylum Kinorhyncha; Ziser Lecture Notes; 2015.9 3 Animals: Phylum Kinorhyncha; Ziser Lecture Notes; 2015.9 4
 Phylum Nematomorpha
(Horsehair Worms)
326 known species; probably many more
a few fossils exist from the Cambrian and Cretaceous
periods
superficially resemble nematode worms – hence their
name
adults are aquatic; larvae are parasites
also called “hairsnakes” or “hairworms”
adults are freeliving and abundant in aquatic and
moist environments
àworldwide distribution
in all types of freshwater habitats with good
oxygen levels
also in damp areas around watering troughs,
swimming pools , puddles and cisterns
also in mats of algae or leaf litter along sides of
ponds and streams; occasionally in damp soil
1 known marine species
Animals: Nematomorpha; Ziser, Lec Notes; 2015.9
covered by cuticle formed from layers of collagen
fibers
secreted by epidermis, no cilia
longitudinal muscles only
mesenchyme forms bulk of the animal
Digestive System
adults are short lived and do not feed
have complete digestive tract but degenerate and
nonfunctional
Nervous System
anterior ganglia surrounding pharynx
ventral nerve cord extends down trunk
some have eyespot on head
no circulatory, respiratory or excretory system
Reproduction & Development
adults usually short lived – up to about a month
long enough to mate and lay eggs, then die
Animals: Nematomorpha; Ziser, Lec Notes; 2015.9
in old days were common in horse troughs
once thought they were literally produced from horses hairs that
fell into the water trough
in days past:
kids would pull hairs out of horses tail and place them in a
jar of water put it in the window watch to see the
“transformation”
adults mostly ~10 cm; range 5-120 cm (up to 3’, a
few 6’) long
but very slender usually ~1 mm dia (but up to
3mm)
reddish or dark brown or black
really do resemble the hair of a horses tail
adults share many roundworm features; once
considered to be nematodes:
adult resemblance
pseudocoelom
body covered with a cuticle
longitudinal muscles only in body wall
nerve ring around pharynx
but larvae are very different
à resemble kinorhynchs (a related phylum)
Body Wall
1 Animals: Nematomorpha; Ziser, Lec Notes; 2015.9 2
dioecious à distinguished by caudal end:
male à bilobed
female à trilobed
internal fertilization
during copulation sperm are released through
anus
adults often found in puddles after rain
sometimes hundreds together contorted into tangled mating
knots of males and females
à also called gordion worms
the name Gordian Worms relates to the fact that the worms often tie
themselves into knots (Gordius of Phyrgia, his ox cart, and an
intractable problem once easily solved by Alexander the Great)
after mating female lays long strands of eggs on
vegetation in water
eggs hatch into larva with spiny proboscis
larval stages are all parasitic in many kinds of
insects, esp. crickets & grasshoppers, also
parasites of centipedes, millipedes and
crustaceans
larvae swim about in water and are usually
ingested as host drinks
(in some species larva can actually bore into host)
3 Animals: Nematomorpha; Ziser, Lec Notes; 2015.9 4
within host, larvae live in the fluid filled body cavity of
their host

absorb nutrients through their body wall from host’s

body fluids

parasitic larvae got through several molts in host

on maturity, parasite acts on host’s brain and

stimulates host to seek water and drown itself

adults emerge only when host is in water and

quickly become sexually active

Animals: Nematomorpha; Ziser, Lec Notes; 2015.9 5

 Animals with a Body Cavity ectoderm à skin, nervous system
mesoderm à muscles, bones, circ sys
endoderm à dig and resp tracts
the animals discussed so far lacked any kind of body
cavity pseudocoelomates:

à organs, when present, were embedded in body wall is lined with mesodermal tissue
mesoglea or parenchyma tissue ie. muscle layers

virtually all other major animal phyla have some kind body cavity is filled with fluid
of body cavity
intestine has no mesodermal tissue
they are “hollow” with organs packed into this therefore no muscle layers
hollow space
tends to be simple, thin, collapsed tube
‘tube within a tube’ body plan:
eucoelomates:
à allows an increase in size

à allows more elaborate lengthening and coiling of internal layers of mesoderm on the inside of the body
organs wall and the outside of the digestive tract
à allows circulation of gasses, food and wastes in the
absence of a circulatory system muscle layers in both places

à provides hydrostatic skeleton digestive system much better developed

2 major kinds of body cavities:

pseudocoelom and true coelom

both have:

three embryonic tissue layers:

Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 1 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 2

externally no distinct head or obvious sense

Phylum Nematoda organs
(roundworms)
most are very small 0.5-1.0mm (100th of an inch to
= eel worms; thread worms 1/5th inch)

~25,000 known species largest is a parasitic species that lives in the

placenta of female sperm whales à 18’ long
specialists estimate that only ~20% of existing
species have been studied and described so far most are colorless and transparent or with
à there may be over 100,000 living species whitish or yellowish tint

a few fossils known; some in amber most abundant of the pseudocoelomate animals
all other pseudocoelomate phyla have relatively few species
very common and diverse group but poorly known and
difficult to identify nematodes may actually be second only to arthropods
in number of species
mostly free living but includes many common parasitic
species over two hundred species have been found in a spoonful of
beach mud

especially notable for their lack of variation in size and

in terms of sheer numbers, nematodes are probably
shape: “they all look alike”
the most abundant animal on earth
species more similar than in any other major à 4 of every 5 animals on planet are nematodes
phylum
à 90,000 nematodes were found in a single rotting apple

very simple and highly adaptable design:

generally; cylindrical, unsegmented worms,
tapered at both ends
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
occur in virtually all habitats from arctic to tropics;
marine, freshwaters, and especially in soil
there is virtually no part of the biosphere that
doesn’t harbor nematodes
3 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 4

à anywhere there is organic matter
have been found in deep ocean trenches and in
hot springs & ice
the deepest living animal known is a nematode
that lives in fractured rock 0.8 miles deep
à almost 3 times deeper than any other animal
(its DNA was found >2 miles below ground)
common as interstitial fauna
nematodes are especially common in soil
as numerous in soil as arthropods
eg. est 6 M individuals in 1 ft3 of soil
eg. upper 1” of soil may contain <1 Bill/acre
eg. 3.5M/m2 in tundra soils to 9M/m2 in grassland soils
à virtually every soil sample will yield new
species
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
nematodes also parasitize virtually every type of
animal and plant
~60% of all known nematode species are parasitic
virtually every species of vertebrate and many invertebrate
groups are hosts to nematode parasites
human parasites are the best known of the nematodes
but make up only a small percentage of total
species
a study done in 1947 found 99% of people around the world
were infected with nematode parasites
“everyone in the world has either had a threadworm
infection, has it now, or will have it in the future”
some of the largest roundworms are parasites,
eg. largest nematode is a parasite of a whale à 27 feet long
Body Organization
elongated, wormlike body; mostly small
few external features
difficult at first glance to distinguish front from back end
“tube within tube” design
animals with simple fluid filled body cavity around
internal organs=pseudocoelom
persistant blastocoel
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
“If all the matter in the universe except the
nematodes were swept away, our world would still
be dimly recognizable, and if, as disembodied
spirits, we could then investigate it, we should find
its mountains, hills, vales, rivers, lakes, and
oceans represented by a thin film of nematodes.
The location of towns would be decipherable, since
for every massing of human beings there would be
a corresponding massing of certain nematodes.
Trees would still stand in ghostly rows
representing our streets and highways. The
location of the various plants and animals would
still be decipherable, and, had we sufficient
knowledge, in many cases even their species could
be determined by an examination of their erstwhile
nematode parasites.”
-N. A. Cobb, 1914, Yearbook of
the US Dept of Agriculture, p. 472
enormous ecological importance
living species feed on a variety of organic material
à aerate soil
à recycle nutrients
à decompose toxins and wastes
there is no sharp distinction between aquatic and
terrestrial species
all nematodes including soil nematodes are
essentially aquatic
à live in water film around soil particles
5 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 6
mesoderm present only on external face of cavity
gut lacks muscle layer
three true tissue layers (=triploblastic)
ectoderm à skin, nervous system
mesoderm à muscles, bones, circ sys
endoderm à dig and resp tracts
eutely is common
à fixed number of cells in adult of each species
Body Wall
body wall a syncytial epidermis
adhesive glands usually present,
no cilia
secretes tough, flexible cuticle containing collagen
à protects worms from abrasion in soil and
sediment
à protects parasites from digestive enzymes
some have an elaborately sculpted cuticle
probably helps them move through soil or
sediment
cuticle is sometimes molted as animal grows
7 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 8
 cuticle sometimes shows superficial segmentation
their cuticle is highly resistant to fairly extreme
environments and conditions
à some can survive pH’s from 1.5-11.5
à some can survive mercuric chloride solutions that would
kill most other animals
à only living organisms to survive a space shuttle explosion
eg. 6 canisters of C. elegans survived the Columbia
disaster
allows them to survive in many unusual habitats
including:
eg. as parasites of both plants and animals
eg. in hot springs
eg. vinegar eel can live in concentrated mercuric
chloride that would kill most other animals
a very common soil animal
feeds on rotting fruit; can thrive in a wide pH range
from 1.5 to pH=11.5
before vinegar was pasteurized it was usually found in
commercial vinegars (cider vinegar from
fermented apples)
eg. a related species is found in pitcher plants
eg. another species has only been found in the felt
coasters under beer mugs in German pubs
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
eat small or microscopic animals
2. some are phytophagous
many marine worms feed on diatoms and other
algae
3. many are parasitic in plants and animals
roots of practically all plants are attacked by
some kind of nematode worm
~20-35% of nematodes found in soil are actually
plant parasites
all the root eaters have a syringe-like stylet
that injects digestive juices into root to liquify
meal
4. a very few may be saprobes:
eat dead or decaying matter
however, more recent studies indicate that most
of these “saprobes” are actually feeding on
live bacteria and fungi and are typically found
on or in dead organic matter such as dung or
decomposing bodies
complete digestive tract:
mouth is at front end surrounded by three “lips”
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
body wall with longitudinal muscles only
fluid filled pseudocoelom
à important as a transport medium for oxygen,
foods and wastes
à pressure created by tough cuticle and muscle
layer creates hydrostatic skeleton
Movement
unlike most wormlike animals they have only
longitudinal muscle in body wall
hydrostatic pressure in fluid filled pseudocoelom
maintains internal pressure and keeps body wall
from collapsing
(circular muscle does this in other worm phyla)
produces characteristic whiplike or snake-like
thrashing motion; “S”
Feeding and Digestion
nematodes feed on a wide variety of foods:
but almost all nematodes eat living cells
1. some are predatory carnivores
9 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 10
often with retractable piercing stylet
muscular pharynx which is able to produce a
suction to draw in food
food passes into a long straight intestine where
it is digested and absorbed
no muscles lining intestine - collapsed thin
tube
intestine only 1 cell layer thick
almost all digestion is extracellular
undigested material exits through anus near (but
not at) posterior end
they have a “postanal tail”, unusual in inverts
Nervous System
“brain” = nerve ring with ganglia around pharynx
dorsal and ventral nerve cords
àmainly controls dorsal and ventral muscle layers
muscles send processes to nerve cord
(opposite more common structue where nerve cells extend to
muscle cells)
11 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 12
 (this also occurs in some flatworms, gastrotrichs, and others)
muscle cells are interconnected so muscles on each
side contract together
à produce whip-like or thrashing contractions
characteristic of these organisms
Senses
visible sense organs generally absent
has mainly chemoreceptors sometimes in head
or tail
Excretion
unique excretory system:
excretory system a series of canals or tubules or
interconnected glandular cells (=renette cells)
sometimes with protonephridia
tubules form lateral line along sides of animal
visible from the outside
empties through excretory pore near front of
animal
no circulatory or respiratory system
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
àcan survive for months or years until
conditions improve
cryptobiosis
some nematodes can enter a state of arrested
activity
à makes them successful in seemingly
unfavorable environments
eg. some have been dried for several years then
rehydrated
eg. some have been placed in liquid air (-194ºC
(= -317ºF)) and revived afterwards
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
pseudocoelom fluids circulate nutrients, oxygen
and wastes
Reproduction
all nematode species show incredible reproductive
abilities
most species have separate sexes (dioecious)
and show sexual dimorphism
most with internal fertilization
sperm lack flagella or cilia à they are amoeboid
after mating, females lay 100,000’s eggs/day
eggs often extremely resistant to environmental
extremes
eg. Ascaris eggs remain viable in 5% formalin
development is usually direct; no larval stage
usually 4 juvenile stages; but resemble adult
juveniles grow by shedding (molting) old cuticle
in some species one of the juvenile stages
becomes an inactive, resistant stage
13 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 14
Ecological Effects of Nematodes
while rarely studied and relatively unknown,
nematodes occur in large numbers in virtually
every habitat
1. Nematodes are of enormous ecological importance:
they feed on a variety of organisms and are an
important part of all food webs especiall in soil
they are fundamental in recycling nutrients in all
ecosystems
eg. they play an important role in the nitrogen cycle by
helping to mineralize nitrogen
they also help to decompose toxins and wastes
soil nematodes help to aerate the soil
nematodes help regulate soil bacterial populations
and overall community composition
eg. one nematode can eat up to 5000 bacteria /minuite
since many nematodes parasitize plants and
animals they play key roles in population
and community ecology within most
ecosystems
2. Some nematodes play unique roles in the life cycles
of some animals
15 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 16
eg. one species of nematode spends its entire life on fig wasps
the wasps spend their entire lives inside the fruit and flower of
the fig tree
wasps are essential for fertilization of the fig flower
the worm attaches to the wasp as it is born in the fruit of the fig
it feeds on them until the fig flowers along with the wasp dies
the worms’ offspring await the birth of the next generation of
wasps as the fruit ripens
eg. the adult of a species of parasitic nematode, Myrmeconema
neotropicum, is found in ‘fruit-eating’ birds
its eggs are passed in the birds feces and collected by foraging
ants to feed their larvae
the infected ant larvae develop large bright red abdomens as
they mature
the ants tend to walk with their abdomens conspicuously
elevated
they also walk more slowly than uninfected ants
the birds apparently mistake the ants for berries and eat them
completing their life cycle
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
also damage ornamental plants, turf grasses and
greenhouse plants
other nematodes attack bark and forest trees
eg. pine wood nematode
in Asia, America and recently found in Europe
2. Other predatory nematodes are beneficial to
agriculture
eg. kill garden pests like cutworms and corn earworm moths
some of these beneficial nematodes are cultured
and sold as an organic form of pest control
3. livestock & pets also suffer heavy losses
eg. Dog Heartworm (Dirofilaria immitis)
a major global pest that affects dogs, cats, wolves, coyotes,
foxes among others
they can also infect humans
in US common in all 50 states, but especially common in SE
US in which ~45% of pets are infected
requires two hosts
mosquito is the intermediate host
ingests juveniles when it bites dog
transfers it to uninfected dog when another is
bitten
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
Human Impacts of Nematodes
while their primary value is their ecological roles, we
are most directly affected and aware of the
numerous parasitic species of nematodes:
1. Plant Parasitic Nematodes
cause extensive crop damage (eg potatoes,
soybeans, etc.) and billions in food & fiber
damage each year
up to 15% of our agricultural crops are damaged by
nematodes each year
eg. root knot nematodes alone cause over $100
billion/yr worldwide in crop damage
most plants can tolerate these parasites to some
degree
but when the balance is tipped in favor of the
parasite large scale damage may ensue
eg. Golden nematode (Heterodera)
parasite of potato plants
has caused crop failures worldwide
chemical in soil diffusing from plant stimulates
emergence of larva from cyst and attracts it
toward plant
17 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 18
in final host (dog) the juveniles undergo further
development
dogs can have juveniles circulating in their
blood and lungs without symptoms
once the number of worms exceeds a certain
number based on size of host the adult
worms move to the heart (usually 6-7 mo.)
and establish themselves in the right side of the
heart
adult worms can reach 12” long and live for
several years
after mating, females bear live juveniles
(=microfilariae) into the blood
the microfilariae can circulate for up to 3 years
“waiting” to be picked up by a mosquito
symptoms begin years after initial infections
begin as soft cough which worsens as the
infection increases
leading to congestive heart disease
untreated dogs die; treatment is difficult and takes several
weeks of discomfort to the dog to rid it of the
parasites
a few human cases are known
eg. Toxocara canis
common intestinal helminth infection in dogs and kittens
virtually all puppies and 20% of kittens are infected
19 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 20
 untill wormed common in tropics; in some countries over 50% of
children are infected
relatively mild symptoms in pet
even in US infections are not uncommon
can infect children but wont complete life cycle
à usually killed in liver or lungs main cause of infection is fecally contaminated food
à but can wander through various tissues and organs
causing inflammation Ascaris eggs are resistant to concentrated bleach and
formalin
human infections are fairly common; 3-20% in children,
especially in SE US they are also coated with an extremely sticky coating to
especially urban children adhere to almost anything

similar life cycle to Ascaris egg can survive >7 yrs after any trace of feces is gone

may be associated with neuropsychological effects after ingestion the juvenile burrows into blood/lymph vessels

4. Human Parasites à circulates into lungs

arrives in lungs ~2 months after initial infection

some species are important human parasites
à enters alveoli and ascends trachea or is
about 50 species of nematodes are able to coughed up and reswallowed

parasitize human hosts become adults in the intestine

virtually every human is host to some parasitic adult worms can survive for 25 years
nematode at some time in their life
they are not really parasites, they feed on material in
eg. Ascaris sp. intestine

the largest human nematode parasites; ~ 10 - 12” (up to 30 cm) if another worm of opposite sex is there they mate
long
female can release ~200,000 eggs/day (~8 Mil/lifetime)
found exclusively in humans
symptoms of infection:
1-1.4 Billion people in world are infected
juveniles:
20,000 die each year
local inflammation if they get into wrong tissue

Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 21 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 22

if large numbers in lungsà fever, spasms, coughing, severe

pneumonia-like symptoms, adult feeds on bacteria and wastes; not on hosts tissues
allergic rxns
adults: dioecious

a few adults à minor effects after copulation male dies

many adults à if “worm burden” is to great may cause
blockage
adults also have a tendency to “wander”; particularly if
living conditions become unfavorable
eg. fever, anaesthetics, worming tablets
they may exit the anus or out the mouth
eggs are not released into feces
instead female crawls to anus to deposit ~1500 eggs when
host is asleep
day or night à worm keys on sleep physiology of host
infections not dependent on fecal contamination for
spread

occasionally may perforate intestine or enter bile ducts à eggs are spread directly

eg. Pinworms (Enterobius vermicularis) à eggs are very resistant

one of the most common nematode infections of humans eggs are highly contageous
worldwide
can cause intense itching
500 Million are infected worldwide
eggs spread on sheets and in the air
small; ~12mm (0.5-.75”)
infections easily transferred to entire household
unusually, its more common in temperate areas than in tropical
areas eggs can also hatch and reenter the intestine

the most common helminth (flukes, tapeworms, (not easily detected by fecal exam since
roundworms) infection in US female lays eggs on skin outside anus;
use ‘scotch tape test’)
30% children infected
16% adults infected eg. Trichina worm (Trichinella spiralis)
more common in US in Caucasians than African
Americans - don’t know why probably the most serious roundworm disease of humans

seldom a health problem causes trichinosis: a potentially lethal disease

Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 23 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 24
 smaller than a pinworm à barely visible
in US an estimated 2.5% of the population is infected each year;
~750,000/yr
infections are also common in other parts of world
often appear in small sporadic outbreaks
each worm requires two separate hosts to complete its life
cycle:
occurs in several hosts: humans, pigs, rats, bears and
other carnivores
each may serve as intermediate or final host
juveniles travel through blood and encyst mainly in muscle
tissue of intermediate host
when raw or poorly cooked meat is eaten, juveniles mature
into adults in intestine of final host
after mating, the female burrows into the wall of the
intestine and releases juveniles into the blood
juveniles circulate to all parts of body
but coil up and encyst only in skeletal muscle cells
(eg. diaphragm, chest & abdominal wall, tongue,
biceps, deltoid)
humans are infected by eating infected meat
often due to undercooked pork, bear, sausage
symptoms of infection
from encysted juvenile
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
female can produce 25000-3000 eggs/day for up to 5 yrs
eggs released in feces
juveniles hatch in soil
feed on bacteria, 2-3 days
juveniles climb to the top of a blade of grass and wait for a host
to walk by
when juvenile contacts skin it immediately burrows into blood
vessels to lungs, then climb up trachea
àswallowed and reattach as adults in intestine
eg. Filarial Worms
8 species (include Wuchereria bancrofti, Loa loa, river blindness
(=onchocercosis))
females up to 10 cm (4”) long
250 Million humans are infected
common in tropical countries
highest rates of infection are in Sri Lanka
female worms release live young (=microfilariae) into blood
mosquito or fly is an intermediate host and vector of spread
microfilariae move to peripheral blood on periodic basis
corresponds to “biting hours” of local vector
eg. Loa loa à diurnal vector
eg. Wulcheria à nocturnal vector = mosquito
but in S Pacific vector bite in
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
range from mild to life threatening
light à soreness; achy muscles
heavy à esp dangerous if in heart
juveniles can remain viable for up to 2 years, but are slowly
calcified
eg. Hookworms (Necator sp. & Ancylostoma)
named for its hooklike anterior end where the head is bent into a
curved shape
up to 11mm long
found in tropics and subtropics
one of the most dangerous roundworm parasites
> 600 Million infected worldwide
adults live in intestine; blood feeders
large plates in mouth cut into intestinal lining to suck blood
often cause excessive blood loss while feeding
eg. a “mild” infection of ~1000 worms can cause a
loss of 100 mL/day (3.3 oz)
heavy infections can cause anemia and weakness
in children can cause retarded physical and mental growth
Reproduction & Life Cycle
dioecious: male smaller than female
25 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 26
day so are diurnal here
eg. in some places, no cycling of vector so no cycling
of movement to peripheral blood
eg. several species of filarial worms, including Wuchereria sp.,
cause Elephantiasis
infects 120 Million/yr in Africa & Asia
the juvenile worm is carried by mosquitoes
in host adult worms live in lymphatic system
symptoms associated with inflammation (fever & skin
lesions) and obstruction of the lymphatic ducts causing
swelling
results in excessive enlargement of affected parts
esp in arms, legs, scrotum
eg. Onchocerca (causes river blindness)
occurs in 38 countries worldwide especially in Africa, South
America, and the Middle East
the worm migrates through the body to victims’ eyes
is a major cause of blindness
18 Million infected worldwide
of those 270,000 are blind
500,000 have severe visual impairment
some villages have 100% infection rate
27 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 28
 or rectal prolapse (protrusion of the rectum out through the
anus
eg. Human Whipworm (Trichuris)
treatment is difficult since Trichuris is resistant to commonly
500 Million are infected worldwide; esp subsaharan Africa used drugs

a medium sized worm (3-4cm) eg. Guinea Worm (Dracunculus medinensis)

1st 2/3rds of animal is thin and hairlike, last one third is thick; an ancient disease
animal resembles a whip
a water borne parasite
found throughout the world, especially tropics
the juvenile lives in small aquatic crustaceans (Cyclops)
almosts 1 Billion humans infected
the adult only lives in humans
other species infect many other kinds of vertebrates
drinking water with infected crustaceans or juvenile worms
lives in large intestine in it

after mating eggs are passes with feces lives beneath skin

can release 7000 eggs/day up to 3’ long

eggs must lie for 3 weeks in soil before becoming infective lives under the skin and creates ulcerations

eggs swallowed the worm creates ulcerations and emerges painfully through the
skin to release its eggs into the water (up to 1.5 M
juveniles in intestine mature into adult eggs/day

new data shows the eggs are triggered to hatch by gut
bacteria
à prevents them hatching in stomach
adults feed on living tissue resulting in more severe symptoms:
providing clean drinking water has greatly reduced incidence of
the disease
the disease has almost been eliminated throughout the world
except for Southern Sudan

normal: diarrhoea, abdominal pain, nausea
heavy infection may lead to intestinal bleeding and anemia
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9
in 1986 there were 3.5 M cases of the disease in Africa and Asia
in 2012 only 542 cases were reported
on track to become only the 2nd human disease after smallpox to
be completely eliminated from the planet
29 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 30

5. Other Human Impacts used to study: genetics, nervous physiology, cell

physiology, aging, etc.
a. evidence is mounting that some parasitic infections
may have benefit in decreasing allergies and we know:
asthma
its complete wiring diagram of nervous system
à it dampens an overactive immune system origin and embryological lineage of all 959 cells making up
its body
proteins secreted by some parasites dampen our
immunity entire genome of 19,820 genes has been mapped

eg. 2007-a study of 1600 vietnamese children infected

with hookworm only 60% were allergic to dust mites

helminths are able to survive in hosts because

they can suppress the host’s immune system

light infections of flukes and other helminths are

used to control allergies and some
autoimmune diseases

nematodes are often able to suppress the immune

system of the host to produce a more
favorable environment

b. some parasitic nematodes show promise as

biological controls against insect crop pests

c. Caenorhabditis elegans important biological

“model” in research (one of a handful; eg. lab rat,
fruit fly, E. coli, etc
Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 31 Animals: Phylum Nematoda; Ziser Lecture Notes; 2015.9 32
 Phylum Acanthocephala proboscis with rows of recurved hooks is used
(spiny-headed worms) to pierce the gut wall of the final host for
attachment
1150 species the size and arrangement of these hooks and spines are used to
identify species
all are parasites in intestines of fish, birds & mammals
proboscis can be retracted into a specialized pouch
body wormlike, somewhat flattened
show eutely (constant number of cells/species
most are less than a couple inches long; up to 1 M
long Body Wall

especially common in the intestines of freshwater syncytial epidermis

fishes
secretes thin cuticle on outer surface
arthropods are intermediate hosts
covered in crypts 4-6 microns deep
one of the most completely parasitic organisms in
the animal kingdom lacunar system of fluid filled canals in body wall

bands of circular and longitudinal muscle fibers

à the digestive tract is completely lost

à only the reproductive system is well developed body wall muscles tube-like and filled with fluids

Body Form body cavity a large pseudocoelom

long, unsegmented, wormlike body with eversible Digestion

proboscis at front end
no digestive system in larva or adult
look like roundworms but with retractable
proboscis exchange of gasses, nutrients and wastes across body
Animals: Phylum Acanthocephala; Ziser Lecture Notes, 2015.10 1 Animals: Phylum Acanthocephala; Ziser Lecture Notes, 2015.10 2

wall by diffusion
gonads contained within thin walled sacs
Respiration
males with pair of testes and penis
no respiratory system
also contain a conspicuous cement gland
absorbs oxygen through tegument
secretes ‘cement’ into the vagina after mating
Circulation which hardens into a plug which prevents
other males from copulating with her
no heart or circulatory system
sperm ejected into vagina and end up in
à but muscle contractions pump fluids through pseudocoel
canals and within body cavity
female has no well defined ovaries, instead the
Excretion ovary fragments as she matures forming
numerous “ovarian balls” that float freely in
if present, excretory system consists of 2 the fluid filled ovarian sac
protonephridia
fertilized egg develops in pseudocoel of female
Nervous System
mature eggs enter the uterus
nervous system with central ganglion near saclike
proboscis receptacle shelled embryo is released in host’s feces
up to 100,000’s/day
but overall greatly reduced
eggs hatch when ingested by intermediate host
Reproduction
Life Cycle
dioecious; females larger than males
requires two hosts to complete its life cycle:
intertnal fertilization
Animals: Phylum Acanthocephala; Ziser Lecture Notes, 2015.10 3 Animals: Phylum Acanthocephala; Ziser Lecture Notes, 2015.10 4

Adult
endoparasite in vertebrates: especially fish, birds
and mammals
also in amphibians and reptiles
only occasional, rare reports of infections in humans
(children)
can be several 1000 in a single host
attach to intestine by spiny proboscis
a host may contain 1000’s of worms
Juvenile
the juvenile cannot complete development and
infect a vertebrate without first going through
an intermediate host
in arthropods (crustaceans & insects)
usually an insect for a terrestrial host
usually a crustacean for an aqutic host
arthropods eat feces with eggs to get infected
in intermediate host the juvenile hatches and
bores through the intestine and into the fluid
Animals: Phylum Acanthocephala; Ziser Lecture Notes, 2015.10
filled body cavity where it completes
development
life cycle may include several “transport hosts” to
complete its life cycle
eg. crustaceanà snailà fish
juvenile can modify the insects behavior to make
it more likely to be eaten by final host
eg. adult in birds (ducks); larva in amphipods
Amphipods (fw crustaceans) typically hide in dark
vegetation during the day to avoid predation
when infected with acanthocephalan worm,
amphipod becomes highly photophilic and
conspicuous near water’s surface
Evolution
recent molecular evidence indicates that
acanthocephala are highly derived rotifers and
should be placed in that phylum
5 Animals: Phylum Acanthocephala; Ziser Lecture Notes, 2015.10 6
 Phylum Rotifera
(Rotifers, Wheel Animals) most aquatic forms are benthic or interstitial fauna

2200 living species some are planktonic

very few fossils have been found – since few hard some are epizoic or parasitic
parts
a few are sessile, living inside tubes or gelatinous
a group of microscopic animals discovered when secretions
microscopes were first being developed
a few rotifers are colonial
1st described in 1696 by Rev John Harris as “an animal like a
large maggot which could contract itself into a spherical most rotifers are 0.1-0.5 mm; but some up to 3 mm
figure and then stretch itself out again; the end of its tail long
appeared with a forceps like that of an earwig.”

von Leewenhoek himself describe a few species in the early some even smaller than some protozoa
1700’s
most are transparent, a few are brightly colored
“wheel bearers” = characteristic ciliated crown
= corona; resembles rotating wheels
great diversity in shape within the phylum
of all invertebrates, the rotifers are most characteristic
à somewhat correlated with their mode of life:
of freshwater habitats
floaters à globular and saclike
most species are common in freshwaters
creepers & swimmers à elongated and wormlike

some (<5%) are found in other habitats: sessile à vaselike

marine or brackish waters some species are cosmopolitan with a worldwide

distribution
a few are terrestrial, in moist soils &
damp mosses & lichens others with very restricted ranges
Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 1 Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 2

Body Form in some: cuticle is very thick, caselike,

fibrous layer
body consists of: head, trunk, and foot
= lorica
head
ciliated corona (or crown) foot
narrow, often tapered, with 1 to 4 toes
gives impression of spinning wheel
often arranged in plates or rings
often with sensory bristles or papillae
often teloscopically retractile (external
used both for feeding segmentation)

draws a vortex of water into mouth for foot is an attachment organ

feeding
toes contain cement glands (=pedal glands)
mouth is inside corona
pedal glands secrete adhesive material
corona can also be used for locomotion
used by both sessile and creeping forms
trunk
foot reduced in pelagic forms
the trunk forms the major part of the body
many species secrete a wide variety of protective
encloses most of the internal organs tubes in which they live:

often elongated or sac-like gelatinous

often has sensory antennae constructed of small pieces of debris

often with false segmentation visible sometimes colored yellow, green or brown
Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 3 Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 4
eutely as in nematodes eating anything small enough to fit in their mouths

Body Wall many are carnivores: feed on protozoa and small

animals
syncytial epidermis (= hypodermis)
some are cannibalistic
secretes thin flexible cuticle
others feed on dead organic matter
in some species some or all of the cuticle is
thickened into a rigid casing (= lorica) most feed by sweeping particles into mouth with
corona
bands of muscles below epidermis
can filter 100,000 x’s its own volume/hr

some circular; some longitudinal à rotifers are sometimes used in fish tanks to clear up
water clouded by organic particles
body cavity a fluid filled pseudocoelom
some are predatory and seek out their prey;
amoeboid cells circulate in fluid probably by touch or chemical stimuli

Movement complete digestive tract

some swim using coronal cilia which pull the animal inside mouth food is directed to a uniquely modified
through the water when unattached pharynx called a mastax that is constantly
working back and forth
some with creeping or leechlike movement
lots of variation in size & shape
some sessile and remain attached to the substrate
operated by bands of muscles
Feeding & Digestion
contains hard chitinous jaws (=trophi) that
most rotifers are omnivores suck in and grind up food
Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 5 Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 6

Excretion
the pharynx leads to the esophagus which opens into
the stomach pair of protonephridial tubes with flame cells

tubes empty into bladder, then to cloaca

both digestion and absorption occurs in the
stomach Nervous System

short intestine leads from stomach to anus in foot bilobed brain dorsal to mastax

some species of rotifers have symbiotic zoochlorellae sends pairs of nerves to sense organs and viscera
within the cells of the stomach wall
senses:
Respiration
eye spots (up to 5)
rotifers have no specific organs for respiration
sensory bristles especiall around the corona
àtheir small size allows them to exchange (touch)
respiratory gasses by diffusion across their
body wall sensory pits lined with cilia in head region

rotifers have a relatively high metabolic rate conpared papillae

to other aquatic invertebrates
usuall 2 pairs of short dorsal antennae
generally have high O2 requirements
Life Cycle
others are capable of withstanding anaerobic or near
anaerobic conditions for extended periods rotifers are notable in that they have a much faster
metabolism than many other cold blooded
eg some live in anoxic mud or in sewage treatment plant
filters
invertebrates

they can grow and reproduce very quickly

Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 7 Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 8
 they can grow from an egg to a reproducing adult in 18 hrs
rotifers even have poor ability to regenerate parts
rotifers can have up to 40 generations/yr
sexual reproduction predominates
in some species, females undergo
cyclomorphosis rotifers are dioecious

=a cyclical change in form of offspring however, females predominate in most

throughout the year populations

rotifer species are also noted for variations in size in some species males are unknown
and appearance in different habitats
males, when present, are minute, degenerate and/or
many rotifers are quite tolerant to drying or short-lived
desiccation (=anhydrobiosis)
commonly only ~1/3rd as long as females
under harsh conditions some can cease
metabolism and dehydrate in some species males are only found for a
few weeks each year
some rotifer eggs can also withstand drying
in others males are degenerate (no digestive
can survive for years (up to 9) and then be tract)
rehydrated and active within a few hours
males are ready for mating within an hour
some can form true cysts that are even more after hatching
resistant to extreme cold and heat
rotifers have internal fertilization
Reproduction
the male uses sensory receptors on its corona
asexual reproduction does not play a major role in this to find a female
phylum as it does in most other animal phyla
Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 9 Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 10

the male attaches its penis to the coronal if a female with mictic eggs is not impregnated
region of the female and transfers sperm she immediately lays the eggs and they
in to her body cavity hatch as males

the male dies soon afterwards resting (winter) eggs:

in some species the males are unknown heavy and thick shelled

à females can reproduce by parthenogenesis overwinter on sediment

recent research has shown that the DNA of these rotifers is
loaded with genes from bacteria, fungi and plants
extremely resistant to drying and extreme
temperatures
this apparently provides adequate mutations to shuffle
genes as an alternative to sexual reproduction in some rotifers, the eggs are retained in the female
until they hatch (ovoviviparous)
in some species, females can produce different
kinds of eggs:
à rotifers bear live young
amictic eggs:
Ecology
diploid eggs produced by parthenogenesis
rotifers are an important part of aquatic foodchains
produced during most of the year
particularly planktonic food chains
mictic eggs:
à reproduce quickly into large populations
haploid eggs
important as food for larger zooplankton
and fish in aquatic ecosystems
capable of being fertilized by male sperm
rotifers affect the species composition of algae in
produced only at certain times of the year
ecosystems through their choice in grazing
Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 11 Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 12
they also compete for the same food sources of
microcrustaceans common in the same habitats

also many species contribute to the decomposition of

organic matter in the soil

Evolutionary Relationships

while rotifers are typically considered with the

pseudocoelomate phyla, new molecular evidence
indicates that they are more closely related to
coelomates than pseudocoelomates

Animals: Phylum Rotifera; Ziser Lecture Notes, 2015.10 13

 Phylum Micrognathozoa cilia on the head create a current that directs food
particles toward the mouth
phylum is represented by a single species collected
from Greenland in 1994 inside the mouth are a complex jaw apparatus

not formally described until 2000 the jaws are made up of up to 15 elements each
controlled by separate muscle strands
tiny interstitial animal, found among mosses eating
diatoms, bacteria and other algae mouth leads to simple gut then to anus

appears to prefer cold temperatures Nervous System

Body large cerebral ganglion in head with a pair of ventral

nerve cords extending to the tail
2 part head
with patches of cilia the whole body has scattered sensory bristles made of
cilia
thorax
resembles sensory bristles of gnathostomulida
abdomen
Excretion
Movement
2 pair of protonephridia
moves using cilia located over the entire body
cilia are arranged in discreet patches or groupings Reproduction

Digestion only females are known and apparently reproduce by

parthenogenesis
digestive system consists of a mouth, pharynx,
esophagus and gut recent study indicates that males could be present for
short periods
Animals: Phylum Micrognathozoa; Ziser Lecture Notes; 2012.9 2

other studies indicate that the animals are

protandrous

the young hatch as males then become females as

they grow older

females produce 2 kinds of eggs; summer eggs and

winter eggs somewhat similar to those of rotifers

Evolutionary Relationships

the complex jaw structure links the rotifers,

gnathostomulids and micrognathozoa

the acanthocephala are also genetically related to

these groups, though because of their specialized
life style as parasites their jaws are greatly
reduced.

Animals: Phylum Micrognathozoa; Ziser Lecture Notes; 2012.9 3

 Phylum Gnathostomulida
(jaw worms) Body Wall

100 species; no known fossils ciliated epidermis used to crawl between sand grains
epithelium is not quite the same as planarians à 1 cilium/cell
à probably many more living species
longitudinal muscle loose in parenchyma in body wall
discovered in 1928 Baltic Sea
acoelomate body – no body cavity
not formally described until 1956
Digestive System
almost all are marine
feed on bacteria, fungi and protists
living in anoxic sand and mud sediments in
shallow coastal waters the ventral mouth is found just behind the head

members of interstitial fauna (meiofauna) pharynx not eversible

between sand grains
pharynx with strong “jaws” made of cuticle
densities up to 3000 jaw worms/lb of sand

all are microscopic sometimes with small teeth

à most less than 1mm long jaws supplied with strong muscles

slender to threadlike worms with transparent body
some have a distinct head, trunk and tail
phylum characterized by distinctive forceps-like jaws
the mouth opens into a blind ended tube (intestine) in
which digestion takes place
no anus but may have small anal pores connecing the
intestine to the epidermis

thought they were small turbellarians until 1969 no circulatory or respiratory system
Animals: Phylum Gnathostomulida; Ziser Lecture Notes; 2015.10 1 Animals: Phylum Gnathostomulida; Ziser Lecture Notes; 2015.10 2

have simple brain and nerve net

the only identifiable sense organs are modified cilia

especiall found in the head region

excretory system consists of protonephridia

but
have solenocytes, not flame bulbs in protonephridia

Reproduction

all are monoecious (hermaphrodites)

gonads are not well defined

egg and sperm in parenchyma

no gonopore

one egg matures at a time

internal fertilization

some species are parthenogenetic

some species are protandric

direct development, no larval stage

Animals: Phylum Gnathostomulida; Ziser Lecture Notes; 2015.10 3

 Phylum Nemertea
(ribbon worms or proboscis worms) larger species use muscular waves to crawl

translates as “unerring one” some swim by undulating body

à refers to animals ability to very accurately shoot its long
proboscis to capture prey some species construct mucus or cellophane-like tubes
(formerly, Rhynchocoela or Nemertini)
others curl up into balls and secrete a covering of
~1150 species mucus around themselves
3 fossils might be nemerteans
almost all are free living, benthic marine worms
first described in the 1700’s; until 1850’s considered
platyhelminthes found in every ocean; from surface to abyssal
zone
share many basic characteristics
most are benthic; a few pelagic species
slender, very fragile ribbon-like worms
à resemble tangled mass of slimy string fairly common on beaches; often found inside
shells, under stones or in tangles of algae;
most < ~8” (20 cm) some burrow into mud and sand

but longest animal in existence is a nemertean arctic ribbon worms commonly wash ashore by the
billions
Lineus longissimus = 197’ (60 M) long
about a dozen species are found in freshwaters
regardless of length most are <1” (23mm) wide
about a dozen species are terrestrial in moist tropical
often very brightly colored soils
some white, red, yellow, green, purple
a very few are parasitic
most use cilia to glide on a trail of slime
Body Plan
Animals: Phylum Nemertea; Ziser Lecture Notes; 2015.9 1 Animals: Phylum Nemertea; Ziser Lecture Notes; 2015.9 2

proboscis (pharynx) is unusual because it is not

general body plan similar to platyhelminthes: connected to GVC in most

triploblastic, acoelomate, bilaterally symmetrical have complete digestive tract with anus
à one way path from mouth to anus
ciliated epidermis
few with rhabdites
like snakes they can devour animals larger than
themselves
Feeding
without food some can live up to 1 yr by self digesting
carnivores like planarians
eat earthworms, sea worms, small mollusks and any
small soft bodied animal, clams and crabs most can shrink at will to < 1/3 their ordinary length
use eversible proboscis (pharynx) to capture prey
Circulatory System
proboscis everted and retracted by muscle bundles
true blood vascular system
in rhynchocoel (a space around proboscis)
Nervous System
often at front of proboscis is sharp pointed spear-like
stylet that impales prey
have a 4 lobed “brain” and two nerve cords extending
down the length of the body
often tipped with poison
nervous system is similar to but somewhat more
sometimes impales with such force that it breaks
elaborate and complex than flatworms
off
writhes for hours
Excretion
animal can grow a new one
excretory system of protonephridia with flame cells
in some the proboscis is a sticky lasso that coils
like flatworms
around its prey
Reproduction
Animals: Phylum Nemertea; Ziser Lecture Notes; 2015.9 3 Animals: Phylum Nemertea; Ziser Lecture Notes; 2015.9 4

Asexual
some reproduce asexually
fragmentation
some can break into 100’s of fragments
makes them difficult to collect whole
each fragment can grow into a complete worm
regeneration
some with great powers of regeneration
Sexual
most are dioecious (unlike flatworms)
fertilized egg develops into ciliated larva
some fragment in warm weather and reproduce
sexually in colder weather
Ecology
nemerteans have few predators: a few bottom feeding
fish, some sea birds, a few invertebrates such as
horseshoe crabs and other nemerteans
Animals: Phylum Nemertea; Ziser Lecture Notes; 2015.9
nemertean skin secrete toxins that deter predators
some crabs “clean” the skin with one claw before
eating them
Evolutionary Relationships
molecular evidence shows them more closely related
to lophophorates than to turbellaria
eg. Cerebratulus
best known example of phylum
may reach lengths up to 2 meters
eg. Prosoma
tiny worm
is common in freshwater ponds
Human Impacts of Nemerteans
one species found near San Francisco is blamed for
the collapse of the dungeness crab fishery
it eats about half of the egg production of the crab
population
other coastal nemerteans have devastated clam beds
In South America a couple of species are sold as fish
bait (locals call them “tapeworms”)
5 Animals: Phylum Nemertea; Ziser Lecture Notes; 2015.9 6
 Phylum Acoelomorpha
sexual reproduction: most are monoecious with
group formerly placed in with flatworms internal fertilization

small flattened worms, <2 mm

all are aquatic; in marine or brackish waters

most are benthic and live in sediments as interstitial

fauna; a few are pelagic

most are free living; some are symbiotic or parasitic

Like flatworms:
3 three tissue layers
double layer of muscles around body
ciliated epidermis
gut sac-like or absent

no well defined digestive system

amoeboid cells function as endoderm to take in

food

very diffuse nervous system

not ladderlike like in flatworms

à resembles nerve net of cnidaria

sense organs: ocelli & statocysts

asexual reproduction by fragmentation

Animals: Phylum Acoelomorpha; Ziser Lecture Notes, 2015.9 1 Animals: Phylum Acoelomorpha; Ziser Lecture Notes, 2015.9 2
 “Mesozoa”
larvae are parasitic in the body spaces of molluscs and
100 species polychaete worms, brittle stars and other
invertebrates
small, worm-like parasites of marine invertebrates
the larvae reproduce asexually by fragmentation
once considered the “missing link” between protozoa
and animals the larva causes destruction of the hosts gonads

unclear if they are degenerate flatworms or truly at some point sexual reproduction is triggered and the
primitive animals larval parasites becomes male or female

recent molecular analysis shows that this group should the males are smaller than the females
be classified into two separate phyla
the males and females leave the host and mate
small wormlike animals (.5-.7MM)
each egg developes into a ciliated larva that makes its
most are parasitic; others are free-living way to another host

made of only 20-30 cells arranged in 2 layers Subphylum Rhombozoa

but no true tissues or organs one group is commensal in kidneys of squid, octopi,
and cuttlefish
ciliated “epidermis”
each species of cephalopod has its own species of
only structures resembling organs are gonads mesozoan

have complex, but poorly understood life cycle only a few millimeters long with 20-30 cells

Subphylum Orthonectida long central reproductive cell (axial cell)

Animals: Mesozoa; Ziser Lecture Notes 2016.9 1 Animals: Mesozoa; Ziser Lecture Notes 2016.9 2

the axial cell can develop asexually into wormlike

juveniles

or it may produce eggs and sperm that self-fertilize to

produce a top-shaped, ciliated larva that leaves
the host and finds a new host

typically it will reproduce asexually until the hosts

kidneys get too crowded then it will reproduce
sexually

there are 3 genera in this group

Evolutionary Relationships

controversial,

some possibilities:

1. very primitive metazoans

may be evolutionary intermediates between protozoa and
metazoa

but from a different stock than choanoflagellates that

are thought to have given rise to animals

2. share ancestry with platyhelminthes

degenerate flatworms due to parasitic lifestyle endoderm
may be mesoderm derived

DNA analysis supports this idea

Animals: Mesozoa; Ziser Lecture Notes 2016.9 3

 Acoelomate Animals à organs are embedded in tissue,
not in any body space (like us)
several phyla including Platyhelminthes share the
following characteristics: like cnidaria and ctenophora

1. have 3 true embryonic tissue layers à only 1 “internal space” =digestive cavity
(=triploblastic):
ectoderm 4. in terms of development these organisms are
mesoderm protostomes
endoderm

between epidermis and digestive cavity is filled à mouth develops first in embryo during
with a 3rd tissue layer = mesoderm gastrulation

à mesoderm allows development of muscle 5. most have bilateral symmetry

layers in body wall
such design allows for a “front end”
à mesoderm allows more elaborate organs = cephalization
more specialization and greater division of
labor than in Cnidaria head contains sense organs, simple brain

2. have true organs was a major new design

each organ is a combination of several tissues à more efficient search for food, mates, etc
specialized for a particular function most animals before this were sedentary filter feeders
sponges have various specialized cells but no true
tissues or organs

jellyfish and corals have 2 true tissue layers and a

few simple organs

3. acoelomate=without body cavity

Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 1 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 2

Phylum Platyhelminthes
diverse array of parasitic species that parasitize
(Flatworms) members of virtually every other animal phylum

[helminth = worm] more specialization and division of labor among

greater variety of tissues and organs
simplest phylum at “organ level” of complexity
acoelomate = no body cavity around digestive
very diverse group but most are poorly known system

includes flatworms, flukes, tapeworms have three true tissue layers (primary germ
layers) = triploblastic
25,000 living species, few fossils à no hard parts
embryonic adult
tissues tissues
poorly known in fossil record but possible trails have ectoderm à epidermis
been found from 565MY mesoderm à parenchyma
endoderm à gastrodermis
may be first animal to have a head & tail
may be first animal to have bilateral symmetry mesoderm makes more elaborate organs possible
may be first animal to show directed movement
à it differentiates into different kinds of
wide variety of body forms all with bilateral symmetry muscle layers

body is usually elongated & slender, leaf-like or organ systems are better developed
long & ribbon-like
only major phylum that is mostly parasitic species
à flattened body allows them to still rely on
diffusion for exchange of gasses, nutrients of 4 classes, 3 are made up of entirely parasitic
and wastes species

Body Wall
range in size from few mm à 10 M long
A. Epidermis
free living in ocean and freshwater habitats, moist soil
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 3 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 4
 free living forms have single layer of ciliated cells
eversible in some à can be extended to find food
parasitic forms have syncytial layer that lacks
cilia secretes enzymes to partially digest food before
“eating” then suck in liquified food
B. Muscle Layers
once ingested enzymes are secreted into GVC
two layers around body wall:
mostly extracellular digestion
circular muscle some intracellular after phagocytosis
longitudinal muscle
Respiration
no rigid skeleton for muscles to act on
no respiratory system
thick muscle layers in pharynx (=feeding tube)
make it “prehensile” flatworms have high surface/volume ratio

in some primitive species these muscle cells gas exchange through flattened body wall
resemble the epitheliomuscular cells of cnidarians

Feeding & Digestion Nervous System

free living forms are mainly carnivores beginnings of cephalization

most species are endoparasites ie. at least some members have distinct head

incomplete digestive tract in most flatworms were probably the first creatures to
have a “brain”
in some planarians digestive tract is highly
branched to distribute food throughout the animal head with cephalic ganglia (~ simple brain)
some parasites (eg. tapeworms) completely lack a
digestive system have pair of ventral nerve cords

pharynx = muscular “throatlike” tube connected by ladder like interconnections

Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 5 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 6

with true synapses between nerve cells

some times produces a chain of zooids
Excretion
à superficial resemblance to segmentation
they are the simplest major phylum with an excretory
system regeneration

some wastes like ammonia are eliminated by diffusion flatworms have considerable powers of
through the body wall regeneration

others have primitive excretory system replacement of lost parts

= protonephridia (tube closed at one end and
exiting body at other end) also to recover from long food shortage

à in most takes the form of “flame cells” budding

cupshaped area with tuft of flagella beat of flagella tapeworms bud off proglottids (reproductive
resemble candle flame under microscope sacs)
wastes and excess water diffuse into bulb
polyembryony
flagella create current to send wastes through tube
which opens to outside of the body flukes
Reproduction
one egg can produce 100’s of larvae
many reproduce both sexually and asexually
increases chances finding a host
Asexual
Sexual
fission
almost all flatworms are monoecious
(hermaphrodites)
pinch in half
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 7 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 8
 cross fertilization not self fertilization Classification

some with internal fertilization Class Turbellaria (planarians)

free living
ventral mouth
locomotion by creeping on cilia and mucous

Class Monogenea
ectoparasites on skin or gills of fish
simple life cycle – no intermediate host

Class Trematoda (flukes)

all are parasitic
endoparasitic flukes
parasites in blood or digestive system
up to 5 developmental stages in 2 or more hosts

Class Cestoda (tapeworms)

all are parasitic
endoparasites in intestine
bud long chains of proglottids

Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 9 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 10

Class Turbellaria Feeding & Digestion

(Planarians)
incomplete digestive tract in most
almost all are free living
àmouth, no anus
mostly bottom dwelling aquatic forms
some have more than one mouth & more than one
mostly freshwater pharynx
some are marine
a few are even terrestrial (6 sp in US) mainly carnivorous
flattened, slender, ribbonlike or leaflike bodies
feed on small crustacea, nematodes, rotifers,
insects
à still rely on diffusion for much exchange of
gasses, nutrients and wastes
can detect food at a distance by chemoreceptors
often brightly colored
entangle their prey in mucous
some marine forms have warning coloration
wraps its body around prey
Body Covering
some marine flatworms prey on molluscs using a
neurotoxin produced by symbiotic bacteria
epidermis is ciliated
they engulf the whole animal or cover the opening of its
secretes mucous trail and uses cilia to glide on it shell then produce the toxin to kill it

land planarians can glide ~6’/hr pharynx = muscular “throatlike” tube

contains rhabdites extend prehensile proboscis to get prey

discharge into water
swell and form protective mucous sheet around body secretes enzymes to help “predigest” prey
may also release toxins to subdue prey and escape
predators sucks up bits of prey
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 11 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 12
in intestine secrete enzymes which further digest prey auricles also contain chemoreceptors

phagocytic cells in gastrodermis take up bits of pieces some turbellaria have statocysts for reacting to
of prey and complete digestion intracellularly gravity

GVC extends to most parts of body Excretory System

à also serves as distribution system
protonephridia with “flame cells”
undigested food is egested through mouth
in freshwater forms this is mainly a way to get rid of
Nervous System & Senses excess water

distinct head with cephalic ganglia metabolic wastes still excreted through body wall

pair of ventral nerve cords connected by ladder- reduced or absent in marine forms
like interconnections
Reproduction
sense organs concentrated on head (vision, smell,
touch, taste) Asexual Reproduction

paired sense organs allow brain to discern the a. Regeneration

direction of the stimulus
turbellarians have considerable powers of
2 eyespots (=ocelli) regeneration
can’t form images, only detect light àreplacement of lost parts
auricles contain tactile cells
àalso to recover from long food shortage
tactile cells are also distributed over body
some can survive for months by self
surface
digesting up to 90% of their body
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 13 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 14

cross fertilization not self fertilization

early researchers also reported learning by cannibalism

taught planarian to run a maze

some with internal fertilization
cut in half and allowed to regenerate
both new worms learned the maze quicker some with vaginas & penises - usually open
through common genital pore
à has never been repeated???

in some monoecious flatworms mating ritual resembles a fight:

b. transverse fission the male organ consists of two dagger-like penises.
during mating, two flatworms “penis fence”
pinch in half: anterior end splits from each trying to get penis in genital pore of the other
posterior end
some without vagina or genital pore use hypodermic
impregnation:
sometimes produces a chain of zooids each tries to stab the other with its penis while trying to
avoid getting stabbed by the other
the one who gets stabbed absorbs the sperm and
à superficial resemblance to segmentation fertilizes its eggs

c. fragmentation fertilized egg is enclosed in cocoon which is

attached by stalks to underside of stones or
when alarmed, some land planarians can plants
break up into dozens of “blobs of slime”
most have no larval stage
in a few hours each piece will become a
new worm embryos emerge as juveniles that grow
into adults
Sexual Reproduction
a few marine species produce planula-like larva
most are monoecious (hermaphrodites) (=Mullers larva)

during breeding season each individual develops àciliated ball of cells (has 8 ciliated lobes)
both male and female organs

Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 15 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 16
 Examples of Turbellarians eg. some marine planarians have symbiotic
zoochlorella (algae) or feed on algae
eg. Dugesia
common in fw streams
some marine forms also have “kleptonematocysts”
eg. Phagocata
they eat cnidarian polyps and keep the
up to 20 pharynxes each with a mouth stinging cells to use for defense
eg Terricola = land planarians

fraction of an inch to almost 1’ long

creatures of dark or dim light

no eyes, 2 eyes or 100’s of eyes

hunters and scavengers

some terrestrial planarians are fast enough to

catch fruit flies (Drosophila)

eg. one Brazilian species pursues earthworms into their

burrows

enfolds it

mouth exudes digestive enzymes that liquifies it

then sucks up liquified worm

eg. Bipalium

terrestrial planarian common here

it also can eat small earthworms

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Animal Parasites Costs to Parasite:

the other classes of Platyhelminths contain only host is a small “discontinuous” habitat
parasitic species parasite must locate and infect new hosts to
propagate its species
Parasitism à most common form of symbiosis
must be able to overcome hosts defenses:
20-50% of all animal species are parasitic inflammation
immune response
1/4th of all animal families are parasites
but can’t kill host
ectoparasites
à parasite lives on outside of host à the most successful parasites do as little
some can use gut for food storage and expand harm as possible to their hosts
to many times their normal size
eg. leeches, ticks, fleas Some Typical Endoparasitic Adaptations

endoparasite 1. Structures for penetration and attachment to

à parasite lives on the inside of host host
digestive system often very simple or gone hooks, suckers, teeth, enzymes
altogether most common point of entry to host is through mouth
eg. tapeworms, flukes, roundworms
2. Usually have a resistant stage in life cycle
Benefits to Endoparasitic Lifestyle:
for getting from one host to another
which is often in a different kind of environment
constant environment if endoparasite - needs to survive trip through digestive system

gets easy access to food 3. Reduction in “unnecessary” structures

protection from predators reduced sense organs

reduced nervous system
reduced locomotion
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 19 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 20
 reduced digestive system
some endoparasites have lost gut entirely
some ectoparasites use gut mainly for food storage
(eg. leeches, ticks)
4. Tendency toward being Hermaphrodite
only need any two, not male and female
some can even self fertilize if necessaryà but usually don’t
5. Enhancement of reproductive capacity
host is a small “discontinuous” habitat
à need extraordinary powers of reproduction to insure
survival
reproductive organs are often the largest, most apparent organ
systems present
often able to produce of large #’s of eggs
Liver fluke(F. hepatica) à 20,000 eggs/day
Ascaris à 200,000 eggs/day
Tapeworm (Diphyllobothrium)
à 1M eggs/day for 15 years
(=5.5 trillion eggs/lifetime)
6. Use of intermediate larval stages on
intermediate hosts
à to enhance chances of getting to final host
Even with large numbers of eggs chances of success are
relatively small
eg. F. Hepatica
in most favorable situation
3-4 out of 20,000/day will actually
hatch
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10
larvae (microfilariae) move to peripheral blood
on periodic basis
corresponds to “biting hours” of local
vector (flies & mosquitoes)
eg. Guinea worm
(nematode: Dracunculus medinensis)
occur in tropical areas; lots of rice fields
eggs must be laid in water to be able to get to its
intermediate host
female may contain up to 1 Million eggs
each with a developing larva inside
larvae must be released in water to
complete life cycle
to do this female moves to part of body
likely to be immersed in water
à lower legs
creates an ulcer
at moment limb enters water the female
protrudes anus and discharges 1000’s of
infective larvae
3. Host Modifying Behaviors
an alternative to modifying the parasites own
behavior is to alter the hosts behavior to
make it more likely to complete parasites
life cycle
eg. Echinococcus (Tapeworm)
sheep infected with hydatid cysts lags behind healthier
members of heard à more easily caught and
eaten by coyote
b. Conspicuous Behavior
eg. Acanthocephalans: adult in birds (ducks); larva in
amphipods (small aquatic crustaceans)
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10
simplest life cycle:
adult parasite à eggs à ingestion by new host
more complex life cycle:
adult parasite à eggs à intermediate host à
definitive host
most complex life cycle:
flukes have several intermediate states that reproduce
7. Behavioral Adaptations
behavior is an important tool for animal survival
this is also true for parasites: behavior can be
used to enhance their chances for success
Examples:
1. Simple host finding behaviors
eg. Entobdella (Monogenea)
skin parasite of a stingray
eggs are released and settle to bottom
larvae emerge from eggs within 3 seconds of sudden
darkness
then swim vertically upwards
2. Periodic Behaviors
parasite keys in on cyclic stimulus
eg. Filarial Worms
live in blood
transmitted by mosquito or fly
21 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 22
Amphipods (fw crustaceans) typically hide in dark
vegetation during the day to avoid predation
when infected with acanthocephalan worm which as
adult infects birds, became highly photophilic and
conspicuous
eg. burrowing clams infected with a fluke,
rather than burrowing into sediment,
remain closer to surface where they are more likely to
be preyed upon
eg. Fluke (Leucochloridium)
adult in birds; larva in snail
when infected, snails tend to crawl to tips of
vegetation instead of hiding like normal
in snail, larvae migrate to tentacles of snail
larvae are brightly colored with red and green bands
they pulsate
makes snails very conspicuous in daytime
at night the larvae withdraw into the snails body
eg. Sacculina
one of best adapted parasites known
Sacculina is a highly modified barnacle that has
become a parasite of crabs
as it matures it sheds all appendages, becomes an oval
sac and penetrates a crab host
develops an extensive system of branches extending
into every appendage
a saclike growth appears under the crabs abdomen
where eggs and sperm form (Sacculina is a
hermaphrodite)
the crabs metabolism is completely altered:
if crab is female:
changes are not as extensive but egg
development is inhibited
if crab is a male:
body assumes shape of a female
23 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 24
 reduced length of some segments persistent poverty
broadening of abdomen
testes reduced or converted to ovaries decreased productivity

à both male and female resemble mature female poorer birth outcomes
bearing eggs: physically and behaviorally
decreased cognitive abilities, epilepsy and neurological disorders
Helminths malnutrition & anorexia

The most common human endoparasites are flukes,

tapeworms (classes of Platyhelminths) and
roundworms

These three are grouped together as “helminths”

(=parasitic worms)

over 25% (1.5-2 Billion) of the world’s population is

infected at any one time

that’s orders of magnitude greater infection rate

as HIV

yet proportionally, helminth diseases receive only

a small fraction of the research dollars

as such most are listed as “neglected tropical

diseases”

over 135,000 die each year from helminth infections

helminth infections also have considerable effects on:

overall health and tissue damage & inflammation, anemias

Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 25 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 26

Class Trematoda they feed aggressively on body fluids

can clog ducts and trigger gall stones and excessive enlargement
over 3x’s more species than any of the other classes of liver
of flatworms
Body Wall
class includes some of our most serious parasites
body covered by thin flexible cuticle
almost all are endoparasitic
protects from hosts digestive enzymes
adults mainly found in vertebrates
integument is syncytium (not divided into individual
cells) with no cilia
leaf-like body shape (=flukes)
muscle layers are embedded in “tegument”
two suckers for attachment:
Feeding & Digestion
one around mouth
the other further down on the body
like turbellaria, they have well developed, incomplete
digestive tract
adults & larvae inhabit a wide variety of sites in
hosts:
digestive tract àbut with mouth at anterior end
respiratory tract
circulatory system gut usually divided into two branches
urinary system
reproductive system
some dissolved nutrients can also be absorbed directly
flukes tend to inflict greater harm to their hosts than through skin
do tapeworms
excretory and nervous systems similar to planarians
flukes can be found in veins of intestines, bladder, bile ducts
and lungs
Reproduction
in high numbers they can cause blockages and damage
most are monoecious and capable of self fertilization
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 transforms into …

sporocyst
trematodes typically have a complex life cycle with sporocyst reproduces asexually (polyembryony)
1 or more larvae occurring in intermediate hosts yeilds more sporocysts or…
and adults in definitive host
redia
also reproduces asexualy
adults are typically parasites of fish or other produces more redia or…
vertebrates
cercaria
emerge from snail
1 to 5 larvae occur in intermediate hosts, usually a penetrate second intermediate host
mollusc or encyst in vegetaton to become…

metacercaria
polyembryony occurs usually in several larval these are juvenile flukes
stages allowing a single egg to develop into when eaten by definitive host develop into adult fluke
100’s of potential adults
Some larval flukes are able to change hosts behavior
typical life cycle: to make them more likely to get to final host:
eg. burrowing clams infected with a fluke,
adult
rather than burrowing into sediment,
remain closer to surface where they are more likely to
metacercaria egg be eaten

eg. Dicrocoelium (Trematode, Fluke)

cercaria miracidium
has 3 host life cycle:
adult in vertebrate, eg sheep
redia sporocyst 1st larva in terrestrial snail
2nd larva in ants
eggs released by host are eaten by snail
egg cercariae emerge entangled in slime of snail forming a
usually passes in feces sticky ball
must reach water to develop ants eat these slimy balls
ants behavior changes so they are more conspicuous
miracidium and more likely to be eaten by vertebrate:
free swimming larva
penetrates tissue of snail à when infected they crawl up blade of grass;
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 29 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 30

seize the grass in their jaws and remain Examples of Flukes

there until the next morning
sheep are early grazers and eat the ant
eg. Clonorchis sinensis (Chinese Liver Fluke)
eg. Fluke (Leucochloridium)
adult in birds; larva in snail the most important human liver fluke
when infected, snails tend to crawl to tips of
vegetation instead of hiding like normal
in snail, larvae migrate to tentacles of snail one of most complex life cycles:
larvae are brightly colored with red and green bands
they pulsate primary host and two intermediate hosts
makes snails very conspicuous
at night the larvae withdraw into the snails body
during the day they are easy prey for birds 5 larval stages

serious problem in China, Asia, Japan

humans are final host (also cats, dogs, pigs)

in which the adult lives

transmitted to humans by eating raw fish

Adult

10-20 mm long
oval, with 2 ventral suckers
simple digestive sacs
hermaphrodite
almost 80% of body is devoted to
reproduction

flukes mature in intestine then move to bile ducts

in liver

typically lives 15-30 years (up to 50 yrs)

Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 31 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 32

light infection are asymptomatic or may
produce light abdominal pain
heavy infection (up to 20,000) can cause liver
damage, cirrhosis and death
1000’s of eggs released each day in feces into
water
eggs can survive weeks in water
if snail eats egg miracidium larva hatches from
egg and developes into sporocyst then redia
then cercaria
through polyembryony, a single egg can produce
250,000 cercaria
cercariae burst out of snail and burrow into fish
and encyst in muscle as metacercariae
if fish is eaten by mammal the metacercaria cyst
dissolves in intestine and fluke moves to liver
of their adult host
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10
eg. Schistosoma (blood fluke, schistosomiasis)
doesn’t occur in US; but >400,000 immigrants
are infected
one of the worlds major helminth infections
à affects 200M worldwide:
esp Africa, S. America, Mid East, Far East
150,000 die each year from direct effects of
the parasite; especially renal failure
another 200,000 die from related causes
snail is intermediate host, humans are final host
for 3 mainly human species
other species of Schistosoma infect birds and
mammals
mature adults live in portal vein of liver, feeding
on blood
differ from most other flukes by being
dioecious (separate sexed)
males larger
females smaller, stay in groove (=gynecophoric canal) in
males body
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10
eg. Fasciola hepatica (Sheep Liver Fluke)
adult lives in liver and bile ducts of liver of sheep,
other ruminants and can live in humans
feed on blood
can produce up to 50,000 eggs/day for several
years
eggs passed in feces
if pasture is wet the eggs hatch into free living
miracidium
miracidium ingested by snail or penetrates land
snail and becomes sporocyst, then redia,
then cercaria
through polyembryony, a single miracidium larva can
produce up to 4000 cercaria larvae
cercaria leaves snail and encysts as
metacercaria on vegetation
when vegetation is eaten by sheep or
ruminants, the adult hatches and moves to
liver
33 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 34
female penetrates wall of blood vessel in liver and
release eggs into bile ducts
eggs move from bile ducts to intestine and
passed in feces
eggs might also enter bladder and be released in urine
many eggs can lodge in liver and cause
abdominal pain, fever and bloody diarrhea, ulcerations,
etc
eggs may also be carried to lungs causing inflammation
if eggs reach water they hatch into ciliated
miracidium
have only a few hours to find snail host or they die
if eaten by snail they develop into sporocyst,
then cercaria (no redia stage)
if humans are in contaminated water:
cercariae bore directly thru skin to get into
blood
cercaria are one of few parasites that can bore through skin
rice farmers are easily infected
in North America some blood flukes of birds may
atempt to bore into humans
they don’t survive and cannot infect us
35 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 36
 but their burrowing and death in skin can eg. Paragonimus (Lung fluke)
cause swimmers itch
lives in lungs of host

many mammals are hosts

found in East Asia, SW Pacific and parts of S.

America

eggs coughed up, swallowed then eliminated in

feces

metacercaria develop in fw crabs

infection is acquired by eating uncooked crab meat

infection causes breathing difficulties and chronic

cough

fatalities are common

one species found in N America infects minks with

its larvae in crayfish

only 1 human case reported

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Class Monogenea
egg hatches into ciliated larva
A group of small (rarely >.75”) mostly ectoparasitic
flatworms larva and adult have large posterior attachment organ
with hooks
widespread and common

1000’s of species

once placed with trematodes

all are parasites

àmainly ectoparasites on gills or skin of fish

some species infect other cold blooded animals

one species infects the eye of hippos, no other warmblooded
birds or mammals

a few are found in urinary bladder of frogs and turtles

have anterior and posterior attachment organs

seem to cause little damage to their hosts

all are hermaphrodites

direct life cycle with single host

they have no intermediate hosts

Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 39 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 40
 Class Cestoda (Tapeworms)
proglottids bud off the scolex
>1000 species
(a chain of proglottids = strobila)
most specialized class of flatworms
bud from scolex with oldest ones furthest
all are endoparasites away

à not same as segmentation since each proglottid acts

can grow up to 10 M (30’) long as an individual animal

1991: doctors removed 37’ tapeworm from Mississippi woman

max tapeworm length ever recorded is over 90’ taken from some individuals can produce a dozen
a sperm whales intestine proglottids/day
adult can live up to 20 years
some tapeworms have up to 3000 proglottids
Body Plan
Body Wall
very different from other classes of flatworms
tegument is syncytial (not subdivided into separate
cells) with microvilli (microtrichs) to increase
àno head
surface area for absorption
front end of the animal is not a head, it’s a
tegument secretes a protective cuticle
special organ for attachment (=scolex)
no external cilia
à has suckers and hooks
well developed muscle layers beneath body wall
use for attachment, not for feeding or sensing
the environment
Feeding & Digestion
à“body” consists of a long chain of reproductive
completely lack digestive system
sacs = proglottids
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 41 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 42

à absorb food through skin (=tegument)

typically the intermediate host is the prey of the
Nervous System adult host

simple nervous system all are monoecious (hermaphrodites)

proglottids are united by nerve cords, the most mature proglottids are those furthest away
from the scolex
but no special sense organs
unlike most hermaphrodites, tapeworm proglottids can
Excretion cross fertilize in same animal

somewhat similar to other flatworms testes usually develop before ovaries so generally can self-
fertilize a single proglottid

protonephridia continuous throughout proglottids eggs or mature proglottids are shed in feces

Reproduction & Life Cycles once egg is released must be ingested by intermediate
host (another vertebrate)
each proglottid acts as “individual”
à usually a vertebrate “prey” of a final host
à any two proglottids can exchange sperm
once eggs ingested larva hatches and bores through
when gravid each proglottid may contain up to intestines of host and into blood
100,000 eggs each
travels to skeletal muscle, heart and other organ
Life Cycle
secretes a protective cyst
almost all tapeworms require at least 2 hosts; mainly
vertebrates in some, cyst develops into a “bladder-worm”
or cysticercus
but same host can bear either the adult or the
larval parasite
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 43 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 44
humans can get infected with eggs by unsanitary
habits with feces, not washing hands, kissing pets,
etc
but humans make poor intermediate hosts from
the parasites perspective
à nothing eats them
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10
larva burrows through intestine and into blood
reach skeletal muscles where they encyst as
bladderworms
= “measly meat”
In US infections are not uncommon:
34 M cattle in US; ~1% of US cattle are infected
20% not federally inspected
1/4th of the infections are missed in inspection of
meat
à 5 in 10,000 or ~150,000 in US infected
humans become infected by eating ‘rare’ or poorly
cooked roast beef, steaks and barbecues
when measly meat is eaten bladderworm hatches
and adult develops in intestine of final host
takes 2-3 weeks for mature worm to develop
if just a few the infection will be mild or
asymptomatic
light infections may cause weight loss
à “diet pills”
heavier infections may cause diarrhea and
vomiting
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10
Examples of Tapeworms
eg. Beef Tapeworms (Taenia saginata)
>50 Million infection worldwide; especially South
America, SE Asia, Africa
adult in human intestine
mature adult may reach 10 M (30 ft) or more
scolex buries itself in intestinal wall
à has 4 suckers to attach (no hooks)
can bud over 2000 proglottids
numerous proglottids are released each day:
gravid proglottids break off and pass with
feces
sometimes they crawl out anus
they crawl out of feces into nearby vegetation
proglottids dry and release eggs
à can remain viable on grass up to 5 months
picked up by grazing cattle
when eaten by cattle the eggs hatch
45 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 46
eg. Pork Tapeworm (Taenia solium)
pork tapeworm is more dangerous to humans
since the larval stage can more easily develop
in humans
adults usually live in human small intestine
juvenile in muscles of pigs
adult can live up to 10 years and grow to 10’ long
generally doesn’t occur in US but thrives in Mexico
and Central America
occasionally shows up along border.
WHO estimates that 2.5 Million are infected with
adult worm and many more with larvae
worldwide
scolex has hooks and suckers
life cycle is similar to beef tapeworm
each proglottid can release 50,000 eggs
eggs eaten by pigs and larva migrates to skeletal
muscles
humans usually infected by eating poorly cooked
pork and adult develops in intestine
47 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 48
 eg. Echinococcus (dog tapeworm)

if humans ingest eggs rather than the larva about 1 Million are infected worldwide.

the eggs will develop into a bladderworm that one of the most dangerous tapeworms
encysts in body tissues
a group called “tissue tapeworms”
=cysticercosis
adult is very small: only a few mm
can cause serious problems by lodging in:
adults occur in dogs, coyotes, wolves and other
eyes à blindness
canines
brain à neurological symptoms or death
juvenile develops in >40 species of mammals
muscle à pain and weakness, inflammation (eg. monkeys, sheep, reindeer, cattle)
and other visceral organs including humans

treatment usually involves surgery sheep infected with juvenile lag behind healthier
members of heard à more easily caught and
eaten by coyote

humans can become intermediate hosts by

fecal-oral route; eg. kissing pets
(humans are dead end choice for parasite since few
eat humans)

once ingested, juvenile moves to various

tissues; eg. liver, lungs, brain

juvenile stage is special kind of cysticercus

= hydatid cyst

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eg. Fish Tapeworm (Diphyllobothrium latum)

grows quickly; 1 cm/month, for up to 20 years humans and other animals are definitive host

may be no symptoms for years ocurs wherever fish are an important food
source and the water supply is easily
can reach size of basketball contaminated with sewage

à up to 4 gallons endemic in Europe, Asia, US & Canada

within main cyst daughter cysts bud off 2 intermediate hosts: copepods & fish

each daughter cyst contains 1000’s of in humans, adult attaches to intestinal lining by
scolices scolex (no hooks)

symptoms and signs depend on the cyst’s eggs are released in feces
location and size
if feces enters water eggs may be eaten by tiny
in humans, growth of cyst can cause damage to crustacean, copepod
organ
fish eats copepod and bladderworm encysts in fish
if cyst ruptures the fluid itself can produce muscle
anaphylactic shock, even death
if fish are improperly cooked, or eaten raw (sushi)
only treatment is surgical removal the infection is transferred to humans

thorough cooking or freezing (-10º C) for 24

hrs kills the parasite

Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 51 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 52
eg. Diphylidium caninum Human Costs of Parasitic Flatworms
adult in small intestine of dog or cat 250-300 Million people worldwide are infected with
some type of parasitic flatworm
up to 6” long
(some put that number much higher)
fleas are intermediate host
results in Billions of dollars in healthcare costs and lost
fleas eat tapeworm eggs released in pet feces productivity

egg hatches and encysts in flea also affects livestock and pets

dog eats fleas and bladderworm hatches into

adult Beneficial Effects of Parasitic
Flatworms
1. weight loss

light infections of adult tapeworms cause little

damage and may cause a loss in weight

à larvae once sold as weight loss pills in US

today the treatment is only available in Mexico,

the approximate cost is $1500.

tapeworm infestation can result in a loss of one to

two pounds per week.

once the target weight loss is reached, a

deworming agent is given
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 53 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 54

However, Tapeworm infestation can result in:

à as hygiene has improved allergies, asthma and
The formation of cysts in the liver, eyes, brain, other autoimmune diseases have dramatically
and spinal cord with potentially lethal increased
consequences.
in developed countries, where improved sanitation
if pork tapeworm much more serious has largely eliminated helminth infections
there has been an increase in such
intestinal blockages
autoimmune disease in the past 100 years

malnutrition these same diseases are rare in poor countries

where intestinal parasites are endemic
2. Helminth Therapy
there is a large “underground market” in helminth
a type of immunotherapy to treat autoimmune parasites fueled by these findings – medicine is
diseases and immune disorders by deliberately just now catching up
infecting patient with intestinal parasites

research has found that intestinal parasites,

particularly roundworms have the ability to temper
the immune system and prevent the overreactions
that cause allergies, asthma, ulcerative colitis,
Chron’s disease, etc

helminthes have thrived in mammals for millions of

years

over that time they have adapted to survive the

onslaught of the hosts immune responses to the
infection

“Hygiene Hypothesis”
Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 55 Animals - Animal Phyla: Phylum Platyhelminthes; Ziser Lecture Notes, 2015.10 56
 Phylum Ctenophora (Comb Jellies,
sea walnuts) a few are compressed and elongated ribbon
shapes; some of these are up to 7’ long
ctenophore = “to bear a comb”
some are pink, orange, olive
~150 species
all are bioluminescent; resemble translucent glowing
include comb jellies and sea walnuts balloons in the surf

first described in late 1600’s as a kind of jellyfish à flashes at night or when prodded

only designated as a unique phylum of animals in most are planktonic

1800’s
some species are common in coastal waters
apparently an ancient group
many occur in very large “schools”
515 M yr old fossils have been found in China
Stromatoveris Texas species sometimes wash up onto beach in large
numbers called “sea snot”
à clearly related to Ediacaran creatures of 542 MY
ago a few are pelagic, esp in deeper waters
new (2014) research has found that this group is the oldest
animal group; even older than sponges some wormlike forms are benthic and creep along
the sediment
all are marine
like Cnidaria, at “tissue” level of complexity
most spherical and transparent; .5” – 2” diameter
no stinging cells; instead have adhesive cells
not bell or polyp shaped as cnidaria (=colloblasts) for getting food

no polymorphism chief identifying feature is 8 rows of comb plates of

cilia extend from mouth to aboral end
Animals: Phylum Ctenophora; Ziser Lecture Notes, 2015.9 1 Animals: Phylum Ctenophora; Ziser Lecture Notes, 2015.9 2

are biradial Feeding

2 long tentacles in most most are carnivores

a few simple organs they eat small crustaceans and other small
zooplankton
Body Wall
no nematocysts
similar to Cnidaria
instead have 2 long tentacles with colloblasts
outer epidermis, inner gastrodermis
=adhesive cells and lasso cells
gelatinous layer of collenchyme (similar but not
some tentacles are relatively long
identical to mesoglea of cnidaria)
eg. Pleurobranchia ~1/2 “ diameter has 6” tentacles
Support and Locomotion
surface of body is sometimes covered with papillae
use vertical ciliated plates (=combs) for locomotion containing colloblasts and suckers

in some the cilia are 200 x’s the length of cilia in other animals eg. Leucotheca

à each comb beats in succession like a wave
starting at aboral (upper) end
a few elongated forms crawl on bottom
comb jellies apparently have muscle layers derived
from mesoderm
a trait not found in sponges or cnidaria but in
higher organism like flatworms
Animals: Phylum Ctenophora; Ziser Lecture Notes, 2015.9
colloblasts are stimulated by movement of prey
some pursue their prey
others cast out tentacles like fishing nets
as tentacles accumulate prey they are wiped
across the mouth and food is extracted
a few are parasitic
3 Animals: Phylum Ctenophora; Ziser Lecture Notes, 2015.9 4
comb jellies have a complete digestive tract: some benthic forms reproduce by fragmentation

most have pharynx (throat) inside mouth sexual reproduction

GVC branches throughout body into jelly layer all are hermaphrodites

has tiny “anal canals” that expel undigested in most sperm and egg discharge through mouth
materials
external fertilization
Excretion
unique larval form = cydippid

have specialized excretory cells (= rosette cells) some northern species are able to reproduce
which line GVC while still larvae

they extend cilia into mesoglea to collect wastes perhaps due to predator pressure

Nervous System Ecology

unlike cnidarians’ nerve net, comb jellies have a more midlevel in food-chain
sophisticated nervous system with a rudimentary
brain and synapses for cell to cell communication they eat plankton

primary sense organ is statocyst for balance in turn eaten by some jellyfish and sea turtles

Classification
Reproduction
Class: Nada
asexual reproduction
no tentacles

remarkable powers of regeneration eg. Beroe

any half can regenerate Class: Tentacula

Animals: Phylum Ctenophora; Ziser Lecture Notes, 2015.9 5 Animals: Phylum Ctenophora; Ziser Lecture Notes, 2015.9 6

tentacles

diverse forms

Phylogeny

origin is obscure

probably arose from radially symmetrical planula-like

ancestor

however, new (2013) genetic research indicates that they

are the earliest animal group to evolve, even
before the sponges

that would mean that comb jellies evolved nerves,

muscles, and other complex features that
were later lost in sponges and cnidaria

may be only surviving remnant of ancient ediacaran

biota (635-542 MY ago)

Human Impacts

sometimes cause serious bioinvasions

eg. introduced by cargo ships from North America into the Black
Sea

ate all zooplankton à fish starved

in 10 years caused complete collapse of fisheries

Animals: Phylum Ctenophora; Ziser Lecture Notes, 2015.9 7

 Coral Reefs
sponges and corals are familiar to most people as main
components of coral reefs

Reefs are unique ecosystems:

1. their structure is created by biological activity

massive deposits of calcium carbonate

 esp by corals
 also coralline algae, mollusks, and a few
other groups

2. reefs are the largest biological structures on the

earth

3. coral reef communities have survived for 1000’s

or 10,000’s of years

relatives of corals appeared over 500 MY ago

 mainly solitary individuals

modern colonial reef building forms appeared

and diversified in the last 25 M years
eg. one reef (Eniwetok) is ~ 4000’ thick (1283 m)
and estimated to be over 60 M years old

oldest reefs are in the pacific; youngest in the atlantic

(10-15,000 years old)

Where are Coral Reefs

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 1

individual corals are found in all oceans from the poles
to the equator

but coral reefs are only found in warm, clear

equatorial waters

 waters >68º F (20º C)

tropical reefs are most common in the western Pacific

and Indian Oceans

coral reefs cover 0.1% of earth’s surface area

all kinds of reefs cover 1.5 M sq miles (=568,600 km2 )

(the most productive shallow water reefs, ie. reefs in <30m

of water, cover ~0.75 M sq. miles (=284,300 km2 )
= area ~ size of Italy)

Coral Reef Diversity

coral reef communities are the most luxuriant,

complex and diverse of all aquatic communities

while they are dominated by coral species practically

all animal phyla are represented

an abundance of sponges, clams, snails, worms,

fish, eels, sea stars, sea urchins, shrimp, crab, etc

also seaweeds, algae, bacteria, protists, etc

coral reefs contain about 200,000 known species

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 2
 (~15% of all species)

tropical rainforests
6% of earth’s surface; 14 M sq mi, support ~50% of all species

coral reefs
0.1% of earth’s surface; 1.5 M sq mi, support ~15% of all species

Diversity per unit area: coral reefs are 400-500 times more
diverse than rain forests

 but estimates range from 600,000 to 9 M

species worldwide
eg. 32 of the 34 animal phyla are found on coral reefs
compared to only 9 of the 34 found in the rainforests

eg. >1/4th of all marine fish species are associated with

coral reefs

many are brightly colored

numerous symbioses occur between reef organisms

yet most reefs grow in areas of ocean with fewest

nutrients

clear clean water = nutrient poor water

Reef Requirements & Structure

to become established a reef has some essential

requirements:

1. hard substrate
Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 3
 initial growth requires a hard surface (firm base)
on which to start construction

reef forming organisms are mainly sessile,

benthic animals

= animals that live in or on a substrate

(don’t swim in open ocean)

2. warm tropical temperature

reef communities are also restricted by water

temperature

 most occur only in tropical and

subtropical seas (±30º latitude)
where average water temperature ~23º - 25º C

none are found below 18º C

few on W coast of N America or Africa due to upwelling of
cold water

3. shallow

most reefs grow depths of 75 ft (25 M) or less

limit is 50-70 M

they are therefore restricted to coastal areas or

seamounts
Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 4
 most reef building corals contain symbiotic
photosynthetic algae that require sunlight

 form basis of reef food chain

not too deep (to 60M)

 light is quickly filtered out
 depth of active reef is restricted by light penetration

the growth and health of the coral community is

directly dependent on the amount of light
reaching the reef

4. salinity near 33ppt

normal salinity of sea water

 can’t withstand lower salt concentrations
eg. don’t see any near E coast of S America because of
outflow of Amazon River

5. clear

reef organisms require clear waters to allow their

photosynthesis

 low amounts of dissolved materials and few

nutrients
 not at mouth’s of large rivers

if the water is shallow, but murky (turbid) sunlight

will not get through for photosynthesis

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 5

 also, too much sediment will smother the polyps
another reason why they are not usually found near outlets
to large rivers

6. Prefer areas with Strong Wave Action

wave action oxygenates waters, brings in

nutrients, and reduces sedimentation

Established Reefs

coral colonies form the main framework of a reef

 may be over 100 species of corals alone

the coral colonies are able to extract calcium

carbonate from sea water to form the reef
structure

 they use sugar produced by the algae that

live inside their tissues to do this

 without the algae the corals cannot grow

most reef building corals contain symbiotic

photosynthetic algae (=zooxanthellae)
present in enormous populations
provides a vital energy source for the reef organisms
base of reef food chain

this symbiosis is beneficial to both organisms:

corals algae
Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 6
 provide CO2 provide O2
N, P remove wastes
make organic nutrients

some corals also have symbiotic nitrogen fixing

cyanobacteria

most corals are hermaphrodites

take 7-10 years to reach sexual maturity

but they can reproduce asexually much more

quickly

many reef animals are colonial

virtually all the nutrients the algae create are

cycled to corals and the rest of the food web

 very efficient, short, direct, quick nutrient

cycling

 prevents nutrients from sinking out of

productive sun lit zone

once established, reefs create their own environment:

numerous crevasses and holes provide excellent

hiding places

 create numerous habitats

any exposed surface created when organisms die,

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 7
 is quickly attacked by boring organisms
especially sponges. worms and clams

as organisms live and die get build up of coral

skeletons, encrusting algae, shells, etc

waves also break up and destroy old reef material

fine materials settles into crevasses and holes

 fills spaces
 cements reef together

reef ecosystems are characterized by:

a. high diversity

 lots of competition especially for space &

food
eg. algae, sponges and corals are constantly growing
over and killing each other

eg. most reef fish are very localized with specific feeding
preferences

eg. reef fish even differ between day and night

b. rapid recycling of nutrients (similar to

rainforests)

(produce several times more organic material/area

than phytoplankton communities)

c. numerous symbioses and interactions

eg. zooxanthellae, sponge symbionts, crabs, molluscs, etc
Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 8
 eg. one common characteristic of many reef organisms is
mass spawning events

 100’s of species synchronize their reproduction

the extensive vertical growth of reefs is the result of

changes in sea level

 virtually all modern reefs have grown

upward due to recent sea-level rise
beginning ~18,000 BP
(3-15 M (10-40’)/1000yrs)

 some of the thickness may also be due to subsistence

(especially at atolls and some barrier reefs)

Kinds of Reefs

Two general types of reefs:

1. Fringing Reefs or Barrier Reefs

most common type
surround islands and border continents
grow in shallow waters and border the coast closely or may be
separated by a shallow stretch of water
project seaward directly from shore
subdivided into several zones:
reef crest – part of reef the waves break over
forereef – medium energy
buttress (spur & groove) – rows of coral with sandy
canyons or passages between rows

eg. Great Barrier Reef is longest in world ~1000 miles

2. Atolls
Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 9
 at summits of submerged volcanoes (seamounts)
usually circular or oval with a central lagoon

Reef Zonation

Both reef types show similarities in profile

(vertical zonation)

these differences due mainly to differences in wave

energy and water depth

a. Reef Face
seaward side
inclined from gentle to steep slopes
often with terraces creating more zonation

10-20M: high energy – help to dissipate wave energy

(30-60’) grooves drain off sand
masses of large dome shaped and columnar
corals, large fish

20-30M: little wave energy

(65-100’) only 25% of surface light reaches here
more delicately branched corals

30-40M: gentler slope

(100-130’) very reduced light
sediments accumulate here
corals become patchy

>50M: slope drops off sharply

(>165’)

b. Reef Crest
highest point of reef front
exposed at low tide, covered by waves at high tide
elkhorn coral and shelf coral

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 10

 c. Reef Flat (back reef)
sheltered, lagoon side
highly variable
short to several 100 meters
lowest energy, coral sand
delicate corals, eg. staghorn
becomes shallower and supports sea grasses

Reef communities are characterized by a coordinated

reproductive frenzy at specific times of the year
often late spring: “spawning”

 one species after another will discharge reddish

clouds of eggs and milky white sperm into the
water

 described as an underwater ‘snowstorm’

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 11

Economic Impacts of Coral Reefs:

reef communities have significant impacts on human

economies and activities:
tourists
coastal protection
fisheries
pharmaceuticals

1. Fisheries

eg. worldwide, coral reefs provide 1/4th of the annual

commercial fish catch and feed over 1 Bil people in asia
alone.

eg. US reefs support millions of jobs and a $200 M annual

fishery

on global basis
1/2 sq mile of reef:
 can sustainably yield 15 tonnes of fish
and other seafood/yr
 $8.6 M in revenue/yr

2. Tourism

eg. reefs of the florida keys generate $1.2 Bil/yr in tourist

dollars

3. Biochemicals

many marine animals produce biologically active

compounds
the earliest known use of marine resources was
Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 12
 for medical uses:
2700 BC – China – medical text

scientists have extracted over 20,000 new

biochemicals from marine life, mainly from
coral reef organisms over past 20 yrs(04)

perhaps 10% of all marine organism could yield

medically important compounds

since the greatest marine diversity is in coral

reefs, they offer the greatest possibilities for
potential uses

scientists first began looking at softbodied sessile organisms of

coral reefs because they thrived under highly competitive
conditions with no apparent claws, teeth, etc for defense

 must use chemical weapons

Some examples:

a. Sponges
antibiotics, antitumor drugs, antifungal drugs

eg. Acyclovir
from Caribbean sponge
1s t antiviral compound approved for human use
fights herpes infections
used since 1982

eg. Vidabarine
may attack AIDS virus

eg. a species of S Pacific sponge produces chemicals

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 13
 that can kill Candida  a human pathogen that causes
thrush and vaginal infections

b. Corals
antiinflammatories, painkillers for arthritis,
antimicrobials

cardiac stimulant from sea anemone

c. Segmented Worms
eg. Padan – a powerful insecticide produced from a
polychaete worm
eg. dolastatins from sea hare (Dolabella auricularia) has
potential anticancer properties

d. Snails & Other Molluscs

muscle relaxants, painkillers
adhesives

e. Bryozoa
potent anticancer chemicals

f. Tunicates
antiviral, antitumor
including possible treatment for malignant
melanoma
 the most dangerous form of skin cancer

by some estimates, coral reefs provide over $30 Billion

in benefits (direct and indirect) , worldwide per
year

destroying 1/2 sq mi of reef costs $137,000-$1.2 M in

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 14
 loss of fisheries, tourism and shoreline protection
over a 25 year period.

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 15

Threats to Reefs:

Coral Reefs are among the most endangered

ecosystems in the world

recent (‘03) assessments of world’s reefs show they are

globally threatened

 there are no “pristine” reefs left:

all reefs are impacted by human activities
only reefs in remote areas are generally healthy

 30% of reefs are damaged

up to 30% have been lost in last 50 years(06)
another 16% are severely damaged

 60% may be completely dead by 2030

generally, coral reefs are very resilient

 have existed for 1000’s to 100,000’s of years

but today are being degraded in a matter of

decades

the greatest threats to reefs are from human activities

eg. ~1/2 of world’s population live in coastal regions

eg. in SE Asia, 70% of population is in coastal areas

Coral Bleaching

one of earliest signs of stress is coral bleaching

 when water gets too warm algae “flee” their

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 16
 coral hosts
therefore lose their color

triggered by disease, pollution, elevated

temperatures, salinity changes, increased UV
radiation, etc

bleaching is a normal response to short term stresses

while bleached, corals stop growing

 leaves reef vulnerable to erosion

after one bout the reef can recover,

but frequent episodes may kill the coral
polyps

what is significant about bleaching today is its

frequency, severity and extent

Coral Reefs are associated with 109 countries,

those in 93 countries show significant damage

reefs at highest risk:

Japan Singapore Taiwan
Sri Lanka India Indonesia
Asia

eg. Phillipines
only 5% of reefs are pristine
30% are dead
39% are still healthy

all are areas with dense coastal populations and

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 17
 heavy coastal development

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 18

Human Causes of Coral Reef Decline:

while natural events, eg diseases and hurricanes

can cause extensive damage to specific reefs

humans are having a global impact on reefs

human causes of reef decline:

1. sedimentation
2. eutrophication
3. shipping and oil spills
4. exploiting for food (overfishing)
5. collecting
6. mining
7. tourism

1. Sedimentation
by far the greatest impact
increase in suspended silt, clay, dirt
mainly due to deforestation esp. mangroves
due to logging, farming, mining, dredging
doesn’t have to occur near coast to have and impact
sediment blankets coral reef
initial plume blocks sunlightreduces photosynthesis
smothers polyps
as they produce mucus to remove it, depletes their
energy reserves; makes them more susceptible to
disease
impedes larval settling

2. Eutrophication
food and nutrients usually limit the growth of most organisms
eg. N & P plants, algae; organics bacteria, heterotrophs
reef ecosystems are especially susceptible since they are found in
nutrient poor waters
too much food can upset the balance between organisms in the
Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 19
 community:
some grow much faster than others and can become
toxic
sometimes a new predator gains upper hand
eg. crown of thorns starfish  can clean out
entire reefs when its predators are eliminated

some algal infestations caused by eutrophication cause

algae to release sugars that fertilize the symbiotic
bacteria making them pathogenic and killing their coral
hosts

3. Shipping and Oil Spills

eg. oil tankers pollute and kill reefs
eg. 1s t gulf war oil release (10M BBL’s)
caused extensive damage to reefs in arabian sea
eg. in Mid East a phosphate tanker ran aground on a reef,
releasing phosphates into the water
killing 500 mi2 of reef

4. Exploiting for Food (overfishing)

reef fish are prone to overfishing because many are slow
growing, long lived fish (K-selected; low natural fertility)
when depleated they are slow to repopulate

historical record shows that over the last several 1000 years,
large fish and animals have been hardest hit of reef
community

blast fishing
use explosives to kill or stun fish
eg ~1/6th of reefs in Phillipines have been damaged this
way since 1945
cyanide fishing
some use cyanide and poisons to fish
 kills other organisms as well
child labor
in Phillipines 40 ships carry 300 children to reef each day
children pound reef with rocks to scare fish into nets
can destroy up to 1 km2 of reef/day
Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 20
 children killed by needlefish, sharks, barracuda,
poisonous snakes, etc

as fish become more scarce, fishermen earn extra income

collecting turtles, clams, etc

5. Collecting
1.5 Million kg’s (15 tonnes; 3M lbs) of coral & shells/year are
harvested
mainly for “shell shops” around the world
~1/3rd from the Phillipines
most is exported
most goes to US gift shops and aquarium shops

live corals were collected and sold in Florida until 1989 when it
was outlawed
but some is still traded on black market

shells etc collected by malacologists: prefer killing live specimens

rather than dead shells from beach

exotic fish collected from reefs feed a $4 Billion/yr aquarium

industry

6. Use as Building Material

in Sri Lanka and parts of India entire sections of reef have been
removed to make cement
there is no other source of rock nearby

7. Tourism (Ecotourism)
walking on reef and touching it kills polyps and kicks up sediment
many break off souvenirs of live reef
beauty of reef stimulates beach front developments

eg. On S Pacific Island of Palau

they mined an area of reef to build a new airport runway
 to accommodate an increasing number of tourists
coming to see the reef

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 21

 eg. in Grand Caymans a 525’ cruise ship dropped a 5 ton anchor
and
dragged its chain across 150M of reef
creating a 3M wide path 150M long
uprooted 8M diameter blocks of coral
destroyed an area 1/2 the size of a football field

Indirect Human Effects:

8. Global Warming
global temperatures are increasing 1/2 – 1 degree every decade
this rate is 100x’s faster than natural rate at end of last
glaciation
most of this accelerated warming is due to human activities
global warming will
alter weather patterns
alter ocean circulation
warm ocean surface waters
cause significant sea level rise up to 6 cm/decade
 but reefs can grow up to 10 cm/decade

9. Ozone Depletion
will continue into next century
ozone levels decrease .5-5% over the tropics
this causes a 1-10% increase in UV radiation

shallow marine communities are particularly susceptible to

damage from this additional radiation

Ecology: Marine Ecology – Coral Reefs; Ziser Lecture Notes, 2006 22

 Phylum Cnidaria
(=Jellyfish & Corals) most members of the phylum (eg. corals) are sessile

9,000 living species, 9,300 fossil species often beautiful and graceful “plant-like” or “flower-
like” forms with one or more rows of large
include: tentacles extending from body
jellyfish
sea anemones like sponges, ancient scholars considered them some kind of
corals plant
sea fans
sea whips not considered animals until 1700’s

another very ancient group with lots of fossil but some (eg. jellyfish) swim weakly as part of the
representatives zooplankton

simplest living animals with true tissues
longest fossil history of any animal
known fossils are even more ancient than
sponges
going back at least 700 M years
though sessile almost all are extremely effective
predators
jellyfish are among the longest of animals
up to 9’ diameter with 120’ tentacles
some colonial forms can grow up to 150’ long

(plenty of hard parts – corals)
st
in terms of evolutionary history they were the 1
animals to appear that had a definite shape
as a colony, they rank as some of the longest – lived
animals on earth
eg. a gold coral colony off the coast of Hawaii was recently
(2009) dated at 2742 years old

radial symmetry eg. a black coral in the same area was dated at 4265 years old

all known animals at that time were sessile all are aquatic
organisms; cnidaria (jellyfish) may have been
the first animals to swim widespread in marine habitats
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 1 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 2

à especially shallow waters, warmer oceans

polyp (=hydroid)
a few found in freshwaters
tubular body
all but 1 species of fw cnidarians are polyps usually sessile – though some can move
upward facing mouth surrounded by tentacles
but there is one small fw jellyfish: Craspedacusta
medusa (=jellyfish)
many are colonial
umbrella shaped
à groups of individuals usually living together and mouth facing downward
interconnected eg corals often, thick jelly-like layer in body wall
à jellyfish
eg. a single coral colony can contain millions of individuals motile: contractions of “bell”
free floating, pelagic planktonic
tissue level of organization
polyp medusa
more complex than sponges but still very simple sessile motile
asexual sexual
do have true tissues benthic pelagic

body wall is made of 2 layers of tissue Cells & Tissues

only a few very simple organs two true tissue layers, not the 3 typical of
animals
Body Forms
=diploblastic
many cnidarians are polymorphic
epidermis & gastrodermis
à with 2 or more separate body forms
2 well defined embryonic layers:
ectoderm
with an alternation between forms endoderm

become two adult tissues

àthe same species has 2 distinct forms epidermis
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 3 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 4
 gastrodermis most epidermal cells contain contractile fibers and
act like muscle cells to produce movement
between the two tissues is a jelly layer called
mesoglea Cells of Epidermis:

à very thick in “jellyfish” a. epitheliomuscular cells

covers outside of body
nontissue layer of mesoglea in between tall T shaped, columnar cells
base elongated with myofibrils
in a few species this mesoglea is replaced by 3rd muscular contractions
true tissue:
b. interstitial cells
mesoderm à connective tissue undifferentiated cells
can form cnidocytes, nerve cells, sex cells, etc
but not epitheliomuscular cells
Body Wall
c. gland cells
epidermis – tissue layer that lines outer surface around basal disc and mouth
secretes mucus and adhesives
those in basal disc can secrete gas bubble for floating
mesoglea – jellylike middle layer; not tissue layer
d. cnidocytes
gastrodermis – tissue layer that lines GVC stinging cells, more later

e. sensory cells
1. Epidermis scattered but especially near mouth and tentacles
respond to chemical and tactile stimuli
outer “skin” of the animal
f. nerve cells
most multipolar (3 or more processes)
consists of cells that cover and protect form synapses with sensory cells and other nerve cells
connect to epitheliomuscular cells and cnidocytes
also contain special stinging cells
2. Mesoglea
some areas also have gland cells for attachment
not really a tissue layer, just a layer of jelly-like
secretions
contain nervous and sensory cells

Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 5 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 6

very thin layer in polyps; much thicker in medusa cavity in which it lives
thus “jellyfish”
muscle layers in body wall contract against
3. Gastrodermis hydrostatic skeleton

inner lining of the digestive sac some polyps of noncolonial forms are motile

mademostly of cells that digest and absorb food eg. fw hydras are not permanently attached
à can glide on pedal disc
à inchworm movements using tentacles
these cells also contain contractile fibers for à gas bubbles and float to surface
movement
medusae are more mobile
Cells of Gastrodermis:
have hydrostatic skeleton
a. nutritive muscular cells
tall T shaped, columnar cells
ciliated nerve net controls contractions of bell for
base elongated with myofibrils swimming
lines GVC
in some freshwater species cells contain green algal
symbionts Feeding and Digestion
in some marine species cells contain dinoflagellate algal
symbionts all are carnivores
b. Interstitial cells
scattered most species have one or more rings of tentacles
transform into other cells as needed, see above surrounding mouth
c. gland cells
in hypostome and scattered throughout armed with cnidocytes (=stinging cells) for
some secrete digestive enzymes capturing prey
mucous glands around mouth

Stinging Cells (cnidocytes)

Movement
one of the most characteristic features of the
typically polyp is sessile and often secretes a cup like
phylum
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 7 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 8
 most are not harmful to humans
used for feeding and defense
eg. most sea anemones stings are harmless
inside each cell is harpoon-like nematocyst
but a few are very painful
à highly coiled tubular thread
eg. Portuguese Man-O-War and some corals
àcontained within a capsule like organelle
à triggerlike cnidocil (tactile trigger) a few can be fatal

when triggered can fire in a fraction of a second eg. cubomedusae (box jellies)

discharge due to: digestive system is a mouth that opens into a saclike
cavity
high osmotic pressure within (140 atm; 10x’s sea level)
when stimulated to discharge water rushes in = gastrovascular cavity lined with gastrodermis
forces thread out with great force – turns inside-out as it
extends at 2m/sec
causes barb to flick out like tiny switchblades to impale prey single opening = mouth

each cell operates independently incomplete digestive tract à mouth only

digestion mostly extracellular, but some intracellular

can differentiate between animate and
inanimate objects most are predatory

à doesn’t just fire at anything use cnidocytes to capture and paralyze prey
use tentacles to move prey toward mouth
cnidoblast must grow new nematocyst after firing engulf prey with mouth

over 20 different kinds inside GVC gland cells secrete digestive enzymes
some wrap around prey or are sticky
nutritive muscular cells take in particles by
some with tiny barbs that impale prey & inject poison pseudopodia

Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 9 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 10

intracellular digestion completes the process at margins of bell

often between lappets
indigestible material is expelled through mouth contains
ocelli à detect light
statocysts à balance organs
sensory pits à chemoreceptors
No respiratory or excretory system
Life Spans
Coordination and Control
little is know of lifespans of cnidaria
no head, no cephalization, no CNS
but one sea anemone kept in an aquarium lived
very simple nervous system, no brain for 80 years until the tank was accidentally
drained
= nerve net
some jellyfish can live up to 10 years
mostly for coordinating contractions in body
Reproduction
diffuse network of nerve fibers connect to:
sensory cells both sexual and asexual reproduction
cnidocytes
epitheliomuscular cells asexual:
nutritive muscular cells asexual reproduction usually by budding

some simple sense organs: if buds remain connected = colonial

statocysts à balance
ocelli à light fission
sea anemones only
polyp pedal laceration
simple sensory cells scattered in epidermis
sexual:
medusae most are dioecious
clusters of sense organs = rhopalium
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 11 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 12
 Classification
gonads are epidermal in hydrozoa
gonads are gastrodermal in other groups
Class: Hydrozoa
most shed gametes into water, often mass most are marine, a few are freshwater
spawnings individuals usually small and inconspicuous
polyp is dominant stage, some completely lack medusa
but one species of box jellyfish actually have a “wedding dance” medusa when present has velum around margin
no septae in GVC, no pharynx (=throat), no cells in mesoglea
begins as male takes hold of female tentacle and pulls her most are colonial - small plant-like appearance
around in the water most have polymorphism with alternation of generations

he then draws her close, male and female become
entwined, so that their manubria touch
male deposits a spermatophore on one of her tentacles and
releases her
the female ingests the spermatophore which then fertilizes
her eggs
embryo in marine species is usually a planula
in many members of the group there is an
alternation of generations between
polyp which reproduces asexually and the
medusae which reproduces sexually
some Cnidaria can switch genders
males require less food and energy and can more easily survive
harsh conditions
Class: Scyphozoa (true jellyfish)
most of the larger jellyfish belong to this group
medusae without velum, cells in mesoglea
all are marine
solitary polyp stage reduced or completely absent
thick jelly layer (=mesoglea)
Class: Cubomedusa (box jellyfish, sea wasps)
cubical jellyfish with extremely potent toxins - some lethal
Class: Anthozoa (Corals and Sea Anemones)
=”flower animals”
all are marine
polyp only; no medusa stage
many cells in mesoglea
polyp with septae and pharynx
some are solitary = sea anemones, usually larger
most are colonial = corals, polyps usually small
most secrete skeleton of calcium carbonate or protein

when conditions improve more will turn into females to insure

their success in producing offspring
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 13 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 14

A. Class Hydrozoa eggs usually mature 1 at a time

eggs fertilized by sperm, then shed
cyst forms around embryo – overwinters
most are marine, a few are freshwater no larval stage
young hydras hatch from cyst in spring
some are colonial
eg. Obelia
common in nearshore marine habitats
most have polymorphism with alternation of
generations more representative of class

have both polyp and medusa stage

polyp is dominant stage
colonial hydrozoan à interconnected hydroid colony
some, eg Hydra, lack medusa stage
attaches to substrate by rootlike hydrorhiza

colonial species often have more than two body forms branching body = hydrocaulus
in same organism
living tissue = coenosarc

different forms act like separate organs and chitinous protective covering = perisarc
are specialized for feeding, stinging,
reproduction attached to hydrocaulus are individual polyps

two types of polyps:

eg. Hydra (hydra)
freshwater species
1. hydranths = feeding polyps
very common in ponds and creeks
tubular or vaselike
feeds on small crustaceans
mouth surrounded by tentacles
seems to “prefer” Daphnia
capture and ingest prey: worms, crustaceans,
no medusa stage
larvae
à polyp reproduces both asexually and sexually
provide nutrition for whole colony
asexual:
à digested broth passes thru common GVC
budding as outpockets of body wall
of whole colony
continuous GVC
cilia of nutritive muscular cells move it
eventually detach
sexual:
2. gonangia = reproductive polyps
dioecious ovaries or testes are temporary
no tentacles
organs on side of polyp
medusa bud off sides
usually appear in autumn (low temp, low O2)
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 15 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 16
 medusae B. Class Scyphozoa (true Jellyfish)
produced by gonangia
small; 2-3mm
velum surrounds inside of bell margin most of the larger jellyfish belong to this group
mouth at end of manubrium a few up to 2 m in diameter
radial canals extend from GVC to margins of bell and
ring canal
GVC also extends into tentacles from ring canal contains one of the longest animals; lion’s mane
jellyfish à over 120’
eg. Craspedacusta

the only freshwater medusa

all are marine
hydroid colony is microscopic in size; <2mm
medusa .5 – 1” in diameter with few or many tentacles around margin of bell
eg. Physalia (Portuguese Man-O-War)
medusa stage is dominant
colonial hydroid form
irridescent purple color solitary polyp stage reduced or absent
common on gulf coast
can produce painful sting even if dead
but with several different kinds of polyps: thick jelly layer (=mesoglea)
float = swimming bell filled with gas may contain amoeboid cells and fibers
gastrozoid polyps = each with single long tentacle
dactylozoids = fishing tentacles
gonophores = sacs of ovaries or testes medusa has no velum

a symbiotic fish, Nomeus, swims among the tentacles Movement

jellyfish are the most motile members of the phylum

more complicated muscle layers:

muscles arranged in radiating and circular bands

contractions of these muscles allow the organism

to propel itself to some degree
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 17 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 18

still, jellyfish are considered part of the zooplankton (the

largest members) since they are not strong enough
swimmers to go against the current
Nervous system & senses

watching some jellyfish swim looks like they’re not since jellyfish are motile their sense organs are
going anywhere better developed than other members in the
phylum
but contractions of bell creates water currents that draw food
through tentacles and toward mouth
à its not ‘trying to go anywhere’ jellyfish in this class have a greater variety of
sense organs than other jellyfish
in a few species the medusa is sessile and spends its
life laying upsidedown on the sediment scalloped margins of bell with indentations bearing
lappets and rhopalia
Feeding
with ocelli, statocysts and sensory pits
mouth hangs down under umbrella on the end of a
throat-like manubrium reproduction & life cycle:

GVC extends into radiating canals or pouches medusa stage is the sexual stage

all jellyfish are carnivorous polyp is asexual stage

buds off small medusa
they eat mostly zooplankton, smaller fish and
other jellyfish jellyfish life cycle:
medusa

larger ones may eat shrimp and other ephyra planula larva
crustaceans
strobila scyphistoma

jellyfish are eaten by spadefish, sunfish and

loggerhead turtules many jellyfish live less than a year

only a few dozen of the 500 or so species of but a few arctic species live up to 10 years
jellyfish are dangerous to humans
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 19 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 20
eg. Aurelia (moon jelly) the most poisonous sea creatures known

common off Texas coast since 1884, it has killed more people along the
northern Australian coast than have sharks in the
7-10 cm diameter; some up to 2 feet area
scalloped margin with indentations bearing lappets and
rhopalium with ocelli, statocysts and sensory pits
mouth on manubrium drawn out into 4 frilly oral arms
feeds on small planktonic organisms

male releases sperm threads into water

female collects and eats them to fertilize eggs

eg. Cannonball jellyfish

eg. Lion’s mane jellyfish

one of largest
can be 8’ in diameter
tentacles can extend to 200’

C. Class Cubozoa (box Jellyfish, sea wasps)

poorly known group

once considered as a group of scyphozoa

medusa is dominant form, polyp is inconspicuous

most are relatively small; <1”

tentacles are at each corner of cubical bell

eg. sea wasp (Chironex fleckeri)

ranges from Indian ocean to coral sea

Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 21 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 22

D. Class Anthozoa (Corals and Sea Anemones) pores in body wall to help catch
prey
=”flower animals”
3 major groups in class:
all are marine 1. sea anemones and stony corals
2. sea fans, sea pansies, sea pens, soft corals
àrange from deep to shallow water 3. tube anemones and thorny corals

some are solitary = sea anemones, usually larger eg. anemones

especially common in tropical waters

most are colonial = corals, polyps usually small
much larger than their coral relatives; some 3’ dia
polyp only; no medusa stage
mostly sessile, but some can glide on pedal disc

GVC large
muscular infolding of mouth = pharynx
GVC partitioned by septa (=mesenteries)
although anemone’s move in slow motion, some group living
species actually battle each other for territory using
specialized tentacles for battle
reproduce by fragmentation; they leave pieces behind as
they move

eg. hard corals (“stony corals”)

mesenteries can be complete or
incomplete colonial polyps

produce rock-like calcareous cups (=theca)

free edge of incomplete septae form
septal filaments with nematocysts secreted by lower half of polyp

in some, lower septal filament prolonged =exoskeleton for support and protection

into acontia form extensive reef structures in warm shallow waters

also with nematocysts
à can be extruded thru mouth or
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9
reef structure consists of compressed & welded together:
calcium carbonate coral skeletons
encrusting coralline algae
23 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 24
 foraminiferan shells Ecological Interactions of Cnidaria
bivalves
sea urchin plates
1. most are aggressive predators
continually destroyed by:
sponges, worms & clams bore into reef eg. a single lions mane jellyfish was found with >200 fish within
waves reduce it to white sand its tentacles
crown of thorns starfish (Acanthaster) feeds on polyps and
decimates populations eg many jellyfish species congregate into large swarms of up to
highly resilient communities à regenerate quickly 1000’s of individuals

can devastate prey in an area

eg. soft corals (octocorallia)
can clog fish nets
secrete a flexible endoskeleton of spicules or keratin-like
protein
swarms often triggered by eutrophication
eg. sea pens, sea pansies, sea fans, whip corals, pipe corals
climate change has caused many swarming species to
sea whips & sea pens
expand their range
eg. tube anemones
2. Some are prey for a variety of specialized
secrete tubes predators
eg. parrot fish, butterfly fish, tangs eat coral polyps

eg. sea turtles like jellyfish

unfortunately many are suffocated by eating plastic bags

floating in the ocean

eg. a number of sea slugs (nudibranchs) eat them but store

unfired nematyocysts in their skin and use them for defense

3. numerous symbioses within this phylum:

eg. The “upside-down jellyfish” is not a predator

it harbors symbiotic algae in its tissues and spends its time
in shallow water laying upside-down ‘sunning’ its algae
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 25 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 26

this community acts in lieu of an immune system to protect

in deeper waters it moves up to the surface during the day the animals which is lacking in most invertebrates,
for photosynthesis including corals

eg. many anthozoa live as commensals on shelled eg. some corals call for help when being smothered by
animals seaweeds
eg. attach to shells of hermit crabs corals compete for light with seaweeds

eg. decorator crabs a new (2012) study found that when a staghorn coral species in
Fiji is being smothered by mats of seaweed it signals gobies
eg. almost every sea anemone is a host to a to come eat the seaweed
variety of fish and other reef animals (shrimp, gobies don’t touch the seaweed unless signalled by the
crab, fish) coral

over 50 species of fish associated with anemones in return gobies get shelter from the coral
(also some shrimp)
5. Coral Reefs
eg. clown fish

fish symbionts are stung on first contact (on tail or nonvital “cold water corals”
body part)
some live below 0ºC
à then body mucous or slime is chemically altered so no light
the fish is not affected by further stings no zooxanthellae for extra nutrients
almost 700 species
must continually refresh mucus layer
“tropical coral reefs”
eg. most corals are mutualistic with dinoflagellate
algae (zooxanthellae)
base of food chain in coral reef communities

eg. most coral species produce large amounts of

slime which harbors a diverse community
of bacteria and other microbes.

Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 27 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 28
 Tropical Coral Reefs
corals are familiar to most people as main components
of coral reefs
Coral Reef Diversity
coral reef communities are the most luxuriant,
complex and diverse of all aquatic communities
while they are dominated by coral species practically
all animal phyla are represented
an abundance of sponges, clams, snails, octopi, squid,
worms, fish, eels, sea stars, sea urchins, shrimp,
crab, etc
also seaweeds, algae, bacteria, protists, etc
each species of life has developed ways to cooperate
and compete within a myriad of nooks and
crannies
reef ecosystems are characterized by high diversity
à lots of competition especially for space & food
eg. algae, sponges and corals are constantly growing
over and killing each other
eg. most reef fish are very localized with specific feeding
preferences
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 29
à also coralline algae, mollusks, and a few
other groups
3. coral reef communities have survived for 1000’s
or 10,000’s of years
relatives of corals appeared over 500 MY ago
à mainly solitary individuals
modern colonial reef building forms appeared
and diversified in the last 25 M years
eg. one reef (Eniwetok) is ~ 4000’ thick (1283 m)
and estimated to be over 60 M years old
oldest reefs are in the pacific; youngest in the atlantic
(10-15,000 years old)
the extensive vertical growth of reefs is the result
of changes in sea level &/or subsistence of the
seabed
à virtually all modern reefs have grown upward due to
recent sea-level rise beginning ~18,000 BP
(rate of 3-15 M (10-40’)/1000yrs)
à some of the thickness may also be due to subsistence
(especially at atolls and some barrier reefs)
Where are Coral Reefs
individual corals are found in all oceans from the poles
to the equator
but coral reefs are only found in warm, clear
equatorial waters
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 31
eg. reef fish even differ between day and night
coral reefs cover 0.1% of earth’s surface area
all kinds of reefs cover 1.5 M sq miles (=568,600 km2)
(the most productive shallow water reefs, ie. reefs in <30m
of water, cover ~0.75 M sq. miles (=284,300 km2)
= area ~ size of Italy)
coral reefs contain about 200,000 known species
(~15% of all species)
tropical rainforests
6% of earth’s surface; 14 M sq mi, support ~50% of all species
coral reefs
0.1% of earth’s surface; 1.5 M sq mi, support ~15% of all species
àDiversity per unit area: coral reefs are 400-500 times more
diverse than rain forests
eg. 32 of the 34 animal phyla are found on coral reefs
compared to only 9 of the 34 found in the rainforests
eg. >1/4th of all marine fish species are associated with
coral reefs
Reefs are unique ecosystems:
1. reefs are the largest biological structures on
the earth
2. their structure is created by biological activity
à they create their own ecosystem
massive deposits of calcium carbonate
à esp by corals
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 30
à waters >68º F (20º C)
tropical reefs are most common in the western Pacific
and Indian Oceans
many are brightly colored
numerous symbioses occur between reef organisms
yet most reefs grow in areas of ocean with fewest
nutrients
àclear clean water = nutrient poor water
Reef Requirements & Structure
to become established a reef has some essential
requirements:
1. hard substrate
initial growth requires a hard surface (firm base)
on which to start construction
reef forming organisms are mainly sessile,
benthic animals
= animals that live in or on a substrate
(don’t swim in open ocean)
2. warm tropical temperature
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 32
 directly dependent on the amount of light
reef communities are also restricted by water reaching the reef
temperature
4. salinity near 33ppt
à most occur only in tropical and
subtropical seas (±30º latitude) normal salinity of sea water
à can’t withstand lower salt concentrations
where average water temperature ~23º - 25º C
eg. don’t see any near E coast of S America because of
none are found below 18º C outflow of Amazon River

few on W coast of N America or Africa due to upwelling of 5. clear

cold water
reef organisms require clear waters to allow their
3. shallow
photosynthesis
most reefs grow depths of 75 ft (25 M) or less
à low amounts of dissolved materials and few
limit is 50-70 M nutrients
à not at mouth’s of large rivers
they are therefore restricted to coastal areas or
seamounts if the water is shallow, but murky (turbid) sunlight
will not get through for photosynthesis
most reef building corals contain symbiotic
photosynthetic algae that require sunlight also, too much sediment will smother the polyps
another reason why they are not usually found near outlets
à form basis of reef food chain to large rivers

not too deep (to 60M) 6. Prefer areas with Strong Wave Action
à light is quickly filtered out
à depth of active reef is restricted by light penetration
wave action oxygenates waters, brings in
the growth and health of the coral community is nutrients, and reduces sedimentation

Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 33 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 34

Established Reefs
àany exposed surface created when organisms die, is quickly
attacked by boring organisms especially sponges. worms
once established, reefs create their own and clams
environment:
àas organisms live and die get build up of coral skeletons,
coral colonies form the main framework of a reef encrusting algae, shells, etc

àwaves also break up and destroy old reef material

à may be over 100 species of corals alone
àfine materials settles into crevasses and holes
the coral colonies are able to extract calcium à fills spaces
carbonate from sea water to form the reef à cements reef together
structure
coral reefs show very rapid recycling of nutrients
à they use sugar produced by the algae that (similar to rainforests)
live inside their tissues to do this
à virtually all the nutrients the algae create are cycled to
à without the algae the corals cannot grow corals and the rest of the food web

most reef building corals contain symbiotic à prevents nutrients from sinking out of productive sun lit
zone and lost
photosynthetic algae (=zooxanthellae)
àproduce several times more organic material/area than
present in enormous populations
phytoplankton communities
provides a vital energy source for the reef organisms
base of reef food chain
reef communities show numerous symbioses and
this symbiosis is beneficial to both organisms: interactions
corals algae eg. zooxanthellae, sponge symbionts, crabs, molluscs, cleaner
provide CO2 provide O2 fish, etc
N, P remove wastes
make organic nutrients a common characteristic of many reef organisms is
some corals also have symbiotic nitrogen fixing mass spawning events
cyanobacteria
most corals are hermaphrodites
ànumerous crevasses and holes provide excellent hiding places
take 7-10 years to reach sexual maturity
à create numerous habitats
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 35 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 36
 corals don’t have complex nervous systems or
sense organs yet many coordinate their
release of sperm and eggs within the same
few evenings each year, sometimes timed to
within minutes of each other
Both reef types show similarities in profile
(vertical zonation)
these differences due mainly to differences in wave
energy and water depth

each year, at late-spring full moon, 100’s of a. Reef Face

coral species simultaneously spawn seaward side
inclined from gentle to steep slopes
à synchronized by a light sensitive pigment in often with terraces creating more zonation
the coral animals
10-20M: high energy – help to dissipate wave energy
(30-60’) grooves drain off sand
Kinds of Reefs masses of large dome shaped and columnar
corals, large fish
Two general types of reefs:
20-30M: little wave energy
(65-100’) only 25% of surface light reaches here
1. Fringing Reefs or Barrier Reefs more delicately branched corals
most common type
surround islands and border continents 30-40M: gentler slope
grow in shallow waters and border the coast closely or may be (100-130’) very reduced light
separated by a shallow stretch of water sediments accumulate here
project seaward directly from shore corals become patchy
subdivided into several zones:
reef crest – part of reef the waves break over >50M: slope drops off sharply
forereef – medium energy (>165’)
buttress (spur & groove) – rows of coral with sandy
canyons or passages between rows b. Reef Crest
highest point of reef front
eg. Great Barrier Reef is longest in world ~1000 miles exposed at low tide, covered by waves at high tide
elkhorn coral and shelf coral
2. Atolls
at summits of submerged volcanoes (seamounts) c. Reef Flat (back reef)
usually circular or oval with a central lagoon sheltered, lagoon side
highly variable
Reef Zonation short to several 100 meters
lowest energy, coral sand
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 37 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 38

delicate corals, eg. staghorn Economic Impacts of Coral Reefs:

becomes shallower and supports sea grasses

Reef communities are characterized by a coordinated reef communities have significant impacts on human
reproductive frenzy at specific times of the year economies and activities:
fisheries
often late spring: “spawning” tourism
coastal protection
à one species after another will discharge reddish pharmaceuticals
clouds of eggs and milky white sperm into the
water 1. Fisheries

eg. worldwide, coral reefs provide 1/4th of the annual

à described as an underwater ‘snowstorm’ commercial fish catch and feed over 1 Bil people in
asia alone.

eg. US reefs support millions of jobs and a $200 M annual

fishery

on global basis
1/2 sq mile of reef:
à can sustainably yield 15 tonnes of fish
and other seafood/yr
à $8.6 M in revenue/yr

2. Tourism
eg. reefs of the florida keys generate $1.2 Bil/yr in tourist
dollars

3. Biochemicals

scientists have extracted over 20,000 new

biochemicals from marine life, mainly from
coral reef organisms over past 20 yrs(04)

Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 39 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 40
 since the greatest marine diversity is in coral
reefs, they offer the greatest possibilities for
potential uses
scientists first began looking at softbodied sessile organisms of
coral reefs because they thrived under highly competitive
conditions with no apparent claws, teeth, etc for defense
à must use chemical weapons
by some estimates, coral reefs provide over $30
Billion in benefits (direct and indirect) , worldwide
per year
destroying 1/2 sq mi of reef costs $137,000-$1.2 M in
loss of fisheries, tourism and shoreline protection
over a 25 year period.
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9
coral hosts
therefore lose their color
triggered by disease, pollution, elevated
temperatures, salinity changes, increased UV
radiation, etc
bleaching is a normal response to short term stresses
while bleached, corals stop growing
à leaves reef vulnerable to erosion
after one bout the reef can recover,
àbut frequent episodes may kill the coral
polyps
what is significant about bleaching today is its
frequency, severity and extent
Coral Reefs are associated with 109 countries,
those in 93 countries show significant damage
reefs at highest risk:
Japan Singapore
Sri Lanka India
Asia
eg. Phillipines
only 5% of reefs are pristine
30% are dead
39% are still healthy
all are areas with dense coastal populations and
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9
Threats to Reefs:
Coral Reefs are among the most endangered
ecosystems in the world
recent (‘03) assessments of world’s reefs show they are
globally threatened
à there are no “pristine” reefs left:
all reefs are impacted by human activities
only reefs in remote areas are generally healthy
à 30% of reefs are damaged
up to 30% have been lost in last 50 years(06)
another 16% are severely damaged
à 60% may be completely dead by 2030
generally, coral reefs are very resilient
à have existed for 1000’s to 100,000’s of years
but today are being degraded in a matter of
decades
the greatest threats to reefs are from human activities
eg. ~1/2 of world’s population live in coastal regions
eg. in SE Asia, 70% of population is in coastal areas
Coral Bleaching
one of earliest signs of stress is coral bleaching
à when water gets too warm algae “flee” their
41 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 42
heavy coastal development
Human Causes of Coral Reef Decline:
while natural events, eg diseases and hurricanes
can cause extensive damage to specific reefs
humans are having a global impact on reefs
human causes of reef decline:
1. sedimentation
2. eutrophication
3. shipping and oil spills
4. exploiting for food (overfishing)
5. collecting
6. mining
7. tourism
8. Climate Change
9. Ozone Depletion
1. Sedimentation
by far the greatest impact
increase in suspended silt, clay, dirt
Taiwan mainly due to deforestation esp. mangroves
Indonesia due to logging, farming, mining, dredging
doesn’t have to occur near coast to have and impact
sediment blankets coral reef
initial plume blocks sunlightàreduces photosynthesis
smothers polyps
as they produce mucus to remove it, depletes their
energy reserves; makes them more susceptible to
disease
impedes larval settling
43 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 44
2. Eutrophication
food and nutrients usually limit the growth of most organisms
eg. N & P àplants, algae; organicsà bacteria, heterotrophs
reef ecosystems are especially susceptible since they are found
in nutrient poor waters
too much food can upset the balance between organisms in the
community:
some grow much faster than others and can become
toxic
sometimes a new predator gains upper hand
eg. crown of thorns starfish à can clean out
entire reefs when its predators are
eliminated
some algal infestations caused by eutrophication cause
algae to release sugars that fertilize the symbiotic
bacteria making them pathogenic and killing their coral
hosts
3. Shipping and Oil Spills
eg. oil tankers pollute and kill reefs
eg. 1st gulf war oil release (10M BBL’s)
caused extensive damage to reefs in arabian sea
eg. in Mid East a phosphate tanker ran aground on a reef,
releasing phosphates into the water
killing 500 mi2 of reef
eg ~1/6th of reefs in Phillipines have been damaged
this way since 1945
cyanide fishing
some use cyanide and poisons to fish
à kills other organisms as well
child labor
in Phillipines 40 ships carry 300 children to reef each day
children pound reef with rocks to scare fish into nets
can destroy up to 1 km2 of reef/day
children killed by needlefish, sharks, barracuda,
poisonous snakes, etc
as fish become more scarce, fishermen earn extra income
collecting turtles, clams, etc
5. Collecting
1.5 Million kg’s (15 tonnes; 3M lbs) of coral & shells/year are
harvested
mainly for “shell shops” around the world
~1/3rd from the Phillipines
most is exported
most goes to US gift shops and aquarium shops
live corals were collected and sold in Florida until 1989 when it
was outlawed
but some is still traded on black market

shells etc collected by malacologists: prefer killing live

4. Exploiting for Food (overfishing) specimens rather than dead shells from beach
reef fish are prone to overfishing because many are slow
growing, long lived fish (K-selected; low natural fertility) exotic fish collected from reefs feed a $4 Billion/yr aquarium
when depleted they are slow to repopulate industry

historical record shows that over the last several 1000 years,
large fish and animals have been hardest hit of reef 6. Use as Building Material
community in Sri Lanka and parts of India entire sections of reef have been
removed to make cement
blast fishing àthere is no other source of rock nearby
use explosives to kill or stun fish
7. Tourism (Ecotourism)
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 45 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 46

walking on reef and touching it kills polyps and kicks up àthis causes a 1-10% increase in UV radiation
sediment
many break off souvenirs of live reef shallow marine communities are particularly susceptible to
beauty of reef stimulates beach front developments damage from this additional radiation

eg. On S Pacific Island of Palau

they mined an area of reef to build a new airport runway
à to accommodate an increasing number of tourists
coming to see the reef

eg. in Grand Caymans a 525’ cruise ship dropped a 5 ton anchor

and
dragged its chain across 150M of reef
creating a 3M wide path 150M long
uprooted 8M diameter blocks of coral
destroyed an area 1/2 the size of a football field

Indirect Human Effects:

8. Climate Change
global temperatures are increasing 1/2 – 1 degree every decade
this rate is 100x’s faster than natural rate at end of last
glaciation
most of this accelerated warming is due to human activities
global warming will
alter weather patterns
alter ocean circulation
warm ocean surface waters
cause significant sea level rise up to 6 cm/decade
à but reefs can grow up to 10 cm/decade
cause acidification (lowering of pH) of ocean waters
à diversity decreases
à individuals are less healthy
à dissolves coral skeletons; reefs don’t grow
à reduces reproductive success of coral eggs and
larvae

9. Ozone Depletion
will continue into next century
àozone levels decrease .5-5% over the tropics
Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 47 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 48
Economic Importance of Cnidaria (excluding
coral reef ecosystems):
1. in orient a few jellyfish are eaten
eg. people in China and Japan eat the mushroom jellyfish; fresh
or pickled
2. stinging cells of some cnidaria are lethal to humans
eg. box jelly or sea wasp (Chironex fleckeri)
from Indian ocean to coral sea - esp around coast of
Australia
can have up to 60 tentacles as long as 15 feet.
eg. treatment for multiple sclerosis from coral venom is in
clinical trials
eg. in development (2009) is a process developed to
harvest stinging cells, remove their venom and then
use them to inject painkillers or insulin into the skin
4. a green fluorescent protein extracted from jellyfish
(also found in fireflies) and used to build tiny fuel
cells which could be useful in powering
nannodevices used to diagnose and treat diseases
in the body
5. space travelers: in 1991 2500 moon jellies flew
aboard the Columbia space shuttle

most poisonous sea creature known to study how their balance organs develop under
weightlessness
stings can kill a human in 5 minutes

each has enough toxin to kill 50 humans

since 1884 at least 5,567 deaths have been

attributed to these creatures.

3. Pharmaceuticals
eg. anti-inflammatories, painkillers for arthritis,
antimicrobials

eg. cardiac stimulant from sea anemone

eg. toxins from soft corals, Palython used as antitumor

medication

Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 49 Animals: Phylum Cnidaria; Ziser Lecture Notes, 2015.9 50
 Phylum Porifera (Sponges)
body consists of just 4 kinds of cells arranged
~9,000 living species; >2200 fossil forms around a system of pores and canals

abundant fossil record though multicellular, no true organs or even

eg. first signs of multicellular life are in 750 MY old rock, in this tissues
same rock are unique chemical signatures of sponges
sponges are closely related to the group of
eg. newly discovered(2010) fossils that may be sponges have
been found in 635-659 MY old rocks
protozoan protists called the
choanoflagellates whose cells very closely
eg. 400 MY ago sponges dominated the oceans as reef builders resemble the collar cells of sponges
eg. some fossil sponge reefs are much larger than the great
~ 1/4th of their genes are shared by all other animals
barrier reef
à covered an arc across most of N Europe 200 MY ago
about 1000 of those are absent from protozoa
and other protists
fossilized into hard rock used to build castles and other
buildings in the Middle ages
à may hold key to origin of multicellularity

genetic analysis indicates that sponges are the most while multicellular their structure is unlike any other
primitive animal group alive today animal group

ancient spongelike ancestors were the first early biologists thought they were some kind of plant
animals
all are aquatic, mostly marine
all other animal groups descended from this
ancestor found at all depths from intertidal to the abyssal
zone
- the earliest sponges in symbiosis with bacteria may have
significantly helped to aerate the ancient oceans making them
more suitable for animal diversification a few (~150 sp./27 US) occur in freshwater

sponges are some of the simplest animals most range from <1/2 inch to over 6 feet tall
(=loggerhead sponges)
all sponges are sessile
Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 1 Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 2

some are round, flat, grow as crusts or vaselike

but only of its own species (if 2 blended together)
most are assymetrical
sponges need only very small amts of oxygen and, as some of the
some are radially symmetrical earliest animals, may have played a role in helping to raise
earths oxygen levels by feeding on dead or dying bacteria that
would deoxygenate the water
often brightly colored: yellows, reds, greens,
oranges, lavenders
Body Plan
à pigments in surface cells
body is a network of pores, canals and passageways
most are colonial – colonies formed by budding
no “mouth”
all are sessile (non motile) , most are filter feeders
water is pumped through these passageways and the
but larvae are free swimming (motile) animals filter nutrients from the water currents

very simple in structure small openings are pores or ostia where water is
drawn into the sponge
though multicellular, they function largely like a
colony of unicellular organisms water exits the sponge through larger openings
= oscula (sing. = osculum)
Aristotle thought they were an intermediate between plants and
animals
natural selection favored an increase in surface area
cellular level or organization leading to more complex “folding” of the sponge:

à no true tissues or organs; loose aggregate of cells simple to more elaborate canal systems
à masses of cells in gelatinous matrix= mesophyll
à only a few cells have specialized for a particular function
a. asconoid
(6 kinds of cells in sponges; simplest type
humans have >250 kinds of cells) very small tube shaped sponges
large central cavity = spongocoel
eg. can force sponge through fine sieve to separate cells in via small openings called ostia
and individual cells will reform a sponge out through single osculum
Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 3 Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 4
 functions
b. syconoid
derived from asconoid pattern by folding
more branching
most cells are “totipotent” (ie. can change form
has incurrent canals and side passages and function – probably an important key to
still have main spongocoel their success)
single osculum

1. Choanocytes (= collar cells)

c. leuconoid
most complex
consists of many small flagellated chambers probably the most distinctive and most important
no longer a central spongocoel but almost unlimited of sponge cells
ability for sponge to grow in size
generally larger colonial forms
each mass has its own osculum each collar cell has a flagellum
incurrent and excurrent canals
surrounded by a sieve-like collar that acts as a
a typical sponge can pump water equivalent to its own strainer
volume in ~ 8 seconds
“collar” is made of microvilli and microfibrils

a sponge must pump >1 ton of water to get 1 oz of

the flagellum beats to draw the water currents into
food
the sponge and then to strain particles
through the collar
they can also control the flow by constricting osculum
at night and opening in day when food is more
food is absorbed (phagocytosis) by the collar cell
plentiful
and then sent to other cells in the sponge
also can reverse the flow to clean out canals after a choanocytes line major cavities depending on canal system:
storm a. spongocoel on asconoid types
b. radial canals on syconoid types
like cleaning pool filters c. flagellated chambers on leuconoid types

Cell Types
collar cells are almost identical to the cells of
choanoflagellates
only a few cells have been specialized for certain
Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 5 Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 6

genetic analysis indicates a very close

relationship between these protists and can differentiate into any other type of cell
sponges
different kinds
à choanoflagellates are ancestors of
animals sclerocytes à secrete spicules
spongocytes à secrete spongin
2. Pinacocytes collencyotes à secrete collagen

form outer “epithelium” and sometimes lines Support

inner passages
the cells in gelatinous matrix of a sponge are arranged
nearest thing to tissues around a skeleton of spicules

thin flat cells spicules maintain its shape and keep pores and
canals open
some are contractile = myocytes in circular bands
around oscula to regulate water flow spicules may be composed of:

3. Porocytes a. calcium carbonate

tubular cells form the pores of asconoid sponges b. silica

à spicules often united to form a rigid network that looks
4. Archaeocytes like fiberglass

amoeboid type cells (eg. Venus Flower Basket)

move about in mesophyll matrix c. spongin (a form of collagen – a protein only

found in animals)
receive particles from choanocytes for digestion à flexible protein fibers related to keratin

phagocytize old cells (eg. common commercial sponge and most sponges
Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 7 Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 8
 normally encountered on reefs)
within days other “tentacles” grow around victim and engulf
Feeding and Digestion and digest it

1. all but a few sponges are filter feeders 3. a small number of sponges are “parasites”

feed on detritus, plankton, bacteria = boring sponges (demospongiae)

pinacocytes, archaeocytes and choanocytes can all phagocytize excavates hollow tubes and passageways into shells and
food corals (living or dead host shells) or limetone rock
archaeocytes can eat larger particles
choanocytes can eat smaller particles the animal grows into the canals and holes it creates

when boring into live animal shells the host will either die
sponges can also absorb dissolved nutrients outright or be much more susceptible to predation
directly from the water
may have significant impacts on coral reefs and oyster reefs
digestion is all intracellular
important in recycling shells and corals = “bioerosion”
each cell is responsible for getting its own food
in some areas bridge supports are no longer constructed of
limestone because it is attacked by these sponges
2. one sponge is a predator
No respiratory or Excretory Systems
until 90’s all sponges were thought to be filter
feeding omnivores take in O2 and get rid of wastes and CO2 by simple
diffusion
one sponge from Mediterranean is now known to
be a predator = Cladorhiza corona a few have contractile vacuoles in choanocytes
found in Mediterranean caves
and archaeocytes

lives in stagnant water à not much to filter No Nervous System or Sense Organs
has developed a tentacle like appendge covered with
velcro-like hooks sponges can react to local stimuli

the hooks snag shrimplike crustacea some produce electrical signals

Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 9 Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 10

specialized sex cells in mesenchyme form egg

sponges do produce some hormones for chemical or sperm
control
sperm released into water
Reproduction & Development
eggs are retained in mesenchyme
reproduce both sexually and asexually
sperm are drawn into female sponge
Asexual through ostia or pores and fertilize
a. regeneration egg

b. asexual buds: most sponges are viviparous

may break off or remain attached to form =retain and nourish embryo
colonies
free swimming ciliated larvae hatch and are
c. fw & a few marine forms produce internal released
buds = gemmules
unique larval form = amphiblastula

à dormant masses of encapsulated swims in plankton for a while then turns

cells inside-out and settles to become a
sessile adult
usually produced during harsh conditions

may be retained inside “parent” sponge or

as original sponge dies, they fall to
bottom

Sexual
some sponges are monoecious,
some are dioecious

Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 11 Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 12
Sponge Classification [not current taxonomy] Phylogeny of Sponges

A. Class Calcarea (calcareous sponges) origin dates to cambrian

small, vase shaped, primitive group
mostly drab coloried; a few are yellow, red, green, lavender
related to flagellate protozoans
all marine, especially shallow waters à choanoflagellates
show all 3 types of canal systems; mostly asconoid canals
spicules of CaCO3, needle shaped or 3-4 rayed sponges diverged early

new genetic analysis (2010) indicate that sponges

B. Class Hexactinellida (glass sponges)
all marine; mostly deep colder waters are the most primitive animal group
body often cylindrical or funnel shaped
most vaselike or tubular; 7-10 cm to 1 m tall and should be divided into at least 2 separate
all with siliceous spicules, 6 rayed or long hairs like fiberglass
usually with large spongocoel and large osculum
phyla; Calcarea and Silicaria
synconoid and leucon canal systems

C. Class Demospongiae (spongin sponges)

largest class: ~90% of all living species,
most diverse group
including most large sponges
includes common bath sponges
mostly marine
1 small family of freshwater sponges
spicules of silica (but not 6-rayed), spongin fibers, or both
some tall and fingerlike, some encrusting
includes 1 group of “boring sponges”; burrow into shells and
corals

Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 13 Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 14

eg. some snails and clams have specific species of

Ecological Interactions sponges encrusting their shells

many sponges have mutualistic associations with

1. Mutualism & Commensalism bacteria

the greatest ecological role of sponges is to eg. heterotrophic bacteria (Pseudomonas, Aeromonas) live
inside the tissue of some sponges
provide homes for a wide variety of organisms
bacteria live in “mesohyl” jelly; a rich growth medium
many commensal organisms live in or on
sponges: sponges, snails, mites, fishes up to half the weight of a sponge is bacteria

sponge eats bacteria

à protection from predators
esp Demospongiae
larger sponges often harbor a larger variety of eg. demospongiae 38% of its volume was bacteria
commensals
eg. some sponges have blue green bacteria or algae
eg. 1 specimen, 2M tall had 16,000 shrimp inside that live inside their tissues
microorganisms get protection
eg. another had >100 species of organisms in and on it sponge gets food

eg. venus flower basket: no other animal has cyanobacterial symbionts

still used as a traditional wedding gift in SE Asia
typically has a male and female shrimp locked 4. Sponges as Prey
inside
=”bonded bliss” or “prisoners of love”
sponges seem to have few predators
Sponges also grow on many other animals;
molluscs, barnacles, corals, crabs some sponges produce chemicals to repel
potential predators
sponges are used by some animals as eg. several sponges are known to be toxic to fish
camoflage
still have some major predators
eg. decorator crabs: mobile substrate

Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 15 Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 16
 à a few bony fish esp from corals and other sponges

àHawksbill turtle produce “dead zone” around sponge

an endangered species associated with tropical reefs
feeds almost exclusively on sponges
these biotoxins can be antimicrobial
only vertebrate known with such a diet
most predators avoid the glass spines and poisonous àmay cause painful skin rashes in humans
secretions of the hawksbill prey

à in freshwaters, spongilla fly larvae feed on

sponges

5. Sponges and Competition

sponges are important components of coral reefs

their distribution is mainly limited by proper

substrate

corals are their chief competitor for space

sponges produce quite a few chemicals that repel

potential predators or other competitors for
space

(often brightly colored to warn others)

à they make a wide range of “biotoxins”

àprevent competition for space on

crowded reefs

Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 17 Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 18

Human Impacts of Sponges may attack AIDS virus

eg. a species of S Pacific sponge produces chemicals

1. bath sponges that can kill Candida à a human pathogen that causes
thrush and vaginal infections
have been used since bronze age; 4000 yrs
a new (2009) chemical derived from a sponge has
holds up to 35 x’s its weight in water the ability to resensitize bacterial pathogens to
antibiotics
takes 5 yrs to reach marketable size
à they lose their resistance to all antibiotics and die
eg. before 1940’s the Florida sponge fleet in Key West had >350
ships and employed 1400 people 3. Material Science
the sponge harvest ceased in 1940’s due to overcollecting,
red tides and a fungal disease that wiped out the the intricate glass skeleton of the venus flower
sponge beds basket is the strongest “glass” structure known
this was also the same time that synthetic sponges
were introduced to the market
it is so sturdy that it is being investigated by
material scientists for the source of its strength
2. sponges produce a wide variety of bioactive
compounds: also, its silica spicules transmit light better than
commercial optic fibers
pharmaceuticals: antibiotics, asthma, arthritis,
anticancer drugs, chemicals that promote efforts are being made to artificially duplicate
wound healing, anti-inflammatories them

eg. antibiotics against bacteria such as E. coli and Staph 3. Aquarium Trade
aureus

eg. Acyclovir
from Caribbean sponge
1st antiviral compound approved for human use
fights herpes infections
used since 1982

eg. Vidabarine
Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 19 Animals –Animal Phyla: Porifera; Ziser Lecture Notes, 2015.9 20
 Phylum Placozoa
has dorsal and ventral; but no front, back or sides
One known species: Trichoplax adhaerens à dorsal of flat ciliated cells and spherical cells
à ventral of columnar ciliated cells and gland cells
but newer genetic analysis indicates there might be up to 8
species between dorsal and ventral layer is fluid filled cavity
with some fibrous cells that can cause animal to
one of simplest animals
contract
à least DNA/cell than any other animal
can make amoeba like movements;
but movement mostly due to ventral cilia
small assymetrical disc-shaped animal
à moves in any direction
barely visible to naked eye 2-3 mm
probably feeds on algae and organic matter by
covering it and secreting enzymes then absorbing
àfew 1000 cells total
the digested material
no tissues or organs,
reproduces
little cell differentiation
asexual: by binary fission & budding
yet all other characteristics place it clearly in the
sexual: (has not been observed)
animal kingdom

phylum is based on a single species; Trichoplax

first discovered in 1883 from marine aquarium

à until 1971 thought it was a larval cnidarian

only known from walls of marine aquaria until

1980’s, when they were found throughout the
Pacific Ocean
Animals: Placozoa; Ziser Lecture notes, 2015.9 1 Animals: Placozoa; Ziser Lecture notes, 2015.9 2
 Taxonomy: the science of naming and describing species
Each distinct species is given a binomial name: The first word is the genus and is capitalized; the second
word is the species epithet; the binomial name is always italicized or underlined; eg. Homo sapiens

Species that are very closely related may be placed in the same genus; eg. Caenis lupus (gray wolf),
Caenis familiaris (domestic dog) and Caenis latrans (coyote); or Felix domesticus (house cat), Felix leo
(lion) and Felix tigris (tiger)

Most binomial names are latinized terms that describe some important characteristic of the species, identify
the location from which it was first collected or honor a particular researcher in the field

In a few instances species have been named after real or imagined celebrities:

Leonardo davincii (moth)

Polemistus chewbacca (wasp)
Polemistus vaderi (wasp)
Aptostichus stephencolberti (spider)
Myrmekiaphila neilyoungi
Aegrotocatellus jaggeri
Agra katewinsletae
Agra schwarzeneggeri
Agathidium bushi (slime mold beetle)
Agathidium cheneyi (slime mold beetle)
Agathidium rumsfeldi (slime mold beetle)
Avalanchurus garfunkeli
Avalanchurus lennoni
Campsicnemius charliechaplini (fly)
Calponia harrisonfordi
Confuciusornis sanctus (bird)
Funkotriplogynium iagobadius — James Brown. Iago is "James" and badius is "brown" in Latin
Hyla stingi (frog)
Orontobia dalailama (moth)
Orectochilus orbisonorum (beetle)
Preseucoila imallshookupis — Elvis Presley. And the specific name for one of his songs
Struszia mccartneyi

[see http://en.wikipedia.org/wiki/List_of_animals_named_after_celebrities for more]

Or sometimes just for fun:

Abra cadabra (clam) La cucaracha (moth)

Agra phobia (beetle) Ohmyia omya (fly)
Agra vation (beetle) Ytu brutus (beetle)
Ba humbugi (snail) Enema pan (beetle)
Gelae donut (beetle) Phthiria relativitae (fly)
Heerz lukenatcha (wasp) Pison eyvae (wasp)

The binomial name with the most letters apparently is:

Parastratiosphecomyia stratiosphecomyioides (fly, with 42 letters in the full name)

B. Dybowski, however, in 1927 proposed the following name for a Lake Baikal amphipod:

Gammaracanthuskytodermogammarus loricatobaicalensis

Which, with 50 letters, may be the longest scientific name ever proposed but this name was later invalidated by the
ICZN.
 History of the Idea of Change Through Time
Before Darwin
Although Darwin has been associated with the idea of “change through time”, such ideas were around
long before his work

1. Aristotle (384 – 322 BC)

saw evidence of “design and purpose” in nature

arranged all organisms into a “Scale of Nature” from simplest to most complex
saw a movement toward a “more perfect state” implied at least some kind of directed change

2. Leonardo da Vinci (15th century)

interpreted fossils as remains of organisms that had become extinct not just “sports of nature”
he doubted the flood story of the Bible

3. James Ussher (1581-1656) & John Lightfoot (1602-1675)

made assumption that the Bible was the only reliable source of chronological information for the
time covered in biblical writings
arrived at the calculation that the earth was created on Sunday, October 24, 4004 BC
Lightfoot, making additional assumptions put the time at 9:00 am

4. Leibnitz, 1690

concluded that the earth was originally a hot liquid; then cooled off forming a hard crust

 by 1700’s most scientists believe that fossils were of organic origin

but most were explained in terms of the Biblical flood

5. Maupertuis (1698-1759)

studied families with hereditary trait of 6 fingers and arrived at some basic rules of genetics
hinted at the concept of mutations and their ability to produce new species

6. Comte de Buffon (1707-1788) “Histoire Naturelle”, 1749

had to deal with dominant religious ideas: “this is what one might think if one did not know what
genesis says”
believed he could get an estimate of the age of the earth based on the rate of heat loss
then he calculated the age of the earth as 74,832 yrs and the origin of life at 40,000 yrs
he recognized 6 geological periods

7. Jean Baptiste de Lamark (1809)

he believed that all organisms were endowed with a “vital force” that drove them to evolve toward
greater complexity
Lamark was 1st to offer a “mechanism” for this change: he was the only biologist before Darwin
to offer a well developed natural theory of how living forms might evolved
 inheritance of acquired characteristics or evolution by “use and disuse of parts”
when scientists discovered the mechanisms of heredity his theories were discredited

8. Chambers, 1844

concluded that life could originate from inorganic substances and electricity

1
 General Ecological Principles

ecology: a study of organisms interactions with the abiotic and biotic components of their
environment •••••
biosphere: the total living world and all aspects of the environment with which it
interacts •••••
ecosystem: self contained group of interacting communities
•••••
community: self contained group of interacting species
•••••
population: self contained group of interacting individuals of the same species
•••••

I. Abiotic Factors
A. Lithosphere - solid: rocks, soil particles, sediments,etc
weathering
mechanical
chemical
B. Hydrosphere - liquid: water in every form; oceans, groundwater, rivers, lakes,
rainfall, ice
C. Atmosphere - gas: primarily nitrogen, oxygen and carbon dioxide gas with
mixture of other gaseous compounds
D. Tolerance Ranges and Limiting Factors
II. Biotic Factors
A. Feeding Strategies: herbivores, carnivores, omnivores, detritivores
B. Production & standing crop
C. Interactions
1. symbioses: mutualism, commensalism, parasitism
2. predator / prey relationships
3. competition
a. intraspecific competition
b. interspecific competition
4. population level interactions (Population Ecology)
a. fluctuations in populations and genetic variation
b. immigration and emigration
c. isolation and evolution
d. local extinctions
5. community level interactions (Community Ecology)
a. concept of the niche
b. foraging theory
c. community structure
III. Matter Cycling in Ecosystems
A. Trophic Levels and Food Webs: producers, consumers, decomposers
B. Pyramids of Numbers and Biomass
C. Biogeochemical Cycles
IV. Energy Flow in Ecosystems
 A. Kinds of energy used in ecosystems
1. solar
2. chemical
3. mechanical
B. Productivity
1. respiration vs production
2. gross vs net productivity
3. primary & secondary productivity
C. Energy Flow & Trophic Structure
1. factors that limit productivity
2. ecological efficiency and the "10% rule"
3. length of food chains
V. Ecosystems
A. Factors which structure ecosystems
1. Climatic Factors & Latitude and Altitude
2. Vegetation
3. Animal Communities
4. Ecosystem Stability
4. species diversity, species richness, and biodiversity
B. Types of Ecosystems (Biomes, Faunal Realms, Biogeographic Provinces)
1. Terrestrial Biomes
a. tundra
b. taiga (boreal forests)
c. coniferous forests
d. desert
e. chaparral
f. grasslands and savannas
g. deciduous forests
h. tropical forests
2. Marine Biomes
a. pelagic (photic vs aphotic zones)
i. neritic = coastal, above continental shelf
ii. oceanic = deep water area beyond continental shelf
b. benthic
(photic zone)
i. supralittoral (= splash zone) = highest tides and waves
ii. littoral (=intertidal) = between normal high and low tide
iii. sublittoral (=subtidal) = along continental shelf
(aphotic zone)
iv. bathyl = continental slope; 200-4000 meters depth
v. abyssal = abyssal plane; 4000-6000 meters depth
vi. hadal = trenches; greater than 6000 meters depth
c. marshes & saltwater swamps
d. coral reefs
3. Freshwater Biomes
i. lentic = lakes, ponds, reservoirs
ii. lotic = streams and rivers
iii. wetlands & freshwater swamps
iv. groundwaters and springs
 Economic Value of Diversity

an analysis by Cornell University (1997) estimated that throughout the world,

the services provided by the myriad species of all forms of life were valued
at almost $3 Trillion dollars annually.

for US alone the value was set at over $300 Billion

Natures services:

Activity United States World

Waste disposal $62 billion $760 billion

Soil formation $5 billion $25 billion
Nitrogen fixation $8 billion $90 billion
Bioremediation of chemicals $22.5 billion $121 billion
Crop breeding (genetics) $20 billion $115 billion
Livestock breeding (genetics) $20 billion $40 billion
Biotechnology $2.5 billion $6 billion
Biocontrol of pests (crops) $12 billion $100 billion
Biocontrol of pests (forests) $5 billion $60 billion
Host plant resistance (crops) $8 billion $80 billion
Host plant resistance (forest)$800 million $6 billion
Perennial grains (potential) $17 billion $170 billion
Pollination $40 billion $200 billion
Fishing $29 billion $60 billion
Hunting $12 billion $25 billion
Seafood $2.5 billion $82 billion
Other wild foods $500 million $180 billion
Wood products $8 billion $84 billion
Ecotourism $18 billion $500 billion
Pharmaceuticals from plants $20 billion $84 billion
CO2 sequestration $6 billion $135 billion

Total $319 billion $2.928 trillion

 The Animal Kingdom
(Metazoa)
Ziser, 2008
Kingdom Metazoa (Animals)
Phylum: Porifera [Sponges]
Phylum: Placozoa
Phylum: Cnidaria [Jellyfish & Corals]
Phylum: Ctenophora [Comb Jellys]
Phylum: Acoelomorpha
Phylum: Platyhelminthes [Flatworms]
Phylum: Mesozoa
Phylum: Nemertea [Ribbonworms]
Phylum: Gnathostomulida
Phylum: Micrognathozoa
Phylum: Rotifera
Phylum: Acanthocephala [Spiny-Headed Worms]
Phylum: Cycliophora
Phylum: Gastrotricha
Phylum: Entoprocta
Phylum: Ectoprocta [Bryozoa]
Phylum: Brachiopoda [Lampshells]
Phylum: Phoronida
Phylum: Mollusca [Molluscs]
Phylum: Annelida [Segmented Worms]
Phylum: Echiura [Spoonworms]
Phylum: Sipuncula [Peanut Worms]
Phylum: Nematoda [Roundworms]
Phylum: Nematomorpha [Horsehair Worms]
Phylum: Kinorhyncha
Phylum: Priapulida
Phylum: Loricifera
Phylum: Onycophora [Velvet Worms]
Phylum: Tardigrada [Water Bears]
Phylum: Arthropoda [Arthropods]
Subphylum: Trilobita
Subphylum: Chelicerata
Class: Merostomata (horshoe crabs)
Class: Pycnogonida (sea spiders)
Class: Arachnida (spiders, scorpions, mites, ticks)
Subphylum: Crustacea
Class: Branchiopoda (tadpole shrimp, water fleas, brine shrimp, fairy shrimp)
Class: Maxillipoda (seed shrimp, copepods, fish lice, barnacles)
Class: Malacostraca (shrimp, crabs, lobsters, crayfish, pill bugs, amphipods,)
Class: Remipedia
Class: Cephalocarida
Class: Pentastomida (tongue worms)
Subphylum: Uniramia
Class: Chilopoda (centipedes)
Class: Diplopoda (millipedes)
Class: Pauropoda
Class: Symphyla
Class: Insecta
Order: Protura
Order: Diplura
Order: Collembola (springtails, snowfleas)
Order: Thysanura (silverfish, bristletails)
Order: Ephemeroptera (mayflies)
Order: Odonata (dragonflies, damselflies)
Order: Orthoptera (locusts, grasshoppers, walking sticks, praying mantis)
Order: Dermaptera (earwigs)
Order: Plecoptera (stoneflies)
Order: Isoptera (termites)
Order: Embioptera (webspinners)
Order: Psocoptera (book lice, bark lice)
Order: Zoraptera
Order: Mallophata (biting lice
Order: Anoplura (sucking lice)
Order: Thysanoptera (thrips)
Order: Hemiptera (true bugs; squash bugs, chinch bugs, stink bugs)
Order: Homoptera (cicadas, aphids, scale insects, leafhoppers)
Order: Neuroptera (dobsonflies, at lios, lacewings)
Order: Coleoptera (beetles, fireflies, weevils)
Order: Strepsiptera
 Order: Mecoptera (scorpionflies)
Order: Lepidoptera (butterflies, skippers, moths)
Order: Diptera (true flies; fruit flies, house flies, mosquitoes, crane flies)
Order: Trichoptera (caddisflies)
Order: Siphonaptera (fleas)
Order: Hymenoptera (ants, bees, wasps, hornets)
Phylum: Chaetognatha [Arrowworms]
Phylum: Echinodermata [Echinoderms]
Phylum: Hemichordata [Acornworms]
Phylum: Chordata [Chordates]
Subphylum: Urochordata invertebrates
Subphylum: Cephalochordata ________________
Subphylum: Vertebrata
Class: Agnatha
Class: Chondrichthyes vertebrates
Class: Osteichthyes
Class: Amphibia
Class: Reptilia
Class: Aves
Class: Mammalia
Order: Marsupialia (opossums)
Order: Insectivora (shrews and moles)
Order: Chiroptera (bats)
Order: Edentata (armadillos)
Order: Lagomorpha (rabbits and hares
Order: Rodentia (rats, mice, squirrels, gophers, beavers)
Order: Cetacea (whales and dolphins)
Order: Carnivora (dogs, bears, raccoons, skunks, cats)
Order: Pinnipedia (sea lions, walruses)
Order: Sirenia (manatees, sea cows)
Order: Artiodactyla (deer, bison, cows, pigs, goats)
Order: Perissodactyla (horses, donkeys)
Order: Primates (lemurs, monkeys, gorillas, baboons, humans)
Suborder: Prosimii (lemurs, bush babies, tarsiers, lorises)
Suborder: Anthropoidea
Superfamily: Ceboidea (new world monkeys)
Superfamily: Cercopithecoidea (old world monkeys)
Superfamily: Hominoidea
Family: Hylobatidae (gibbons)
Family: Pongidae (apes, chimpanzees, gorillas)
Family: Hominidae
Genus: Australopithecus
Genus: Homo
Species: Homo habilis
Species: Homo erectus
Species: Homo sapiens (humans)
Subspecies: Homo sapiens neanderthalis
Subspecies: Homo sapiens sapiens (modern humans)
 Biol 1413: General Zoology
Taxonomy and Classification of Animals

Animals are distinguished from all other kingdoms by the following

characteristics:
♦ Eucaryotic cells that lack cell walls
♦ lack photosynthesis (although some animals harbor symbiotic algae or bacteria that may
carry out photosynthesis or chemosynthesis), instead they are heterotrophs

The taxonomy and classification of the different animal phyla relies

on some basic differences in the following general characteristics:
◊ Type of symmetry:
asymmetrical, radial symmetry, bilateral symmetry
◊ Number of Germ Layers (embryonic tissue layers):
no true tissue layers, diploblastic, triploblastic
◊ Presence and type of body cavity:
acoelomate, pseudocoelomate, coelomate (eucoelomate)
◊ Presence of segmentation:
segmented vs unsegmented
◊ Embryonic formation of body cavity:
schizocoelous, enterocoelous
◊ Embryonic mouth formation:
protostomes, deuterostomes
◊ Embryonic cleavage patterns:
spiral, radial, discoidal & some others

Some other characteristics that are important in the classification of

specific phyla and classes of animals relate to the diversity of
different organ systems found in the animal kingdom:
• Type of digestive system: complete vs incomplete
• Type of skeletal system: exoskeleton, endoskeleton, hydroskeleton
• Type of circulatory system: open vs closed
• Type of respiratory system: gills, tracheae, lungs (and others)
• Type of reproduction: asexual, parthogenesis, hermaphroditic, dioecious
 Animal Behavior
-one of the functions of the nervous and endocrine systems
-everything an organism does
-requires a stimulus

I. Simple Behaviors - require simple neural pathways

A. Tropisms
B. Taxes
C. Kineses
D. Reflexes
E. Other Innate Behaviors
II. Complex Behaviors - more complex neural pathways; require some form of learning
A. Species Specific Behaviors (Intraspecific Behaviors)
1. Cyclic or Rhythmic Behaviors
a. biological clocks
b. circadian Rhythms
2. Learned Behaviors
a. habituation
b. conditioning
i. classical
ii. operant
c. trial and error
d. imprinting
e. insight
3. Communication
a. Visual
b. Chemical
c. Auditory
4. Navigation & Migration
5. Social Behaviors
a. courtship
b. parental & family behaviors
i. licking
ii. feeding
iii. submissive actions
iv. affectional systems
c. group interactions
i. chemical conditioning of the environment
ii. aggressive behaviors
iii. dominance hierarchy
iv. territoriality
v. sociobiology (altruism & reciprocity)
B. Interspecific Behaviors
a. positive releasers
b. negative releasers (avoidance responses)
i. Mullerian mimicry (conspicuous coloration)
ii. Batesian mimicry (deceptive mimicry)
iii. camouflage
III. Behavioral Ecology
IV. Evolution of Behavior
 Animal Reproduction
Most animals can reproduce both asexually and
sexually.
Asexual reproduction produces genetically identical
copies (ie. clones) while sexual reproduction
produces genetically unique offspring.
There are advantages and disadvantages to both
types of reproduction:
Asexual reproduction
makes identical copies (clones) of the parent.
Only a single parent is required
the process is much quicker than in sexual
reproduction.
Asexual reproduction is beneficial when
resources are abundant.
Sexual reproduction
involves the fertilization of a female’s egg by a
male’s sperm
Zoology: Animal Reproduction 8/27/15
In some parasitic animals each different larva is
able to clone copies of itself. This allows a
single egg to produce 100’s of potential
offspring and enhances chances that at least a
few will be able to find a new host to complete
their life cycle. In humans this process occurs
occasionally to produce identical twins.
Regeneration: This process is most commonly used
not as a form of reproduction but to replace
missing or damaged parts.
Some organisms have great powers of
regeneration while others can only regenerate
simple cells and tissues.
Humans for example regenerate all their bone
tissue about every 7 years and all their
blood cells about every 4 months.
In other animals, such as lizards and arthropods,
tails or whole limbs can be replaced. Starfish
can regenerate new “arms” when one is
broken off and sometimes a single arm can
regenerate an entire starfish.
Examples of Sexual Reproduction
Most variations in sexual reproduction depend on
where the gametes come from or whether the
developing egg has been fertilized or not.
Zoology: Animal Reproduction 8/27/15
takes more time to find a mate, requires
courtship rituals, nesting, parenting skills,
etc
results in genetically unique individuals or
offspring
Sexual reproduction provides much of the
genetic variation required for evolution
and adaptation.
Examples of Asexual Reproduction
Budding: A very common type of asexual
reproduction especially in colonial animals
In budding a new offspring begins as an outgrowth
of the parent and may either remain attached
and form a colony or break away and begin an
independent life.
Fragmentation: Some animals spontaneously break
into many separate pieces which then regrow into
a complete animal.
Polyembryony (twinning): A type of asexual
reproduction in which the embryo or larva
(resulting from sexual reproduction) then clones
itself into separate individuals.
For example armadillos typically produce 4
identical offspring from a single fertilized egg.
1 Zoology: Animal Reproduction 8/27/15 2
Monoecious Animals (Hermaphrodites):
Monoecious organisms are those containing both
male and female reproductive organs.
About 15% of animals, especially those that are
sessile (nonmotile) or parasitic, tend to be
hermaphrodites.
Dioecious Animals: These are organisms that
produce either male or female reproductive
organs and gametes but never both at the same
time.
Protandry: is the ability in some animals to switch
their sex based on environmental cues such
as changes in temperature.
Sexual Dimorphism: Sometimes it is impossible to
distinguish the male from the female of a species
by outward appearance.
In other dioecious animals the males and females
are not identical, but differ in appearance,
thus showing dimorphism.
For example, in most invertebrates the male is
usually smaller than the female, while in
vertebrates the female is usually the
smaller one.
3 Zoology: Animal Reproduction 8/27/15 4
 In addition to size, dimorphism may also
result in differences in structures and color
between genders.

In birds the male is usually more brightly

colored.

Parthenogenesis (‘virgin birth’): this process only

regularly occurs in nonhuman animals, both
vertebrates and invertebrates.

In these cases the unfertilized egg is able to

develop even though it has not been fertilized
by a sperm.

Most rotifers, brine shrimp and some social

insects such as bees and ants regularly
reproduce this way.

Some higher animals such as fish and frogs

can also sometimes reproduce in this way.

Zoology: Animal Reproduction 8/27/15 5

 Animal Development
All living organisms exhibit some form of growth and
e gg spe rm
development.

Members of the animal kingdom have the most z yg ote

( fe rtilize d egg )
complex developmental cycles of any living
organism. clea vag e
pr eem br y onic
bl ast ula st ag es
The sequence of discrete, recognizable stages that
ga st rul a
these organism pass through as they develop from
the formation of a zygote (the fertilized egg) to
the sexually mature adult are referred to as its em br y onic s ta ges

developmental cycle.
lar va ny mph fe tus

Animal development can be subdivided into several

sequential processes:
1. Gametes.
gametogenesis
fertilization The gametes are produced by the process of
preembryonic development meiosis which differs from mitosis in that only
embryonic development one of each chromosome ends up in the cells
post embryonic development after division.

Embryonic development includes the processes of
growth, determination, differentiation and
morphogenesis.
The male gamete, the sperm, is small and almost
always flagellated.
The female gamete us usually large since it
contains yolk, and spherical.

2. Fertilization.
1 2

At fertilization only a single sperm penetrates and

fertilizes the egg to produce a zygote. The egg
and sperm each contribute a set of chromosomes
so that the fertilized egg then has a pair of each
chromosomes, one set from the male parent and
the other set from the female parent. To prevent
additional sperm from penetrating the egg a
fertilization cone is produced to produce the original
sperm into the egg quickly. Then a fertilization called a blastomere
membrane expands around the egg and pushes
away and “locks out” other sperm cells.
Slide: starfish early cleavage wm (Fig.
4.2)
-observe the slide of starfish eggs in early
cleavage and be able to distinguish this stage
from those that follow
-note 2, 4, 8 cell stages; each cell is
b. Morula:

Slide: starfish unfertilized egg wm (Fig. 4.2a) Continued division leads to a solid ball of
Slide: starfish fertilized egg wm (Fig. 4.2b) cells called the morula.

-Be able to distinguish between unfertilized and Slide: starfish late cleavage wm (Fig.
fertilized eggs 4.2G)

3. Preembryonic Development. [Ex 4-1] -identify the morula stage. How does its size
compare with that of the fertilized egg?
Almost immediately, the zygote begins to divide to
produce a multicellular embryo. c. Blastula:

a. Cleavage. Cell division continues until the embryo

becomes a hollow ball of cells. If the embryo is
The first identifiable period of development spherical as in starfish it is called a blastula;
occurs as the fertilized egg begins to divide. The cavity inside the blastula is called the
These early divisions are called cleavage blastocoel.
divisions and each cell produced is an identical
genetic copy of the zygote and is called a Slide: starfish blastula wm (Fig. 4.2H)
blastomere. At this stage each blastomere has
the potential to become a complete embryo.
3 4
 -Identify the blastula and distinguish it
from other embryological stages
-hollow ball of cells
-blastocoel
d. Gastrula:
In the gastrula, a depression forms at one
end of the embryo, cells move in to form a saclike
pouch. The embryo is now essentially two
layered. The cavity of this new pouch is called
the archenteron (or gastrocoel) which is
surrounded by the original, now much smaller
blastocoel. The archenteron opens to the
outside through the blastopore. The blastopore
will eventually become either the mouth or
the anus of the adult animal. By this time the
individual cells of the embryo are beginning to
differentiate into 3 embryonic tissue layers;
the ectoderm, the mesoderm and the
endoderm. The ectoderm will eventually become the
skin and nervous system. The mesoderm
will become the skeletal system, muscular system
and circulatory system. And the endoderm
will develop into the respiratory and digestive
system. A cell removed at this stage can no
longer develop into a complete embryo.
Slide: starfish gastrula wm (Fig. 4.3C,D)
-Identify the blastula and distinguish it from
other embryological stages.
5 6
forms that are found in more than one phylum.
Similar larvae imply similar ancestry; indicating
that the taxa are relatively closely related. Many
groups of animals have characteristic larval stages.
A few examples are illustrated below:
i. Planula larva.
A simple multicellular, oval larva with no
discernable organs, its surface is covered with
cilia for movement common in jellyfish and
some flatworms
Slide: Aurelia planula wm (HO)
ii. Nauplius larva.
A triangular larva with three pairs of jointed
appendages, eyespots, and digestive organs.
Nauplii are larvae of crustaceans such as
shrimp and crabs
Slide: nauplius, barnacle wm (HO)
iii. Glochidia larva.
Glochidia are larvae of freshwater clams that
are parasitic on the gills of fish until they mature
and fall to the sediment and begin life as freeliving
clams. Note the large ‘teeth’ that they use to
attach to their hosts.
7 8
-Note the blastocoel, archenteron
(gastrocoel), and blastopore
4. Embryo
After the three embyronic tissue layers are formed
in the gastrula the immature animal is referred to as
an embryo. During embryonic development the
organs and organ systems form. By the end of the
embryonic stage all tissues and major organs have
formed.
Slide: chick embryo 72 hours (HO)
Model: human embryo
-Note on the chick embryo and locate the
brain, heart, limb buds, eye
The animal embryo may next develop into any of
several forms such as larvae, nymphs, or fetuses.
Further development varies considerably in different
animal groups. Some examples are given below.
5a. Larvae.
In many animals, the embryo soon develops into a
free living larval form. Larvae are an immature
stage which move and feed independently and
often have no resemblance at all to the adult of the
species. While each phylum or class often has its
own characteristic larvae, there are a few larval
Slide: mussel glochidia wm (HO)
iv. Caterpillars, maggots, grubs and tadpoles
(HO)
are examples of larvae that look and feed
quite differently than the adults that they become
(butterflies, flies, beetles and frogs,
respectively)
Preserved: Caterpillars, maggots,
grubs & tadpoles (HO)
5b. Nymphs:
Nymphs are immature stages of animals that at
least somewhat resemble the adult of the species
and that live and feed independently
Preserved: mayfly, dragonfly, stonefly
nymphs
5c. Fetus:
In addition to embryonic development, vertebrates
(higher animals) produce an immature stage that
does resemble the adult but that is usually completely
dependent on the mother (or enclosing egg) for
nutrition and protection.
Preserved: misc vertebrate fetuses
Illustrations: human fetal development
Models: Human fetus

9
 Protists – General The Animal-like Protists: The Protozoa

Protists were the earliest and simplest of eukaryotic most are unicellular; a few are colonial
organisms
protozoa share several animal-like traits:
they were the second major form of life to appear lack cell wall
on the earth after the prokaryotes (bacteria) most are motile
heterotrophic nutrition
mostly single celled organisms
! protozoa gave rise to animals
very efficient cells compared to procaryotic cells
most are microscopic (3-300!m)
Protists are not a natural grouping, some divide
but some are relatively large cells than can be seen with the
protists into 5 or 6 separate kingdoms and 50 or unaided eye
so phyla
one species of amoeba (foraminiferan) is 2.5” long
very diverse group of organisms; algae, seaweeds,
some colonial
protozoa, slime molds & water molds
diverse group of organelles with highly developed
we will concentrate on those that have some affinities
division of labor
to the Animal Kingdom
protozoa are found in all aquatic environments
most of the “animal – like” protists are collectively
anywhere there is water or moisture:
called the Protozoa
freshwater ponds, lakes, creeks, rivers
common name for an unrelated group of protists that share at marine environments
least some characteristics with the animal kingdom
some found in terrestrial habitats where moisture is
abundant:
sand
soil
decaying organic matter
Protists - 1413, Ziser Lecture Notes, 2012.9 1 Protists - 1413, Ziser Lecture Notes, 2012.9 2

some can send extensions outward, then “ooze” into

them (eg. “the blob)
some are symbiotic in animals & other organisms
or they can even actually “walk” on these false feet
some are mutualistic
some protozoa are nonmotile (=sessile) but use cilia
many are parasites of animals or flagella to create water currents for feeding
eg. most vertebrates hae protozoan parasites in their
intestinal tract or in their blood most have optimum temperature range of
36 – 40 º C (=96.8 – 104º F)
eg. a few are ectoparasites of fish (Costia
most protozoa are heterotrophs
! must eat organic food:
in aquatic environments they are an important part of
plankton
they have many ways to take in organic food:
= organisms that drift with currents
1. absorbing dissolved organic nutrients
phytoplankton include autotrophic protists
through cell membrane
zooplankton include heterotrophic protists
2. ingest solid particles
most protozoa are motile by
through a mouth-like opening (=cytostome)

cilia eg. eat bacteria

usually many short whiplike filaments that beat in 3. some are herbivores
unison to move protozoan along
eat algae
flagella
4. some are saprophytic
much longer whiplike filaments; usually only one or a
few per cell =eat decaying organic matter in water or sediment

amoeboid motion (false feet; pseudopodia) eg. scavengers, detritus feeders

Protists - 1413, Ziser Lecture Notes, 2012.9 3 Protists - 1413, Ziser Lecture Notes, 2012.9 4
 5. many are predators fission = divide in equal halves
(not same as bacterial fission)
some can eat prey larger than themselves eucaryotic fission involves mitosis
(eg. didinium takes 1 min & digests for 2 hrs)
some split longitudinally (flagellages)
some have long hollow “tentacles” and pierce other
protozoa to suck contents out (suctoria) some split transversely (ciliates)

6. some are parasites budding = unequal fission

once inside the cell: multiple fission = >2 daughter cells

food becomes enclosed in vacuole which sexual: involves some exchange of genes
travels through cytoplasm (endocytosis)
conjugation = exchange of a few genes
digestive enzymes are injected into the (ciliates only)
vacuole to digest the food
syngamy = fusion of gametes (egg & sperm)
undigested material is expelled by a reverse
process (exocytosis) or through an “anal some have alternation of sexual and asexual
pore” generations

a few protozoa are autotrophs and have Life Cycles

chloroplasts
most protozoa exist in a single form which feeds and
do photosynthesis to make organic molecules reproduces

Reproduction and Life Cycles some alternate between two stages in their life
cycle:
protozoa reproduce both asexually and sexually:
troph = active vegetative feeding form
asexual: main form of reproduction
cyst = more resistant stage,
low metabolic rate
Protists - 1413, Ziser Lecture Notes, 2012.9 5 Protists - 1413, Ziser Lecture Notes, 2012.9 6

allows them to be successful even in sexual: involves some exchange of genes

harsh environments between 2 cells

! cilia or flagella are reabsorbed produces genetically unique individuals

! metabolism slows or stops
conjugation
! hard resistant outer covering is secreted
two individuals come together and one
the resistant stage can withstand harsh conditions
and become an active feeding stage again gives a few of its genes to the other
when conditions improve
they separate as genetically different
some cysts have survived for 38 yrs and 49 yrs in
dried soil
individuals and usually then reproduce
asexually
Reproduction
(ciliates only)

protozoa, like all protists, reproduce both asexually

syngamy
and sexually:
two separate cells, acting and male and
asexual: identical copies
female actually join and fuse together
like egg and sperm in a zygote
this is their main form of reproduction
(fertilized egg)
most protozoa divide several times per
combine their genetic material and then
day
divide asexually as a genetically
distinct individual
by: fission
most protists alternate between asexual and sexual
budding
reproduction
multiple fission a few parasitic forms also have several different developmental
stages in more than one host
Protists - 1413, Ziser Lecture Notes, 2012.9 7 Protists - 1413, Ziser Lecture Notes, 2012.9 8
Some major kinds of Protozoa: 1. “Amoebas”
these are just convenient groupings of a considerably larger number
of actual phyla and does not follow current classification schemes
amoeba = “to change form”

1. “Amoebas” includes protozoa that move by pseudopodia

(=false feet)
protozoa that move primarily by amoeboid motion
organism can alternate between solid gel-like and
44,000 living and extinct species liquid cytoplasm to produce pseudopodia

2. “Flagellates” ! false feet used for locomotion

protozoa that move mainly with flagella
! false feet used to engulf food
~1,500 species
! some are long thin tentacle-like for grabbing
3. “Ciliates” food and drawing it in
protozoa that use cilia for movement or for feeding
simplest protozoans !relatively few organelles
~8,000 species
also, some of the largest single celled organisms
4. Apicomplexans
nonmotile, parasitic protozoa with complex life cycles ! some amoebas are up to 4” long (forams)

~ 5,000 species the life cycle of some amoebas involve the alternation
between amoeba and flagellate forms

found in all aquatic environments

many are symbiotic in animals

amoebas are the only group of protozoa that have an

extensive fossil record
Protists - 1413, Ziser Lecture Notes, 2012.9 9 Protists - 1413, Ziser Lecture Notes, 2012.9 10

over 20,000 fossil species

a. Entamoeba gingivalis
some member of the group secrete or construct
protective shells found in the mouth near base of teeth

!the shell may be composed of calcium carbonate found in 95% of people with gum disease and 50% of
or silica secreted by cytoplasm people with healthy gums

!foreign material such as sand grains embedded in parasitic ! feeds on RBC’s and WBC’s at sites of infection
cement like secretion and gum disease

does not form cysts

two most important shelled forms:
! requires direct transmission by kissing, shared
radiolaria secrete a silica shell (SiO2) utensils

found from surface to bottom of ocean b. Entamoeba hystolytica (amoebic dysentery)

foraminiferans produce calcium carbonate shells (CaCO3) intestinal parasite

most live on the ocean floor in incredible numbers infects 400 Million worldwide
esp tropics and areas of poor sanitation
have existed since precambrian times 10% of world population is infected

form thick “oozes” that cover a third of the deep ocean up to 10 Million in US
floor
kills >10,000/yr
both have an extensive fossil record are are
90% hosts are asymptomatic
valuable to geologists as “index fossils”
humans only reservoir
amoebas reproduce mostly asexually
spread by fecal/oral route

a few also reproduce sexually cysts passed in feces

Human Impacts: !ingested with contaminated water

invade intestinal lining and feed on RBC’s

1. some amoebas are common human pathogens:
Protists - 1413, Ziser Lecture Notes, 2012.9 11 Protists - 1413, Ziser Lecture Notes, 2012.9 12
 can cause ulcerations and profuse bleeding in acute pools but does seem to survive in low numbers even in
cases treated water supplies

may spread to liver, lungs, brain, etc may prefer areas where other organisms have been wiped
out by natural or man made disasters (eg Mt. St.
2. Naegleria fowleri Helens)

members of the genus are found in almost all freshwater 3. Acanthamoeba

lakes, rivers, hot springs
one of the most common amoebas in soil
but extremely rare in them
also found in freshwaters
feeds as an amoeba on bacteria
though free living it can occasionally cause severe infections
once most of the food is gone they transform into a of eyes, skin and brain especially in patients with
flagellated cell (<90 minutes) which is better able compromised immune systems
to go in search of food
spread by improperly disinfected contact lens solutions
one species, Naegleria fowleri, is a human pathogen
can damage cells of the cornea
35 cases reported in Texas (2007) including a few in central
Texas have died from infections of this amoeba
parasite

usually infects from getting contaminated water into nose

makes its way to the brain

causes always-fatal primary amoebic meningoencephalitis

or PAM

most die within 2 weeks

mature adults seem to be immune

the parasite prefers warm waters with a high iron content

especially warm stagnant water

usually cannot survive highly chlorinated water of swimming

Protists - 1413, Ziser Lecture Notes, 2012.9 13 Protists - 1413, Ziser Lecture Notes, 2012.9 14

2. “Flagellates” within the colony there is a division of labor with some cells
specializing in feeding and locomotion and larger germ cells
in specialized for sexual and asexual reproduction
includes several major phyla
asexual reproduction includes the formation of daughter
cell membrane surrounded by pellicle that “stiffens” colonies inside the “adult” colony

the cell membrane

most flagellates are symbiotic
move using one or a few long flagella
one cellulose digesting group has a mutualistic
symbiosis with animals
some have “sail-like” undulating membrane
used for food gathering and locomotion animals are not able to produce the enzymes
to break down cellulose or lignin
reproduce by binary fission
eg. cellulose digesting flagellates in the gut of
termites
a few are free living
1/3rd to 1/2 of a termites weight is these symbiotic
eg. Euglena is common in stagnant ponds and creeks protozoa

it usually has chloroplasts and does photosynthesis eg. cellulose digesting flagellates in cow rumen

when sunlight is not available it gets rid of its chloroplasts contains 1 M protozoa/ml (100 l of fluid total
and becomes a heterotroph
they provide cow with ~20% of its protein needs
eg. Volvox is a colonial flagellate that is thought to resemble
what the first truly multicellular animals might have looked
like.
some are parasitic in humans and other animals

Each hollow spherical colony is made up of 50,000 one group of flagellates, the “Choanoflagellates”
individual cells embedded in a gelatinous ball

each cell is similar to Euglena cells and are interconnected are believed to be the protists group most closely
by cytoplasmic strands related to the protozoa that gave rise to
animals and fungi
they are autotrophic

resemble feeding cells (collar cells) of sponges

Protists - 1413, Ziser Lecture Notes, 2012.9 15 Protists - 1413, Ziser Lecture Notes, 2012.9 16

common in freshwaters and salt water
many species are colonial
Human Impacts
many protozoan flagellates are important human
pathogens throughout the world
eg. Giardia (one cause of “traveler’s diarrhea”)
first observed by von Leeuwenhoek in his own feces
worldwide distribution: one of most common intestinal
parasites in the world
! up to 20% of all humans are infected (7% US )
also occurs in cattle, cats, bears, coyotes, bird &
amphibians
transmitted by comtaminated food or water:
cysts shed in feces; fecal/oral transmission
epidemics associated with contaminated water
especially common in poor overcrowded areas with
poor sanitation and lack of clean water
can also be transmitted in ponds and pools
! cysts can survive up to 2 months in water
! chlorine doesn’t always kill cysts
once ingested Giardia infects small intestine
Protists - 1413, Ziser Lecture Notes, 2012.9
begins with aching joints, headache and fever
affects CNS: personality changes, headaches, apathy,
sleepiness, emaciation
usually results in death from coma, malnutrition,
secondary infections
so far, no safe and effective treatment
eg. Chagas disease (T. cruzi)
new world tropics; eg Mexico, Central America, So. America
40-50% of population in So. America; 2-3 Million are
chronically infected
! 45,000 die each year
the most serious cases occur in children <5 yrs old
only a few cases in extreme SW US
also requires 2 hosts in its life cycle:
kissing bug and humans
in kissing bug its an intestinal parasite
in humans it’s a blood parasite
other mammals serve as reservoirs: rodents,
possums, armadillos
contracted when “kissing bug” bites (usually on lips)
bug usually defecates after feeding
when the bite is scratch some of the infected
feces is rubbed into the wound
Protists - 1413, Ziser Lecture Notes, 2012.9
it is not usually a parasite: it usually feeds on dead
organic material; no invasive ability
usually asymptomatic
in large #’s can cause chronic diarrhea, cramping,
dehydration
incidence is increasing in US where it affects 3x’s more
children than adults; esp daycare centers & public
places
eg. Trypanosoma (Africal Sleeping Sickness)
some of the most important protozoan parasites are in this
genus
trypanosomes are blood parasites and occur in all
vertebrate groups
human parasites occur mainly in the tropics of Africa and
the Americas
African Sleeping Sickness occurs in old world tropics; esp in
Africa
about 10,000 new cases occur each year; kills ~5,000
people/yr (2007); many of the rest suffer permanent
brain damage
requires two hosts:
the tse-tse fly is the final host for the sexual stage of
the parasite
humans and other animals are intermediate hosts
humans become infected when fly bites for blood meal
parasite moves into blood and lymphatic system
17 Protists - 1413, Ziser Lecture Notes, 2012.9 18
symptoms somewhat similar to sleeping sickness
chronic and hard to treat
may also affects many organs; eg. brain, heart,
intestines
most dangerous to children
! can affect many organs
eg. Trichomonas
several species; commensal or parasitic
T tenax
lives in mouth, is not a pathogen
5-10% oral infections, esp with poor oral hygeine
T. vaginalis
20-40% infection rate worldwide
one of most common infections in US
(2.5 M inf/yr: 3-15% US infected)
lives in human urogenital tract: likes acidity of female tract
~50% are asymptomatic carriers
no cyst form ! usually requires personal contact (STD)
occasionally spread in communal baths
and mother to child
if acid balance is disturbed, eg. by other infections, can
become more virulent
19 Protists - 1413, Ziser Lecture Notes, 2012.9 20
 esp common in promiscuous young women who are
already infected with other STD’s
in some women infection may produce a frothy,
smelly green discharge & painful urination
not often virulent in men
Protists - 1413, Ziser Lecture Notes, 2012.9
all ciliates have more than one nucleus and
usually two different kinds of nuclei
macronucleus ! vegetative chores
micronuclei (up to 80) ! sexual reproduction
eg. “mouth” (=cytostome) and throatlike area
called a gullet
most feed on microorganisms
have mouthlike cytostome; opens into a throat;
food vacuole forms at end of throat
an unusual group are called the suctoria
which paralyze their prey (other protozoa) and suck
out the cell’s contents with tubelike “tentacles”
eg. food vacuoles
contain digestive enzymes for processing organic food
eg. contractile vacuoles
freshwater species tend to take on water
must constantly pump out excess; like a bilge pump on
a boat
eg. trichocysts
long thread like proteins that the protozoan is able to
shoot out to anchor the cell or to capture prey
eg. myonemes
Protists - 1413, Ziser Lecture Notes, 2012.9
3. “Ciliates”
the most diverse group of single celled 'protozoan'
protists
they also tend to be larger than most protozoans and
some can even be seen without magnification
most are freeliving and solitary
in a wide variety of aquatic habitats, especially in
freshwaters
most are motile by means of cilia
= 1000’s oarlike projections produce coordinated
movements
fastest of all the protozoans
in some bundles of cilia are fused to form rigid spines (=cirri)
that the organism uses to crawl on substrates
a few are nonmotile, and some of these are colonial
live attached to substrate by stalk
use cilia for filter feeding, not for movement
ciliates have the greatest variety of organelles
and internal structures of all the protists:
eg. more than one nucleus
21 Protists - 1413, Ziser Lecture Notes, 2012.9 22
muscle-like fibers that allow stalked forms (eg. Vorticella) to
rapidly contract from danger
“eg. chloroplasts!”
ciliates are heterotrophs but …
some ciliates can steal chloroplasts from the algae
they eat and then use them for photosynthesis
Reproduction:
asexual: binary fission
sexual: conjugation: portion of micronuclei
are exchanged between + and – forms
Ecological Interactions
ciliates play a vital role in food webs, particularly of
freshwater ecosystems
many are part of the zooplankton
others are benthic - spending their lives
crawling about the substrate for food
23 Protists - 1413, Ziser Lecture Notes, 2012.9 24
4. “Apicomplexans” malaria has probably killed more people than any other
disease in history

All members of this group are nonmotile chronic in some parts of world

all are endoparasites worldwide infects 300 - 5000M each year and kills 1-3 M/yr
(90% of cases in Africa, also in Asia & Latin America)

most have fairly complex life cycles every 12 seconds someone dies from malaria

unlike many other parasitic diseases it is NOT a disease of

!same species exists in lots of different forms poor sanitation and contamination

alternating between forms that reproduce its distribution and incidences is closely correlated with its
sexually and those that reproduce mosquito host

asexually relatively rare in US (usually travelers)

sometimes in two hosts single most important disease hazard for people
traveling to foreign lands

Human Impacts: requires two hosts to complete life cycle:

Human parasites include: Anopheles mosquito has sexual stages in its salivary
glands

eg. Texas fever (Babesia) humans harbor the asexual stages in blood, especially
vessels in liver
killed 1000’s of cattle in US in late 1800’s and early 1900’s
transmitted by mosquito bite
spread by tick
symptoms of infection:
destroy RBC’s ! causes red urine ! “red water fever”
7-14 days after infection cold chills and shaking begin
today almost completely eliminated by dipping cattle to
kill ticks uncontrollable deep tremors take over the body (can
propel a bed across a room)
eg. Plasmodium (malaria); several species
next comes fever (up to 106º F) with profuse sweating

cyclic chills/fever, headache every 3-4 days

Protists - 1413, Ziser Lecture Notes, 2012.9 25 Protists - 1413, Ziser Lecture Notes, 2012.9 26

generally a nonhuman parasite that can occasionally

can produce irreversible damage to liver, spleen, infect humans
kidneys and brain
humans can become intermediate hosts
many succumb by way of delerium and coma
humans contract by contaminated soil, cat feces (litter
if not treated may be self limiting but host may be box), infected meat
a reservoir for up to 3 years
generally no human-human transfer
most effective prevention is elimination of mosquito
! 16% of US adults are infected
WHO has been trying to eliminate it but with little success
often asymptomatic in adults; children sometimes get rash
mosquitoes have developed resistance to insecticides (‘macropapular rash’)

the parasite has developed antibiotic resistance in humans can invade blood and multiply in WBC’s and
various organs
experimental vaccines are being tested
if contracted by pregnant woman (especiallly in the first
some living in endemic areas have developed genetic trimester) the parasite can cross placenta and cause
resistance to disease (sickle cell) retardation blindness and convulsions in embryo, fetus
or newborn
eg. Toxoplasma
! 2% of all mental retardation in US may be due to
requires two hosts to complete life cycle: prenatal Toxoplasma exposure

cats are primary host, new info indicates that though there are usually no
symptoms in most infected adults there seems to be a
prey species such as rodents serve as intermediate correlation with more risky behaviors in humans the
hosts mimmic the results of the parasite in rodents

infected cats release cysts in feces ! 1000’s of years ago would increase the chances of
humans falling prey to large cats
rodents, cattle, sheep are intermediate hosts
in another study 3900 drivers were monitored for 18
to spread toxoplasma manipulates rodents brains months
making them reckless and more likely to be
caught by cats those who were infected with Toxoplasma were 2.5X’s
more likely to have an accident
toxoplasma is an example of a zoonosis

Protists - 1413, Ziser Lecture Notes, 2012.9 27 Protists - 1413, Ziser Lecture Notes, 2012.9 28
 ! based on current rate of world infections, 0.4-1
million of world’s annual road deaths
might be due to toxo infections Protists -
Toxoplasma has also been implicated in the mental
Slime Molds & Water Molds
disorder; obsessive-compulsive disorder, but results
are not yet conclusive ~1100 species

eg. Cryptosporidium sp. two distinct groups of fungus-like protists:

first reported in people in 1976
slime molds and water molds
is now recognized as a major cause of diarrheal disease
worldwide both superficially resemble fungi at some stage in life
cycle
especially in children in tropical countries

occasional outbreaks occur in US !heterotrophs

can be life threatening in AIDS patients ! some produce chitinous cell walls at some
stage in their life cycle

!body of threadlike filaments = hyphae

! many produce a fruiting body with spores

for reproduction

but differ from fungi:

! most are motile by false feet or flagella at some

point during life cycle; fungi are NEVER motile

!produce flagellate reproductive cells; fungi

produce nonmotile spore

Protists - 1413, Ziser Lecture Notes, 2012.9 29 Protists - 1413, Ziser Lecture Notes, 2012.9 30

!some have cellulose cell walls or no cell walls; all 1. Slime Molds
fungi have cell walls, usually made of chitin
this group is probably more closely related to amoebas
than to fungi
! sometimes referred to as “social amoebas”

common in cool, moist shady places

most easily found in summer and early fall

eg. crevasses of rotting wood

two basic stages to its life cycle:

a. a relatively large motile feeding stage

b. the reproductive stage in the form of a

fungus- like fruiting body that produces spores

a. feeding stage (‘plasmodium’):

for most of a slime molds life it exists as a thin,

free-living amoeba-like mass of protoplasm

essentialy a large single cell up to several

inches across that

can cover an area of several square yards

(to 30 g = ~ 1oz)

Protists - 1413, Ziser Lecture Notes, 2012.9 31 Protists - 1413, Ziser Lecture Notes, 2012.9 32
 creep along in amoeboid fashion and feeds ! biologists “saved the world!”
on decaying organic matter, bacteria and
protozoa b. reproductive stage:

it is thick and slimy to the touch when food supply dwindles reproduction is
initiated
feeds and grows as long as there is food and
moisture it moves out of its normal habitat and goes to a
drier, more exposed location to produce a
some species form extensive growths on lawns, fruiting body
croplands
often seen crossing roads, lawns, climbing trees, etc
! do little, if any, damage
fruiting bodies can also be produced by absence of food, changes
in moisture, pH, temperature
! may appear in the same locations, year after year
as patches of purple, gray, white & cream
plasmodium divides into numerous mounds

some species found on lawn are mistaken for dog each mound forms cells surrounded by cell walls
vomit
at this stage the slime mold more closely resembles fungi
some pet owners find them then rush their dogs to than amoebas
the vet to find out why their pet is sick
produces multicellular fruiting body (= sporangium)
eg. Fuligo septica plasmodium (shades of war of
the worlds)
! very small (~1-2mm); look like tiny mushrooms
1973 found in Dallas suburb & reported in paper
appeared on lawns as bright yellow masses ! goblets, globes, plumules
spread over large areas
described in paper as a “pulsating yellow blob” ! with or without a stalk
blobs broke apart when sprayed with hose
!but pieces continued to crawl around ! often colored yellow, orange, red
caused local panic:
!must be indestructible aliens from space ! produces very resistant reproductive spores
! or mutant bacteria that might take over
the earth some slime molds can produced a hardened resistant
excitement soon dissipated once identified
sclerotium to survive adverse condition
Protists - 1413, Ziser Lecture Notes, 2012.9 33 Protists - 1413, Ziser Lecture Notes, 2012.9 34

2. Water Molds
new (2010) research indicates that some slime molds
1000 species ( ~100 genera) described
show traits usually encountered in more complex
organisms: most primitive group of fungi
eg. slime molds can be taught to “run mazes for food”
molecular evidence suggests that they are a direct link between
protists and fungi
eg. some slime molds “farm” the bacteria they eat

they stop grazing on bacterial while there is still some left some are unicellular, some multicellular

then mix uneaten bacteria into the spores they produce to have chitin in cell wall
make a “starter kit” for the next generation

fossils of this group has the distinction of being the mostly aquatic, a few are terrestrial
first true fossil that actually shows an organism
caught in the act of sexual reproduction (65MY) extremely abundant
a teaspoon of water from virtually any freshwater habitat should
Economic Importance of slime molds: yield samples

1. slime molds are eaten in Veracruz Mexico: most are saprobes –absorptive
some are collected, fried and eaten by indigenous
peoples others are parasites of plants, animals and other
called “caca de luna” fungi

most commonly seen as the fuzzy filaments

growing on skin or eggs of fish & amphibians
eg. Saprolegnia is common parasite of aquarium fish;

causes lesions

sometimes becomes a problem in fish farms

other species infect rotifers, nematodes,

arthropods and diatoms
Protists - 1413, Ziser Lecture Notes, 2012.9 35 Protists - 1413, Ziser Lecture Notes, 2012.9 36
 virtually the entire Irish potato crop was wiped out in
Economic Impacts of Water Molds: one week

> 1 million deaths from starvation

1. some (chytrids) are part of the microorganism began large scale emmigration of Irish to US
community in the stomachs of most farm animals
and grazing animals within a decade the population of Ireland
dropped 50%: 8M -> 4M

they are anaerobic and produces cellulases to help eg. other Phytophthora species
digest plant material along with other protists
have caused widespread destruction of many crops
and bacteria throughout the world:
pineapples, tomatoes, rubber, onions,
therefore all products coming from these animals strawberries, apples, soybeans, tobacco,
(beef, milk, dairy products, leather, wool, etc) citrus

are in part a product of these protists

3. Animal Pathogens
2. some are serious plant pathogens
a primitive water mold pathogen
(Batrachochytrium dendrobatidis (chytrid)) is
eg. root rotting fungi, blister rusts, white rusts
at least partly responsible for current decline
and downy mildews
in amphibians around the world
eg. Downy Mildews
today one third of the worlds 6,000 amphibian species are
infect grapes, lettuce, corn, cabbage and many other threatened
crop plants
! one of largest extinction spasms in vertebrate
introduced into France in late 1800’s history

almost destroyed the wine industry unsure of exact causes of declines:

problem was accidentally solved using copper possibly caused by acid precip, deforestation
sulfate and lime urbanization, climate change

eg. Potato Blight (Phytophthora infestans) more recently noted deformities pollutants in water

Cause of Irish Potato Famine (1845-7) in Ireland most recently has been tied to worldwide spread of
(including in and around central Texas)
Protists - 1413, Ziser Lecture Notes, 2012.9 37 Protists - 1413, Ziser Lecture Notes, 2012.9 38

the fungus spreads very rapidly;

don’t know how it kills frogs

Barton springs salamander and some other amphibians

have natural antibiotics in its skin that seem to
protect it from the pathogen)

2008-probiotic treatment with normal amphibian skin

bacterium, Janthinobacterium lividum, seems to
protect frogs from the chytrid.

It apparently produces an antibiotic that is deadly

to the chytrid.

Treatment is now being tested on wild

populations

Protists - 1413, Ziser Lecture Notes, 2012.9 39

 Origin and Evolution of Animals the main difference in multicellular organisms is
that in them cells clump together, specialize
For the first 3 Billion years of earth’s history life and become more dependent on each other in
existed only as single celled bacteria and protists multicellular organisms

3/4ths of all geologic time was dominated by Multicellular Life

primitive, slowly evolving microbes
~1 BY ago multicellular organisms appeared and
virtually the entire basic organization of the biological began to diversify
world dates from this time
multicellularity was not a major step but a natural
! bacteria and protists set the stage for animals: progression toward increased competitive
interaction and specialization
1. locomotion: flagella, cilia, amoeboid motion

2. skeletons and shells for support while we tend to think of bacteria and protists as
single celled organisms they more typically
3. cell to cell communication:
form groupings, films, filaments, etc
chemical messengers leading to hormones and
neurotransmitters bacteria don’t really function as single cells in
nature but as microbial communities:
4. metabolism:

virtually all but a handful of chemical reactions that teams of several kinds of bacteria live
animal cells use already existed together responding to and creating
eg. aerobic respiration each others environment

eg. digestion ! wastes of one becomes another’s food

! complementary enzymes to share the process
eg. lethal toxins for defense of digestion
! they communicate chemically with each other
eg. hemoglobin to store O2 ! protect each other
! essentially become interdependent
5. asexual and sexual reproduction
Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 1 Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 2

! in other words they were functionally

multcellular 2. allow much more variation in size & shape

a multicellular body is the only way that a living most single cells take in food and oxygen and get
organism can achieve sizes larger than a few rid of wastes by simple diffusion in and out of
millimeters the cell

! cells can’t get too large without breaking in larger organisms there are more ways to solve
these problems
better to make lots of small cells than a eg. getting food: herbivores, carnivores, fluid feeders,
few very large cells insectivores, etc

multicellularity evolved many separate times among eg. getting oxygen: gills, book gills, lungs, tracheae,
etc
eukaryotes:
seaweeds
3. multicellular organism becomes more resilient
slime molds
fungi damage doesn’t kill you
plants
animals
!it can often be repaired
!must have been a clear advantage
but there are some tradeoffs:
Advantages of Multicellular Life:
Disadvantages of Multicellular Life:
1. allows individual cells to become more specialized
for specific functions: 1. individual cells become completely dependent on
each other
all cells don’t have to do everything
! can no longer survive on their own
eg. epithelial cells: cover, protect, support, etc
eg. muscle cells: movements
eg. vascular tissues: transport of nutrients and wastes and
2. larger organisms need MUCH MORE food and
hormones oxygen
Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 3 Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 4
 molecular evidence puts animal origins at ~625 MY ago
! lots more cells to feed
poor fossil record of animals during most of this
3. as size increases need more elaborate ways to
earliest time
supply each cell with nutrients and remove wastes
elaborate organ systems to collect and distribute nutrients mostly soft bodied forms ! leave few fossils
and get rid of wastes

eg. vascular systems, etc

4. need method of coordination and control to get

all parts on the “same page”
eg. hormones, nervous systems

5. need some kind of support, especially on land, to

counteract gravity

6. Dispersal becomes more of an issue

eg. most bacteria and protists have worldwide distributions

relatively few large animals and plants have worldwide

distributions

of the three multicellular kingdoms (fungi, plants &

animals; animals appeared earliest in the fossil
record

earliest fossil evidence of animals is 575 MY ago

but indirect evidence of animal life are found over 1 billion
years ago

! not true fossils, trace fossils (burrows, etc)

Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 5 Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 6

The Cambrian “Explosion” earliest fossils of land animals are arthropod fauna:
eurypterids, scorpions, centipedes ~440 MY
(Ordovician)
once multicellularity originated the more complex
lifeforms evolved fairly quickly What caused the Cambrian ‘Explosion’?

!evolution of multicellularity is correlated with an the convergence of several factors may have
explosion of diversity in the fossil record produced this dramatic increase in fossils

only 3 of the major animal phyla alive today have a 1. rapid melting of snowball earth
fossil record before the 530 MY (during Cambrian)
before this time the surface of the earth was completely
eg. 700MY Ediacaran fossils frozen = ”snowball earth”

just before Cambrian it began to thaw very quickly (possibly

after 530 MY, fossils of all major animal phyla are in less than 1000 years)
found
! may have lead to an explosion of diversity
eg. sponges, corals, molluscs, arthropods, echinoderms,
vertebrates, etc 2. some suggest that a threshold level of O2
induced animal evolution
!virtually all major animal groups (phyla)
appeared in a geologic instant (~10 M yrs) before this time there was not enough oxygen in the
atmosphere or surface waters to meet the much higher
= Cambrian Explosion aerobic energy demands of animals

3. warm shallow seas may have promoted

all modern animal phyla originated as marine diversification
organisms
eg. much of North America was covered by a warm
only a relatively few animal groups successfully shallow sea rich in nutrients, rising oxygen levels
adapted to land
4. evolution of hard teeth and skeletons
! no animal phylum is completely terrestrial
almost all fossils BEFORE the Cambrian
Explosion were soft bodied forms
Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 7 Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 8
 Arthropods quickly became the dominant lifeforms
! indicated that predators had not yet and have dominated the fossil record since
evolved
one of the oldest animal species on earth (has remained
unchanged) is Triops cancriformis
only after animals began trying to eat each
other ! 180 M yrs ! requires no males

! evolved larger sizes, harder coverings

hard bodied invertebrates (animals) are much

more likely to leave fossils

shells may have been a way to recycle

abundant calcium in sea water

in marine cells the [Ca++] is 10,000 x’s less than in sea

water

! Calcium tended to diffuse into the cells

in large amounts this could be toxic

animals began to use this excess toxin for defense,

support, teeth, spines

! transformed a pollutant into something useful

[Nabokov – the greatest enhancements in nature

involve deception]

by the middle of the Cambrian age virtually every

major invertebrate group was well
represented

Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 9 Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 10

History of Animal Life in the past 600 MY paleontologists estimate that 100’s
of times the number of species alive today have
existed and become extinct
The Animal Kingdom has dominated life on earth for
the past 600 MY (at least in terms of ‘visible’ species) they also estimate that individual species only last 5-
10 MY before they have adapted to such a degree
we have a fairly good fossil record as evidence of life’s as to become new species
history over this time
! the ‘lifespan’ of a species is usually 5-10 MY
Fossils
the fossil record reveals two major trends in the past
actually, only a relatively few organisms actually leave history of animals (and all life on earth):
a record of their past existence
fossils are usually preserved as molds or casts or when 1. the oldest species are very different from the
tissue is replaced with harder minerals species in the world today

fossils of aquatic forms are much more common (even though they represent most of the same phyla)
than fossils of terrestrial organisms
they are more likely to get buried in sediment before 2. the total number of species has generally
they are eaten or decompose increased over the history of life on earth

fossils are especially common when an organism ! today there are more species alive at one
produces some kind of hard part time than at any other time in hearth’s
eg. teeth, shells, bones
history

only occasionally do we find impressions of soft

bodied forms or tissues
eg. jellyfish or impressions of skin, feathers or hair

aware of these limitations, paleontologists are adept at

interpreting the past history of life on our planet

Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 11 Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 12
 Classification of Animals assymetrical, radial symmetry, bilateral symmetry

2. degree of tissue and organ differentiation:

Animals can be classified into 34 phyla based mainly
on: embryonic: ectoderm, mesoderm, endoderm

adult: epithelial, connective, muscle, nervous

1. overall body form:
sac like 3. nature of body cavity or coelom
tube like
segmented this is the actual space within the body in which most
solid body or with body cavity organs are found

2. level of complexity of structure acoelomate

cellular, tissue, organ, organ systems no body cavity – most primitive

a. some have no true tissues or organs pseudocoelom – space, but missing tissue layers

true coelom
b. some have some tissues but few or no organs
additional tissue layers enclosing the coelom allow
c. simple organs vs relatively complex organ for much more elaborate developent of skin
and internal organs, circulatory system and
systems nervous sytem

3. complexity of the nervous system and sense 4. presence or absence of segmentation

organs
(=metamerism)
eg. distinct head with sense organs, mouth, etc

eg. nerve net, nerve cords and ganglia or brain and spinal cord serial repetition (segmentation) of body parts is an ancient
feature of animal design
4. developmental and life cycle characteristics
5. cephalization
Additional characteristics used to help classify animals: the formation of and the development of a nervous
system
1. type of symmetry:
leads to sense organs and mouth at anterior end of animal
Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 13 Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 14

Brachiopoda (Lampshells)
Animal Classification Ectoprocta (bryozoa)
Deuterostomes
Echinodermata (starfish & sea urchins)
Cellular level of organization: Chaetognatha (arrow worms)
Porifera (sponges) Hemichordata
Mesozoa Chordata
Placozoa Urochordata
Tissue level of organization: Cephalochordata
Vertebrata
Cnidaria (Jellyfish & corals) Agnatha
Ctenophora (comb jellies) Chondrichthyes
Osteichthyes
Organ level of organization: Amphibia
Acoelomates Reptilia
Aves
Acoelomorpha Mammalia
Platyhelminthes (flatworms)
Nemertea (ribbonworms)
while animals are classified into 34 different body
Pseudocoelomates plans or phyla
Nematoda (roundworms)
Rotifera
Gnathostomulida (jaw worms) 95% of all species belong to only ~8 different
Micrognathozoa phyla
Gastrotricha
Kinorhyncha
Loricifera the animal kingdom can also be considered as 2 great
Priapulida groups:
Nematomorpha (horsehair worms)
Acanthocephala (spiny-headed worms)
Cycliophora invertebrates vs vertebrates
Entoprocta
Eucoelomates 95% of all animals are invertebrates
Protostomes
Mollusca (clams & snails)
! animals without backbones
Annelida (segmented worms)
Arthropoda
Sipuncula (peanut worms) represented by all 34 phyla
Echiura (spoon worms)
Onycophora (velvet worms)
Tardigrada (water bears)
5% of all animals are vertebrates
Phoronida
Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 15 Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 16
 ! animals with backbones

found in only one phylum: Chordates (including

vertebrates: fish, amphibians, reptiles, birds &
mammals)

Origin & Evolution of Animals; Ziser Lecture Notes; 2010.9 17

 Taxonomy and Classification
Taxonomy
= the science of naming and describing species

“Wisdom begins with calling things by their right names”

-Chinese Proverb

museums contain ~ 2 Billion specimens worldwide

about 1.5 M different species of life have been

described

each year ~ 13,000 new species are described

most scientists estimate that there are at least

50 to 100 Million actual species sharing our planet
today

most will probably remain unknown forever:

 the most diverse areas of world are the most remote
 most of the large stuff has been found and described
 not enough researchers or money to devote to this work

Common vs Scientific Name

many larger organisms have “common names”
 but sometimes >1 common name for same
organism
 sometimes same common name used for 2 or
more distinctly different organisms
eg. daisy eg. moss
eg. mouse eg. fern
eg. bug

Taxonomy and Classification, Ziser Lecture Notes, 2004 1

without a specific (unique) name it’s impossible to
communicate about specific organisms

What Characteristics are used

how do we begin to categorize, classify and name all

these organisms

there are many ways to classify:

form
color
size
chemical structure
genetic makeup

earliest attempts used general appearance

ie anatomy and physiological similarities
plants vs animals
 only largest animals were categorized
everything else was “vermes”
 but algae, protozoa

today, much more focus on molecular similarities

proteins, DNA, genes

History of Classification
Aristotle
was the first to try to name and classify things based on
structural similarities

described ~520 species of animals; esp around Greece

Taxonomy and Classification, Ziser Lecture Notes, 2004 2
 believed a name should reflect hidden reality or essence

used phrases or single words, not consistent

Theophrastus classified plants into herbs, shrubs and trees

after invention of the printing press in 1400’s “Herbals” were printed

they emphasized plants with medicinal uses

Carolus Linnaeus,
a Swedish botanist,

developed and published the first comprehensive and consistent

classification system for both plants and animals:

plants: Species Plantarum 1753

(described and classified all
plants known in his time
= 7300 species)

animals: Systema Naturae 1758

Linnaeus

categorized and classified ~8000 different plants and

~10,000 animals

(including 828 mussels & molluscs; 2100 insects; 4777 fish, birds &
mammals)

‘God designed life; Linnaeus did the filing’

offered the first comprehensive, consistent

and much simpler method of naming and
organizing species into a collection:
Taxonomy and Classification, Ziser Lecture Notes, 2004 3
 1. emphasized morphological characteristics as
the basis for arranging specimens in a
collection

2. each species is given a unique scientific name

 while some species can have 100’s of common names
each has only one binomial name

3. each unique name is a binomial

binomial name: Genus + species epithet

before – species consisted of up to 12 words

eg. tomato
was: Solanum caule inerme herbaceo foliis
pinnatis incisis racemis simplicibus
(= Solanum with a smooth herbaceous stem,
incised pinate leaves and simple
inflorescence)
Linnaeus: Solanum lycopersicum

“God Created, Linnaeus arranged”

most names are descriptive, latinized or greek forms, often

to honor someone

but there are also some irreverent names:

eg. Verae peculya  arthropod (?insect)
eg. Heerz lukenatcha  arthropod (?insect)
eg. Pison eyvae  arthropod (?insect)
eg. Agra phobia  beetle
eg. Phthiria relativitae  fly

sometimes “overdo” the descriptive value of a name:

eg. Gammaracanthuskytodermogammarusloricatobai
calensis
Taxonomy and Classification, Ziser Lecture Notes, 2004 4
  was later disapproved

4. species were arranged in an ascending series of

inclusive categories or ‘taxa’

ie a hierarcy:
kingdom
phylum
class
order
family
genus – a typical genus contains about 10-12 species
species

in this classification scheme only “species” is a

real category

the species is the basic unit of classification

the only real unit
 higher taxa are purely mythical creations to
help us understand relationships between
organisms

and sometime change as our knowledge of

the group increases

eg. “lumpers and splitters”

biological species concept:

species = group of organisms capable of
interbreeeding
but fossils
asexual reproduction

Taxonomy and Classification, Ziser Lecture Notes, 2004 5

 ‘type’ species is collected and described

when a new species is discovered a representative sample is

collected and used as the “type”

this becomes a permanent part of a museum collection

it is the type that is described and named

if there are any future questions it can be examined further

when Linnaeus 1st proposed his system

Evolution had not yet been formally proposed

after Darwin and Wallace

Taxonomy took on a new role
was also used to reflect evolutionary
relationships

systematics = determining phylogeny of a species

phylogeny = evolutionary relationships

based on phenotypic similarities and differences

the original hierarchy was originally used just to categorize

degrees of similarity between organisms

after Darwin it was interpreted to show phylogenetic relationships

 to construct evolutionary trees

1. look for anatomical and physiological similarities

2. compare with similar fossils that might exist
3. study its embryological development
4. ecological characteristics

Taxonomy and Classification, Ziser Lecture Notes, 2004 6

 eg. classes older common ancestor
eg. genera younger common ancestor

New criteria and terminology for classification:

Primitive vs Advanced
Primitive  more similar to hypothetical ancestor
Advanced  considerable change from ancestor

eg. magnolia vs orchid

eg. scorpion vs spider
eg. bat or primate vs mouse or shrew

Generalized vs Specialized
Generalized  one structure can be used for lots of different
things
Specialized  structure is modified to perform a specific
function

eg. insect ovipositor to lay eggs

becomes specialized to sting, drill holes,
parasitize hosts, etc
eg. leaf for photosynthesis
becomes specialized as thorns, tendrils, food or water
storage, flower parts, etc

Homologous vs Analogous
Homologous  structures with same origin
implies evolutionary relationship

eg. bird wing, human arm, whale flipper

Analogous  structures with similarity in function and

sometimes appearance
with no evolutionary connection

eg. bird wing, insect wing

eg. octopus eye, human eye

Taxonomy and Classification, Ziser Lecture Notes, 2004 7

 but analogous structures can indicate
convergent evolution

eg. cacti in America, euphorbs in Africa

eg. placentals in west, marsupials in australia
eg. insect wing arose several times in evolution

Taxonomy and Classification, Ziser Lecture Notes, 2004 8

Numerical Taxonomy

studies and opinions on these criteria are used to

construct phylogenetic trees

but each decision is a value judgement

 some traits get more emphasis or more
importance than others

numerical taxonomy is an attempt to remove some of

the subjectivity

uses a large number of characteristics

all characteristics have equal value

the more features shared between organisms the

more closely related they are assumed to be

Cladistics

an even more specific method of comparison

tries to remove even more of the subjectivity

uses only homologous structures in comparing

orginisms
 completely new traits are not used

each change in “primitive” structure is given equal

weight
Taxonomy and Classification, Ziser Lecture Notes, 2004 9
those with the most changes in the structure are the
most distantly related

Molecular Taxonomy
more recently, molecular evidence is used
provides a more objective way to determine
relationships

the variations in structure of proteins or genes on

strand of DNA can be used to calculate how close of
a relationship there is between several organisms

can also be used to estimate how long ago new groups

evolved

generally parallels other information

but sometimes provides new insight into

evolutionary relationships

Taxonomy and Classification, Ziser Lecture Notes, 2004 10

Phylogeny

each species is included in a hierarchy of classification

each level of the hierarchy is more general than the

one below

each level = a taxon

before Darwin
species names were given based on
eg. physical characteristics or
eg. to honor a researcher in field,
eg. location where found, etc
classification into higher taxa was by their
structural similarities (morphology)

after Darwin classification became closely tied to

evolutionary relationships (= phylogeny)

each taxon must be monophyletic

 all members must be from same original
ancestor

Phylocode [www.ohio.edu/phylocode]

a new type of taxonomy seeks to more closely reflect

these phylogenetic relationships in the naming of a
species

propose renaming many species to reflect evolutionary

Taxonomy and Classification, Ziser Lecture Notes, 2004 11
 position

an alternative to traditional binomial name:

a species name might be shortened or

hyphenated with its former genus name

or given a numerical designation

instead of being grouped into ranks such as genus,

family, order, etc
 organisms would be assembled into “clades”

clade = any set of organisms with a common ancestor

Taxonomy and Classification, Ziser Lecture Notes, 2004 12

 Introduction to Ecology
as far as we know; only earth harbors life
one of the most basic characteristics of life is that
living organisms are constantly interacting with
their environment and with each other
can’t study the diversity of life on earth without
knowing something about a few ecological
principles that affect them
even on earth, life is contained within a thin veneer
near the earth’s surface
Biosphere:
=the total living world and all aspects of the
nonliving environment with which life directly
interact
from ~ 7 miles (11,000M) below the surface of ocean
to ~ 6 miles (9,000M) above sea level
eg. if earth were the size of an apple the biosphere would
be the thickness of its skin
these general conditions have molded animals
along similar patterns & designs throughout
earth’s history
Animal Ecology, Ziser, Lecture Notes, 2012.4
! also freezing kills because sharp ice crystals
expand and break the cells open
yet some animals are able to survive below
freezing as long as they can keep some of the
water liquid
water is kept liquid by
! high salts (eg. making ice cream or salt
melting snow ! keeps water liquid below
freezing temperatures)
eg. there are a few brackish pools in Anarctica
where water remains fluid at –5º F (-20.6º C )
! “antifreeze” compounds like glycerol (car
antifreeze) or other alcohols
eg. numerous insects survive freezing in tundra by
producing glycerol in their cells
eg. human, after 7 hrs exposure to freezing
temperatures whose body temp fell to 77ºF, were
revived; such survivors were often drunk
but high temperatures can also be dangerous
some of the large, complex organic molecules
(proteins and DNA) are very sensitive to
higher temperatures:
!above ~ 120ºF (50º C) proteins are destroyed and
Animal Ecology, Ziser, Lecture Notes, 2012.4
yet in this range, each kind of animal has developed a
different set of tolerances for a variety of
environmental conditions
eg. Temperature of liquid water
most animals are adapted to temperatures between
68-104ºF (20º-40ºC)
life requires liquid water
all life is made mostly of water
eg. humans 60-70 %
life is basically a series of chemical reactions
=metabolism
and you can’t have chemical reactions unless the
chemicals are dissolved in liquid
!no water no metabolism
however, some animals can temporarily stop
metabolism when there is no water yet still
survive
eg. tardigrades, nematodes
frozen water is the same as no water at all
! there can be no metabolism
1 Animal Ecology, Ziser, Lecture Notes, 2012.4 2
cell membranes begin to break down
!DNA melts above 150ºF (65-75º C)
yet some fish are able to survive in hot springs
and some marine invertebrates thrive at hot
thermal vents
eg. Salt /Water Concentration at ~3%
the relative concentration of salt in water is referred to
as its salinity
most animal cells have a salinity of 3%
in environments with too little or too much salt
animals will die without special adaptations
eg. marine animals generally can’t tolerate
freshwaters and freshwater animals generally
cannot tolerate ocean water
the salinity of the open ocean is a very constant 3%
invertebrates that live in the ocean usually have
no problems with salinity variations since their
bodies are the same salinity as the ocean
water
marine vertebrates (mainly fish) have cells that
are slightly less salty than seawater
3 Animal Ecology, Ziser, Lecture Notes, 2012.4 4
 ! since they naturally absorb lots of salts they
must continually remove salts (usually
from gills or kidneys) to survive
very few animals can live in extremely high salt
concentrations such as those found in the great
salt lake or the dead sea
eg. brine flies
animals that live in freshwaters and on land often
have trouble getting salt
eg. freshwater fish have perfected the ability to conserve salts
from the food they eat while constantly getting rid of excess
water
eg. large land animals such as deer tend to congregate at
natural “salt licks”
hunters put out salt to attract deer
eg. pH Range near Neutral
pH is a measure of the balance between acids and
bases
examples of acids: lemon juice, carbonated drinks, coffee,
battery acid, etc
examples of bases: bleach, drain cleaner, oven cleaner
cement,
a pH of 7 is neutral
!above 7 means more bases, fewer acids
Animal Ecology, Ziser, Lecture Notes, 2012.4
only a few animal species live regularlyabove
22,000 ft elevation
the main effect of higher pressures (deep in the
ocean) is felt on gasses trapped in lungs and air
sacs of animals
! too much pressure pushes extra gasses into blood
nitrogen narcosis
! too quick of a return to lower pressures produces gas
bubbles in blood
deep diving mammals can collapse their lungs to
prevent these problems
Ecosystems
variations in the above factors (and others),
throughout the biosphere produce distinctive
sets of environmental conditions and results in
distinctive living communities
ie, ecosystems
an ecosystem is a portion of the biosphere with
similar environmental conditions supporting a
characteristic and distinctive group of species
other terms for ecosystems: biomes, ecoregions,
life zones
different ecosystems are usually easily recognizable
Animal Ecology, Ziser, Lecture Notes, 2012.4
!below 7 means more acids, fewer bases
the fluid inside most animal cells is near neutral
like salts, too much or too little acidity can affect large
organic molecules
eg. destroys proteins
eg. humans body fluids pH=7.4; <7.0 and >7.8 ! death
a few fish can tolerate pH’s near 4, but cannot
reproduce in such environments
eg. acid rain in northern lakes kills fish
animals can rarely grow in environments above pH of
10.5
eg. Pressure Range Near 1 atm
most animals live at a pressure near that at sea level
but mountain and deep sea pressures vary greatly
the main effects of lower pressures (at higher
elevations) relates to the amount of O2 available
to air breathing animals
! too little pressure ! not enough O2
eg. the lowest pressure humans can survive is about 1/5th
of an atmosphere (~22,000’ above sea level)
!would become starved for oxygen
5 Animal Ecology, Ziser, Lecture Notes, 2012.4 6
eg. rainforest, desert, tundra, etc
the study of these distinctive patterns throughout the
biosphere is called “ecology”
ecology: the study of organisms’ interactions with
their environment & with each other
General Kinds of Ecosystems
all the world’s ecosystems can be grouped into just a
two broad categories that share many similar
characteristics that life in them must adapt to:
A. Aquatic Ecosystems (~73% earth’s surface)
B. Terrestrial Ecosystems (27% earth’s surface)
A. Aquatic Ecosystems
water based
most stable overall (most of ocean is 2º C)
buoyancy of water reduces need for support
less oxygen in water than in air
!larger animals need more efficient extraction
eg. gills
7 Animal Ecology, Ziser, Lecture Notes, 2012.4 8
 eg. streams, rivers, lakes, ponds
heavy dependence on chemical senses and ability to
detect vibrations in water (<2% earth’s surface = less than the area of Europe)

water is an ideal medium for reproduction more variable in temperature, amount of light,
spawning nutrients, etc than marine
motile larvae for dispersal
doesn’t require internal fertilization
very few salts in water
two different kinds of aquatic ecosystems:
1. Marine FW systems are disproportionately rich in species and
2. Freshwater disproportionately imperiled
FW ecosystems encompass <2% of earth’s surface
Marine Ecosystems
!they contain 12% of all animal species
eg. oceans, seas, bays, estuaries, intertidal shores,
deep trenches ! including 41% of all fish species

oceans dominate the biosphere: but a much greater proportion of fw species are now
endangered, threatened or at risk
71% of area of earth; 99% of volume of biosphere
eg. 20-36% of all fw fish species
salt concentration roughly the same as most cells eg. 67% fw clams
eg. 64% crayfish species
eg. 35% amphibians
! no need for salt/water regulation
[compare to terrestrial:eg. 17% of mammals; 11% of birds
are at risk]
most of the ocean gets NO light from the sun
B. Terrestrial Ecosystems (~27% earth’s surface)
algae, seaweeds, and blue green bacteria
are only found in the upper layers eg. forests, marshes, deserts, rainforests, savannahs, praries,
etc
organic material rains down from above
harshest, most variable environment
Freshwater Ecosystems
Animal Ecology, Ziser, Lecture Notes, 2012.4 9 Animal Ecology, Ziser, Lecture Notes, 2012.4 10

animals living on land must be resistant to drying or animals must be able to get sperm cells to egg
be able to store water cells

oxygen more freely available animals must find mates

eg. air contains 20x’s more oxygen than water
! often more elaborate mating behaviors
but respiratory organs must be protected inside than in aquatic animals
body to avoid drying out
Community Interactions
eg. lungs, book lungs, tracheae
in addition to interactions between the living and
on land animals need considerably more support nonliving parts of an ecosystem, there are also
numerous interactions between the living
water is 800x’s more dense than air organisms themselves
water is harder to move through but does buoy
plants vs herbivores
up the body
the largest animals that ever existed are aquatic animals some animals have very specfic food needs

land animals need strong skeleton & muscular eg. Panda eat only bamboo
system to get around eg. many insects eat only a certain species of plant

land based life must adapt to extreme changes in predators vs prey

temperature throughout the seasons
symbiosis = when two organism are usually found
water fluctuates little in temperature
together and are interdependent on each other
ocean temperatures are constant
not the same as predator/prey or food chain relationships
land has harsh cycles of freezing and drying
is a closer more specific kind of interdependence
on land reproduction become more complicated

Animal Ecology, Ziser, Lecture Notes, 2012.4 11 Animal Ecology, Ziser, Lecture Notes, 2012.4 12
ALL living organisms including all animals form
symbioses with other animals and other lifeforms ie. the other is harmed in some way
eg. tapeworms, liver flukes, fleas, ticks, etc
Kinds of Symbioses
a. mutualism
b. commensalism
c. parasitism

a. Mutualism

both organisms benefit from the relationship

eg. symbiotic algae in corals and sponges

eg. protozoa in gut of termite

eg. some gut bacteria protect us from disease and

pathogens

b. Commensalism

one organism benefits, the other neither benefits

nor is harmed (neutral effect)
eg. follicle mites

eg. many gut bacteria

c. Parasitism

most common form of symbiosis

eg. 20-50% of all animal species are parasitic

one organism benefits at the other’s expense

Animal Ecology, Ziser, Lecture Notes, 2012.4 13 Animal Ecology, Ziser, Lecture Notes, 2012.4 14
 Introduction to Evolution eg. same species of plant or insect may have 1 generation in
northern part of its range or 2, even 3, generations in the
Evolution is the process by which nature selects, southern part of its range

from the genetic diversity of a population, those traits eg. virtually every bacterial pathogen has become at least
that would make an individual more likely to survive and somewhat resistant to antibiotics over the past 60 years

reproduce in!a continuously changing environment. ! over time, these populations may change in their
appearance and other visible characteristics and
Over many years and many generations the full will surely change in their genetic structure
diversity of life on earth is expressed.
eg. many unrelated species often adapt in similar was when
Evolution is one of the most fundamental organizing subjected to the same environmental conditions

principles of the biological sciences and as such is

the single most dominant theme in biology today over long periods of time these changes could be
significantly different from what you started with
evolution stresses the relatedness of all life
rather than its differences yet, no one has ever witnessed the origin of a
major new animal or plant group
! it provides a framework (=unifying principle) ! takes 10,000’s or millions of years
for the way that we study and understand the
living world we do however have an increasing amount of fossil data
that shows the evolution of one species from another, step
by step
! it’s a way of bringing together many diverse
aspects of life’s tremendous complexity and today with molecular techniques we can actually observe
and measure the rate of evolution in many species today

Adaptation vs Evolution there is no controversy surrounding evolution within the scientific

community itself;
One of the “characteristics of life” is that organisms
adapt to their environment as it changes from year the “controversy” is fabricated by those who seek to inject
to year nonscientific beliefs into a very powerful scientific concept

eg. same species of plant adapts to dryer conditions in one part

of its range and wet conditions in another
Genetics & Evolution: Introduction to Evolution & Natural Selection, Ziser, Lecture Notes, 2010.6 1 Genetics & Evolution: Introduction to Evolution & Natural Selection, Ziser, Lecture Notes, 2010.6 2

The Theory of Evolution by eg. in Darwin’s time scientists were beginning to

realize that the world was much older than
Natural Selection previously thought
the theory of evolution was developed by Charles a. before Darwin the accepted age of the earth was
Darwin, in the mid 1800’s, after a lifetime of determined by James Ussher (1581-1656) & John
travel, observation, experimentation and Lightfoot (1602-1675)
discussion made assumption that the Bible was the only reliable
source of chronological information for the time
in his 3 year voyage on the Beagle, he collected and catalogued covered in biblical writings
1000’s of plants and animals and made numerous
observations arrived at the calculation that the earth was created on
Sunday, October 24, 4004 BC
Darwin collected copious notes on species variations and their
relationship to fossil forms Lightfoot, making additional assumptions put the time
at 9:00 am
he also studied breeds of domesticated animals and plants and
pondered how we could produce such variations by so the earth was believed to be ~6000 years old
selective breeding
b. in the next century, Comte de Buffon (1707-1788;

eg. Dogs today consist of >300 breeds “Histoire Naturelle”, 1749) believed he could get an
! all were created by humans within the last estimate of the age of the earth based on its
200 years rate of heat loss

eg. cats, cattle, sheep he calculated the age of the earth as 74,832 yrs
eg. corn, brassicas (and the origin of life at 40,000 yrs)

= human directed “evolution”: humans did the selecting he also recognized 6 geological periods
instead of nature
much of western science at this time was still
if humans can do it in 100’x or 1000’s of years dominated by Church beliefs and he was heavily
surely nature can do it given Millions of years pressured by the Church to reconsider his
calculations
many of Darwin’s ideas were stimulated by an
explosion of new scientific information his solution: “this is what one might think if one did
not know what genesis says”

Genetics & Evolution: Introduction to Evolution & Natural Selection, Ziser, Lecture Notes, 2010.6 3 Genetics & Evolution: Introduction to Evolution & Natural Selection, Ziser, Lecture Notes, 2010.6 4
 c. by Darwin’s time geologists were beginning to realize The Theory of Evolution by Natural Selection
that the earth was 100’s of millions or even billions of
years old
at its core is a relatively simple idea:
eg. paleontologists were learning that fossils were
representatives of previous forms of life from the a. all living things consist of a unique
ancient past combination of chemicals organized in
unique ways
much earlier, fossils were thought of as “sports of nature”

by 1700’s most scientists believe that fossils were of organic ! variations occur in every species
origin
no two individuals of a species are alike
but most were explained in terms of the Biblical flood
b. species’ populations are able to adapt to
as geologists were realizing the extreme age of the earth that gradually changing environments
that would mean that fossils trapped in these ancient layers
were also millions of years old
the same species in different parts of the world
eg. previous biologists had already suggested that have different tolerances and slightly
different characteristics to survive the local
all species are interrelated
conditions in which it lives
species change through time
eg. live oak in Austin, vs live oak in Baton Rouge
and the environment is a factor in that change
eg. flower and gardening catalogues vs local growers
Jean Baptiste de Lamark (1809) produced the first
“evolutionary tree” to illustrate “change through time” still they are the same species:

but he could not offer a reliable explanation or “mechanism” they interbreed naturally where they come nto contact
for how these processes could occur
c. Most of these variations have a genetic basis

! they can be passed on to their offspring

Darwin was not aware of Mendel’s work, He didn’t know

HOW traits were passed on, just observed that some
were
Genetics & Evolution: Introduction to Evolution & Natural Selection, Ziser, Lecture Notes, 2010.6 5 Genetics & Evolution: Introduction to Evolution & Natural Selection, Ziser, Lecture Notes, 2010.6 6

variations are selected over 100’s or 1000’s of

took another 50-60 yrs before hereditary information was
added to Darwin’s original theory
generations new forms will arise

! made it even more powerful

Additional evidence supporting evolutionary
d. each species produces more offspring than theory
will survive into maturity
1. today the layers of rock can be accurately
eg. if not, 1 bacterial cell ! 36 hours would cover earth 3-4
ft deep dated by strata and by radioactive decay
methods
eg. fruit fly ! in 7 months would produce enough offspring
to equal the mass of the earth
we can see that species have been altered
over geologic time
e. those individuals whose variations best fit
their environment will be more likely to
the fossil record shows clearly that all
survive and reproduce
organisms did not appear at the same time
fittness = ability to reproduce
many that once existed have become
extinct
organisms with less favorable variations will be
less likely to survive ~99% of all life that ever existed on earth is now
extinct
"There is a “struggle for existence”
the fossil record also shows there has been an
orderly sequence species change and
" with “survival of the fittest”
replacement over billions of years from the
simplest forms of live to the most
f. by a process of natural selection, evolution sorts
complex:
through these numerous variations within a
population and “chooses” the most fit combination 3.5 BY bacteria
1.5 BY eucaryotes
as the environment slowly changes and certain 500 MY animals
400 MY plants
190 MY 1s t flowering plants
Genetics & Evolution: Introduction to Evolution & Natural Selection, Ziser, Lecture Notes, 2010.6 7 Genetics & Evolution: Introduction to Evolution & Natural Selection, Ziser, Lecture Notes, 2010.6 8
 65 MY modern plants, primates the more closely related an animal is the more
15 MY hominids
similar its biochemistry:
2. also, we have found fossils of many eg. DNA
“intermediate forms” between major groups in eg antibodies
the fossil record: eg. protein structure

eg. between fish and amphibians eg. nematode worm shares 40% of its DNA with us

eg. between dinosaurs and birds eg. chimpanzees and humans share 98% of their DNA

eg. evolutionary stages of the horse, elephant, etc eg. all humans share 99% of our DNA

eg. human ancestors eg. closer relatives 99.5%

in a few cases we have essentially every major we can even quantify the degree of difference
step in the evolutionary process from one and the evolutionary timelines for virtually
species to another all forms of life
eg. 1 snail species into 2 in So American Lake
sediment (year by year evolution) virtually everything we know about the natural world
biology
3. we have learned the science of genetics and geology
can explain how mutations occur and how they chemistry
are passed on (this process was completely physics
unknown to Darwin) astronomy

contribute to our current understanding of the

4. In modern times we have added a massive
process of evolution
amount of molecular evidence that supports
evolutionary theory

similarities and differences in biochemistry

correlates with assumed evolutionary
relationships

Genetics & Evolution: Introduction to Evolution & Natural Selection, Ziser, Lecture Notes, 2010.6 9 Genetics & Evolution: Introduction to Evolution & Natural Selection, Ziser, Lecture Notes, 2010.6 10
 What is Science?
Many people think of science as a bunch of facts, a
giant dictionary
but it is more: a process, a way of looking at the
world; a way of uncovering knowledge
there are many ways to uncover knowledge and to
better understand our world and ourselves:
music
art
poetry
philosophy
theology
history
scientific investigation
etc
each these disciplines can provide valid
information
all teach us something important
but each addresses different questions or different
aspects of a question
eg. science can show us how to clone animals & plants,
how to improve crops, how to produce stem cells, etc
eg. philosophy & religion can highlight the moral and
ethical implications of those advances and help us to
decide whether we should really be doing these things
or not, whether we have a right to do them
Defining Science –What is Science, Ziser, Lecture Notes, 2011.3
to use scientific methods correctly we must make
certain assumptions and follow certain rules
otherwise we are NOT “doing science”
1. explanations must be guided by Natural Laws
the world works as a natural machine
mechanism (Descarte, 17th century)
mechanistic view !no “vital spirits”
reductionism: complex processes can be reduced to
simpler components
In science, we assume that everything is
reducible to simple understandable
natural explanations:
body functions
smile of a baby
even a persons philosophical and religious beliefs
(eg temporal lobe seizures)
there is no room in science for magic or
supernatural
eg. life or health is not the result of “vital forces”
eg. ”creation science” is an oxymoron
eg. “intelligent design” assumes unnatural or
supernatural forces control the universe, it is
therefore not a scientific concept
Defining Science –What is Science, Ziser, Lecture Notes, 2011.3
eg. politics, government tells us how we can make fair laws
to protect all who are faced with such difficult decisions
eg. music and art can help to express our emotional
connections to the world
all are just different languages or ways to describe
the world
science is one process of understanding how the world
works
science is a language, a tool, a method for searching
for knowledge
(but only one of many sources of knowledge)
we use them all and need them all to produce a well
rounded understanding or our world
Each method of describing the world requires that we
follow a specific set of assumptions or criteria:
music ! tonal qualities, tempos, harmonies
art ! mix of colors, lines, perspective, depth
philosophy ! criteria of logic, inductive and deductive
reasoning
eg flower, a sunset, our “place” in the world
1 Defining Science –What is Science, Ziser, Lecture Notes, 2011.3 2
2. explanations must be falsifiable
scientists begin by making hypotheses
(=educated guesses) about what is going
on
scientific hypotheses are not necessarily more
likely to be true
but it must be able to be disproven
or its not a scientific explanation
eg. observation: my car wont start
possible hypotheses:
battery is dead
out of gas
aliens zapped car with an energy
sucking ray
the last one is NOT a scientific hypothesis
statements below are NOT valid subjects of
scientific investigation because they
cannot be disproven
eg. your neighbor insists that he was abducted by
aliens
eg. a friend believes that her astrological sign
accurately describes her personality or
future events
eg. the belief that “God created the universe”
all explanations must be testable by
3 Defining Science –What is Science, Ziser, Lecture Notes, 2011.3 4
 experimentation
an experiment is essentially an attempt to
disprove your hypothesis
eg. virtually ALL strides in medical treatments
have come from scientific knowledge
! from testing and retesting various
hypotheses or explanations
almost all rigorous testing of alternative
medical practices have shown them to
be ineffective
3. the process of science requires logical
reasoning
“human nature” generally tends to cause most
people to actually form an opinion first and
then collect information that seems to
support or make sense of the idea
eg. there are today people who firmly believe that the
earth is flat or that it is only 6000 years old
despite massive amounts of data that disprove
thess concepts
eg. politics: we tend to like or hate someone and see
all that happens through that lens. So much so
that we essentially dehumanize that person, and
its often hard to resist even when we know better
experimentation requires careful unbiased
observations
Defining Science –What is Science, Ziser, Lecture Notes, 2011.3
a given question may have 1000’s of
logical answers but only one is correct,
therefore the chances are high that a
random guess will be wrong
eg. observation: my car wont start
hypotheses: battery is dead
wiring is fried
out of gas
someone stole the
distributor cap
must choose each one at a time and test it
until you
c. experiments must be rigorously controlled
! but must be aware of your assumptions
4. statistical analysis is a powerful tool to sort
through large numbers of observations
(data) and likely conclusions
but must be used correctly; one can lie with statistical
methods that are misused
eg. from huge amounts of data collected we can
logically conclude that consumption of
pickles is dangerous & can cause:
cancer
war
communism
auto accidents
Defining Science –What is Science, Ziser, Lecture Notes, 2011.3
! can’t set out to “prove” something
eg. if you already believe in bigfoot or flying
saucers you are biased and cannot make
very effective scientific observations
eg. Loch Ness assumes something is there then
builds elaborate theory to “explain” data or
lack of data
eg. Bermuda Triangle
percentage of vessels lost in triangle are no
higher than anywhere else
but percentage of false losses are much
higher for this area
it is very difficult, even for scientists to
eliminate bias; to keep an open mind
a. one way to reduce bias is that the same
observation must be repeatable by
others
eg. one (or a few) persons claim of UFO or
abduction is not a scientific observation
! testamonials are not scientific data
when many people see the same UFO and
their observations are unrelated in any way
and can produce unambiguous physical
evidence then UFO’s will be accepted as
scientific fact
b. in designing an experiment it is best to
pose question (hypothesis) as an
“either/or” statement
5 Defining Science –What is Science, Ziser, Lecture Notes, 2011.3 6
data:
those born in 1839 who ate pickles
have sufferend 100% mortality
rate
rats force fed 20 lbs of pickles a day for
a month ended up with bulging
abdomens, poor health and loss of
appetite
99.9% of cancer victims had eaten
pickles sometimes in their lives
100% of all soldiers
96.8% of communist sympathizers
99.7% of those involved in car
accidents
5. the simplest logical explanation is the
preferred one (=Occam’s Razor)
Extraordinary hypotheses require
extraordinary evidence
eg bigfoot, ufo’s, astrology, etc
6. scientific explanations must have predictive
value
eg. if an apple falls to the ground today, you can
predict that it will do so tomorrow and the day
after
the scientific method is a self-correcting
process
it is the ONLY source of knowledge that is self
correcting
scientists think like everyone else but with
more awareness of the possibility that they
may be wrong!
7 Defining Science –What is Science, Ziser, Lecture Notes, 2011.3 8
 versus a politician who must act like he can never
be wrong !they are supported by massive amount of
or theologian who accepts a concept on faith
evidence
alone, with no experimentation
a theory is a scientific fact
7. the strongest explanations that have survived
numerous attempts to disprove them its as close to the truth as a scientist can get
become theories
but we haven’t “proven” a thing
A theory is an hypothesis that has been
repeatedly verified, not disproven !nothing has been (is ever) proven
in science
the word “Theory” in general use is used for any idea
someone has on a topic; a hunch
but only need one disproof
in science an hypothesis is a “hunch”
theories do not last indefinitely
a theory is an hypothesis that has been repeatedly tested,
1000’s of times, in many different ways by many
different people and the results always support the most theories have a very short life span as
hypothesis; never disprove it. originally proposed
we sometimes hear the rationalization for not valuing a
scientific theory that “its just a theory”
! they are only temporary truths

but then; few survive more than a couple of years

without being revised or discarded
its just a theory that the sun rises every morning

its just a theory that the heart pumps blood in
the body
its just a theory that penicillin can be used to
effectively treat some bacterial infections
theories are not speculation
Defining Science –What is Science, Ziser, Lecture Notes, 2011.3
as more and more tests are done the theory is
revised, remolded, “tweaked” to be even
more precise and more accurate
but only rarely does the entire theory get
discarded
9 Defining Science –What is Science, Ziser, Lecture Notes, 2011.3 10

eg. spontaneous generation are usually discovering new factors that might
eg. earth centric solar system
influence a particular outcome

they don’t necessarily become incorrect, just sometimes a theory becomes so powerful at
obsolete for some situations predicting that it becomes a natural law

as theories are refined they become more !implies there are NO exceptions
inclusive and are able to make stronger
eg. an apple always falls to the ground
predictions until the ‘50’s anyway

replacing a theory that is wrong with one that is more

“subtly wrong”
! one exception even a “law” must be altered
or trashed
eg. “theory of agriculture”
eg. “Law of Gravity” became just a small part of a
seed sprouts more complete “Theory of Relativity”
seed sprouts better with water
seed sprouts better under soil with water Science can never explain everything
add manure
prevent disease
root nodules only those things that can be investigated using
mycorrhizae correct assumptions
etc.
eg. science will never be able to:
as theory is refined it becomes more and more
accurate at predicting future events define right and wrong?
why we are here?
eg. Heredity is there life after death?
something in cell transmits traits to offspring (Mendel) is there a “god”
it is in nucleus
it is in chromosomes
its either proteins or DNA these are not scientific questions
its DNA
specific genes identified they are best studied in other disciplines
when we find exceptions to a current theory we

Defining Science –What is Science, Ziser, Lecture Notes, 2011.3 11 Defining Science –What is Science, Ziser, Lecture Notes, 2011.3 12
 The Animal Kingdom à each year ~ 13,000 new species are
described
Animals are the largest most diverse ‘kingdom’ of
life on earth à some of the most diverse areas on
earth are just beginning to be
all life on earth can be categorized into 5 major investigated
groups
eg. vertebrates
although more recent information has now expanded that to
6 to 9 major groups; for our purposes we’ll consider 5 here birds: 3 new bird species/yr described

over 2 million individual species have been fish: estimate only 40% of those in South
identified so far in all life America have been described

eg. invertebrates
known species in each group:
insects: ~700 new species of insects are
# of species described each year
Bacteria 10,000
Protista 65,000 eg. of 19 trees in one Panama study 1200
Fungi 100,000 species of beetles were collected and 80% of
Plants 300,000 them were new species
Animals 1,700,000
eg. marine invertebrates of the deep ocean
Animals cleary dominate
only 1.5% of the deep ocean have been
investigated
à over 3/4ths of all know species belong in
animal kingdom on an area the size of a parking lot off New
England coast
Estimates of the total number of living animal
à 800 different species of inverts found
species, known & unknown, are difficult to
make recently discovered whole new kind of
community of 100’s of new species of
organisms
yet all indications are that there are many
more species yet to be discovered:
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 2

marine biologists estimate there may be up Animal Records

to 10M invertebrate species in the deep
ocean alone
A. Largest Animal
à there are probably ~25 x’s more animal
species that have not yet been named =longest
than those that have been named longest animal in existence is a nemertean
= 60 M (180’) long
Also, all animals alive today comprise <10% of all
"Lions Mane" jellyfish à ~150’ long.
animals that have ever lived
= tallest living animal
perhaps 500 Million species of animals have
lived on this earth in the last billion years giraffe, Giraffa camelopardalis à 19’ (5.8 m)

Even in terms of total numbers of individuals, =most massive

animals are second only to bacteria
a. blue whale, Balaenoptera musculus, would be bigger in
eg. estimate 10,000 Trillion ants on planet every other dimension (weight, girth, displacement)

à all the earth’s ants weigh ~ same as all earth’s eg. Mature blue whales typically measure anywhere
human population from 75 feet (23 m) to 100 feet (30.5 m) from
head to tail
eg. there are more animals on earth than stars in our
galaxy and can weigh as much as 150 tons (136 metric
tons).

The largest blue whale on record is a 110’ female that

weighed 195 tons (177 tonnes).

b. largest of all reptiles: Seismosaurus hallorum

("Earth-shaking lizard")

120+ feet long (37 m); 30-80 tons

à largest animal ever to have walked on land

b. the sauropod dinosaur, Argentinosaurus, weighed

Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 3 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 4
 about 90 tons (82 tonnes).
àThat's little more than half the size of an adult blue
whale.
It makes a lot of sense that the world's largest
animal would be a sea creature.
Land animals have to support their own weight, whereas
sea creatures get some help from the water.
c. The largest land animals today are male Savannah
Elephants, up to 11’ tall with one known example
weighing around 7.25 tons(6.7 tonnes)
d. Largest land mammal ever was Baluchitherium
B. Smallest Animal
some unusual invertebrates are microscopic being made up of
<100 cells
C. Longest Lived Animal
Confirmed: The ocean quahog (Arctica islandica), a type of
clam, lived in the deep water off iceland before being
dragged onto a ship, age was confirmed by growth rings on
shell
Invertebrates (unconfirmed estimates):
eg. biochemical and nuclear evidence indicates that some
sea urchins may live up to 200 years
eg. some crinoids may live for 1000’s of years
eg. some sea anemones are essentially immortal, can live
1000’s of years
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9
What exactly is an Animal?
1. ALL animals are multicellular organisms
the CELL is the basic unit of life
à ALL living things are composed of cells
an organism can consist of one or more cells:
unicellular = solitary cell
eg. bacteria, some protists
multicellular = always consist of many
cells
different cells have become specialized for
different functions
cells are interdependent on each other
àcells never found alone
eg. most fungi, all plants, all animals
animals show the best development of
multicellularity of all kingdoms
2. Most animals are motile; they move around in
some way; walk, run, swim, climb, dig, etc
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9
Cold Blooded Vertebrates:
eg. Chilean sea bass is claimed to live over 100 yrs
eg. Orange roughy up to 150 years
eg. A Madagascar radiated tortoise (Geochelone radiata)
à ~188-192 years
presented to the Tongan royal family by the British
explorer Captain Cook in either 1773 or 1777, died in
1965, lived to the age of at least 188 years old! The
animal was called Tui Malila.
eg. A Giant Galapagos Tortoise Harriet, collected in
1835, lived in captivity until she died in 2006
at about 177 years old.
Warm Blooded Vertebrates
eg. some Macaw birds may live up to 100 years
eg. unconfirmed report of a bowhead whale living to be 245
yrs old; dated from stone and metal harpoon points
found in the whales & chemical testing
eg. primates are generally the longest lived as a group
and man is the longest lived of the primates: 122
years (Jeanne Louise Calment, 1875 - 1997).
(mammals with shortest lifespan are tiny shrews: maybe 1
to 1.5 years.
àThe faster you live, though, the shorter you live)
5 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 6
only a few, mostly simple animals are sessile;
they don’t move around
eg. corals, barnacles, etc
but even they are motile at some point in their life cycle
3. most animals have true tissues, organs and
organ systems
4. ALL animals are heterotrophs
a few marine species carry chloroplasts in their skin cells
and can do photosynthesis
à but these chloroplasts are taken from the algae they eat
a newly discovered species of sea slug actually contains some of
the genes for photosynthesis in its own genome
à but still needs to steal chloroplasts to do photosynthesis
all life requires a set of nutrients
à chemicals that are essential to keep the
organism alive
these nutrients may be used as building
blocks or as an energy source
nutrients are used as building blocks to
make cells, tissues, skeletons, blood, etc
organisms also need an energy source to
“power” everything that an organism does
7 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 8

the direct source of a cell’s energy is the
breakdown of organic molecules (mainly
sugar)
à ie. they use chemical energy
energy is stored in “chemical bonds”
when you break bonds you release energy
organic molecules have lots of bonds and store
lots of energy
ie. they don’t need organic molecules
in their diet
eg. plants, algae and some bacteria
5. Most animlas produce energy by aerobic
respiration
(all but 3 newly discovered animal species)
sugars and organic molecules can be broken down
for energy with or without O2 gas.

organisms have developed two basic ways to get producing energy without the use of O2 is called
these energy molecules: anaerobic respiration and is common in
bacteria and some fungi
Heterotrophs
aerobic respiration extracts much more energy
=organisms that must eat organic from organic food
molecules and then break them down
eg. anaerobic respiration of sugar à 2 units of energy
for energy
aerobic respiration of sugar à 36-38 units of energy
eg. us and other animals (19x’s more)

Autotrophs
= organisms that extract energy from
sunlight or some other source to make
organic molecules
then they can break them down as
needed for energy
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9
some animals can use anaerobic respiration
temporarily or in specific tissues such as
muscle tissue but primarily rely in aerobic
respiration for most energy
recently, 3 new species of loriciferans have been found to
completely lack mitochondria and get all their energy
from anaerobic respiration
6. most animals store extra energy as fats or oils
9 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 10

most efficient way to store energy for active 10. most animals show complex development, with
animals extended embryonic phase, often with
free living larval stages
eg. twice as efficient as storing extra energy as starch as
plants do
all organisms (even bacteria) pass through a
7. most animals have a head with distinct sense characteristic life cycle
organs and some kind of brain
changes in size and shape
8. most animals have outgrowths, processes or eg. Frog: embryoà larvaàadult
appendages for sensory functions, collecting food eg. Human: embryoàfetusàjuvenileàadult
eg. Butterfly: embryoàcaterpillar(larva)àpupaàadult
&/or for movement eg. Fly: embryoàmaggot(larva)àpupaàadult
eg. Clam: embryoàlarva(glochidium)àadult
eg mouthparts
antennae some involve alternation of two completely
tentacles
fins
different body forms; one that reproduces
legs asexually, one sexually
wings
the immature forms may be self sufficient or
9. most reproduce both sexually and asexually completely dependent on mother for
nourishment and protection
asexual: exact copies, clones
eg. embryos, larvae, nymphs, etc
easier, quicker, don’t need a mate
the life cycle often involves metamorphosis of
sexual: unique combination of traits one free living form into another

produces variation which allows in some animals the embryonic or immature stage
adaptation and evolution is a dormant or resistant stage

involves specialized sex cells: 11. behavior is an important tool for animal survival
sperm and egg
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 11 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 12

behavior plays a major role in how they interact
with their environment and with each other
behaviors play a major roll in food gathering,
social interactions, mating and care of young
all behavior has a genetic basis
à follows Darwinian evolution to some degree
predictable
programmed
adaptive (reproductive advantage)
behaviors are either:
Instinctive
taxes
reflexes
fixed action patterns
mimicry, camoflage
the most basic type of behavior:
stimulus à response
may or not be aware of the stimulus
stimulus may be internal or external
à perceived by sensory organ or cell
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9
behaviors in invertebrates are usually highly rigid,
stereotyped, patterns
àalmost all are genetically preprogrammed
in more complex animals (vertebrates) learning
plays a larger role
12. Probably some of the most unique chemicals
produced by animals are a huge variety of
venoms
venomous creatures are found throughout the
animal kingdom
estimated 100,000 venomous animals from
corals and sponges to spiders, scorpions,
snakes, etc
each carries its own unique coctail of toxins,
mostly, various kinds of proteins
13. all living organisms (including animals) constantly
adapt to their environment in many ways through
physiological, anatomical or behavioral changes
some adaptations can be in the form of changes in
growth and physiology
eg. fur color of the same species may vary across habitats
adapting to temperature or amount of sunlight
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9
response is controlled or modified by nervous or
endocrine system
Taxes à response to single stimulus by
motile organism
eg. attraction to light, food, etc
Reflexes àmore complex but unlearned,
unmodifiable response in organisms with
well developed nervous systems
involves a complete functional circuit
of nervous system:from receptor to
effector
or Learned
imprinting
eg. blinking as a reflex arc
habituation
eg. touching hot skillet
conditioning
social:
courtship Learned Behaviors --> from simple to very
reproductive complex
family
group eg. predator avoidance
eg. abstract thought & planning
but what is learned vs. innate
eg. Newborns don’t blink when object is brought close
to their eyes
à learned
à maturation of pathways for reflex
eg. right or left thumb on top when folding hands
13 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 14
eg. same species of bird may have harder stronger beak in
one habitat than another depending on type of food
available
some adaptations involve behavioral changes and
learning
eg. tool use among animals
14. adaptations also occur across generations as the
most useful traits are selected for and passed on
while harmful traits are eliminated
à =evolution by natural selection
eg. same species in different ecosystems
eg. reproductive displays
eg. niches and evolution à marsupials in australia
15 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 16
Animal Cells eg. lysosomes à cellular digestion

animals are generally much more active, have a

animal cells are “eukaryotic” cells
much higher metabolism than the members
of other kingdoms
much larger, much more complex,
they therefore require much more energy than
most genetic material is contained in prominent
species in any other kingdom
nucleus
almost all this energy is the product of
lots of internal and surface structures including
aerobic respiration inside mitochondria
organelles
àanimal cells have many more
animal cells lack cell walls mitochondria than those of other
organisms
the thin flexible cell membrane forms the
outer boundary of each cell animal cells make much more use of proteins for
structure, movement, nerve impulses and
not rigid, but flexible metabolism

also acts as a gateway for things moving into and àmany more ribosomes (protein
out of the cell factories)

organelles and other internal structures provide a

division of labor and allow the cells to work much
more efficiently
eg. mitochondria à energy factories

eg. vacuoles à storage containers

eg. ribosomes à energy factories

Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 17 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 18

Animal Tissues
1. epithelial
single celled organisms like protozoa are “jacks of all
2. connective
trades”
move
feed 3. muscular
produce energy
respond to stimuli 4. nervous
reproduce
etc
1. Epithelial Tissues
in multicellular organisms such as animals the large
size and complexity dictates that individual cells this is the most primitive animal tissue
specialize àpresumably the first true animal tissue to evolve.

rather than performing all tasks each group of It forms the outer coverings of animals
cells becomes specialized to do one or a few
tasks very well ( much more efficiently): lines the inner and outer surfaces of all organs.
eg. muscle cells à contraction & movement
eg. bone cells à support
consists of cells fitted tightly together
eg. blood à circulation
Functions of Epithelial Tissues:
while they become more efficient at one or a few
specific jobs, they lose their independence and 1. protection
can no longer exist on their own
from microbes, physical injury, water loss, etc

tissues are groups of similar cells performing similar

2. absorption
functions
of food, water etc in the intestine
includes cells and any secretions (=matrix) they
produce 3. transport
sometimes have cilia (tiny hairlike processes)
There are 4 basic animal tissues: that move things along a tube
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 19 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 20
 the nature of the matrix and the fibers it
eg. oviducts à move egg toward uterus
contains identifies the specific kind of
eg. respiratory tract à move dust and bacteria connective tissue
out of lungs
fibers composed mainly of collagen
4. filtration
a uniquely “animal” protein
in kidneys

(esp. skin, tendons, ligaments, cartilage)

5. gas exchange
lungs Functions of Connective Tissue:

6. secretion 1. Glue
eg. form glands that secrete various substances eg. areolar tissue

eg. mucous, sweat, digestive juices 2. Support & movement

2. Connective Tissues eg. bone & cartilage

the most widespread and abundant type of tissue 3. Nutrient Storage

in animals eg. bone, adipose

the most diverse in structure and function 4. Temperature Homeostasis

most connective tissues are heavily vascularized eg. fat for heat production and cold insulation

5. Transport
connective tissues have an abundance of matrix
eg. blood, lymph
àthe noncellular matrix often comprise the majority of the
tissue volume 3. Muscle Tissues

elongated cells, spindle shaped, up to 1 ft long

Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 21 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 22

= muscle fibers
4. Nervous Tissues
highly contractile and elastic cells
nerve cells are also elongated into long fibers
all cells contract to some degree, but muscle
cells are much stronger and contract much typically large cell body with one or more long
more efficiently fibers extending from it
eg. our calf muscles can support 1 ton
grouped together to form extensive interconnected
muscle cells generally stop dividing at birth network of “wires” that extend throughout the
body
(# fixed at birth)
nerve cells are able to conduct impulses to send
but each cell can expand greatly in volume signals throughout the animal body

Functions of Muscle Tissue: Functions of Nervous Tissue

1. movement 1. to sense internal and external

environmental changes,
both voluntary movements such as
swimming or running 2. coordination and control of muscles
and glands.
and internal involuntary movements of
various organs such as the pumping Stem Cells
heart, and peristalsis of the digestive
organs. Most adults retain some kinds of “embryonic
cells” called stem cells
2. posture
can later differentiate into replacement cells
3. heat generation and tissues.

used for movement

Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 23 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 24
 Animal (Including Human) à in some skin is hardened to offer support
Organ Systems àin some animals the skin color is important in
The greater specialization of cells and tissues behaviors:
communication
increases the efficiency by which animals can camoflage
carry out life’s basic processes and allows for reproductive behaviors
almost limitless opportunities for evolutionary etc
variations and adaptations to numerous kinds of
many animals can quickly change the color of their
habitats and environmental conditions.
skin
à chromatophores
to understand and appreciate the complexity of
different kinds of animals we’ll focus on the most 2. Skeletal System
familiar (and most complex) animal à US
especially terrestrial animals
1. Skin (integumentary system)
(in land plants support was also an important
outer covering of the animal consideration à cellulose & lignin)

(plants also have outer covering, epidermis, but it’s much 3 main kinds of support system in animals:
simpler in structure and function)
exoskeleton
àin some animals (usually relatively small
ones) is a simple covering that allows on the outside à especially good protection
food, gasses and waste products to easily (eg. clams, snails, insects)
secreted by the skin
diffuse in and out of the animal grows at edges (clams & snails)
or must be shed periodically for growth
à in terrestrial animals it may serve as a (insects and other arthropods)
waterproofing layer to keep animal from
drying out endoskeleton
internal à grows with the body
à skin often contains various sense organs eg. vertebrates including us

Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 25 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 26

hydrostatic skeleton à animals are much more active than any

muscles of body wall control fluid pressure other kingdom
eg. most worms, jellyfish, octopus

General Functions of muscles:

eg. human skeleton(endoskeleton)
1. movement
is made of over 200 bones
most animals are motile
grows continuously throughout life
à very active tissue à walk, run, crawl, swim, fly, climb, etc
à recycled every ~7 years
a few animals are sessile
subdivided in axial (skull, vertebrae, rib cage)
& appendicular skeleton (arms & legs) but even these have internal muscles
that circulate blood, move food
functions in through digestive tract, etc

1. support some muscle are voluntary

strong and relatively light; 10% body weight
some are involuntary
2. movement
framework on which muscles act
àact as levers and pivots 2. Heat Production

3. protection important for warm blooded animals like

some of our most delicate organs are well
us
protected by being encased in bone
à muscle generate lots of heat
eg. brain, lungs, heart, reproductive system
warm blooded vs coldblooded
3. Muscular System
all animals alive today except birds and mammals
are “cold blooded”
unique to animals:
bird & mammals are warm blooded
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 27 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 28
 à much more active in most animals the organs of digestive system
à require much more food to maintain
heat production
form essentially a long continuous tube that
is open at both ends
4. Digestive System
à alimentary canal (GI tract)
like fungi, and many protists and bacteria, animals are
heterotrophs à take in organic food mouthà pharynxà esophagusà stomachà
small intestineà large intestineà anus
animal food needs are much more complex
near the beginning of the system food is
physically and chemically digested
digestive system functions to break down the food
so that it can be absorbed and used by the eg. typically the mouth is armed with the appropriate
body tools to rip and tear the food into smaller pieces

most animals digest the food after it is eaten, eg. the stomach and beginning of the small intestine
produces enzymes and other chemicals to break
not before as in fungi or some plants large proteins and starches into smaller molecules

but a few (eg. spiders) predigest their food the rest of the system is used to absorb the
nutrients released by digestion and to get
lots of specialization in structures depending on rid of undigestible wastes
how an animal gets its food & what kind of
food it prefers eg. most absorption occurs in the small intestine

a few things (water, alcohol) can be absorbed by

eg. predator, herbivore, parasite, filter feeder, fluid feeder
the stomach; and the large intestine can absorb
additional water and nutrients released by
in some animals the digestive system is a simple bacterial action
sac, opened at one end
our small intestine is greatly modified for absorption
surface area is greatly increased for more
à food is eaten, digested and the wastes efficient absorption of nutrients:
are “spit out” the mouth
1” diameter x 10’ long
eg. corals, jellyfish, flatworms à if smooth tube= 0.33 m2 (3 sq ft)

Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 29 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 30

but: interior is folded aquatic animals

also has fingerlike projections = villi &
microvilli Gasses diffuse much slower in water than in air

àTotal Area = 200m2 (1800 sq ft) water contains 20 times less oxygen than air
à aquatic organisms must have more efficient
in us, once the nutrients are absorbed they go to the respiratory systems
liver for processing and storage
high surface area provided by gills, book
5. The Respiratory System gills, etc

like plants, all animals require O2 to produce numerous flaps or feather-like

energy and release C02 as a waste product structures exposed on the sides of
the animal
oxygen gas is needed as a nutrient;

carbon dioxide gas is a waste product of must keep water moving across gills
respiration (energy production) àgills in constant motion
àwater is constantly pumped over
since animals are more active than plants they gills
require more efficient ways to get oxygen
(plants just used simple pores: stomata or lenticels, air breathers:
or pneumatophores)
easier to extract O2 from air
Respiratory system functions as this gas exchange air contains 20 times more air than water
system in animals
but air dries respiratory surface
in very small animals there is no specific
“organ” àrespiratory organs must be
à breath through their skin protected and kept moist

air breathing animals have different lungs, trachea, book lungs, etc
requirements than those that extract oxygen
from water
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 31 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 32
 often the respiratory system is closely associated =blood vessels: arteries, capillaries, veins
with some kind of circulatory system to more
effectively collect and distribute the oxygen “pumps”
= heart
eg. Human lungs
can be “open” or “closed” system:
some of the most efficient
open system (eg. insects)
à lots of area for gas exchange fluid sloshes around in body cavity
pumping heart keeps fluid in motion

contain millions of microscopic alveoli

surrounded by capillaries closed system (vertebrates; us)
2 2
total surface area ~ 70M (=760 ft ~20’x38’)
blood flows in closed system of vessels
6. Circulatory System over 60,000 miles of vessels (mainly capillaries)

the circulatory system is the major connection arteries à capillaries à veins

between external and internal environment
7. The Endocrine System
àeverything going in or out of body must go
through the circulatory system to get to virtually all multicellular organisms use chemicals
where its going to coordinate activities and communicate

in small organisms gas exchange and food and wastes
enter and leave by simple diffusion
in large, multicellular organisms some kind of
circulatory system is needed to move things
around
typically, the circulatory system consists of
“plumbing”
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9
in animals, chemicals (= hormones) are used to
help control long term activities such as
growth, development, reproductive cycles, etc
virtually all organs produce various hormones but
in some organs hormone production is their
main job
eg. thyroid gland, pituitary gland, pancreas, etc
33 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 34

O2
8. Nervous System
many of an animals coordination is hard wired into
animals are much more active than members of circuits that produce predictable responses to
the other two multicellular kingdoms stimuli

animals move much more quickly, must reflex = a rapid, automatic, predictable motor
respond to things much quicker response to a stimulus
chemicals may take minutes or hours to unlearned
unplanned
produce a response involuntary
à “hard wired” into our neural anatomy
animals need a system to control quick reactions:
movements, emergencies, etc Kinds of Nervous Systems

à only members of the animal kingdom have animal nervous systems range from very
an additional systems of control simple to increasingly complex:

all major animal groups except sponges have eg. Nerve net:
some kind of nervous system
no brain

cells of the nervous system are highly specialized simple coordination of swimming or
for receiving stimuli and conducting impulses feeding movements
to various parts of the body eg. jellyfish & corals

made up mainly of neurons eg. Ganglia and nerve cords

long thin fiber like cells up to 4 ft long
very minimal “central processing”
very high metabolic rate (highest of any cells in body)
nerve cords can be paired; dorsal, ventral,
require glucose, can’t use alternate fuels lateral, etc

require lots of O2 – only aerobic metabolism eg. flatworms, segmented worms, arthropods
can’t survive more than a few minutes without
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 35 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 36
 eg. True brain and spinal cord à receptors convert one form of energy
into nerve impulses that the brain can
in vertebrates only
interpret
nervous system is organized into 2 major
subdivisions: our body has millions of sensory receptors
à some we are consciously aware of
CNS: brain and spinal cord
PNS: cranial nerves and spinal nerves à most are internal, and help maintain body
at an unconscious level

9. The Senses kinds of transducers:

monitor and allow organism to respond to its photoreceptor - light

chemoreceptor - chemicals
environment mechanoreceptor - bending, pressure, touch
thermoreceptor - temperature
senses provide direct contact between animal and osmoreceptor – salt/water conc
its surroundings nocioceptor (“to injure”) – pain

no animal is completely aware of its environment
à only selectively aware
eg. those that live in caves depend more on smell and
sound
eg. those that live on surface of land rely heavily on sight
10. Excretory System (Urinary System)
excretory wastes = metabolic wastes
à chemicals & toxins produced by cells during
metabolism

eg. those that live in water use smell, currents and all organisms must get rid of excess materials and wastes
vibrations fungi, protists, bacteria à diffusion;
plants àstomata, converted to “secondary plant
sensory receptors are transducers products” for defense or support or stored
in woody tissue)

information presents itself in different energy having greater metabolism, animals generate
forms more wastes

Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 37 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 38

à need more effective way to get rid of only major human system that doesn’t work
wastes continuously

à only activated at puberty

main job of excretory system is to collect and
eliminate toxic wastes

may also help to stabilize salt and water balance

in body

12. Reproductive System

most animals reproduce both asexually and

sexually

à most vertebrates reproduce only sexually

animals typically go through more complex stages

of development

sometimes spending years in immature forms

some animals go through an alternation of

generations

contains ovaries and testes for sexual

reproduction

sometimes contains organ for development of

young
Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 39 Animals – Introduction to Zoology; Ziser, lecture notes, 2016.9 40