07/03/2023

Gnathiferans and Smaller

Lophotrochozoans
Chapter 15

Pseudocoelomates

⚫ Pseudocoelomates have a body cavity

(the pseudocoel) between the gut
(derived from endoderm) and body wall
(derived from mesoderm).
⚫ Derived from the blastocoel.

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Advantages of a Body Cavity

⚫ A body cavity, pseudocoel or coelom,

has several advantages. It provides:
⚫ Greater freedom of movement.
⚫ Space for development of organ systems.

⚫ A simple means for circulation of materials

around the body.
⚫ Storage place for waste products.

⚫ A hydrostatic organ.

Common Features of
Pseudocoelomates
⚫ All have a body wall of epidermis, a
dermis, and muscles surrounding the
pseudocoel.
⚫ Almost all have a complete digestive
tract.
⚫ The epidermis of many secretes a
nonliving cuticle with bristles & spines.

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Pseudocoelomates

⚫ Pseudocoelomates do not form a clade.

⚫ Some are part of superphylum
Lophotrochozoa, others are in
superphylum Ecdysozoa.
⚫ All share the pseudocoelomate body plan.

Lophotrochozoa (10 Phyla)

⚫ Ancestors possessed complex cuticular jaws – Clade
Gnathifera
⚫ Gnathostomata
⚫ Micrognathozoa
⚫ Rotifera
⚫ Acanthoephala
⚫ 6 other lophotrochozoan phyla
⚫ Gastrotricha
⚫ Tiny aquatic animals that may be closely related to gnathiferans
⚫ Molecular characteristics place the following with
Lophotrochozoa
⚫ Cycliophora
⚫ Entoprocta
⚫ Ectoprocta
⚫ Brachiopoda
⚫ Phoronida

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Clade Gnathifera

⚫ Possess small cuticular jaws with a

homologous microstructure.
⚫ Numbers of pairs of jaws vary.
⚫ Gnathostomulida, Micrognathozoa, and
Rotifera are tiny, free-living, aquatic
animals.
⚫ Acanthocephalans are worm-like
endoparasites living as adults in fish or
other vertebrates.

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Clade Gnathifera

⚫ Rotifera and Acanthocephala

⚫ Presumed sister taxa.
⚫ Form a clade called Syndermata.

⚫ Have eutelic syncytial epidermis.

⚫ Constant number of nuclei.
⚫ Grouping is controversial.

Phylum Gnathostomulida

⚫ Phylum Gnathostomulida includes the jaw

worms.
⚫ Very small - <2mm.
⚫ Live in interstitial spaces of fine coastal
sediments.
⚫ Can endure low O2.

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Phylum Gnathostomulida
⚫ Feed by scraping bacteria and fungi from the
substratum with a pair of jaws on the pharynx.
⚫ Acoelomate with a poorly developed parenchyma
layer.
⚫ Sexual stages include males, females, and
hermaphrodites.
⚫ Fertilization is internal.

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Phylum Micrognathozoa

⚫ Micrognathozoans are tiny animals that

live interstitially (between sand grains).
⚫ Body consists of a two-part head, a
thorax, and abdomen with short tail.
⚫ Move using cilia and have a unique
ventral ciliary adhesive pad that
produces glue.
⚫ Three pairs of complex jaws.

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Phylum Micrognathozoa
⚫ Simple gut
⚫ Anus opens to outside
only periodically.
⚫ Reproductive system is
not well understood:
⚫ Only female reproductive
organs have been
identified.
⚫ May reproduce
parthenogenetically.
⚫ Cleavage and subsequent
development have not
been studied.

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Phylum Rotifera

⚫ Members of the
phylum Rotifera are
pseudocoelomate
and have three
embryonic germ
layers
(triploblastic).
⚫ Complete digestive
system.

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Phylum Rotifera
⚫ Dioecious (separate sexes)
but some species
parthenogenetic (females
produce diploid eggs).
⚫ Some are parthenogenetic
during part of the year,
depending on environmental
conditions.
⚫ Thick shelled eggs that can
withstand harsh conditions
are sometimes produced.

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Phylum Rotifera

⚫ Rotifers have a ciliated crown, the corona,

that is characteristic of the phylum.

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Phylum Rotifera
⚫ Rotifers come in a wide
range of colors and
shapes.
⚫ Shapes often
correspond to lifestyle
(floaters, swimmers,
sessile).
⚫ They may be individual
or colonial.
⚫ Mostly freshwater.
⚫ Benthic and pelagic
forms.

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Phylum Acanthocephala

⚫ All spiny-headed worms (Phylum

Acanthocephala) are parasites in the
intestines of vertebrates.
⚫ Over 1100 species known.
⚫ Occur worldwide and parasitize fish,
birds, and mammals.
⚫ Larvae develop in crustaceans or
insects.

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Phylum Acanthocephala

⚫ Proboscis has rows of recurved spines that

penetrate and may rupture host intestines.
⚫ Proboscis with hooks can be inverted into a
proboscis receptacle by retractor muscles.

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Phylum Acanthocephala

⚫ Body somewhat flattened.

⚫ About 80% of tegument is a lacunar
system of fluid-filled canals that may
distribute nutrients and remove wastes
from muscles.
⚫ No heart - function provided by lacunar
fluid.
⚫ Both longitudinal and circular body wall
muscles are present.

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Phylum Acanthocephala

⚫ No respiratory system.
⚫ Protonephridia with flame cells, if
present, perform excretory functions.
⚫ Nutrients are absorbed across the
tegument, which bears some enzymes -
no digestive tract.
⚫ Dioecious
⚫ No species normally parasitizes humans.

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Phylum Acanthocephala

⚫ Acanthocephalans penetrate the

intestinal wall with spiny proboscis.
⚫ Remarkably little inflammation on host wall,
but pain of infection is intense.
⚫ Larval acanthors burrow through beetle
intestine.
⚫ Develop into juvenile cystacanths in the
insect hemocoel.

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Phylum Gastrotricha
⚫ Gastrotrichs appear similar
to rotifers, but without the
ciliated corona and have a
bristly looking body.
⚫ Members of the phylum
Gastrotricha are
pseudocoelomate and
have three embryonic germ
layers (triploblastic).
⚫ Complete digestive system.
⚫ Hermaphroditic or
parthenogenetic.

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Phylum Entoprocta

⚫ About 150 species in the phylum

Entoprocta occur worldwide.
⚫ Usually in marine environments.
⚫ Less than 5 mm long and mostly
microscopic, resembling hydroid
cnidarians.

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Phylum Entoprocta
⚫ Urnatella gracilis is a common freshwater species
in North America.
⚫ Body or calyx is cup shaped and bears a circular
crown of ciliated tentacles.
⚫ Attaches by a stalk with adhesive glands.

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Phylum Entoprocta

⚫ Tentacles (3-30) and stalk are

continuations of the body wall.
⚫ Tentacles on lateral and inner surfaces
can roll inward but cannot be retracted
into the calyx.
⚫ Gut is U-shaped with both mouth and
anus opening within the circle of
tentacles.

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Phylum Entoprocta

⚫ Long cilia on sides generate current bringing in

particles.
⚫ Short cilia on inner surfaces capture food and
direct it to mouth.
⚫ Pair of protonephridia embedded in gelatinous
parenchyma.
⚫ Well-developed nerve ganglion on the ventral
side of stomach.
⚫ No circulatory or respiratory organs.

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Phylum Entoprocta

⚫ Some are monoecious, some dioecious, and

some first produce sperm and later eggs.
⚫ Fertilized eggs develop in a brood pouch.
⚫ Modified spiral cleavage leads to trochophore-
like larva.

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Lophophorates
⚫ Phylum Ectoprocta
⚫ Phylum Brachiopoda
⚫ Phylum Phoronida
⚫ Belong within the
lophotrochozoan
subgroup of
protostomes.
⚫ Grouping very
controversial.
⚫ Evidence comes from
sequence analysis of
genes encoding small-
subunit ribosomal
RNA.
⚫ Some aspects of
development place
these taxa within
Deuterostomia.

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Lophophorates
⚫ Members of these 3 taxa possess a feeding device
called a lophophore.
⚫ Unique arrangement of ciliated tentacles borne on a ridge or
fold on the body wall.
⚫ Tentacles are hollow and contain an extension of the
mesocoel.

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Lophophorates
⚫ Ciliated tentacles are also a respiratory device
permitting gas exchange between surrounding water
and internal coelomic fluid.
⚫ Gut is U-shaped:
⚫ Mouth opens inside the lophophore ring, and the anus
opens outside the ring.

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Phylum Ectoprocta

⚫ Phylum Ectoprocta contains aquatic animals

that often encrust hard surfaces (bryozoans).
⚫ Approximately 4500 living species.
⚫ Inhabit both shallow freshwater and marine
habitats.
⚫ Most are sessile, some slide slowly, and
others crawl actively across surfaces.
⚫ Mostly colony builders.
⚫ Each member is less than 0.5 mm in length
and is called a zooid.

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Phylum Ectoprocta

⚫ Zooids feed by
extending
lophophores into
surrounding water to
collect tiny particles
⚫ Zooids secrete
exoskeleton in which
they live in.

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Phylum Ectoprocta
⚫ Exoskeleton may be gelatinous, chitinous, or stiffened with calcium
and possibly impregnated with sand.
⚫ Shape may be boxlike, vaselike, oval, or tubular.
⚫ Some colonies form limy encrustations on seaweed, shells, and
rocks.
⚫ Others form fuzzy or shrubby growths or erect branching colonies.
⚫ Freshwater colonies may form mosslike colonies on stems of plants
or on rocks.

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Phylum Ectoprocta
⚫ To feed, the lophophore is
extended and tentacles
spread out into a funnel.
⚫ Cilia on tentacles draw
water into funnel.
⚫ Food particles caught by
cilia in the funnel are
drawn into the mouth.
⚫ Digestion begins
extracellularly in the
stomach and is completed
intracellularly within the
intestine.

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Phylum Ectoprocta
⚫ Respiratory, vascular, and excretory organs
absent.
⚫ Gas exchange is through body surface.
⚫ Ganglionic mass and a nerve ring around the
pharynx.
⚫ No sense organs.

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Phylum Ectoprocta

⚫ Reproduction - most hermaphroditic.

⚫ Some species shed eggs into seawater, but most
brood their eggs.
⚫ Brooding occurs within coelom and some have an
external chamber called an ovicell.
⚫ Sometimes embryos proliferate asexually from the
initial embryo.
⚫ Cleavage is radial but mosaic.
⚫ Larva of nonbrooding species have a functional gut
and swim for a few months before settling.

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Phylum Ectoprocta
⚫ Larva of brooding species do not feed and settle after
a brief free-swimming existence.
⚫ Attach to substratum by secretions from an adhesive
sac, then metamorphose to adult form.
⚫ New colonies begin from this single metamorphosed
primary zooid, called an ancestrula.
⚫ Ancestrula undergoes asexual budding to produce many
zooids of a colony.

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Phylum Ectoprocta
⚫ Freshwater ectoprocts
undergo budding that
produces statoblasts.
⚫ Hard, resistant
capsules containing a
mass of germinative
cells.

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Phylum Brachiopoda

⚫ Brachiopods appear
similar to bivalve
molluscs because they
have two calcareous
shell valves secreted
by a mantle.
⚫ Dorsal/ventral instead
of left/right.
⚫ Pedicel – a fleshy
stalk used for
attachment.

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Phylum Brachiopoda

⚫ Brachiopods are an
ancient group they
were prolific during the
Paleozoic and
Mesozoic eras.
⚫ One living species,
Lingula, is considered
to be a living fossil
since it has changed
little since the
Ordovician (505 mya).

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Phylum Brachiopoda

⚫ Characteristics of both protostomes &

deuterostomes:
⚫ Cleavage is radial (deuterostome)
⚫ Coelom formation enterocoelous at least in
some brachiopods. (deuterostome)
⚫ The relationship of the blastopore to the
mouth is uncertain.

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Phylum Phoronida

⚫ Species in the phylum

Phoronida are small
wormlike animals.
⚫ Secrete tubes to live in.
⚫ Tentacles of the
lophophore are
extended for feeding.
⚫ U-shaped digestive
tract.

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Phylum Phoronida

⚫ Characteristics of both protostomes &

deuterostomes:
⚫ Blastopore becomes mouth (protostome).
⚫ Cleavage is radial (deuterostome).

⚫ Coelom formation – highly modified

enterocoelous (deuterostome).

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Phylogeny
⚫ Analysis of rRNA gene sequencing suggests
that after the ancestral deuterostome diverged
from ancestral protostomes:
⚫ Protostomes split into two large groups:
⚫ Ecdysozoa that molt.
⚫ Lophotrochozoa that exhibit lophophore feeding and
trochophore-like larvae.
⚫ Most lophotrochozoans share some
developmental features:
⚫ Spiral mosaic cleavage and formation of mouth
from embryonic blastopore.
⚫ No common body plan.

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Phylogeny
⚫ Lophotrochozoan protostomes are a heterogeneous
group for which evolutionary branching order remains
to be determined.
⚫ DNA sequence analysis has led to the conclusion that
acanthocephalans are highly derived rotifers.
⚫ Sequences put Acanthocephala and Rotifera together as
clade Syndermata, sharing a eutelic syncytial epidermis.
⚫ Syndermata is placed with Micrognathozoa and
Gnathostomulida in clade Gnathifera.
⚫ Controversial placement of Cycliophora, Gastrotricha, and
Platyhelminthes close to Gnathifera.

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Phylogeny

⚫ Entoprocts and ectoprocts were once

considered both Bryozoa, but ectoprocts have
been broken out as true coelomate animals.
⚫ Sequence analysis now places them both in the
lophotrochozoan phyla.

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Phylogeny

⚫ Ectoprocts, brachiopods, and phoronids share

a lophophore and a tripartite coelom.
⚫ Other features are mixtures of developmental traits
from both protostomes and deuterostomes.
⚫ Debate continues on whether the lophophorates
form a clade or whether the group members, either
individually or collectively belong within Protostomia
or Deuterostomia.

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