STF1053 BIODIVERSITY

LU3:
Species Diversity
PART I

SPECIES

SPECIES are the different kinds of

organisms found on the Earth.
A more exact definition of species is a
group of interbreeding organisms
that do not ordinarily breed with
members of other groups.

SPECIES (cont.)

If a species interbreeds freely with

other species, it would no longer be a
distinctive kind of organism.

This definition
animals.

works

well

with

SPECIES (cont.)

However, in some plant species

fertile crossings can take place
among
morphologically
and
physiologically different kinds of
vegetation.

In this situation, the definition of

species given here is not appropriate.

SPECIES (cont.)

There are several competing theories, or

"species concepts

The
most
widely
accepted
are
the
morphological species concept, the biological
species concept, and the phylogenetic
species concept.

Although the Morphological Species Concept

(MSC) is largely outdated as a theoretical
definition, it is still widely used.

SPECIES (cont.)

According to this concept:

species are the smallest groups that are

consistently and persistently distinct, and
distinguishable by ordinary means.

Cronquist (1978): states that "a species is a

community,
or
a
number
of
related
communities, whose distinctive morphological
characters are, sufficiently definite to entitle it,
or them, to a specific name"

Cronquist, A. (1978). Once again, what is a species? In L.V. Knutson (Ed.),Biosystematics in Agriculture. (pp. 3-20). Montclair, New
Jersey, U.S.A: Allenheld Osmin.

SPECIES (cont.)

The Biological Species Concept (BSC), states

that "a species is a group of interbreeding
natural populations that is reproductively
isolated from other such groups".

According to the Phylogenetic Species Concept

(PSC), is defined a species : "is the smallest
diagnosable cluster of individual organism (that
is, the cluster of organisms are identifiably
distinct from other clusters) within which there
is a parental pattern of ancestry and descent".

SPECIES (cont.)

These concepts are not in agreement,

and considerable debate exists about
the advantages and disadvantages of
all existing species concepts.

In practice, systematists usually

group specimens together according
to
shared
features
(genetic,
morphological, physiological).

SPECIES (cont.)
When two or more groups show
different sets of shared characters,
and the shared characters for each
group allow all the members of that
group to be distinguished relatively
easily and consistently from the
members of another group, then the
groups are considered different
species.

SPECIES (cont.)

This
approach
relies
on
the
objectivity
of
the
phylogenetic
species concept (i.e., the use of
natural, shared, characters to define
or diagnose a species) and applies it
to
the
practicality
of
the
morphological species concept, in
terms of sorting specimens into
groups.

Species Diversity

Species diversity is the number of

different species in a particular area
(species richness) weighted by some
measure of abundance such as number of
individuals or biomass.

However, it is common for conservation

biologists to speak of species diversity
even when they are actually referring to
species richness.

Species Diversity (cont.)

Another measure of species diversity

is the species evenness, which is the
relative abundance with which each
species is represented in an area.

An ecosystem where all the species

are represented by the same number
of individuals has high species
evenness.

Species Diversity (cont.)

An ecosystem where some species are
represented by many individuals, and
other species are represented by
very few individuals has a low
species evenness.

Species Diversity as a Surrogate for

Global Biodiversity

Global biodiversity is frequently

expressed as the total number of
species currently living on Earth, i.e.,
its species richness.
Between about 1.5 and 1.75 million
species have been discovered and
scientifically described thus far
(Cracraft, 2002).

Cracraft, C. (2002). The seven great questions of systematic biology: an essential foundation for conservation and the
sustainable use of biodiversity.Annals of the Missouri Botanical Garden,89, 127-144.

(cont.)

Estimates
for
the
number
of
scientifically valid species vary partly
because of differing opinions on the
definition of a species.

For
example,
the
phylogenetic
species concept recognizes more
species than the biological species
concept.

(cont.)

Also, some scientific descriptions of

species appear in old, uncertain, or
poorly circulated publications.

In these cases, scientists may

accidentally overlook certain species
when preparing inventories of biota,
causing them to describe and name
an already known species.

(cont.)

More significantly, some species are very

difficult
to
identify.
For
example,
taxonomically "cryptic species look very
similar to other species and may be
misidentified (and hence overlooked as being
a different species).

Thus, several different, but similar-looking

species, identified as a single species by one
scientist, are identified as completely
different species by another scientist

(cont.)

Scientists expect that the scientifically described

species represent only a small fraction of the total
number of species on Earth today. Many additional
species have yet to be discovered, or are known
to scientists but have not been formally described.

Scientists estimate that the total number of

species on Earth could range from about 3.6
million up to 7.7 million, with 13 to 20 million
being the most frequently cited range (Cracraft
2002).

Cracraft, C. (2002). The seven great questions of systematic biology: an essential foundation for conservation and the
sustainable use of biodiversity.Annals of the Missouri Botanical Garden,89, 127-144.

(cont.)

Species are grouped together according to

shared characteristics (genetic, anatomical,
biochemical, physiological, or behavioral)
and this gives us a classification of the
species based on their phylogenetic, or
apparent evolutionary relationships.

We can then use this information to assess

the proportion of related species among the
total number of species on Earth.

(cont.)

Most public attention is focused on the

biology and ecology of large, charismatic
species such as mammals, birds, and certain
species of trees (e.g., mahogany, sequoia).

However, the greater part of Earth's species

diversity is found in other, generally
overlooked groups, such as mollusks,
insects, and groups of flowering plants.

Regional patterns of species

diversity
(i) Species diversity or richness
A basic "statistic" of zoogeography is species
diversity or "richness" which describe the
numbers of species within a prescribed area,
without regard to their numerical abundance or
ecological importance in a given community.
There are three basic kinds of richness indices:
1. Alpha diversity or the numbers of species within
a generally small area (e.g., a hectare, sq. km, or
a defined habitat such as a single lake).

(cont.)
2. Beta diversity

the change in the species richness

between
communities
across
an
environmental gradient over a relatively
small distance (e.g., the change across
a salinity gradient or across a mountain
slope). Beta diversity is often estimated
by calculating a species turnover rate

(cont.)

(cont.)
3. Gamma diversity

the numbers of species across a very large area such

as a biome or a continent. Gamma diversity tends to
be a function of both alpha and beta diversity, i.e., it
will trend to be high in areas of high alpha diversity
and particularly across a diversity of habitats.
Gamma diversity is simply the sum of all species
across each area (without double counting shared
species). In the table below, an "X" means that a
species (1-9) is present in one of three areas (Forest,
grassland, and scrub ecosystems).

(cont.)

(cont.)
(ii) The latitudinal gradient in species
diversity
Described by some as the only true
"law" of ecology is the observation
made even by early zoogeographers
that there tend to be more species
(higher alpha, beta, and gamma
diversities) in the tropics than at
higher or lower latitudes.

(cont.)
(iii) What can explain the latitudinal gradient in species
diversity?

The gradient in species diversity is clearly not just an

distinctive characteristic of a few species. Its broad
observance suggests a general shared process that
consistently "produces" more species of any given taxon in
the tropics than the poles.

There have been up to 35 different hypotheses proposed

to explain the gradient in species diversity, many of which
are not mutually exclusive (because many things co-vary
with latitude, e.g., temperature, productivity, etc).

(cont.)

There are, however, several ways in which hypothese

can be grouped. The so-called "mid-domain effect of
Colwell and Hurtt (1994), for example, states that
random distribution of geographic range boundaries
between the "hard edges" of the poles of the earth
results de facto in the greatest range overlaps mid-way
between these edges, i.e., the equatorial regions, and,
consequently, greater species diversity at the equator
than at either pole.

Hubbell's (2001) "neutral theory of biodiversity and

biogeography" is another null model against which
predictions
can
be objectively
tested.
Colwell,
R. K., and Hurtt, G. C. 1994.
Nonbiological
gradients in species richness
and a spurious Rapoport effect.American

Naturalist144:570-595.

Hubbell, S.P. (2001) The Unified Neutral Theory of Biodiversity and Biogeography. Princeton University Press, Princeton, NJ.

(cont.)

Two other general classes of hypotheses:

equilibrium and non-equilibrium hypotheses.
There have been alternative classification
schemes
proposed
(time-based,
habitat
heterogeneity-based, evolutionary-based, etc ).

Equilibrium hypotheses suggest that a "steady

state has been reached among the processes
that influence species diversity and that the forces
that increase diversity are exactly balanced by
those that reduce diversity such that species
diversity remains fixed through time.

Willig, M. R., D. M. Kaufmann, and R. D. Stevens. 2003. Latitudinal gradients of biodiversity: pattern, process, scale and synthesis.
Annu. Rev. Ecol. Syst. 34:273-309.

(cont.)

In other words, if we revisited an area in 100,000 years

it would have the same species diversity (although the
species composition may have changed).

The differences among areas in observed species

diversity simply reflect the different relative
magnitudes of forces that increase or decrease species
diversity in a given area - imagine two containers each
with a hole at a different height from the bottom.
Water enters each container at the same rate, but the
one with the lower hole holds less water (fewer
species).

(cont.)

Non-equilibrium hypotheses involve the idea that a given

community has not yet reached a "steady state and
species diversity is still in the process of increasing (or
decreasing) after some historical disturbance.

In other words, if we came back 100,000 years later, the

species diversity may have changed to reach its "true"
level. For example, Pleistocene glaciations eliminated most
life from North America north of about 46 degrees latitude.
After the ice sheets left, animals and plants re-colonized the
vast area and perhaps the gradient in species diversity is
simply due to the fact that not enough time has elapsed to
allow all species to reach or establish populations in the
newly-exposed habitat.

(cont.)

Such habitats are considered as

"unsaturated" and given enough time
the species diversities will increase to
be the same as at more tropical
latitudes.

Six major sub-hypotheses have been

proposed to account for the latitudinal gradient
1.

Historical shift (non-equilibrium): as

explained above

2.

Productivity: tropical areas are more

productive and there is more
useable energy available to be
subdivided amongst more niches
and
hence
more
species.
Opportunities
for
resource
specialization
increase
species

(cont.)

Note
that
in
the
productive
environment, more species are able
to achieve some critical value of
population
size
although
each
occupies a narrower niches. This way,
more productive environments may
promote greater species diversity by
allowing the persistence of more
species that show a greater degree of
resource specialization.

(cont.)

Although productivity is typically correlated positively

with increase species diversity, it has its limits.

In fact, the productivity-species diversity relationship

is often "dome-shaped", i.e., maximal species
diversity is found in areas of intermediate
productivity, that is, at intermediate latitudes.

Some of the most productive environments on earth

are not the most speciose (rich in species). E.g.,
estuaries, salt marshes, hydrothermal vents, shallow
eutrophic lakes).

3. Harshness:

small isolated or otherwise "harsh environments (like extreme

temperature or moisture) have higher extinction rates, lower
colonization potential, and less opportunity for resource
specialization than more benign tropical environments.

This may explain why some of the most productive

environments are not the most species in many cases, they are
too stressful (e.g., salt marches [salinity stress], hot springs
[heat stress]).

Stressful environments may simply increase extinction rates or

reduce the potential for specialization and place a limit on the
numbers of species that exist in such environments because
only a few possess the requisite adaptations to extreme
conditions.

4. Climate stability

more variable climates prevent resource specialization

and hence are able to support fewer species.

Again, problems with this hypothesis include the many

exceptions. for instance, many regions that are quite
stable (high mountain tops, deep abyssal regions of the
ocean) have low species diversity. By contrast, many
variable environments (e.g., the "dry" and "flood"
seasons in Amazonia) are incredibly species rich.

Climate stability in terms of seasonality of solar

radiation, like the tropics, are more productive

5. Habitat heterogeneity

diverse physical environments promote

isolation, and resource specialization, and
hence speciation, and the coexistence of
more species.

Such relationships have been documented for

desert rodents and some aquatic habitats.
Coral reefs, for instance, are outstanding
examples of complex physical environments
that support some of the highest diversities of
marine species.

6. Interspecific interactions

(2-6 are equilibrium hypotheses): More

species in tropical areas create a positive
feedback through increased interspecific
interactions such as competition, predation
or parasitism.

The intensity of these interactions prevents a

few species from dominating the resources,
promotes specialization, and, therefore,
potential coexistence of more species.

Rapoport's Rule

Rapoport
in
1982
formalized
the
observation
that
subpopulations
of
mammals in North America (i.e., within
species) tend to show larger average
geographic range sizes with increasing
latitude.

This trend has been observed in several

other taxa and is known as "Rapoport's
Rule.

Species Dominance

Also noted in community comparisons is another latitudinal

trend. Tropical areas tend to have more species, but these
species also tend to be rarer, i.e., any individual species
accounts, in terms of numerical abundance, for a lower
proportion of the total individuals summed across all
species in that area, than the average species in more
temperate areas.

In other words, there are fewer species in temperate areas,

but those fewer species tend to be more abundant (a
relatively few species "dominate" the landscape) and they
have larger geographic ranges than those in tropical areas
(where species are more numerous, but each one is less
abundant and has a smaller average range size).

STF1053 BIODIVERSITY
LU3:
Species Diversity
PART II
SPECIES CONCEPT &
SPECIATION

What is a species?
1. Biological species concept

A species is a population or group of

populations whose members have
the potential to interbreed with one
another and produce viable offspring,
but who cannot produce viable
offspring with other species.

(cont.)

Speciation is the process by which a new species

originates and involves the creation of a population
of organisms that are novel enough to be classified
in their own group. There are two processes by
which this can occur:
Anagenesis is the accumulation of heritable traits in a

population, that transforms that population into a new species

Cladogenesis is branching evolution, in which a new species

arises as a branch of from the evolutionary tree. The original

species still exists. This process is the source of biological
diversity

(cont.)

For a new species to form, there

needs to be isolation of some
members of a species as a separate
population. Forms of isolation, that
interfere with breeding include both..

2. Prezygotic and postzygotic

barriers

Prezygotic barriers prevent mating or egg

fertilization if members of different species try to
mate. Examples:

a. Habitat isolation
- Two species that live in the same area, but occupy
different habitats rarely encounter each other.
b. Behavioral isolation
- Signals that attract mates are often unique to a
species (e.g. different species of fireflies flash
different patterns).

(cont.)
c. Temporal isolation
- Two species breed at different times of the day or during
different seasons.
d. Mechanical isolation
- Closely related species attempt to mate, but are
anatomically incompatable. (Example: flowering plants with
pollination barriers; some plants are specific with respect to
the insect pollinator, often occurs with butterflies/moths)
e. Gametic isolation
- Gametes must recognize each other. (Example: fertilization
of fish eggs, chemical signals between sperm and egg
allows sperm to recognize the correct egg)

Postzygotic barriers

prevent a hybrid zygote from developing into a fertile adult.

Examples:

a. Reduced hybrid viability

- Abort development of hybrid at some embryonic stage.
b. Reduced hybrid fertility
- Meiosis doesnt produce fertile gametes in vigorous
hybrids.
c. Hybrid breakdown
- First-generation hybrids are fertile, but they cannot produce
fertile offspring in the next generation (e.g. different
species of cotton).

Alternative concepts of species

a.

Ecological species concept

Species are defined by their use of environmental

resources; their ecological niche (e.g. species that

are defined by their food source such as
butterflies with certain flowers)
b.

Morphological species concept

- Takes into consideration factors such as body
shape, size, etc.

(cont.)
c.

Paleontological species concept

- Species in the fossil record are characterized

according to a unique set of structural features.

d.

Phylogenetic species concept

- Recognizes species are sets of organisms with
unique genetic histories. This idea is based
often on molecular analyses such as DNA
sequences.

Modes of speciation
1. Allopatric speciation
- Allopatric speciation describes speciation that takes place in
populations with geographically separate ranges. Gene flow is
interrupted and new species evolve.

2. Sympatric speciation
-

Sympatric speciation describes speciation that takes place in

geographically overlapping populations. Chromosomal changes
and nonrandom mating reduce gene flow.

Remember: Species arise when individuals in a

population become isolated one from the other.

Modes of speciation (cont.)

Modes of speciation (cont.)

Sympatric speciation

Sympatric
speciation
describes
speciation that takes place in
geographically
overlapping
populations. This can occur by
chromosomal
changes
and
nonrandom mating. Both can reduce
gene flow between organisms and
cause populations to evolve to new
species.