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What determines the distribution and abundance of species in space and time?
All organisms have the inherent potential to increase their numbers exponentially.
However, because resources limit growth, and many environments are unsuitable, no
organisms actually increase indefinitely. This basic principal of biogeography is called as
Malthusian Theory.
The niche concept- the species utilize certain specific resources from the surrounding
environment for their survival,habitat,water. Species niche in the absence of any
interspecific competition and predation is called fundamental niche while species niche in the
presence of competing and/or predator species is its realized niche.
These fundamental and realized niches are reflected in geographic distributions of species
Also, some favorable habitats may be uninhabited just by chance (historical factors)
Additionally, temporal fluctuations in the environment can determine species
presence/absence in a typical habitat.
Biogeographic patterns emerging from speciation & extinction on a dynamic
Earth/biodiversity of a region-
species distribution can be of three types-
1) Cosmopolitan: same/similar species widely distributed around the world.
but no species or family is truly cosmopolitan Certain higher taxonomic levels are nearly
cosmopolitan (e.g. Chiroptera)
2) Endemic: same/similar species occurring in one geographic location and nowhere
else.Endemism actually depends on the taxonomic level and spatial scale. Example-Devils Hole
pupfish near las vegas; mountain lion in southern north america,mid america, and south
America; galapagao tortoise on gelapagao island;Tuatara from islands off New Zealand.
causes of species endemism-
1)Lack of dispersal
2)Shrinking range
3)Influences of historical events
Endemism can be studied with help of Relicts which may be a)Taxonomic b)Biogeographic. both
Usually occur together living fossils
one vareity of endemism is Provincialism which is coincident occurrence of large numbers of
well-differentiated endemic forms in an area.
3) Disjunction- Closely related organisms that occur in separated areas.
Disjunction occur because of 1. Vicariance events 2. Long-distance dispersal 3. Extinction in the
intervening areas
Examples-Nothofagus forest of New Zealand-also found in southern tip of south
americai.e.chile coast; Ratite birds-Rhea (from latin america),Ostrich(from sudan,ethiopia,south
africa),Cassowary(from papua new guinea),kiwi(new zealand),emu( from australia). Distribution
and phylogeny of ratite birds reflect: -plate Tectonics -subsequent Extinctions.
discontinuous range of a taxon in which at least two closely related populations are separated
by a wide geographic distance
2 possible explanations 1. Dispersal 2. Geological events(vicariance)
Biogeographic regions/realms-
It is a region that shares species with similar biogeographic history. It helps to understand
evolutionary and geological factors that contributed to regional patterns of biodiversity
Early biogeographers defined them subjectively (Sclater , Wallace). Modern methods use
mathematical classification techniques (e.g. Simpson Index), phylogenetics .
Major biogeographic regions (Wallace-Sclater)

Recent classification- incorporates phylogenetic analysis and plate tectonics.

Factors determining the distribution of plants and animals-
As per liebig's law of minimum, Population is limited by single most limiting factor,i.e.yield is
proportional to the amount of the most limiting nutrient but is a oversimplified
concept,multiple of factors which determines.
Most Important Factors are -
Physical environment
Biological interactions
1) Physical limiting factors
Saguaro distribution limited by low temperatures
Bark beetles overwintering temperature limits
Climate change
Eastern phoebe winter range (food limited)
B)Precipitation (moisture)
Orographic precipitation and vegetation gradients
Soil Moisture
Xerophytes Full sunlight in dry soils
Mesophytes Wetter and more shaded environments
C) Solar radiation light saturation eg.lightward side of slope will have more vegetations and
variety than leeward side of the slope.
Solar radiation light limitation eg. In tropical rainforest due to thick canopy of tall trees,
sunlight cant reach land surface so many different plant species requiring less sunlight grow
D)Wind--Timberline and wind
Determined by temperature at broad scales
Wind critical local limiting factor.
e) Edaphic factors- soil related factors.
Based on soil structure ,nutrients presence vegetations differes.
2) Disturbance-
A)Fire, volcanism, floods, hurricanes, etc.
Great impact on species distributions by killing individuals, changing resource availability &
environmental conditions , altering species interactions
B)Also biotic disturbances: insect outbreaks, pathogens, herbivore grazing, etc.
Example: Fire suppression in the Patagonian steppe allows Austrocedrus chilensis expansion
3) Biological interactions-
A)Competition (negative-negative)
Competition may be
Exploitative : using resources and therefore making them unavailable for other species
Interference: aggressively denying the use of resources to other species
Diffuse competition: one species is affected by multiple species that collectively diminish a
shared resource
Example-Kangaroo rats in the southwest
B) Predation (positive-negative)
Predation - herbivory-Predator distribution influenced by prey (resource)
Parasitism-Prey distribution influenced by predator/s
Parasitism Alcon blue butterfly larva Fools ants with chemistry by mimicking the surface
chemicals that the ants have on their own brood Reduces ant reproductive output
C) Mutualism (positive-positive)
Can be obligate or nonobligate
Obligate mutualisms have stronger effects on species distributions
Example-mistletoe and its seed dispersers ,Yellow fruits disperser: mockingbird, Green fruits
disperser: marsupial
D) Facilitation (positive-zero or positive-positive)
Presence of one species aids another
Silene acaulis increases plant species richness & abundance
Principle of Competitive Exclusion
Species cannot coexist in a community if their niches are identical
BUT Exclusion takes time Only applies if the ecological factors are constant
4) Interacting factors-
Interacting factors may have more extreme consequences on species distributions than any
factor alone
Most commonly the interaction among multiple factors determine species distributions (and
not a single factor alone)
Relative importance of physical and biological limiting factors-
On harsh physical environments competition is less important than on resource rich
environments as a factor affecting species distributions.
5) Vicariance Vs dispersal-
Vicariance geographical range of a taxa is split into parts by the formation of a barrier
Events such as plate tectonics, orogeny , climate change, sea level rise or glacial movement
split populations
After vicariance events, evolutionary divergence can give rise to different (but related) species
in the disjunctions
1)Ecological dispersal movement of individuals away from an existing population or parent
organism Reduce intra-specific competition But, habitat similarity decreases with distance
2)Biogeographic dispersal species shift their ranges by moving over long distances across
large barriers (broader spatiotemporal scales) Infrequent but very important Mostly historic
Why disperse?
Reduce intraspecific competition Find more suitable habitats
Mechanisms of dispersal
Active: movement of an organism from one location to another by its own means Passive:
movement of an organism from one place to another by means of a stronger force, such as
water flow, wind or another organism
examples-Active dispersal - plants- Himalayan Balsam,Squirting cucumber, Monarch
Butterfly,Golden Plover,Ruby-throated hummingbird
Passive dispersal-by wind,Diaspore is a plant dispersal unit (consists of seed plus additional
tissues adapted for dispersal) e.g.Pappus,Wings,
Exozoochory seeds dispersed on outside of animal
Endozoochory seeds dispersed on inside of animal
Types of dispersal
1). Jump dispersal 2. Diffusion 3. Secular migration
Jump dispersal Long distance dispersal accomplished during a relatively short period Occurs
infrequently, large effect Some species are better at jump dispersal than others e.g. Hawaii
and Galapagos
1. Explains wide and often discontinuous distributions of many taxa
2. Accounts for similarities/ differences among biotas inhabiting similar environment in
different geographic areas
3.Human-aided jump dispersal increasing
2).Diffusion- Slow dispersal of individuals spreading out from the margins of the species range
(accomplished over generations)-Often follows jump dispersal
example-Cattle egret;Africanized honey bees-Brought from Africa Escaped from Sao Paolo
Dispersed ~ 100 mi/yr
3)Secular migration- Very slow dispersal (e.g. hundreds of generations) that commonly
involves evolutionary changes in the dispersing populations
6)Types of Barriers-
It can be of three types --1)Physiological, 2)Ecological,3)Behavioral
1)Physiological Barriers- Physical environments outside the range an organism normally
encounters so it cannot survive and disperse across the barrier. Examples- Land-water
Salinity Temperature.
like oceans, Saharan deserts, Himalayan mountains, cold polar weather.

2)Ecological Barriers- Dispersing organisms must be able to withstand ecological hazards Like
3)Behavioral Barriers- Ability of some organisms to select suitable habitat can limit ability to
disperse across barriers
Example bird species capable of flying long distances will not cross open areas.
7)Dispersal Routes-
Dispersal routes can be --1)Corridors 2)Filters 3)Sweepstake routes
1)Corridor dispersal route that permits movement of many related species from one region
to another Similar environment to that of the two regions Balanced taxonomic composition.
E.G.Beringia - Pleistocene
2)Filters - selectively impede the movement of certain taxa while allowing others to disperse
freely (biotic or abiotic) Colonist-biased subset Often form transition zones.E.G. Two-way
filter for reptiles between Australia and Asia
3)Sweepstake Routes severe barrier that permits rare dispersal events (jump dispersal)
Rare on ecological timescale, likely and important on geological timescale. It can be followed
by dispersal and adaptive radiation.
E.g. Hurricane Luis Vegetation mats Green iguanas (Iguana iguana) , sweepstakes route from
Guadeloupe to Anguilla
8) Resistance to invasion-
More diverse communities may be more resistant to invasion(controversial not proved).
when barriers are crossed or disappear, then two or more distinct biotas come into contact
and biotic interchanges occured. There is Little evidence to reconstruct what happened during
these events in the past
Example- The Great American Interchange
Mammals evolved on Pangaea about 220 MYears Ago. e.g.Monotremes, marsupials, placentals
South America drifted from Gondwana about 160 MYA, Shift in mammal-dominance 65 MYA,
about 2.5 - 3.5 MYA formation of the Central American land bridge, and many species
croosed the two landforms k/as great american interchange.
Of those who crossed the landmasses-only Better migrators and better competitors survived as
per Habitat theory and Darwins survival of the fittest.
9) Pleistocene Glaciations- is one of the important factors that affected the distribution of flora
and fauna significantly.
Effects of glaciations on the physical environment
1) Temperature changes 2) Sea level changes 3) Climatic zones changes.
Temperature-falled significantly so organisms migrated to lower latitudes and altitudes.
Sea level changes-During glaciations sea level dropped (up to 160 m compared to current sea
level!)-That leads to Creation of corridors and barriers.
Climatic zones- Climatic zones shifted towards the equator-Novel combinations of temperature,
winds and precipitation generated non-analogue climatic zones
Biogeographic Responses to Glaciation
Changes in location, extent, and configuration of prime habitat
Changes in the nature of climatic and environmental zones
Formation and dissolution of dispersal routes
Biogeographic consequences-1)Migration 2)Refugia 3)Pleistocene extinctions
Altitudinal migration of entire biomes
Changes in single species geographic ranges
Species with low migration capabilities had to adapt to the changing environment
Many went extinct or remained isolated from other populations
The ability to migrate did not guarantee persistence in the new area (mutualisms, disease,
as new species invaded Biotic interchanges occured as explained earlier.
Example-Eurasian species crossed Beringia.
1)GLACIAL REFUGIA-Isolated fragments of non-glaciated land
Provided opportunities for divergence and diversification of species
Pleistocene MEGAFAUNA
Teratorn (Teratornis incredibilis);Giant bison (Bison latifrons) ;Ground sloth (Megatherium
sp);Sabertooth (Smilodon fatalis);Cave Bear (Ursus spelaeus) ;Giant deer(Megaloceros
giganteus )
3) Pleistocene extinctions
1) Climate based hypothesis- Climatic changes increased aridity and decreased food for
Evidence in support- Climatic changes modified vegetation leading to less food to large
herbivores; Large animals are more affected during environmental changes and food scarcity
than smaller animals
Drawbacks- Megafauna survived aprox. 20 glacial cycles, Large species in Africa and Eurasia did
not suffer the extreme extinction rates that the American mega fauna did.
2)Overkill hypothesis--Humans crossed Beringia from Asia, were Skillful hunters while prey was
Evidence in support- Megafauna and humans coexisted and fossil record shows hunting
evidence; Most affected terrestrial animals by the Pleistocene extinction were large herbivores
and their predators; Few extinctions in Eurasia and Africa (animals coexisted for longer with
human hunters); Extinctions occurred in a north to south fashion
Drawbacks- Hunting skills and technology of humans not as good as to bring down large prey;
Only archeological evidence that mammoths and bison were killed by human hunters
3)More comprehensive explanations- Combination of humans and environmental changes:
Fire use by humans Competition for resources with humans Introduction of exotics
Disease Hunting.
Scales of biodiversity-
Alpha Diversity Species richness of a local ecological community (e.g. defined as a study plot
such as a hectare)
Beta Diversity Change (or turnover) in species composition between two distinct communities
Gamma Diversity Total species richness over a large geographic area such as a biome.
Gradients in biodiversity-
The Latitudinal Gradient Increasing diversity towards the equator
The Peninsula Gradient Decreasing diversity away form the mainland
The Elevation Gradient Species diversity decreases with elevation
The Aridity Gradient Species diversity decreases with diminishing water availability
Aquatic Environments Similar patterns
Ecogeographic rules-
Bergmanns Rule (1847) Animals with larger body forms occur at high latitudes. Larger
animals have a lower surface area to volume ration.
Allens Rule (1877) Endothermic vertebrates that live in warmer climates have longer
Glogers Rule (1883) Coloration of related forms is correlated to humidity with darker
coloration occurring in more humid regions. Most likely driven by crypsis (camouflage).
Biotic communities are not static, they change through time.simplest change is the growth,
interaction and death of individual organisms. But there are other levels of community
change that act over longer time spans and that account for much larger community
composition and structure. These include biotic succession and community evolution.
biotic succession is an orderly and progressive replacement of one community by another
more suitable community called as climax community,also called as ecosystem development.
in a long span ,series of communities grow there and replace one another.
example- When a cropfield is deserted or a forest is severely burnt over, it is just like a plot of
bare ground and a series of plant communities grow there and replace one another-first
annual weeds, then perennial weeds and grasses, then shrubs, and trees until a forest
ends the development.
1)It is an orderly process of community development that involves changes in species
structure and community process with time. It is reasonably directional and, therefore,
2)It results from modification of the physical environment by the community; that is,
succession is community-controlled even though the physical environment determines the
patern and the rate of change and often sets limits as to how far development can go,
3)It culminates in stabilised eco-system in which maximum biomass and symbiotic
function between organisms are maintained per unit of available energy flow.
4)With succession, the following changes occur diversity of species increases production per
biomass decreases energy flow decreases new habitat niches are created climax or stable
community controls or becomes a buffer against the physical forces, such as, temperature,
moisture, light, wind, etc.
5)The first organisms to become established in an eco-system, undergoing succession, are
known as pioneers.
The stable community that ends the succession is termed the climax community.
6)The whole series of communities which are involved in the ecological succession in a
given area, for instance, from grass to shrub to forest, and which terminates in a final stable
climax community, is called a sere and seral stage.Each seral state is a community, although
temporary, with its own characteristics and it may remain for a very short time or for many
The successions may be of two types, in any of the basic environments such as terrestrical,
fresh water or marine.
It is the process of species colonization and replacement on sites not occupied
previously by any other community, such as sand beach, sand dune, fresh lava flows, volcanic
ash plans, etc. The sere involved in primary succession is called presere. Initially, only
those species which are resistant to extreme conditions flourish and add to the humus.
Thus ground is prepared for higher order species with broad foliage. Initial species are
called the pioneer communities (lichens on bare rocks, for instance). Colonisation of
beaches can be cited as an example of a primary succession.
Sand Beach Beach Grass Woody Shrubs Pine Trees Dry Oak Moist Forest Beach
Maple Forest ComplEX.
The bog successions of Canada are an example of a primary succession.
It is a process of change that occurs on sites previously occupied by well developed
communities, for instance, an old field succession where an abandoned field acts as the site.
secondary succession is more rapid than primary. The sere involved in secondary succession is
called subsere.
Bare FieldGrasslandPine ShrubsPine ForestOak Forest Climax
The complete process of primary ecological succession involves the following sequential
1. NUDATION--the formation of a bare area or nudation which could be due to volcanic
eruption, landslide, flooding, erosion, deposition, fire, distance or some other catastrophic
agency.New lifeless bare areas are also created by human activity, for instance, walls, quarries,
burning, digging, flooding large land areas under reservoirs.
2. INVASION--The next stage is invasion or the arrival of the reproductive bodies of various
organisms and their settlement in the new or bare area. The plants are the first
invaders (pioneers) in any area because the animals depend on them for food.
3. COMPETITION AND INTERACTIOn--As the number of individuals of species increases by
multiplication, the competition for space and nutrition beings-within different individuals of
the same species (intra-specific competition) and between two or more species
(interspecific competition). These species, in turn, interact with the environment, and the
exchange is a two-way process-the environment gets modified and different species also
modify their behaviour. Increased availability of food allows various kinds of animals to join
the community and the resulting interactions further modify the environment, thus paving
the way for fresh invasions by other species of plants and animals and continuing the process
of succession.
4. STABILISATION OR CLIMAX--Eventually a stage is reached when the final terminal community
becomes more or less stabilized for a comparatively long period of time and it can maintain
itself in the equilibrium or steady state with the climate of that area. This terminal community
is characterized by an equilibrium between gross primary production and total respiration,
between the energy captured from sunlight and energy released by decomposition,
between the intake of nutrients and the return of nutrients by litter fall. It has a wide
diversity of species, a well developed spatial structure, and complex food chains; and its living
biomass is in a steady state. This final stable community of the state. This final stable
community of the sere is the climax community, and the vegetation supporting it is the climax
5)CONTINUUM CONCEPT-- According to this concept, the vegetation undergoes gradual and
continuous changes, and cannot be differentiated into distinct communities.