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HISTORICAL PHYTOGEOGRAPHY C. C. Amadi

Phytogeography: Floral Biogeography

Biogeography studies the past and present distribution of the world's plant and animal species, and
often relates to the examination of the physical environment and how it affects species and shapes
their distribution across the world. As such, biogeography also includes the study of the
world's biomes and taxonomy—the naming of species—and has strong ties to biology, ecology,
evolution studies, climatology, and soil science as they relate to plant populations and the factors
that allow them to flourish in particular regions of the globe. Much of this knowledge has emerged
from the tremendous body of work in biogeography from one scientist, Alfred Russel Wallace
widely regarded as the “Father of Biogeography.” Aside from co-originating the process of Natural
Selection with Charles Darwin, Wallace spent extended periods studying the distribution and
diversity of plants and animals in Amazonia and Southeast Asia in the mid -1800s. Wallace,
originally from England, was a naturalist, explorer, geographer, anthropologist, and biologist who
first extensively studied the Amazon River and then the Malay Archipelago (the islands located
between the mainland of Southeast Asia and Australia). During his time in the Malay Archipelago,
Wallace examined the flora and fauna and came up with the Wallace Line—a line that divides the
distribution of animals in Indonesia into different regions according to the climates and conditions
of those regions and their inhabitants' proximity to Asian and Australian wildlife. Those closer to
Asia were said to be more related to Asian animals while those close to Australia were more related
to the Australian animals. Following Wallace were a number of other biogeographers who also
studied the distribution of species, and most of those researchers looked at history (leading to
historical biogeography) for explanations, thus making it a descriptive field. Continued studies
changed the way biogeographers looked at species and eventually promoted the study of the
environmental features of that time as important to understanding spatial patterns in species
distribution. Initially, island biogeography and the fragmentation of habitats caused by islands
became popular fields of study as it was easier to explain plant and animal patterns on the
microcosms developed on isolated islands. The study of habitat fragmentation in biogeography then
led to the development of conservation biology and landscape ecology.

Biogeography is the study of the distribution and evolution of living organisms and their
interactions with the physical and biological environment. It examines how and why different
species are found in different regions of the world and how they have changed over time.
Phytogeography is a sub-field of biogeography whose other counterpart is zoogeography. It studies
the past and present distribution of plants while zoogeography studies the past and present
distribution of animal species. Phytogeography is the study of the distribution of flora over the
earth and of the principles that govern this distribution. It studies major geographic patterns in
species richness, and the processes and theories that are thought to account for these patterns.
Phytogeography deals with the spatial distribution patterns of plants of different phylogeny
(species, populations) and function, as well as their communities (ecosystems, biodiversity), and
aims to capture, understand and predict these patterns. Why do some places contain more species
than others? Many patterns in the spatial distribution of species have been identified by
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biogeographers, and many mechanisms have been proposed to explain these patterns. A species
occurs in a given place and time because members of this species (or its ancestors) evolved in this
location or dispersed to it at some point in the past. Biogeographers seek to discover new patterns
in the distribution of species across space and use diverse research methods to study the historical
and ecological factors that can explain these patterns. Many of the spatial patterns in biodiversity
are overt, others are subtle and yet additional patterns remain undetected. While the existence of
these patterns may be obvious — and changes in the environment that are paired with these
patterns may also be obvious — the mechanisms that cause the differences in biodiversity along
environmental gradients are under still the subject of scientific debate.

Historical Biogeography in Phytogeography

Historical biogeography is called paleobiogeography and studies the past distributions of species.
It looks at their evolutionary history and things like past climate change to determine why a certain
species may have developed in a particular area. Paleogeographic research uses fossils to show the
movement of species across space via moving continental plates. Paleobiogeography also takes
varying climate as a result of the physical land being in different places into account for the
presence of different plants. For example, the historical approach would say there are more species
in the tropics than at high latitudes (temperate and arctic regions) because the tropics experienced
less severe climate change during glacial periods which led to fewer extinctions and more stable
populations over time. When glaciers advanced in temperate regions during the Pleistocene Epoch,
vegetation zones were pushed lower in elevation or lower in latitude. When glaciers retreated,
vegetation zones rebounded; the climate gradient softened and landscape diversity increased. Each
cycle of advance-and-retreat caused some extinctions, however, so the biota of the temperate zone
became simpler in that era.

Historical biogeography is a branch of biogeography that focuses on the historical processes that
have shaped the distribution of living organisms on Earth. It examines how the movement,
diversification, and extinction of species have been influenced by past events such as continental
drift, climate change, and mass extinctions. Historical biogeographers use a variety of methods,
including phylogenetics, geology, and palaeontology, to reconstruct the evolutionary history of
different groups of organisms and to understand how their distributions have changed over time.
One of the main goals of historical biogeography is to reconstruct the biogeographic history of
different regions and to identify the factors that have influenced the distribution of different groups
of organisms. This includes understanding how land bridges, oceanic barriers, and other
geographical features have affected the dispersal and diversification of species, as well as how
climate change and other environmental factors have influenced the survival and extinction of
different groups of organisms.
Historical biogeography also provides a framework for understanding the current distribution of
biodiversity and the factors that threaten it, such as habitat destruction and climate change.
Understanding the historical context of biodiversity can help inform conservation and management
efforts aimed at protecting endangered species and preserving the Earth's biodiversity
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Conservation Biogeography in Phytogeography

In recent years, scientists and nature enthusiasts alike have further expanded the field of
biogeography to include conservation biogeography—the protection or restoration of nature and
its flora (and fauna), from devastation which is often caused by human interference in the natural
cycle. Scientists in the field of conservation biogeography study ways in which humans can help
restore the natural order of plant (and animal) life in a region. Often times, this includes re-
integration of species into areas zoned for commercial and residential use by establishing public
parks and nature preserves at the edges of cities (e.g., greenspaces, green infrastructure). It can also
be applied in land reclamation or vegetation restoration projects.

Conservation biogeography is a field of study that applies biogeographic principles and methods to
the conservation of biodiversity. It uses an understanding of the historical and current distribution
of species and ecosystems to identify areas of high conservation value, such as biodiversity
hotspots, and to design conservation strategies that aim to protect these areas and the species and
ecosystems they contain. One of the main goals of conservation biogeography is to identify and
rejuvenate areas of high biodiversity that are at high risk of habitat loss or fragmentation and
degradation, such as tropical rainforests, prairies or woodlands. These areas, known as biodiversity
hotspots, are considered a priority for conservation efforts because they contain a large number of
species that are found nowhere else on Earth. Additionally, conservation biogeography uses
biogeographic principles to understand how climate change affects the distribution and survival of
species, and to design conservation strategies that take into account the potential effects of climate
change on biodiversity.

Ecological Biogeography in Phytogeography

Ecological biogeography is a field of study that focuses on how the distribution and abundance of
living organisms are influenced by their interactions with the physical and biological environment.
It examines how the characteristics of different ecosystems, such as climate, geology, topography,
and the presence of other species, affect the distribution and abundance of different groups of
organisms. One of the main goals of ecological biogeography is to understand how different factors,
such as climate, geology, topography, and the presence of other species, interact to shape the
distribution and abundance of living organisms. This includes understanding how different
ecological factors, such as temperature, precipitation, and sunlight, influence the distribution of
different groups of organisms, and how the presence of other species, such as predators,
competitors, and symbionts, affects the survival and reproduction of different groups of organisms.
Ecological biogeography also involves the study of biogeographic regions and biotic provinces.
Biogeographic regions are defined based on the similarity of the biotic components, such as the
presence of certain types of plants, animals, and microorganisms, across a large area. Biotic
provinces are defined based on the similarity of the abiotic components, such as temperature,
precipitation, and geology, across a large area. Ecological biogeography also provides a framework
for understanding how human activities, such as habitat destruction, pollution, etc are affecting the
distribution and abundance of living organisms, and for designing conservation strategies that aim
to protect biodiversity.
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CAUSATIVE FACTORS OF PLANT DISTRIBUTION

Internal and external factors cause plants to become abundantly distributed or vulnerable to
extinction. Internal factors refer to the plant biological characteristics, including failures in
heritability, reproduction, viability, and adaptability. External factors include both natural and
human factors; human factors refer to human disturbances, roads, and land use types. Most of these
have become the harmful factors impacting on rare and endangered plants. Natural factors refer to
the ecological environment and they include climate, topography, soil type, human activity,
associations, water supply, fire, salinity.

Climatic factors

Climate
Plants that are adapted to cold climates will struggle to survive in areas that are consistently warm,
and vice versa. Similarly, plants that require a lot of water will not thrive in areas that are prone to
drought, while those that are drought-tolerant will. Additionally, the amount and duration of
sunlight in an area will influence which plants can grow there. These factors are the main drivers of
plant distribution and can be observed in different biomes across the world. Climate is often
thought to be the chief determinant of vegetation. First, climate varies over larger spatial scales and
acts, therefore, as an environmental filter at these scales, whereas, in comparison, soils show
smaller‐scale variation and act as an additional filter under homogeneous climatic conditions at
smaller spatial scales. Second, our tropical forests are at the drier end of the rainfall gradient
(1100–2200 mm year−1), and water availability is clearly a more limiting factor here than for
forests at the wetter part of the rainfall gradient, where nutrient availability might become a
limiting factor.

Temperature
Tropical tree species should be sensitive to such small temperature variation, given the fact that
they experience – and are adapted to – a very small range in temperatures: within the lowland
tropics, mean annual temperature ranges only from 24 to 27 °C, and seasonal monthly
temperature variation is <4 °C, over a band of 20° latitude around the equator. Small changes in
temperature may have drastic effects on tropical species and thus on their distribution patterns as
many of these species are adapted to low temperature variation and lack populations inhabiting a
wider range of temperatures. Harsh habitats help limit the intrusion of new species and the
abundance of some resident species. The topography and environmental conditions order the
nature of vegetation. Habitats like tundra, alpine tundra, salt marshes, rocky outcrops, seem almost
free of introduced species. Sometimes, the environment, the isolation of a place and the genetic
potential of plants growing there combine to create unique and bizarre vegetation found nowhere
else in the world. E.g. the flora and vegetation of alpine zones on tropical African mountains
(Kilimanjaro) may live for a century.

Microenvironment
This also plays key roles in plant distribution. It is the part of environment close to the surface of a
plant, animal structure or the ground. It is modified by nearby surfaces. Because bare soil absorbs
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heat on a clear day, the soil surface and just below surface air temperature will be higher than that
recorded by meteorology agencies. Prostrate plants with basal leaves will experience warmer
temperatures, greater humidity, less turbulence etc. Plant species that are not favoured by such
environmental conditions migrate and are absent, making the tolerant species to appear abundant.
Plants growing in shallow depressions experience frost more frequently than those growing on
higher ground because cold air flows downhill. These plants have to build up tolerance for
prolonged cool temperatures and the non-tolerant species migrate or are lost. Soils in depressions
may also be wetter (due to melting snow) and anaerobic (due to reduced light penetration); hence,
creating a kind of cold marsh that can support floaters, reeds and swamp species that can withstand
the cold, as well as microorganisms.

Precipitation, as Water and Snow; Water supply

The seasonal timing of precipitation (as rain or snow) can be as critical to vegetation as the annual
amount. Rainfall is generally more than 200cm annually in the tropics; when the precipitation is
divided evenly among 12 months, a 3-storeyed evergreen forest results; but if divided into wet and
dry seasons, a less complex forest with many deciduous trees will result. The nature of precipitation
is also important: at high altitudes, precipitation falls as snow while below 2000 metres, it falls as
rain. Hence, the species of trees, shrubs and herbs will be different in the 2 zones even if the total
and seasonal precipitation are the same. Experimental studies in the shade-house and the field also
conclude that seasonal drought has a stronger effect on growth and survival over other factors, as it
immediately affects cellular processes and plant physiology. Plant water availability and drought
sensitivity are, therefore, important determinants of the distribution of tropical tree species.

Plant distribution is largely affected by moisture content/water availability. Plant-water

relationships have led to the classification of species as xerophytes (plants able to grow in dry
conditions), mesophytes (plants that grow best in moist soil), helophytes (plants that can grow in
saturated soil e.g. marshes and bogs) and hydrophytes (able to grow rooted, submerged, emergent
or floating in standing water). Plants in each category have unique morphological, anatomical and
physiological traits that help them adapt to the moisture-determined environment. For example,
xerophytes may have small, hard leaves, epidermis with thick cuticle, light colour and hair-like
trichomes, sunken stomata that close early during the day, and sometimes succulent tissue with
large cells and water-filled vacuoles. Leaves may be permanently or seasonally absent; they possess
green stems which are photosynthetic. Xerophytes can live in humid environments provided their
roots are in dry microenvironment e.g. cactus. Some of them live in tropical rainforests as epiphytes
attached to dry tree barks and trapping water from rain with their succulent leaves and stem; the
root surfaces have specialised water-storing tissues e.g. orchidaceae (orchid) and bromeliaceae
(Ananas).

Salinity.
Saline soils are said to be physiologically arid because dissolved salts decrease the water potential
of soil water, making the water less available. Salinity occurs in soils where marine water bodies
receded from, leaving behind expansive amounts of salts or when water table rises to a few meters
from the soil surface because of the drainage of water from waterways, thereby bringing out broken
down salts from the bed rocks. It also results when water leaks through unlined channel banks and
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raises the water table, or when water (e.g., from floods) contains broken down salts of carbonate,
sodium and potassium (the salts from the groundwater are raised by hair-like activity to the surface
of the soil). All kinds of water (other than characteristic precipitation) contain some salts. When
plants take up water, salts are left behind in the soil and collect there, making it more troublesome
for plants to absorb soil moisture. Chemicals like gypsum (CaSO4) and iron sulphate encourage
salinity (gypsum reacts with sodium to form NaSO4).

Salinity destroys soil structure; diverse layers of the soil are seriously influenced. It lessens seed
germination; seedling development and yield is extremely poor in many plants. Saline soil contains
chlorides and sulphate of sodium and potassium. These salts are solvent in water and can be
filtered out by flushing of soil with water, e.g., in the wake of planting rice, harvest flushing is
exceptionally useful in controlling salinity. Soybean, spinach, cabbage, cauliflower, sweet potato,
ginger, cucumber, pea sugar beet and so forth can be grown in saline soils, as well as grasses like
Sporobolus arabicus, Cynodon dactylon, Chloris barbata. Herbs like Melilotus albus, Trifolium
fragiferum. Sesbania bispinosa and Sesbania sesban (Jantar) fix nitrogen in saline soils, while some
plants collect salts in crystalline form and adsorb the crystals to the roots, thereby freeing up the
soil e.g., Suaeda fruticosa, Suaeda monoica etc.

Topography (altitude, slope aspect, slope gradient, and exposure of rock).

A study conducted to study the early stage of the relationship between vegetation and topographic
factors has found that landscape variations in vegetation are strongly related to topographic factors.
In a vegetation study, most orchid species were concentrated in high altitudinal habitats. Seed
germination and growth of some orchids are dependent on high altitudinal habitats, which not only
have favorable temperatures and higher humidity, but also avoid human disturbance. Also, it was
found that the peak richness of the small populations always occurred at the medium level across
an environmental gradient, and that it had larger species similarity values. Research found that
many rare and endangered plants are often concentrated at medium altitudes. On the other hand,
slope gradient can reflect the soil moisture and its nutrient condition. The distribution of many rare
plants was found to be concentrated in a medium level of slope gradients with favorable soil
conditions. The peak richness of extremely small populations always appeared at the south aspect,
and that their frequencies were always the largest on sunny or semi-sunny slope aspects, due to
their drought tolerance and sun preference. But several studies have also shown that the species
richness of endangered plants and their individual numbers is higher on shady and semi-shady
slope aspects than those on other aspects, because they prefer high humidity and low irradiance.
Also, since the increasing forest canopy can improve the forest microhabitat conditions, and
regulate the distribution, composition, and growth of forest subordinate layers, the seedlings of
most of the rare and endangered plants thus grew better with high abundance under medium
canopy densities. Topography can also partly affect the accumulation and export of soil nutrients,
thereby indirectly impacting plant distribution. Some studies have indicated that changes in
topography have clear influences on soil physical and chemical properties and soil water
characteristics. Micro-topographic factors (such as slope, slope aspect and slope position/elevation)
exert a strong influence on plant community structure and species distribution.
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Soil Quality
Soil variables, such as the available potassium (K) and total nitrogen (N) have a significant influence
on vegetation growth and development. Research found that the available K was the variable with
the highest explanatory power compared with the other soil nutrient variables, followed by total N
and soil organic matter. Many studies have shown that the soil organic matter presents surface
accumulation and has a significant influence on vegetation growth and development. Because a high
amount of litter was introduced into the soil each year, its decomposition by microorganisms
resulted in a greater amount of humus, and increases in the soil organic matter. It has been found
that soil organic matter was positively correlated with canopy density and negatively correlated
with above-ground biomass and herb coverage, implying that vegetation could increase soil organic
matter and, in this way, soil nutrient availability could influence species distribution and
community composition.

Soil composition
Soil fertility may act as a more important environmental filter at smaller spatial scales. Indeed,
edaphic conditions are found to control species distribution in climatically homogeneous
environments and small spatial scales. Soil texture may modify in an important way plant water
availability through its effect on water‐holding capacity, and it may also constrain species
distribution, by creating waterlogged or anaerobic conditions. Intriguingly, many species that show
a significant response to soil texture have a higher probability of growing in soils of poor texture
given the high sand content and, hence, a lower water‐holding capacity, suggesting that these
species prefer edaphically dry soils or are better competitors there. This enforces the importance of
water availability and soil drainage as drivers of plant distributions.

Soil parent material

Plant distribution may be affected by soil parent material and water availability acting together.
Very coarse soils that are high in sand and gravel permit most rainwater to percolate rapidly,
leaving behind arid topsoil within a climate that might seem adequately wet. This will promote
growth for xerophytes. Similarly, shallow soils and rock outcrops are unable to retain much water
for roots. Clay soils, on the other hand, have low water absorption and so can store water briefly,
but they can also easily lose water if it rains in torrential bursts, forcing the soil to act as a hard
surface that permits runoff and preventing water percolation in the soil.

Human Interference
It was found that this was one of the leading factors changing the appearance frequency of
extremely small populations among plant species. For example, the dominant factors affecting the
appearance frequency of orchid and non-orchid species were altitude and the distance between
population and road, respectively. For many endangered plants, the direct factors making them
susceptible to extinction are human activities. This may be because many of these species have high
economic utilization value or excellent ornamental characteristics and, thus, that they are often
overexploited by local people. Additionally, it was found that the appearance frequencies of three
orchid species (Dendrobium hainanense, Oxystophyllum changjiangense, Dendrolirium tomentosum)
were significantly and positively correlated with roads and agricultural activities. These species
were also concentrated along roadsides and in areas of human activity, suggesting that the
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distribution of some endangered orchid species benefit from human activities or roads. Roads
usually have larger negative effects on the distribution of native plants than positive effects.

Man controls plant distribution through agriculture, migration, importation for horticulture,
foraging etc. The early man gathered and used wild-growing plants, but over time and upon the
discovery of the importance of certain crops, put in extra effort to ensure the continued availability
of those species through cultivation and subsequently, vegetative propagation and fertilization to
encourage greater yield. This simplistic form of agriculture often referred to as Protoagriculture
began this selection/distribution of choice plants. It started as selective harvesting, burning,
pruning, sowing of rhizomes, bulbs and seeds (without geometric farm patterns) in a way that
favours the choice plants. However, because of human-induced selection, some species lose their
natural characteristics over time e.g., the capacity for sexual reproduction, morphological traits,
genetic traits, etc. Also, as man explored new land, he brought along his already domesticated plants
and their companion weeds, and seeds/spores sticking to them. Upon cultivation of domestic
plants, the weeds were also sown, and through fierce competition over time and with favourable
soil and climatic conditions, these weeds edge out the native species from the landscape. A large
percentage of weeds in the US were imported from Europe, Asia, Africa and Australia; the
introduced plants have naturalised in forests, grasslands, wetlands, deserts and coastal strands.

Invasive species
An invasive species is one which is transported outside its original geographic range to a novel
habitat, where it increases in density, and can cause detrimental effects on the indigenous species in
that area, often reducing biodiversity. Most introduced species do not become invasive, indeed they
frequently become extinct because they are less likely to be well adapted to the new environment
than indigenous species, but the subset of introduced species that become invasive is problematic
in terms of the economy, ecosystem services, and biodiversity. The widespread exchange of species
among distant parts of the world as a result of human commerce has clearly resulted in a loss of
biodiversity at many spatial scales.

Ecological Associations
Mutualism is a mutually beneficial relationship between two organisms. In the case of plants,
mutualistic relationships often involve the exchange of nutrients or other resources. For example,
many plants form mutualistic relationships with fungi, where the fungi provide the plant with
nutrients and in return, the plant provides the fungi with a place to live (e.g., in a lichen association).
Other examples of mutualism animals feed on nectar from flowers and fruits, thereby aiding in
pollination and seed dispersal, respectively, influence plant distribution in this way. Also, nitrogen-
fixing bacteria living in the roots of leguminous plant increase yield and distribution of the legumes.
Mutualistic relationships can allow plants to thrive in areas where they otherwise might not be able
to survive, by providing them with the resources they need to grow and thrive.

Parasitism is a relationship where one organism (the parasite) benefits at the expense of the other
organism (the host). In the case of plants, parasitic relationships often involve one plant stealing
resources from another. For example, some plants are able to grow roots that can penetrate the
roots of other plants and steal water and nutrients from them. These parasitic relationships can
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make it difficult for the host plant to survive and can influence the distribution of both the host and
parasite plant species.

Fire.
Some grasslands, chaparral scrub, the boreal forests, mountain conifer forests and desert oases owe
their presence and plant composition ton climates with episodic, natural fires. When man
suppresses these fires due to their supposed catastrophic/destructive nature, they begin to change
the mix of species and the shape of the landscape that has resulted over time. Many species in these
fire-prone zones not only tolerate fire but may depend on it, either to complete their cycle, or to
maintain dominance over other species.
Some plant species are adapted to fire-prone environments and are able to survive and even benefit
from fires. These plants are known as fire-adapted or fire-dependent species. They often have thick
bark, deep roots, or resprout from underground storage organs after a fire. For example, some
species of eucalyptus in Australia, have a thick bark that protects the tree from fire damage, and the
heat from fire causes the tree to release its seeds, which will germinate after the fire. On the other
hand, other plant species are not adapted to fire and are killed or severely damaged by fires. These
plants are known as fire-sensitive or fire-intolerant species. They often have thin bark, shallow
roots, or no underground storage organs. The frequency and intensity of fires, as well as the timing
of fires also influence plant distribution. For example, frequent, low-intensity fires can benefit fire-
adapted species, while infrequent, high-intensity fires can benefit fire-sensitive species.

Herbivory.
Some plant species have their population members thoroughly dispersed to make it difficult for the
foraging herbivores to locate them. Some plant species propagate profusely by means of
underground stem and rhizomes. Explosive mechanism and wind dispersal of seeds aid in achieving
wide distribution in a species. To ward off foragers, some species are able to put up physical
barriers such as spines, sclerophylls, thick barks, hard seed coats and fruits. Other plants
manufacture and accumulate metabolic by-products that are distasteful or that otherwise inhibit
herbivores from grazing; this is referred to as Amensalism. These chemicals include toxins, tannins
and terpenes. Toxins have small molecular weight and are effective in small concentrations. They
repel herbivores by interfering with nerve and muscle activity, hormone function, liver and kidney
metabolism. Tannins and terpenes have large, complex molecules and repel herbivores by their
bitter, unpleasant taste.

More for you…

Ecological biogeography looks at the current factors responsible for species distribution, and the
most common fields of research within ecological biogeography are climatic equability, primary
productivity, and habitat heterogeneity:
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Climatic equability (climate, precipitation)

This looks at the variation between daily and annual temperatures as it is harder to survive in areas
with high variation between day and night and seasonal temperatures. Because of this, there are
fewer species at high latitudes because more adaptations are needed to be able to survive there. In
contrast, the tropics have a steadier climate with fewer variations in temperature. This means
plants do not need to spend their energy on being dormant and then regenerating their leaves or
flowers, they don’t need a flowering season, and they do not need to adapt to extreme hot or cold
conditions. Common factors include Climate, precipitation, temperature.

Primary productivity (temperature, microenvironment)

This looks at the evapotranspiration rates of plants. Where evapotranspiration is high and so is
plant growth. Therefore, areas like the tropics that are warm and moist foster plant transpiration
allowing more plants to grow there. In high latitudes, it is simply too cold for the atmosphere to
hold enough water vapor to produce high rates of evapotranspiration and there are fewer plants
present.

Habitat heterogeneity (succession, herbivory, competition)

All species occurring in at a given place and time either arrived from another place or originated in
that location from ancestral species. This fact applies to extinct species that were ancestors of all
extant species. Species richness in a given location is the result of three factors — the rate of
speciation, the rate of extinction, and the dispersal of species from other locations. In principle, if
biogeographers could understand how the current and past environment has shaped these three
factors, we would then obtain a comprehensive understanding of what generates all biogeographic
patterns of species richness. The study of historical factors shaping species richness and
distribution is often broken into two major categories: vicariance and dispersal.

Vicariance: here, species occur in a location because their ancestors remained there passively as
the environment moved around them. Vicariance describes the disruption of the biogeographic
range of a group of organisms by changes in the environment. Vicariant events can happen when
landmasses move apart through tectonic action, or when mountains emerge to divide the
geographic ranges of species. Vicariance typically leads to the emergence of new species through
allopatric speciation, in which one ancestral species will result in the production of two new species
that evolve apart from one another in geographic isolation, frequently by genetic drift rather than
natural selection.
Dispersal: here, a species or its ancestors may have arrived at a location via movement from
another location. Dispersal biogeography focuses on the movement of species from one location to
another location. Jump dispersal events, when individuals of a species travel a relatively long
distance to a new environment in which they did not previously occur, can result in the adaptive
radiation of one ancestral species giving rise to a broad diversity of new species.

Major Spatial Patterns in Biodiversity

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One major geographic pattern in biodiversity is the latitudinal gradient in species richness. As one
travels further away from the equator, for most taxa, the number of species declines. This general
pattern holds true for most taxa and ecosystem types in both marine and terrestrial environments.
There is broad agreement that this pattern is caused by differences in the abiotic, climatic
environment, but the specific mechanism or mechanisms causing this pattern are a continued topic
of discussion and investigation.

Another recurring pattern in biogeographic theory is the elevational gradient in species richness.
As one travels to higher elevations, the number of species declines, or, in many cases, peaks at mid-
elevations. Aside from the environmental mechanisms driving this diversity gradient, there is a
phenomenon that is based on the geography of species range distributions called the mid-domain
effect. The mid-domain effect predicts a peak of diversity at the midpoint along any domain simply
by the fact that the ranges of more species overlap in the middle of a domain (like a mountain or an
island) than on the edges, and this effect works together with environmental determinants to affect
the net distribution of species along many elevational gradients.

A third recurring pattern in the distribution of species is the area effect on species richness. The
larger a place is, the more species it can support. This applies to actual islands in bodies of
water, as well as habitat islands such as those surrounded by human development.