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Biogeography

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

Biogeography

By Joy Nystrom Mast, Carthage College

Biogeography is the study of the geographical distribution of living and fossil

plants and animals as a result of ecological and evolutionary processes. Biogeography

analyzes organism-environment relations through change over space and time, and often

includes human-biota interactions. The main questions explored by biogeographers deal

with organism patterns in order to understand the underlying processes. Biogeographers

ponder questions such as: Why is a species present in a given area? Conversely, if a

species is not present, then why is it missing from the area? What are the historical and

ecological factors that help determine where a species occurs? What are the effects on

evolution and plate tectonics? How have humans altered geographic distribution of

organisms? The science of biogeography has been revitalized in the past 60 years due to

our understanding of plate tectonics, mechanisms limiting distributions, island

biogeography theories, and mathematical and technological tools.

Current work in biogeography uses spatial patterns of organisms, past and

present, to determine ecological processes. Biogeographers use experimental testing and

quantification of biotic interactions. Vegetation dynamics is the primary focus for

approximately half of the biogeographic research conducted by U.S. geographers. Other

major focuses include ecosystem structure and function, zoogeography, paleoecology,

and development of new biogeographic methodology. In particular, mapping and

modeling spatial patterns of abundance and distribution of species of plants and animals

has greatly advanced with geographic information systems and remote sensing

technology.

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

Development of Biogeography

To better understand the current field of biogeography, it is important to explore

the foundations and history of the science. Biogeography is a synthetic study which is

based in part on the subjects of community ecology, geology, systematics, evolutionary

biology, and paleontology. The development of the subject of biogeography may be

broken into four historical periods.

1600 – 1850: Age of Reason

Early studies of organisms’ geographic distributions were focused on descriptive

studies with historical explorations. These scientists focused on documenting spatial

patterns of organisms, focusing on the effects of climate, latitude, and altitude. Comte de

Buffon (1707-1788), also known as Georges-Louis Leclerc, determined that distant

regions with similar climate and similar-appearing vegetation have different animal

species. This is now referred to at Buffon’s Law. He is also the author of Historie

Naturelle, a 44 volume natural history encyclopedia. Carl Linnaeus (1707-1778) studied

the plants and animals spread from Mount Ararat in Turkey in order to explore the idea of

the biblical flood. As a result of documenting elevational zones of Ararat, he came up

with the idea of biomes defined as major ecological communities. In addition, Carl

Linnaeus is considered the father of the science of taxonomy, which is the science of

classification.

This time period is also known as a great age for exploration. Johann Reinhold

Forster (1729 - 1798) was the naturalist on James Cook's second Pacific voyage in 1778.

He advanced biogeography by creating global biotic regions for plants. Forster noted the

higher species diversity in tropics, as well as species diversity being correlated with

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

island size. Alexander von Humboldt (1769-1859) created a botanical geography that was

foundational to the field of biogeography. He determined that plant vegetation types are

strongly correlated with local climate to create latitudinal belts of vegetation. Moreover,

he developed elevational vegetation Zones for the Andes in South America.

1850 – 1900: Evolution by Natural Selection

The idea of evolution based on natural selection greatly altered the way species

distributions were explained. Charles Darwin (1809-1882) is most famous for publishing,

The Origin of Species, outlining his idea of evolution through natural selection. Natural

selection occurs when individuals in a population either do not survive equally well, do

not breed equally well, or do not survive and breed equally well due to inherited

differences. Evolution in turn can be thought of in two ways: microevolution and

macroevolution. In microevolution, evolution is considered changes in the genetic

composition of a population with the passage of each generation. For macroevolution,

evolution is the gradual change of organisms from one form into another, with the origins

of species and lineages from ancestral forms. For an example, Darwin studied the

adaptation of Galapagos Island finches to specialize in tree versus ground varieties, then

further evolving their bill structure (for grasping, probing, crushing) into seed, insect,

cactus, or fruit eaters. This divergent evolution is a diversification over evolutionary time

of a species into several different species, commonly referred to as adaptive radiation.

Alfred Russel Wallace (1823-1913) is also famous for independently developing

the idea of evolution by natural selection, based on his work in Indonesia. He found that

the species on Sumatra and Java were very different from nearby New Guinea, even

though the climates were similar. Wallace’s study of biota in Southeast Asia showed

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

geographic distance is not equal taxonomic similarity, and the boundary area between

these islands is now referred to as “Wallace’s Line”. Wallace is also considered to be the

originator of zoogeography, which is the biogeography focused on animals. Wallace

integrated geological, fossil, and evolutionary information to consider paleoclimate

influences distributions, developing six great biotic regions.

Other notable contributions to biogeography during this period include mapping

biotic regions and understanding limiting factors. Philip Lutley Sclater (1829-1913)

advanced the subject of biogeography with his defining terrestrial biotic regions for birds

and marine regions for marine mammals. Justus Liebig (1803-1876) changed the way

scientists viewed restrictions on organisms away from a focus on total resources available

with his law of the minimum. The law of the minimum states that the scarcest resource

(or limiting factor) in the environment makes it difficult for a species to live, grow, and

reproduce.

1900 – 1950: Continental Drift and Ecology

Themes in biogeography in the first half of the twentieth century focused on links

to paleontology, centers of species origins, and the biological species concept. The

emphasis in the science of biogeography was on evolution, history, dispersal, and

mechanisms of survival. The greatest impact on biogeography in this period was the

theory of continental drift in 1912 and 1915 by the German geologist Alfred Wegener

(1880-1930). Before the theory of plate tectonics, it was difficult for biogeographers to

explain certain patterns of species distributions with the assumption that land masses

were fixed in their geographic positions. Wegener’s theory was actually not widely

accepted until the 1960s when proof of continental drift came from a series of linear

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

magnetic anomalies on either side of the Mid-Atlantic Ridge. With the acceptance of the

continental drift theory, biogeographers could now explain the disjunct biogeographic

distribution of present day organisms found on different continents but having similar

ancestors. Species can interact as continents collide. Subsequently, when the continents

separate they take their new species with them. Biogeographers now ponder how plate

tectonics may have affected the evolution of life. In turn, biogeographers offer evidence

for plate tectonics such as dispersal of species via corridors such as the Bering land

bridge or widely separated (“disjunct”) species distributions that can’t be explained by

dispersal. For instance, Nothofagus (southern beech) trees which only occur in southern

South America and in New Zealand.

In addition to historical explanations of organism distributions, biogeographers

also examined ecological reasons for spatial patterns. Theories on ecological succession

were formally developed in the late 1800s and early 1900s to show predictable and

orderly changes in the composition or structure of ecological communities. In 1899,

Henry Cowles published his study of stages of vegetation development on dunes along

Lake Michigan. In 1916, Frederic Clements published his famous theory of vegetation

development focusing on gradually changes over time to best fit to the local conditions.

His climax theory of vegetation dominated plant ecology was later largely replaced by

other theories, notably by Henry Gleason’s 1926 concept of distribution of plants

depending on the individual species rather than the Clements’s idea of plant associations.

In 1934, Christen Raunkiaer (1860-1938) helped change the way biogeographers’

classified species with life forms based on ecological rather than taxonomic

classification. In 1935, Sir Arthur Tansley (1871-1955) refined the term “ecosystem” to

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

mean the whole complex natural unit in a system consisting of all plants, animals and

micro-organisms (biotic factors) in an area functioning together with all of the non-living

physical (abiotic) factors of the environment.

1950 – Present: Ecological and Historical Theories

Since 1950, the field of biogeography has been revitalized with advanced in

ecological and historical theories focused on phylogenetic classification to related

different species, mechanisms limiting geographic distribution, and distances and size

influencing number of species in an area. During this period, the concept of new species

arising due to geographic isolation was developed by the Ernst Mayr (1904-2004). Mayr

is also well known for defining the “biological species concept” as potentially

interbreeding to produce fertile offspring. In addition, Mayr helped to define the term

“cladistics” to refer to classifications which only take into account genealogy, based on

evolutionary ancestry. Cladistics, or phylogenetic classification, views a species as a

group of lineage-connected individuals, compared to the traditional Linnaean taxonomy

which focused on the similarities between species. Cladograms are created based on the

order in which different groups branched off from their common ancestors, arranged with

the most closely related species on adjacent branches of the phylogenetic tree.

Theories also expanded during this time period on how a species can occur in

widely geographically separated areas and the mechanisms that limit these distributions.

In 1958, Leon Croizat (1932-1982) published his concept of “vicariance biogeography”

to explain disjunction of multiple species due to the growth of barriers instead of via

dispersal. Croizat’s works include Manual of Phytogeography (1952), Panbiogeography

(1958), Space, Time, Form (1964). Robert Harding Whittaker (1920–1980) proposed a

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

new method to analyze limits to plant distributions by comparing species abundance to

environmental gradients. His gradient analyses approach focuses on abiotic factors such

as light, water, temperature, and soil nutrients in plant communities.

Biogeography during this period moved from observational to predictive studies

with the theory of island biogeography. In 1963, R. H. MacArthur and E. O. Wilson

hypothesized that species richness of an area could be predicted to explain distributions.

The theory states that if one knows the rates of colonization and extinction of an island,

then it is possible to predict number of equilibrium species that area could support. They

based the species richness prediction on two factors: (1) distance of island from a

mainland source of species for colonization pool; and (2) the size of the island for

available habitat and variety of niches. With these two factors, MacArthur and Wilson

predicted the number of species the area could maintain, as well as the turnover rate for

the area. According to island biogeography theory, small and distant islands have a lower

number of species that can be maintained compared to large and near islands. The theory

also states that there would be a turnover of the species as new species colonize and old

species go extinct, but the number of species overall should achieve an equilibrium

number. This theory has been applied to other non-island areas that act like islands due

to habitat fragmentation, such as nature preserves and national parks.

Spatial Distributions of Organisms

Modern biogeography explores spatial patterns in the geographic variation of individuals

and populations, including genetic, physiological, and morphological variations.

Organisms are often studied in biomes, consisting of distinct flora and fauna related to

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

climate, soil, and geological factors. Biogeographers analyze species ecological niches

(both fundamental and realized), defined as the total requirements for resources and

physical conditions. The foundation of biogeographic distribution patterns follows the

first rule of geography, namely that closer equates to more similar (referred to as “spatial

autocorrelation”). When relating species niche to geographic distributions, key

interactions and adaptations to consider include: (1) stress, with regards to climate,

predation, and availability of symbiosis (close association between species), (2)

competition, both between and within species which exclude some species from their

fundamental niches, and (3) disturbance, which occurs less predictably and causes a

greater change in the environment than stress. Biogeography seeks to answer why

species distribution patterns as a response to historical and ecological limiting factors,

dispersal mechanisms, and human influences.

Vicariance Biogeography

Geographic distributions may be separated into disjunct populations by historic

events that create barriers, called “vicariant events”. Disjunct distribution occurs when

two or more closely related taxa live today in widely separated areas. Barriers which split

a continuous distribution into disjunctions are created by many processes, including

changing in the distribution of land via by continental or tectonic shifts, volcanism and

mountain-building, shifts in river patterns, glacial cycles, climate change, and human

alterations to landscapes. Barriers may be physical (like a mountain), physiological (such

as fresh versus salt water blocking for aquatic organisms), and/or ecological-behavioral

(for example predators). For example, one vicariance splitting taxons’ ranges occurred

from historical climate change during glacial periods, resulting in the drying of tropical

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

rainforest in South America into smaller fragments of rainforest refugia surrounded by

grassland. Disjunct distribution from vicariant events are supports by fossil evidence and

by present species can’t be explained by natural dispersal. For example, Nothofagus

(Southern Beech) trees occur today in such widely separated regions as southern South

America and New Zealand, which can be explained by historic plate movement but not

dispersal. In another example, disjunct populations of southern migrations were created

during glacial ice advancements in North America, with isolated populations cut off as

ice retreats (partly because soil was removed by glaciers). Organisms from temporary or

fluctuating environments (such as seasonally variations in temperature) typically are less

separated by barriers. For instance, crossing mountain ranges are less of a barrier to

species from temperate climates (adapted to cold winters) compare to species from

tropical climates.

Geographic isolation from vicariant events leads to reproductive isolation. Even

if the disjunct population in time reunites, the geographic isolation may have already

resulted in the groups no longer being able to interbreeding (especially in animal species).

In other words, when barriers geographically isolate populations over time, a different

evolutionary lineage might occur with new species created (referred to as “allopatric”

speciation). Geographic isolation from vicariant events leads to reproductive isolation

because different geographic regions have different selective pressures, such as

temperature, rainfall, predators and/or competitors. In addition, even if the environments

separated by a barrier are not very different, the populations may differentiate because

different genetic combinations and mutations occur by chance.

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

Dispersal Biogeography

Disjunct biogeographic distributions may also be created by dispersal events.

Dispersal can be defined as movement away from one’s point of origin. Possible results

of dispersal include: extending within current range by colonizing new habitat in range;

or (2) colonizing distant location across a major physical barrier of unfavorable habitat.

Individuals may move great distances through unsuitable habitats by traveling through

corridors, flying over hostile environments, being blown or floating through sweepstake

events (such as a hurricane). Dispersal agents typically are wind, water, rafting, or

animals. Dispersal biogeography studies distribution patterns of organisms, emphasizing

dispersal capabilities as well as ecological properties of species to evaluate origins of taxa

in a biota. Dispersal mechanisms affect rate of species movement across landscape,

efficiency species colonize new areas, and successfully establishments.

Modes of dispersal vary from jump dispersal (such as the movement of gypsy

moth to North America) to diffusion (such as gypsy moth spread within North America).

Jump dispersal includes traveling over long distances across inhospitable habitat, in other

words long distance dispersal mostly by organisms that can fly or swim. Jump dispersal

events are rare, cover large distance, and are considered "surprising" events. These long

distance dispersals can explain large discontinuous distributions of some organisms, as

well as taxonomic similarity of distant biotas and populations. Most animal and all plant

jump dispersals are passive, although occasionally some animals have active long

distance dispersal. Those individuals who succeed at jump dispersal have the ability to:

(1) travel long distances; (2) withstand unfavorable conditions during passage; and (3)

establish viable population on arrival.

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

Pathways of dispersal can be broken into the categories of corridors, filters, and

sweepstakes. Corridors are routes that permit spread of taxa through continuous

favorable habitat with relatively little risk. These routes may in turn serve to link larger

areas of habitat. Famous corridors include past land connections created by sea level

changes, such as the Bering land bridge connecting Asia and North America. In contrast,

a “net” or “filter” is a route that contains patches of suitable habitat interspersed between

larger areas of unsuitable habitat. Filters can act as a barrier for some taxa, blocking or

slowing passage of some organisms. A classic example is the Isthmus of Panama

filtering the dispersal between North and South America starting around 3.5 million years

ago. The “sweepstakes” pathway of dispersal refers to chance dispersal across a major

barrier, in other words a long shot for dispersal which usually involves accidents, low

probability, and unusual means of travel. For instance, sweepstakes dispersal includes

birds caught in hurricanes, seeds traveling in upper atmosphere winds, and animals on

floating on driftwood (called “Noah’s Ark” if there is an assemblage of organisms

deposited in mass).

Human Impacts of Distributions

In addition to barriers and dispersal events, biogeography also explores species

distribution patterns impacted by humans, whether in limiting or expanding ranges.

Humans decrease other species ranges through habitat destruction, hunting and

commercial exploitation, polluting environments, deliberately or accidentally introducing

competing non-native species, change historic disturbance regimes, and causing the loss

of ecological partners. Species most likely to have their geographic ranges limited by

humans include: species who have relatively few offsprings who are nurtured for a long

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

time (“K” species), specialists who are selective about their niche conditions,

economically valuable species, naturally rare species, species with naturally restricted

ranges, species sensitive to pollutants, and species in competition with humans for habitat

or resources. People also increase species geographic ranges, either by introducing

species to a new geographic region or by expanding ranges by favoring some species that

can adapt to human landscapes (weeds, agricultural species, species adapted to urban

areas). Introduction of species by people into a new area may be deliberate or accidental.

Although most introduced species fail to establish viable population, those who succeed

typically have harmful long-term effects, competing with native species or transmitting

diseases.

Conclusion

Biogeographers seek to develop theories to explain past, present, and changing

future composition of plants and animals distribution patterns in order to better

understand the processes of how organisms interact with our planet. Biogeography is

both interdisciplinary within other subfields of geography as well as other disciplines,

such as ecology, geology, and biology. An integrative approach helps biogeographers

provide a holistic understanding of the diversity of life, and make recommendations for

the conservation of biological diversity.

See also in Encyclopedia of Geography: Chapters on individual Biomes, Biota,

Darwinism, Ecosystems, Island Biogeography, and Landscape Ecology

Joy Nystrom Mast

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 Encyclopedia of Geography (2010), Sage Publications; ISBN 9781412956970

Further Readings

Cox, C.B., & Moore, P.D., (2005). Biogeography: An Ecological and Evolutionary

Approach. Malden, MA: Blackwell.

Lomolino, M.L., & Heaney, L.R. (Eds) (2004). Frontiers of Biogeography. Sunderland,

MA: J.H. Sinauer.

Lomolino, M.V., Riddle, B., and Brown, J.H. (Eds.) (2006). Biogeography (3rd Edition).

Sunderland, MA: J.H. Sinauer.

Lomolino, M.L., Sax, D.F., & Brown, J.H., (Eds) (2004). Foundations of Biogeography:

Classic Papers with Commentaries. Chicago, IL: University of Chicago Press.

MacArthur, R.H., & Wilson, E.O. (reprinted 2001). The Theory of Island Biogeography.

Princeton, NJ: Princeton University Press.

MacDonald, G., (2003). Biogeography: Introduction to Space, Time, and Life. New

York, NY: John Wiley & Sons.

Quammen, D. (1997). The Song of the Dodo: Island Biogeography in an Age of

Extinction. New York, NY: Simon and Schuster.

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