Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, isolation and habitat area. Phytogeography is the branch of biogeography that studies the distribution of plants. Zoogeography is the branch that studies distribution of animals.
Knowledge of spatial variation in the numbers and types of organisms is as vital to us today as it was to our early human ancestors, as we adapt to heterogeneous but geographically predictable environments. Biogeography is an integrative field of inquiry that unites concepts and information from ecology, evolutionary biology, geology, and physical geography.
Modern biogeographic research combines information and ideas from many fields, from the physiological and ecological constraints on organismal dispersal to geological and climatological phenomena operating at global spatial scales and evolutionary time frames.
The short-term interactions within a habitat and species of organisms describe the ecological application of biogeography. Historical biogeography describes the long-term, evolutionary periods of time for broader classifications of organisms. Early scientists, beginning with Carl Linnaeus, contributed to the development of biogeography as a science. Beginning in the mid-18th century, Europeans explored the world and discovered the biodiversity of life.
The scientific theory of biogeography grows out of the work of Alexander von Humboldt (1769–1859), Hewett Cottrell Watson (1804–1881), Alphonse de Candolle (1806–1893), Alfred Russel Wallace (1823–1913), Philip Lutley Sclater (1829–1913) and other biologists and explorers.
The patterns of species distribution across geographical areas can usually be explained through a combination of historical factors such as: speciation, extinction, continental drift, and glaciation. Through observing the geographic distribution of species, we can see associated variations in sea level, river routes, habitat, and river capture. Additionally, this science considers the geographic constraints of landmass areas and isolation, as well as the available ecosystem energy supplies.
Over periods of ecological changes, biogeography includes the study of plant and animal species in: their past and/or present living refugium habitat; their interim living sites; and/or their survival locales. As writer David Quammen put it, "...biogeography does more than ask Which species? and Where. It also asks Why? and, what is sometimes more crucial, Why not?."
Modern biogeography often employs the use of Geographic Information Systems (GIS), to understand the factors affecting organism distribution, and to predict future trends in organism distribution. Often mathematical models and GIS are employed to solve ecological problems that have a spatial aspect to them.
Biogeography is most keenly observed on the world's islands. These habitats are often much more manageable areas of study because they are more condensed than larger ecosystems on the mainland. Islands are also ideal locations because they allow scientists to look at habitats that new invasive species have only recently colonized and can observe how they disperse throughout the island and change it. They can then apply their understanding to similar but more complex mainland habitats. Islands are very diverse in their biomes, ranging from the tropical to arctic climates. This diversity in habitat allows for a wide range of species study in different parts of the world.
One scientist who recognized the importance of these geographic locations was Charles Darwin, who remarked in his journal "The Zoology of Archipelagoes will be well worth examination". Two chapters in On the Origin of Species were devoted to geographical distribution.
The first discoveries that contributed to the development of biogeography as a science began in the mid-18th century, as Europeans explored the world and described the biodiversity of life. During the 18th century most views on the world were shaped around religion and for many natural theologists, the bible. Carl Linnaeus, in the mid-18th century, initiated the ways to classify organisms through his exploration of undiscovered territories. When he noticed that species were not as perpetual as he believed, he developed the Mountain Explanation to explain the distribution of biodiversity; when Noah's ark landed on Mount Ararat and the waters receded, the animals dispersed throughout different elevations on the mountain. This showed different species in different climates proving species were not constant. Linnaeus' findings set a basis for ecological biogeography. Through his strong beliefs in Christianity, he was inspired to classify the living world, which then gave way to additional accounts of secular views on geographical distribution. He argued that the structure of an animal was very closely related to its physical surroundings. This was important to a George Louis Buffon's rival theory of distribution.
Closely after Linnaeus, Georges-Louis Leclerc, Comte de Buffon observed shifts in climate and how species spread across the globe as a result. He was the first to see different groups of organisms in different regions of the world. Buffon saw similarities between some regions which led him to believe that at one point continents were connected and then water separated them and caused differences in species. His hypotheses were described by his books, Histoire Naturelle, and Générale et Particulière, in which he argued that varying geographical regions would have different forms of life. This was inspired by his observations comparing the Old and New World, as he determined distinct variations of species from the two regions. Buffon believed there was a single species creation event, and that different regions of the world were homes for varying species, which is an alternate view than that of Linnaeus. Buffon's law eventually became a principle of biogeography by explaining how similar environments were habitats for comparable types of organisms. Buffon also studied fossils which led him to believe that the earth was over tens of thousands of years old, and that humans had not lived there long in comparison to the age of the earth.
Following this period of exploration came the Age of Enlightenment in Europe, which attempted to explain the patterns of biodiversity observed by Buffon and Linnaeus. At the end of the 18th century, Alexander von Humboldt, known as the "founder of plant geography", developed the concept of physique generale to demonstrate the unity of science and how species fit together. As one of the first to contribute empirical data to the science of biogeography through his travel as an explorer, he observed differences in climate and vegetation. The earth was divided into regions which he defined as tropical, temperate, and arctic and within these regions there were similar forms of vegetation. This ultimately enabled him to create the isotherm, which allowed scientists to see patterns of life within different climates. He contributed his observations to findings of botanical geography by previous scientists, and sketched this description of both the biotic and abiotic features of the earth in his book, Cosmos.
Augustin de Candolle contributed to the field of biogeography as he observed species competition and the several differences that influenced the discovery of the diversity of life. He was a Swiss botanist and created the first Laws of Botanical Nomenclature in his work, Prodromus. He discussed plant distribution and his theories eventually had a great impact on Charles Darwin, who was inspired to consider species adaptations and evolution after learning about botanical geography. De Candolle was the first to describe the differences between the small-scale and large-scale distribution patterns of organisms around the globe.
In the 19th century, several additional scientists contributed new theories to further develop the concept of biogeography. Charles Lyell, being one of the first contributors in the 19th century, developed the Theory of Uniformitarianism after studying fossils. This theory explained how the world was not created by one sole catastrophic event, but instead from numerous creation events and locations. Uniformitarianism also introduced the idea that the Earth was actually significantly older than was previously accepted. Using this knowledge, Lyell concluded that it was possible for species to go extinct. Since he noted that earth's climate changes, he realized that species distribution must also change accordingly. Lyell argued that climate changes complemented vegetation changes, thus connecting the environmental surroundings to varying species. This largely influenced Charles Darwin in his development of the theory of evolution.
Charles Darwin was a natural theologist who studied around the world, and most importantly in the Galapagos Islands. Darwin introduced the idea of natural selection, as he theorized against previously accepted ideas that species were static or unchanging. His contributions to biogeography and the theory of evolution were different from those of other explorers of his time, because he developed a mechanism to describe the ways that species changed. His influential ideas include the development of theories regarding the struggle for existence and natural selection. Darwin's theories started a biological segment to biogeography and empirical studies, which enabled future scientists to develop ideas about the geographical distribution of organisms around the globe.
Alfred Russel Wallace studied the distribution of flora and fauna in the Amazon Basin and the Malay Archipelago in the mid-19th century. His research was essential to the further development of biogeography, and he was later nicknamed the "father of Biogeography". Wallace conducted fieldwork researching the habits, breeding and migration tendencies, and feeding behavior of thousands of species. He studied butterfly and bird distributions in comparison to the presence or absence of geographical barriers. His observations led him to conclude that the number of organisms present in a community was dependent on the amount of food resources in the particular habitat. Wallace believed species were dynamic by responding to biotic and abiotic factors. He and Philip Sclater saw biogeography as a source of support for the theory of evolution as they used Darwin's conclusion to explain how biogeography was similar to a record of species inheritance. Key findings, such as the sharp difference in fauna either side of the Wallace Line, and the sharp difference that existed between North and South America prior to their relatively recent faunal interchange, can only be understood in this light. Otherwise, the field of biogeography would be seen as a purely descriptive one.
Moving on to the 20th century, Alfred Wegener introduced the Theory of Continental Drift in 1912, though it was not widely accepted until the 1960s. This theory was revolutionary because it changed the way that everyone thought about species and their distribution around the globe. The theory explained how continents were formerly joined together in one large landmass, Pangea, and slowly drifted apart due to the movement of the plates below Earth's surface. The evidence for this theory is in the geological similarities between varying locations around the globe, fossil comparisons from different continents, and the jigsaw puzzle shape of the landmasses on Earth. Though Wegener did not know the mechanism of this concept of Continental Drift, this contribution to the study of biogeography was significant in the way that it shed light on the importance of environmental and geographic similarities or differences as a result of climate and other pressures on the planet. Importantly, late in his career Wegener recognised that testing his theory required measurement of continental movement rather than inference from fossils species distributions.
The publication of The Theory of Island Biogeography by Robert MacArthur and E.O. Wilson in 1967 showed that the species richness of an area could be predicted in terms of such factors as habitat area, immigration rate and extinction rate. This added to the long-standing interest in island biogeography. The application of island biogeography theory to habitat fragments spurred the development of the fields of conservation biology and landscape ecology.
Classic biogeography has been expanded by the development of molecular systematics, creating a new discipline known as phylogeography. This development allowed scientists to test theories about the origin and dispersal of populations, such as island endemics. For example, while classic biogeographers were able to speculate about the origins of species in the Hawaiian Islands, phylogeography allows them to test theories of relatedness between these populations and putative source populations in Asia and North America.
Biogeography continues as a point of study for many life sciences and geography students worldwide, however it may be under different broader titles within institutions such as ecology or evolutionary biology.
In recent years, one of the most important and consequential developments in biogeography has been to show how multiple organisms, including mammals like monkeys and reptiles like lizards, overcame barriers such as large oceans that many biogeographers formerly believed were impossible to cross. See also Oceanic dispersal.
Biogeography now incorporates many different fields including but not limited to physical geography, geology, botany and plant biology, zoology, and general biology. A biogeographer's main focus is on what environmental factors and what the influence of humans do to the distribution of the specific species of study. In terms of applications of biogeography as a science today, technological advances have allowed satellite imaging and processing of the Earth. Two main types of satellite imaging that are important within modern biogeography are Global Production Efficiency Model (GLO-PEM) and Geographic Information Systems (GIS). GLO-PEM uses satellite-imaging gives "repetitive, spatially contiguous, and time specific observations of vegetation". These observations are on a global scale. GIS can show certain processes on the earth's surface like whale locations, sea surface temperatures, and bathymetry. Current scientists also use coral reefs to delve into the history of biogeography through the fossilized reefs.
Paleobiogeography goes one step further to include paleogeographic data and considerations of plate tectonics. Using molecular analyses and corroborated by fossils, it has been possible to demonstrate that perching birds evolved first in the region of Australia or the adjacent Antarctic (which at that time lay somewhat further north and had a temperate climate). From there, they spread to the other Gondwanan continents and Southeast Asia – the part of Laurasia then closest to their origin of dispersal – in the late Paleogene, before achieving a global distribution in the early Neogene. Not knowing that at the time of dispersal, the Indian Ocean was much narrower than it is today, and that South America was closer to the Antarctic, one would be hard pressed to explain the presence of many "ancient" lineages of perching birds in Africa, as well as the mainly South American distribution of the suboscines.
Paleobiogeography also helps constrain hypotheses on the timing of biogeographic events such as vicariance and geodispersal, and provides unique information on the formation of regional biotas. For example, data from species-level phylogenetic and biogeographic studies tell us that the Amazonian fish fauna accumulated in increments over a period of tens of millions of years, principally by means of allopatric speciation, and in an arena extending over most of the area of tropical South America (Albert & Reis 2011). In other words, unlike some of the well-known insular faunas (Galapagos finches, Hawaiian drosophilid flies, African rift lake cichlids), the species-rich Amazonian ichthyofauna is not the result of recent adaptive radiations.
For freshwater organisms, landscapes are divided naturally into discrete drainage basins by watersheds, episodically isolated and reunited by erosional processes. In regions like the Amazon Basin (or more generally Greater Amazonia, the Amazon basin, Orinoco basin, and Guianas) with an exceptionally low (flat) topographic relief, the many waterways have had a highly reticulated history over geological time. In such a context, stream capture is an important factor affecting the evolution and distribution of freshwater organisms. Stream capture occurs when an upstream portion of one river drainage is diverted to the downstream portion of an adjacent basin. This can happen as a result of tectonic uplift (or subsidence), natural damming created by a landslide, or headward or lateral erosion of the watershed between adjacent basins.
Some fundamental concepts in biogeography include:
The study of comparative biogeography can follow two main lines of investigation:
There are many types of biogeographic units used in biogeographic regionalisation schemes, as there are many criteria (species composition, physiognomy, ecological aspects) and hierarchization schemes: biogeographic realms (or ecozones), bioregions (sensu stricto), ecoregions, zoogeographical regions, floristic regions, vegetation types, biomes, etc.
The Avon Wheatbelt is an Australian bioregion in Western Australia and part of the larger Southwest Australia savanna ecoregion.Biogeochemical cycle
In ecology and Earth science, a biogeochemical cycle or substance turnover or cycling of substances is a pathway by which a chemical substance moves through biotic (biosphere) and abiotic (lithosphere, atmosphere, and hydrosphere) compartments of Earth. There are biogeochemical cycles for the chemical elements calcium, carbon, hydrogen, mercury, nitrogen, oxygen, phosphorus, selenium, and sulfur; molecular cycles for water and silica; macroscopic cycles such as the rock cycle; as well as human-induced cycles for synthetic compounds such as polychlorinated biphenyl (PCB). In some cycles there are reservoirs where a substance remains for a long period of time (such as an ocean or lake for water).Community (ecology)
In ecology, a community is a group or association of populations of two or more different species occupying the same geographical area and in a particular time, also known as a biocoenosis. The term community has a variety of uses. In its simplest form it refers to groups of organisms in a specific place or time, for example, "the fish community of Lake Ontario before industrialization".
Community ecology or synecology is the study of the interactions between species in communities on many spatial and temporal scales, including the distribution, structure, abundance, demography, and interactions between coexisting populations. The primary focus of community ecology is on the interactions between populations as determined by specific genotypic and phenotypic characteristics. Community ecology has its origin in European plant sociology. Modern community ecology examines patterns such as variation in species richness, equitability, productivity and food web structure (see community structure); it also examines processes such as predator–prey population dynamics, succession, and community assembly.
On a deeper level the meaning and value of the community concept in ecology is up for debate. Communities have traditionally been understood on a fine scale in terms of local processes constructing (or destructing) an assemblage of species, such as the way climate change is likely to affect the make-up of grass communities. Recently this local community focus has been criticised. Robert Ricklefs has argued that it is more useful to think of communities on a regional scale, drawing on evolutionary taxonomy and biogeography, where some species or clades evolve and others go extinct.Cosmopolitan distribution
In biogeography, a taxon is said to have a cosmopolitan distribution if its range extends across all or most of the world in appropriate habitats. Such a taxon is said to exhibit cosmopolitanism or cosmopolitism. The opposite extreme is endemism.Ecological niche
In ecology, a niche (CanE, UK: or US: ) is the match of a species to a specific environmental condition. It describes how an organism or population responds to the distribution of resources and competitors (for example, by growing when resources are abundant, and when predators, parasites and pathogens are scarce) and how it in turn alters those same factors (for example, limiting access to resources by other organisms, acting as a food source for predators and a consumer of prey). "The type and number of variables comprising the dimensions of an environmental niche vary from one species to another [and] the relative importance of particular environmental variables for a species may vary according to the geographic and biotic contexts".A Grinnellian niche is determined by the habitat in which a species lives and its accompanying behavioral adaptations. An Eltonian niche emphasizes that a species not only grows in and responds to an environment, it may also change the environment and its behavior as it grows. The Hutchinsonian niche uses mathematics and statistics to try to explain how species coexist within a given community.
The concept of ecological niche is central to ecological biogeography, which focuses on spatial patterns of ecological communities. "Species distributions and their dynamics over time result from properties of the species, environmental variation..., and interactions between the two—in particular the abilities of some species, especially our own, to modify their environments and alter the range dynamics of many other species." Alteration of an ecological niche by its inhabitants is the topic of niche construction.The majority of species exist in a standard ecological niche, sharing behaviors, adaptations, and functional traits similar to the other closely related species within the same broad taxonomic class, but there are exceptions. A premier example of a non-standard niche filling species is the flightless, ground-dwelling kiwi bird of New Zealand, which feeds on worms and other ground creatures, and lives its life in a mammal-like niche. Island biogeography can help explain island species and associated unfilled niches.Geraldton Sandplains
Geraldton Sandplains is an interim Australian bioregion of Western Australia and part of the larger Southwest Australia savanna ecoregion, as assessed by the World Wildlife Fund.Great Sandy Desert
The Great Sandy Desert (GSD) is an interim Australian bioregion, located in the north west of Western Australia straddling the Pilbara and southern Kimberley regions. It is the second largest desert in Australia after the Great Victoria Desert and encompasses an area of 284,993 square kilometres (110,036 sq mi). The Gibson Desert lies to the south and the Tanami Desert lies to the east of the Great Sandy Desert.Great Victoria Desert
The Great Victoria Desert, an interim Australian bioregion, is a sparsely populated desert area in Western Australia and South Australia.Holarctic
The Holarctic is the name for the biogeographic realm that encompasses the majority of habitats found throughout the northern continents of the world, combining Wallace's Palearctic zoogeographical region, consisting of North Africa and all of Eurasia (with the exception of the southern Arabian Peninsula, Southeast Asia, and the Indian subcontinent), and the Nearctic zoogeographical region, consisting of North America, north of Mexico. These regions are further subdivided into a variety of ecoregions. Many ecosystems, and the animal and plant communities that depend on them, are found across multiple continents in large portions of this realm. The continuity of these ecosystems results from the shared glacial history of the realm. The floristic Boreal Kingdom corresponds to the Holarctic realm.Indigenous (ecology)
In biogeography, a species is indigenous to a given region or ecosystem if its presence in that region is the result of only natural processes, with no human intervention. The term is equivalent to the concept of native or autochthonous species. Every wild organism (as opposed to a domesticated organism) has its own natural range of distribution in which it is regarded as indigenous. Outside this native range, a species may be introduced by human activity, either intentionally or unintentionally; it is then referred to as an introduced species within the regions where it was anthropogenically introduced.The notion of indigeneity is often a blurred concept, as is a function of both time and political boundaries. Seen over long periods of time, plants and animals take part in the constant movement of tectonic plates—species appear and may flourish, endure, or become extinct, and their distribution is rarely static or confined to a particular geographic location.
An indigenous species in a location is not necessarily also endemic to that location. Endemic species are exclusively found in a particular place. An indigenous species may occur in areas other than the one under consideration. The terms endemic and indigenous also do not imply that an organism necessarily first originated or evolved where it is currently found.Insular biogeography
Insular biogeography or island biogeography is a field within biogeography that examines the factors that affect the species richness and diversification of isolated natural communities. The theory was originally developed to explain the pattern of the species–area relationship occurring in oceanic islands. Under either name it is now used in reference to any ecosystem (present or past) that is isolated due to being surrounded by unlike ecosystems, and has been extended to mountain peaks, seamounts, oases, fragmented forests, and even natural habitats isolated by human land development. The field was started in the 1960s by the ecologists Robert H. MacArthur and E. O. Wilson, who coined the term island biogeography in their inaugural contribution to Princeton's Monograph in Population Biology series, which attempted to predict the number of species that would exist on a newly created island.Journal of Biogeography
The Journal of Biogeography is a peer-reviewed scientific journal in biogeography that was established in 1974. It covers aspects of spatial, ecological, and historical biogeography. The founding editor-in-chief was David Watts, followed by John Flenley and Philip Stott (1987-2004). The current editor-in-chief is Peter Linder (University of Zurich).Land bridge
A land bridge, in biogeography, is an isthmus or wider land connection between otherwise separate areas, over which animals and plants are able to cross and colonise new lands. A land bridge can be created by marine regression, in which sea levels fall, exposing shallow, previously submerged sections of continental shelf; or when new land is created by plate tectonics; or occasionally when the sea floor rises due to post-glacial rebound after an ice age.Phytogeography
Phytogeography (from Greek φυτόν, phytón = "plant" and γεωγραφία, geographía = "geography" meaning also distribution) or botanical geography is the branch of biogeography that is concerned with the geographic distribution of plant species and their influence on the earth's surface. Phytogeography is concerned with all aspects of plant distribution, from the controls on the distribution of individual species ranges (at both large and small scales, see species distribution) to the factors that govern the composition of entire communities and floras. Geobotany, by contrast, focuses on the geographic space's influence on plants.Tasmanian Southern Ranges
The Tasmanian Southern Ranges is an interim Australian bioregion located in the southern region of Tasmania, comprising 757,228 hectares (1,871,150 acres).Unified neutral theory of biodiversity
The unified neutral theory of biodiversity and biogeography (here "Unified Theory" or "UNTB") is a hypothesis and the title of a monograph by ecologist Stephen Hubbell. The hypothesis aims to explain the diversity and relative abundance of species in ecological communities, although like other neutral theories of ecology, Hubbell's hypothesis assumes that the differences between members of an ecological community of trophically similar species are "neutral", or irrelevant to their success. This implies that biodiversity arises at random, as each species follows a random walk. The hypothesis has sparked controversy, and some authors consider it a more complex version of other null models that fit the data better.Neutrality means that at a given trophic level in a food web, species are equivalent in birth rates, death rates, dispersal rates and speciation rates, when measured on a per-capita basis. This can be considered a null hypothesis to niche theory. Hubbell built on earlier neutral concepts, including MacArthur & Wilson's theory of island biogeography and Gould's concepts of symmetry and null models.An ecological community is a group of trophically similar, sympatric species that actually or potentially compete in a local area for the same or similar resources. Under the Unified Theory, complex ecological interactions are permitted among individuals of an ecological community (such as competition and cooperation), provided that all individuals obey the same rules. Asymmetric phenomena such as parasitism and predation are ruled out by the terms of reference; but cooperative strategies such as swarming, and negative interaction such as competing for limited food or light are allowed (so long as all individuals behave in the same way).
The Unified Theory also makes predictions that have profound implications for the management of biodiversity, especially the management of rare species.The theory predicts the existence of a fundamental biodiversity constant, conventionally written θ, that appears to govern species richness on a wide variety of spatial and temporal scales.World Register of Marine Species
The World Register of Marine Species (WoRMS) is a taxonomic database that aims to provide an authoritative and comprehensive list of names of marine organisms.Zoogeography
Zoogeography is the branch of the science of biogeography that is concerned with the geographic distribution (present and past) of animal species.Zoology
Zoology () is the branch of biology that studies the animal kingdom, including the structure, embryology, evolution, classification, habits, and distribution of all animals, both living and extinct, and how they interact with their ecosystems. The term is derived from Ancient Greek ζῷον, zōion, i.e. "animal" and λόγος, logos, i.e. "knowledge, study".-History-
-ancient history to Darwin-
The history of zoology traces the study of the animal kingdom from ancient to modern times. Although the concept of zoology as a single coherent field arose much later, the zoological sciences emerged from natural history reaching back to the biological works of Aristotle and Galen in the ancient Greco-Roman world. This ancient work was further developed in the Middle Ages by Muslim physicians and scholars such as Albertus Magnus. During the Renaissance and early modern period, zoological thought was revolutionized in Europe by a renewed interest in empiricism and the discovery of many novel organisms. Prominent in this movement were Vesalius and William Harvey, who used experimentation and careful observation in physiology, and naturalists such as Carl Linnaeus, Jean-Baptiste Lamarck, and Buffon who began to classify the diversity of life and the fossil record, as well as the development and behavior of organisms. Microscopy revealed the previously unknown world of microorganisms, laying the groundwork for cell theory. The growing importance of natural theology, partly a response to the rise of mechanical philosophy, encouraged the growth of natural history (although it entrenched the argument from design).
Over the 18th, 19th, and 20th centuries, zoology became an increasingly professional scientific discipline. Explorer-naturalists such as Alexander von Humboldt investigated the interaction between organisms and their environment, and the ways this relationship depends on geography, laying the foundations for biogeography, ecology and ethology. Naturalists began to reject essentialism and consider the importance of extinction and the mutability of species. Cell theory provided a new perspective on the fundamental basis of life.
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