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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. Mycogeography is the branch that studies distribution of fungi, such as mushrooms.
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, taxonomy, geology, physical geography, palaeontology, and climatology.
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.
The scientific theory of biogeography grows out of the work of Alexander von Humboldt (1769–1859),Francisco Jose de Caldas (1768-1816), 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 in his work, the 36 volume Histoire Naturelle, 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 the 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 birth of the 19th 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.
Several additional scientists contributed new theories to further develop the concept of biogeography. Charles Lyell 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 1967showed 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, general biology, and modelling. A biogeographer's main focus is on how the environment and humans affect the distribution of species as well as other manifestations of Life such as species or genetic diversity. Biogeography is being applied to biodiversity conservation and planning, projecting global environmental changes on species and biomes, projecting the spread of infectious diseases, invasive species, and for supporting planning for the establishment of crops. Technological evolving and advances have allowed for generating a whole suit of predictor variables for biogeographic analysis, including 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.
Biogeography is a synthetic science, related to geography, biology, soil science, geology, climatology, ecology and evolution.
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 terms biogeographic unit,biogeographic area or bioregion sensu lato, can be used for these categories, regardless of rank.
In 2008, an International Code of Area Nomenclature was proposed for biogeography.
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Ecology is the study of the relationships between living organisms, including humans, and their physical environment, . Ecology considers organisms at the individual, population, community, ecosystems, and biosphere level. Ecology overlaps with the closely related sciences of biogeography, evolutionary biology, genetics, ethology and natural history. Ecology is a branch of biology, and it is not synonymous with environmentalism.
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 ('animal'), and λόγος, logos.
In ecology, a niche 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 and how it in turn alters those same factors. "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".
Phylogeography is the study of the historical processes that may be responsible for the past to present geographic distributions of genealogical lineages. This is accomplished by considering the geographic distribution of individuals in light of genetics, particularly population genetics.
This glossary of ecology is a list of definitions of terms and concepts in ecology and related fields. For more specific definitions from other glossaries related to ecology, see Glossary of biology, Glossary of evolutionary biology, and Glossary of environmental science.
Biological dispersal refers to both the movement of individuals from their birth site to their breeding site, as well as the movement from one breeding site to another . Dispersal is also used to describe the movement of propagules such as seeds and spores. Technically, dispersal is defined as any movement that has the potential to lead to gene flow. The act of dispersal involves three phases: departure, transfer, settlement and there are different fitness costs and benefits associated with each of these phases. Through simply moving from one habitat patch to another, the dispersal of an individual has consequences not only for individual fitness, but also for population dynamics, population genetics, and species distribution. Understanding dispersal and the consequences both for evolutionary strategies at a species level, and for processes at an ecosystem level, requires understanding on the type of dispersal, the dispersal range of a given species, and the dispersal mechanisms involved.
Endemism is the state of a species being native to a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found elsewhere. For example, the Cape sugarbird is found exclusively in southwestern South Africa and is therefore said to be endemic to that particular part of the world.
Evolutionary ecology lies at the intersection of ecology and evolutionary biology. It approaches the study of ecology in a way that explicitly considers the evolutionary histories of species and the interactions between them. Conversely, it can be seen as an approach to the study of evolution that incorporates an understanding of the interactions between the species under consideration. The main subfields of evolutionary ecology are life history evolution, sociobiology, the evolution of interspecific interactions and the evolution of biodiversity and of ecological communities.
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 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.
Ecology is a new science and considered as an important branch of biological science, having only become prominent during the second half of the 20th century. Ecological thought is derivative of established currents in philosophy, particularly from ethics and politics. Its history stems all the way back to the 4th century. One of the first ecologists whose writings survive may have been Aristotle or perhaps his student, Theophrastus, both of whom had interest in many species of animals and plants. Theophrastus described interrelationships between animals and their environment as early as the 4th century BC. Ecology developed substantially in the 18th and 19th century. It began with Carl Linnaeus and his work with the economy of nature. Soon after came Alexander von Humboldt and his work with botanical geography. Alexander von Humboldt and Karl Möbius then contributed with the notion of biocoenosis. Eugenius Warming’s work with ecological plant geography led to the founding of ecology as a discipline. Charles Darwin’s work also contributed to the science of ecology, and Darwin is often attributed with progressing the discipline more than anyone else in its young history. Ecological thought expanded even more in the early 20th century. Major contributions included: Eduard Suess’ and Vladimir Vernadsky’s work with the biosphere, Arthur Tansley’s ecosystem, Charles Elton's Animal Ecology, and Henry Cowles ecological succession. Ecology influenced the social sciences and humanities. Human ecology began in the early 20th century and it recognized humans as an ecological factor. Later James Lovelock advanced views on earth as a macro-organism with the Gaia hypothesis. Conservation stemmed from the science of ecology. Important figures and movements include Shelford and the ESA, National Environmental Policy act, George Perkins Marsh, Theodore Roosevelt, Stephen A. Forbes, and post-Dust Bowl conservation. Later in the 20th century world governments collaborated on man’s effects on the biosphere and Earth’s environment.
Zoogeography is the branch of the science of biogeography that is concerned with geographic distribution of animal species.
Landscape connectivity in ecology is, broadly, "the degree to which the landscape facilitates or impedes movement among resource patches". Alternatively, connectivity may be a continuous property of the landscape and independent of patches and paths. Connectivity includes both structural connectivity and functional connectivity. Functional connectivity includes actual connectivity and potential connectivity in which movement paths are estimated using the life-history data. The degree to which a landscape is connected determines the amount of dispersal there is among patches, which influences gene flow, local adaptation, extinction risk, colonization probability, and the potential for organisms to move as they cope with climate change.
The following outline is provided as an overview of and topical guide to ecology:
The body size-species richness distribution is a pattern observed in the way taxa are distributed over large spatial scales. The number of species that exhibit small body size generally far exceed the number of species that are large-bodied. Macroecology has long sought to understand the mechanisms that underlie the patterns of biodiversity, such as the body size-species richness pattern.
Plant ecology is a subdiscipline of ecology which studies the distribution and abundance of plants, the effects of environmental factors upon the abundance of plants, and the interactions among and between plants and other organisms. Examples of these are the distribution of temperate deciduous forests in North America, the effects of drought or flooding upon plant survival, and competition among desert plants for water, or effects of herds of grazing animals upon the composition of grasslands.
Island ecology is the study of island organisms and their interactions with each other and the environment. Islands account for nearly 1/6 of earth’s total land area, yet the ecology of island ecosystems is vastly different from that of mainland communities. Their isolation and high availability of empty niches lead to increased speciation. As a result, island ecosystems comprise 30% of the world’s biodiversity hotspots, 50% of marine tropical diversity, and some of the most unusual and rare species. Many species still remain unknown.
Ecological fitting is "the process whereby organisms colonize and persist in novel environments, use novel resources or form novel associations with other species as a result of the suites of traits that they carry at the time they encounter the novel condition". It can be understood as a situation in which a species' interactions with its biotic and abiotic environment seem to indicate a history of coevolution, when in actuality the relevant traits evolved in response to a different set of biotic and abiotic conditions.
Microbial biogeography is a subset of biogeography, a field that concerns the distribution of organisms across space and time. Although biogeography traditionally focused on plants and larger animals, recent studies have broadened this field to include distribution patterns of microorganisms. This extension of biogeography to smaller scales—known as "microbial biogeography"—is enabled by ongoing advances in genetic technologies.
This glossary of evolutionary biology is a list of definitions of terms and concepts used in the study of evolutionary biology, population biology, speciation, and phylogenetics, as well as sub-disciplines and related fields. For additional terms from related glossaries, see Glossary of genetics, Glossary of ecology, and Glossary of biology.
Taxon cycles refer to a biogeographical theory of how species evolve through range expansions and contractions over time associated with adaptive shifts in the ecology and morphology of species. The taxon cycle concept was explicitly formulated by biologist E. O. Wilson in 1961 after he surveyed the distributions, habitats, behavior and morphology of ant species in the Melanesian archipelago.