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116-411: In biogeography , a land bridge is an isthmus or wider land connection between otherwise separate areas, over which animals and plants are able to cross and colonize 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

232-476: 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

348-465: A dramatic effect on community structure. Hunting of sea otters, for example, is thought to have led indirectly to the extinction of the Steller's sea cow ( Hydrodamalis gigas ). While the keystone species concept has been used extensively as a conservation tool, it has been criticized for being poorly defined from an operational stance. It is difficult to experimentally determine what species may hold

464-428: A dramatic impact on the ecology of individual species or whole ecosystems. For instance, the replacement of an ant species by another (invasive) ant species has been shown to affect how elephants reduce tree cover and thus the predation of lions on zebras . A trophic level (from Greek troph , τροφή, trophē, meaning "food" or "feeding") is "a group of organisms acquiring a considerable majority of its energy from

580-412: A flattened body relative to the main populations that live in open savanna. The population that lives in an isolated rock outcrop hides in crevasses where its flattened body offers a selective advantage. Habitat shifts also occur in the developmental life history of amphibians, and in insects that transition from aquatic to terrestrial habitats. Biotope and habitat are sometimes used interchangeably, but

696-421: 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. Two global information systems are either dedicated to, or have strong focus on, biogeography (in the form of the spatial location of observations of organisms), namely

812-419: 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

928-749: 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

1044-612: A higher one." Small scale patterns do not necessarily explain large scale phenomena, otherwise captured in the expression (coined by Aristotle) 'the sum is greater than the parts'. "Complexity in ecology is of at least six distinct types: spatial, temporal, structural, process, behavioral, and geometric." From these principles, ecologists have identified emergent and self-organizing phenomena that operate at different environmental scales of influence, ranging from molecular to planetary, and these require different explanations at each integrative level . Ecological complexity relates to

1160-444: A keystone role in each ecosystem. Furthermore, food web theory suggests that keystone species may not be common, so it is unclear how generally the keystone species model can be applied. Complexity is understood as a large computational effort needed to piece together numerous interacting parts exceeding the iterative memory capacity of the human mind. Global patterns of biological diversity are complex. This biocomplexity stems from

1276-408: A keystone species can result in a community collapse just as the removal of the keystone in an arch can result in the arch's loss of stability. Sea otters ( Enhydra lutris ) are commonly cited as an example of a keystone species because they limit the density of sea urchins that feed on kelp . If sea otters are removed from the system, the urchins graze until the kelp beds disappear, and this has

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1392-741: A legacy niche that was constructed before their time. Biomes are larger units of organization that categorize regions of the Earth's ecosystems, mainly according to the structure and composition of vegetation. There are different methods to define the continental boundaries of biomes dominated by different functional types of vegetative communities that are limited in distribution by climate, precipitation, weather, and other environmental variables. Biomes include tropical rainforest , temperate broadleaf and mixed forest , temperate deciduous forest , taiga , tundra , hot desert , and polar desert . Other researchers have recently categorized other biomes, such as

1508-439: A linear successional route, changes might occur quickly or slowly over thousands of years before specific forest successional stages are brought about by biological processes. An ecosystem's area can vary greatly, from tiny to vast. A single tree is of little consequence to the classification of a forest ecosystem, but is critically relevant to organisms living in and on it. Several generations of an aphid population can exist over

1624-439: A macroscopic view of the system. While the notion of trophic levels provides insight into energy flow and top-down control within food webs, it is troubled by the prevalence of omnivory in real ecosystems. This has led some ecologists to "reiterate that the notion that species clearly aggregate into discrete, homogeneous trophic levels is fiction." Nonetheless, recent studies have shown that real trophic levels do exist, but "above

1740-516: A mix of herbivores and predators). Omnivores do not fit neatly into a functional category because they eat both plant and animal tissues. It has been suggested that omnivores have a greater functional influence as predators because compared to herbivores, they are relatively inefficient at grazing. Trophic levels are part of the holistic or complex systems view of ecosystems. Each trophic level contains unrelated species that are grouped together because they share common ecological functions, giving

1856-420: A new appreciation for the influence that organisms have on the ecosystem and evolutionary process. The term "niche construction" is more often used in reference to the under-appreciated feedback mechanisms of natural selection imparting forces on the abiotic niche. An example of natural selection through ecosystem engineering occurs in the nests of social insects , including ants, bees, wasps, and termites. There

1972-413: A number of methods have been developed to produce arguably more complete "predictive" or "modelled" distributions for species based on their associated environmental or other preferences (such as availability of food or other habitat requirements); this approach is known as either Environmental niche modelling (ENM) or Species distribution modelling (SDM). Depending on the reliability of the source data and

2088-435: A population-level phenomenon, as with the migration routes followed by plants as they occupied northern post-glacial environments. Plant ecologists use pollen records that accumulate and stratify in wetlands to reconstruct the timing of plant migration and dispersal relative to historic and contemporary climates. These migration routes involved an expansion of the range as plant populations expanded from one area to another. There

2204-401: A region), and sites are classed either as sources or sinks. A site is a generic term that refers to places where ecologists sample populations, such as ponds or defined sampling areas in a forest. Source patches are productive sites that generate a seasonal supply of juveniles that migrate to other patch locations. Sink patches are unproductive sites that only receive migrants; the population at

2320-987: 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 (ecozones), bioregions ( sensu stricto ), ecoregions , zoogeographical regions , floristic regions , vegetation types, biomes , etc. The terms biogeographic unit, biogeographic area can be used for these categories, regardless of rank. In 2008, an International Code of Area Nomenclature

2436-517: A seasonal influx of new immigrants. A dynamic metapopulation structure evolves from year to year, where some patches are sinks in dry years and are sources when conditions are more favorable. Ecologists use a mixture of computer models and field studies to explain metapopulation structure. Community ecology examines how interactions among species and their environment affect the abundance, distribution and diversity of species within communities. Johnson & Stinchcomb (2007) Community ecology

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2552-456: A species describes the environment over which a species is known to occur and the type of community that is formed as a result. More specifically, "habitats can be defined as regions in environmental space that are composed of multiple dimensions, each representing a biotic or abiotic environmental variable; that is, any component or characteristic of the environment related directly (e.g. forage biomass and quality) or indirectly (e.g. elevation) to

2668-402: A specific habitat, such as a cave or a pond, and principles gleaned from small-scale studies are extrapolated to larger systems. Feeding relations require extensive investigations, e.g. into the gut contents of organisms, which can be difficult to decipher, or stable isotopes can be used to trace the flow of nutrient diets and energy through a food web. Despite these limitations, food webs remain

2784-446: A valuable tool in understanding community ecosystems. Food webs illustrate important principles of ecology : some species have many weak feeding links (e.g., omnivores ) while some are more specialized with fewer stronger feeding links (e.g., primary predators ). Such linkages explain how ecological communities remain stable over time and eventually can illustrate a "complete" web of life. The disruption of food webs may have

2900-440: A whole suite 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

3016-399: A wide array of interacting levels of organization spanning micro-level (e.g., cells ) to a planetary scale (e.g., biosphere ) phenomena . Ecosystems, for example, contain abiotic resources and interacting life forms (i.e., individual organisms that aggregate into populations which aggregate into distinct ecological communities). Because ecosystems are dynamic and do not necessarily follow

3132-485: Is a central concept in the ecology of organisms and is sub-divided into the fundamental and the realized niche. The fundamental niche is the set of environmental conditions under which a species is able to persist. The realized niche is the set of environmental plus ecological conditions under which a species persists. The Hutchinsonian niche is defined more technically as a " Euclidean hyperspace whose dimensions are defined as environmental variables and whose size

3248-504: Is a function of the number of values that the environmental values may assume for which an organism has positive fitness ." Biogeographical patterns and range distributions are explained or predicted through knowledge of a species' traits and niche requirements. Species have functional traits that are uniquely adapted to the ecological niche. A trait is a measurable property, phenotype , or characteristic of an organism that may influence its survival. Genes play an important role in

3364-414: Is a larger taxonomy of movement, such as commuting, foraging, territorial behavior, stasis, and ranging. Dispersal is usually distinguished from migration because it involves the one-way permanent movement of individuals from their birth population into another population. In metapopulation terminology, migrating individuals are classed as emigrants (when they leave a region) or immigrants (when they enter

3480-424: 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

3596-461: Is an emergent homeostasis or homeorhesis in the structure of the nest that regulates, maintains and defends the physiology of the entire colony. Termite mounds, for example, maintain a constant internal temperature through the design of air-conditioning chimneys. The structure of the nests themselves is subject to the forces of natural selection. Moreover, a nest can survive over successive generations, so that progeny inherit both genetic material and

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3712-502: 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

3828-404: Is another statistical approach that is often used in conservation research . Metapopulation models simplify the landscape into patches of varying levels of quality, and metapopulations are linked by the migratory behaviours of organisms. Animal migration is set apart from other kinds of movement because it involves the seasonal departure and return of individuals from a habitat. Migration is also

3944-438: Is greater than respiration) by photosynthesis or chemosynthesis . Heterotrophs are organisms that must feed on others for nourishment and energy (respiration exceeds production). Heterotrophs can be further sub-divided into different functional groups, including primary consumers (strict herbivores), secondary consumers ( carnivorous predators that feed exclusively on herbivores), and tertiary consumers (predators that feed on

4060-404: Is in the geological similarities between varying locations around the globe, the geographic distribution of some fossils (including the mesosaurs ) on various 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

4176-473: 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

4292-590: Is the Park Grass Experiment , which was initiated in 1856. Another example is the Hubbard Brook study , which has been in operation since 1960. Holism remains a critical part of the theoretical foundation in contemporary ecological studies. Holism addresses the biological organization of life that self-organizes into layers of emergent whole systems that function according to non-reducible properties. This means that higher-order patterns of

4408-460: Is the natural science of the relationships among living organisms and their environment . Ecology considers organisms at the individual, population , community , ecosystem , and biosphere levels. Ecology overlaps with the closely related sciences of biogeography , evolutionary biology , genetics , ethology , and natural history . Ecology is a branch of biology , and is the study of abundance , biomass , and distribution of organisms in

4524-413: Is the biosphere: the total sum of ecosystems on the planet. Ecological relationships regulate the flux of energy, nutrients, and climate all the way up to the planetary scale. For example, the dynamic history of the planetary atmosphere's CO 2 and O 2 composition has been affected by the biogenic flux of gases coming from respiration and photosynthesis, with levels fluctuating over time in relation to

4640-522: Is the study of the interactions among a collection of species that inhabit the same geographic area. Community ecologists study the determinants of patterns and processes for two or more interacting species. Research in community ecology might measure species diversity in grasslands in relation to soil fertility. It might also include the analysis of predator-prey dynamics, competition among similar plant species, or mutualistic interactions between crabs and corals. These ecosystems, as we may call them, are of

4756-556: Is used for managing wildlife stocks and setting harvest quotas. In cases where basic models are insufficient, ecologists may adopt different kinds of statistical methods, such as the Akaike information criterion , or use models that can become mathematically complex as "several competing hypotheses are simultaneously confronted with the data." The concept of metapopulations was defined in 1969 as "a population of populations which go extinct locally and recolonize". Metapopulation ecology

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4872-614: 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

4988-713: The Global Biodiversity Information Facility (GBIF: 2.57 billion species occurrence records reported as at August 2023) and, for marine species only, the Ocean Biodiversity Information System (OBIS, originally the Ocean Biogeographic Information System : 116 million species occurrence records reported as at August 2023), while at a national scale, similar compilations of species occurrence records also exist such as

5104-405: The abundance or biomass at each level. When the relative abundance or biomass of each species is sorted into its respective trophic level, they naturally sort into a 'pyramid of numbers'. Species are broadly categorized as autotrophs (or primary producers ), heterotrophs (or consumers ), and Detritivores (or decomposers ). Autotrophs are organisms that produce their own food (production

5220-436: The logistic equation by Pierre Verhulst : where N(t) is the number of individuals measured as biomass density as a function of time, t , r is the maximum per-capita rate of change commonly known as the intrinsic rate of growth, and α {\displaystyle \alpha } is the crowding coefficient, which represents the reduction in population growth rate per individual added. The formula states that

5336-479: 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

5452-509: 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

5568-551: The Earth and atmospheric conditions within a narrow self-regulating range of tolerance. Population ecology studies the dynamics of species populations and how these populations interact with the wider environment. A population consists of individuals of the same species that live, interact, and migrate through the same niche and habitat. A primary law of population ecology is the Malthusian growth model which states, "a population will grow (or decline) exponentially as long as

5684-597: The Gómez Farias Region, Tamaulipas, Mexico , which has been described as "ground-breaking" and "a classic treatise in historical biogeography". Martin applied several disciplines including ecology , botany , climatology , geology , and Pleistocene dispersal routes to examine the herpetofauna of a relatively small and largely undisturbed area, but ecologically complex, situated on the threshold of temperate – tropical (nearctic and neotropical) regions, including semiarid lowlands at 70 meters elevation and

5800-619: 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

5916-452: 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

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6032-482: 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 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

6148-603: 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

6264-639: The U.K. National Biodiversity Network , the Atlas of Living Australia , and many others. In the case of the oceans, in 2017 Costello et al. analyzed the distribution of 65,000 species of marine animals and plants as then documented in OBIS, and used the results to distinguish 30 distinct marine realms, split between continental-shelf and offshore deep-sea areas. Since it is self evident that compilations of species occurrence records cannot cover with any completeness, areas that have received either limited or no sampling,

6380-414: The anticipated effects of climate change can also be used to show potential changes in species distributions that may occur in the future based on such scenarios. 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

6496-421: 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

6612-521: The concept was first proposed by Jules Marcou in Lettres sur les roches du Jura et leur distribution géographique dans les deux hémisphères ("Letters on the rocks of the Jura [Mountains] and their geographic distribution in the two hemispheres"), 1857–1860. Hypothesized land bridges included: The theory of continental drift provided an alternate explanation that did not require land bridges. However

6728-744: The context of the environment. It encompasses life processes, interactions, and adaptations ; movement of materials and energy through living communities; successional development of ecosystems; cooperation, competition, and predation within and between species ; and patterns of biodiversity and its effect on ecosystem processes. Ecology has practical applications in conservation biology , wetland management, natural resource management ( agroecology , agriculture , forestry , agroforestry , fisheries , mining , tourism ), urban planning ( urban ecology ), community health , economics , basic and applied science , and human social interaction ( human ecology ). The word ecology ( German : Ökologie )

6844-443: The continental drift theory was not widely accepted until the development of plate tectonics in the early 1960s, which more completely explained the motion of continents over geological time. Biogeography 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

6960-420: 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

7076-714: The diversity of life from genes to ecosystems and spans every level of biological organization. The term has several interpretations, and there are many ways to index, measure, characterize, and represent its complex organization. Biodiversity includes species diversity , ecosystem diversity , and genetic diversity and scientists are interested in the way that this diversity affects the complex ecological processes operating at and among these respective levels. Biodiversity plays an important role in ecosystem services which by definition maintain and improve human quality of life. Conservation priorities and management techniques require different approaches and considerations to address

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7192-530: The dynamic resilience of ecosystems that transition to multiple shifting steady-states directed by random fluctuations of history. Long-term ecological studies provide important track records to better understand the complexity and resilience of ecosystems over longer temporal and broader spatial scales. These studies are managed by the International Long Term Ecological Network (LTER). The longest experiment in existence

7308-456: The ecology and evolution of plants and animals. Ecological theory has also been used to explain self-emergent regulatory phenomena at the planetary scale: for example, the Gaia hypothesis is an example of holism applied in ecological theory. The Gaia hypothesis states that there is an emergent feedback loop generated by the metabolism of living organisms that maintains the core temperature of

7424-434: The energy is transferred through a chain of organisms by consumption. The simplified linear feeding pathways that move from a basal trophic species to a top consumer is called the food chain . Food chains in an ecological community create a complex food web. Food webs are a type of concept map that is used to illustrate and study pathways of energy and material flows. Empirical measurements are generally restricted to

7540-460: The environment experienced by all individuals in the population remains constant." Simplified population models usually starts with four variables: death, birth, immigration , and emigration . An example of an introductory population model describes a closed population, such as on an island, where immigration and emigration does not take place. Hypotheses are evaluated with reference to a null hypothesis which states that random processes create

7656-504: 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

7772-451: The equilibrium, r / α {\displaystyle r/\alpha } as K , which is known as the "carrying capacity." Population ecology builds upon these introductory models to further understand demographic processes in real study populations. Commonly used types of data include life history , fecundity , and survivorship, and these are analyzed using mathematical techniques such as matrix algebra . The information

7888-597: 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

8004-589: The flux of energy and matter through an environment. Ecosystems have biophysical feedback mechanisms that moderate processes acting on living ( biotic ) and abiotic components of the planet. Ecosystems sustain life-supporting functions and provide ecosystem services like biomass production (food, fuel, fiber, and medicine), the regulation of climate , global biogeochemical cycles , water filtration , soil formation , erosion control, flood protection, and many other natural features of scientific, historical, economic, or intrinsic value. The scope of ecology contains

8120-458: The formation of regional biotas. For example, data from species-level phylogenetic and biogeographic studies tell us that the Amazonian teleost 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

8236-483: The former Lifemapper project at the University of Kansas (now continued as a part of BiotaPhy ) and AquaMaps , which as at 2023 contain modelled distributions for around 200,000 terrestrial, and 33,000 species of teleosts , marine mammals and invertebrates, respectively. One advantage of ENM/SDM is that in addition to showing current (or even past) modelled distributions, insertion of changed parameters such as

8352-400: The former applies to a community's environment, whereas the latter applies to a species' environment. Definitions of the niche date back to 1917, but G. Evelyn Hutchinson made conceptual advances in 1957 by introducing a widely adopted definition: "the set of biotic and abiotic conditions in which a species is able to persist and maintain stable population sizes." The ecological niche

8468-544: The former relates only to the physical modifications of the habitat whereas the latter also considers the evolutionary implications of physical changes to the environment and the feedback this causes on the process of natural selection. Ecosystem engineers are defined as: "organisms that directly or indirectly modulate the availability of resources to other species, by causing physical state changes in biotic or abiotic materials. In so doing they modify, maintain and create habitats." The ecosystem engineering concept has stimulated

8584-518: The full ecological scope of biodiversity. Natural capital that supports populations is critical for maintaining ecosystem services and species migration (e.g., riverine fish runs and avian insect control) has been implicated as one mechanism by which those service losses are experienced. An understanding of biodiversity has practical applications for species and ecosystem-level conservation planners as they make management recommendations to consulting firms, governments, and industry. The habitat of

8700-655: 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

8816-408: The herbivore trophic level, food webs are better characterized as a tangled web of omnivores." A keystone species is a species that is connected to a disproportionately large number of other species in the food-web . Keystone species have lower levels of biomass in the trophic pyramid relative to the importance of their role. The many connections that a keystone species holds means that it maintains

8932-406: The human and oceanic microbiomes . To a microbe , the human body is a habitat and a landscape. Microbiomes were discovered largely through advances in molecular genetics , which have revealed a hidden richness of microbial diversity on the planet. The oceanic microbiome plays a significant role in the ecological biogeochemistry of the planet's oceans. The largest scale of ecological organization

9048-411: 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. In 1958 paleontologist Paul S. Martin published A Biogeography of Reptiles and Amphibians in

9164-404: The interplay among ecological processes that operate and influence patterns at different scales that grade into each other, such as transitional areas or ecotones spanning landscapes. Complexity stems from the interplay among levels of biological organization as energy, and matter is integrated into larger units that superimpose onto the smaller parts. "What were wholes on one level become parts on

9280-412: The interplay of development and environmental expression of traits. Resident species evolve traits that are fitted to the selection pressures of their local environment. This tends to afford them a competitive advantage and discourages similarly adapted species from having an overlapping geographic range. The competitive exclusion principle states that two species cannot coexist indefinitely by living off

9396-559: 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

9512-402: The latter focuses on materials and energy fluxes. System behaviors must first be arrayed into different levels of the organization. Behaviors corresponding to higher levels occur at slow rates. Conversely, lower organizational levels exhibit rapid rates. For example, individual tree leaves respond rapidly to momentary changes in light intensity, CO 2 concentration, and the like. The growth of

9628-424: The lifespan of a single leaf. Each of those aphids, in turn, supports diverse bacterial communities. The nature of connections in ecological communities cannot be explained by knowing the details of each species in isolation, because the emergent pattern is neither revealed nor predicted until the ecosystem is studied as an integrated whole. Some ecological principles, however, do exhibit collective properties where

9744-432: The lower adjacent level (according to ecological pyramids ) nearer the abiotic source." Links in food webs primarily connect feeding relations or trophism among species. Biodiversity within ecosystems can be organized into trophic pyramids, in which the vertical dimension represents feeding relations that become further removed from the base of the food chain up toward top predators, and the horizontal dimension represents

9860-488: The mid-18th century, improved our classifications of organisms through the exploration of undiscovered territories by his students and disciples. 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

9976-515: The most important and consequential developments in biogeography has been to show how multiple organisms, including mammals like monkeys and reptiles like squamates , 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

10092-558: The most various kinds and sizes. They form one category of the multitudinous physical systems of the universe, which range from the universe as a whole down to the atom. Tansley (1935) Ecosystems may be habitats within biomes that form an integrated whole and a dynamically responsive system having both physical and biological complexes. Ecosystem ecology is the science of determining the fluxes of materials (e.g. carbon, phosphorus) between different pools (e.g., tree biomass, soil organic material). Ecosystem ecologists attempt to determine

10208-505: 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

10324-683: The nature of the models employed (including the scales for which data are available), maps generated from such models may then provide better representations of the "real" biogeographic distributions of either individual species, groups of species, or biodiversity as a whole, however it should also be borne in mind that historic or recent human activities (such as hunting of great whales , or other human-induced exterminations) may have altered present-day species distributions from their potential "full" ecological footprint. Examples of predictive maps produced by niche modelling methods based on either GBIF (terrestrial) or OBIS (marine, plus some freshwater) data are

10440-456: The nature of the unique physical environments that shapes the biodiversity within each. A more recent addition to ecosystem ecology are technoecosystems , which are affected by or primarily the result of human activity. A food web is the archetypal ecological network . Plants capture solar energy and use it to synthesize simple sugars during photosynthesis . As plants grow, they accumulate nutrients and are eaten by grazing herbivores , and

10556-573: The niche is called the ecotope , which is defined as the full range of environmental and biological variables affecting an entire species. Organisms are subject to environmental pressures, but they also modify their habitats. The regulatory feedback between organisms and their environment can affect conditions from local (e.g., a beaver pond ) to global scales, over time and even after death, such as decaying logs or silica skeleton deposits from marine organisms. The process and concept of ecosystem engineering are related to niche construction , but

10672-476: The northernmost cloud forest in the western hemisphere at over 2200 meters. 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

10788-531: The number of nitrogen fixers , can lead to disproportionate, perhaps irreversible, changes in the system properties." Biodiversity refers to the variety of life and its processes. It includes the variety of living organisms, the genetic differences among them, the communities and ecosystems in which they occur, and the ecological and evolutionary processes that keep them functioning, yet ever-changing and adapting. Noss & Carpenter (1994) Biodiversity (an abbreviation of "biological diversity") describes

10904-399: The observed data. In these island models, the rate of population change is described by: where N is the total number of individuals in the population, b and d are the per capita rates of birth and death respectively, and r is the per capita rate of population change. Using these modeling techniques, Malthus' population principle of growth was later transformed into a model known as

11020-406: The organization and structure of entire communities. The loss of a keystone species results in a range of dramatic cascading effects (termed trophic cascades ) that alters trophic dynamics, other food web connections, and can cause the extinction of other species. The term keystone species was coined by Robert Paine in 1969 and is a reference to the keystone architectural feature as the removal of

11136-583: The origins of species in the Hawaiian Islands , phylogeography allows them to test theories of relatedness between these populations and putative source populations on various continents, notably 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

11252-448: 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

11368-468: The published work of George Perkins Marsh ("Man and Nature"). Within an ecosystem, organisms are linked to the physical and biological components of their environment to which they are adapted. Ecosystems are complex adaptive systems where the interaction of life processes form self-organizing patterns across different scales of time and space. Ecosystems are broadly categorized as terrestrial , freshwater , atmospheric, or marine . Differences stem from

11484-502: The rate of change in population size ( d N ( t ) / d t {\displaystyle \mathrm {d} N(t)/\mathrm {d} t} ) will grow to approach equilibrium, where ( d N ( t ) / d t = 0 {\displaystyle \mathrm {d} N(t)/\mathrm {d} t=0} ), when the rates of increase and crowding are balanced, r / α {\displaystyle r/\alpha } . A common, analogous model fixes

11600-420: 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,

11716-424: The same limiting resource ; one will always out-compete the other. When similarly adapted species overlap geographically, closer inspection reveals subtle ecological differences in their habitat or dietary requirements. Some models and empirical studies, however, suggest that disturbances can stabilize the co-evolution and shared niche occupancy of similar species inhabiting species-rich communities. The habitat plus

11832-484: The sea floor rises due to post-glacial rebound after an ice age . In the late 19th and early 20 centuries, vanished land bridges were an explanation for observed affinities of plants and animals in distant locations. Such scientists as Joseph Dalton Hooker noted puzzling geological, botanical, and zoological similarities between widely separated areas, and proposed land bridges between appropriate land masses that allowed species to spread between land masses. In geology,

11948-399: The site will disappear unless rescued by an adjacent source patch or environmental conditions become more favorable. Metapopulation models examine patch dynamics over time to answer potential questions about spatial and demographic ecology. The ecology of metapopulations is a dynamic process of extinction and colonization. Small patches of lower quality (i.e., sinks) are maintained or rescued by

12064-524: The study of ecology into a conceptually manageable framework, the biological world is organized into a nested hierarchy , ranging in scale from genes , to cells , to tissues , to organs , to organisms , to species , to populations , to guilds , to communities , to ecosystems , to biomes , and up to the level of the biosphere . This framework forms a panarchy and exhibits non-linear behaviors; this means that "effect and cause are disproportionate, so that small changes to critical variables, such as

12180-428: 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

12296-432: The sum of the components explain the properties of the whole, such as birth rates of a population being equal to the sum of individual births over a designated time frame. The main subdisciplines of ecology, population (or community ) ecology and ecosystem ecology , exhibit a difference not only in scale but also in two contrasting paradigms in the field. The former focuses on organisms' distribution and abundance, while

12412-416: 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

12528-670: The tree responds more slowly and integrates these short-term changes. O'Neill et al. (1986) The scale of ecological dynamics can operate like a closed system, such as aphids migrating on a single tree, while at the same time remaining open about broader scale influences, such as atmosphere or climate. Hence, ecologists classify ecosystems hierarchically by analyzing data collected from finer scale units, such as vegetation associations , climate, and soil types , and integrate this information to identify emergent patterns of uniform organization and processes that operate on local to regional, landscape , and chronological scales. To structure

12644-415: The underlying causes of these fluxes. Research in ecosystem ecology might measure primary production (g C/m^2) in a wetland in relation to decomposition and consumption rates (g C/m^2/y). This requires an understanding of the community connections between plants (i.e., primary producers) and the decomposers (e.g., fungi and bacteria). The underlying concept of an ecosystem can be traced back to 1864 in

12760-439: The use of a location by the animal." For example, a habitat might be an aquatic or terrestrial environment that can be further categorized as a montane or alpine ecosystem. Habitat shifts provide important evidence of competition in nature where one population changes relative to the habitats that most other individuals of the species occupy. For example, one population of a species of tropical lizard ( Tropidurus hispidus ) has

12876-447: 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,

12992-593: Was coined in 1866 by the German scientist Ernst Haeckel . The science of ecology as we know it today began with a group of American botanists in the 1890s. Evolutionary concepts relating to adaptation and natural selection are cornerstones of modern ecological theory . Ecosystems are dynamically interacting systems of organisms, the communities they make up, and the non-living ( abiotic ) components of their environment. Ecosystem processes, such as primary production , nutrient cycling , and niche construction , regulate

13108-400: 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

13224-529: 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,

13340-776: Was proposed for biogeography. It achieved limited success; some studies commented favorably on it, but others were much more critical, and it "has not yet gained a significant following". Similarly, a set of rules for paleobiogeography has achieved limited success. In 2000, Westermann suggested that the difficulties in getting formal nomenclatural rules established in this field might be related to "the curious fact that neither paleo- nor neobiogeographers are organized in any formal groupings or societies, nationally (so far as I know) or internationally — an exception among active disciplines." Ecology Ecology (from Ancient Greek οἶκος ( oîkos )  'house' and -λογία ( -logía )  'study of')

13456-502: 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

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