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Fauna

Fauna refers to the collective animal life inhabiting a specific region, time period, or environment, encompassing all species of animals such as mammals, birds, reptiles, amphibians, fish, insects, and other invertebrates present in that context.^[1] This term, derived from Fauna, the name of a Roman goddess associated with wildlife and fertility, is often contrasted with flora, which denotes the plant life in the same area, together forming the biotic components of an ecosystem.^[2] Fauna can vary widely by habitat, from the diverse marine species in coral reefs to the specialized insects in soil layers, and is studied to understand biodiversity patterns and ecological dynamics.^[1] In ecology, fauna plays a critical role in maintaining ecosystem balance through processes like pollination, seed dispersal, predation, and nutrient cycling.^[3] For instance, herbivorous animals such as deer and insects regulate plant populations, while predators like birds and mammals control pest species, preventing overpopulation and supporting food web stability.^[4] Wildlife, as a key component of fauna, also contributes to broader environmental services, including carbon sequestration via animal-mediated decomposition and habitat engineering by species like beavers that create wetlands.^[5]^[6] Additionally, fauna influences pathogen transmission and pollination, which are essential for agricultural productivity and human health.^[7] The study and conservation of fauna are vital amid global threats like habitat loss, climate change, and invasive species, which have led to declining biodiversity and ecosystem disruptions.^[3] Efforts to protect fauna often focus on preserving habitats and monitoring populations, as diverse animal communities enhance resilience against environmental stresses and provide cultural, recreational, and economic value to societies.^[8] Notable examples include the role of large mammals in forest regeneration through seed transport^[9] and the importance of soil fauna in decomposition processes that sustain soil fertility.^[10]

Fundamentals

Definition and Scope

Fauna refers to the animal life occurring within a particular region, geological period, or environment, encompassing multicellular animals such as vertebrates and invertebrates.^[11] This term typically excludes plants, which are classified under flora, and microorganisms, which are addressed separately in microbial ecology unless explicitly included in broader biotic studies.^[11]^[12] The scope of fauna includes both wild and domesticated animals, though ecological analyses often emphasize native, wild species that form the natural assemblages of an area.^[13] For instance, the fauna of the Amazon rainforest highlights this focus, featuring diverse native groups such as over 400 mammal species including jaguars and sloths, more than 1,300 bird species like scarlet macaws, numerous reptiles such as anacondas, and approximately 2.5 million insect species.^[14] These examples illustrate how fauna represents the collective animal populations adapted to specific ecosystems, contributing to biodiversity assessments. Fauna can be examined at various scales, from local assemblages like the unique species on isolated islands—such as the endemic reptiles and birds of the Galápagos—to global distributions spanning continents and oceans.^[12] As a dynamic assemblage, fauna is shaped by factors including geography, which determines species ranges; climate, which influences habitat suitability; and evolutionary processes like speciation and extinction that drive changes over time.^[15]^[16] This fluidity underscores fauna's role as an evolving component of biotic communities.

Importance in Ecology and Biodiversity

Fauna play pivotal roles in ecological systems by occupying diverse positions within food webs, acting as predators, prey, pollinators, and decomposers, which collectively maintain ecosystem stability and nutrient cycling. As predators, animals such as wolves regulate herbivore populations, preventing overgrazing and promoting vegetation recovery; for instance, the reintroduction of gray wolves in Yellowstone National Park in 1995 led to a trophic cascade that reduced elk numbers, allowing willow and aspen trees to regenerate and benefiting beaver and bird populations.^[17] In turn, many animals serve as prey, supporting higher trophic levels and ensuring energy transfer across the web. Pollinators like bees and butterflies facilitate plant reproduction by transferring pollen, enabling the persistence of approximately 87.5% of flowering plant species that rely on animal pollination.^[18] Decomposers, including insects such as dung beetles and millipedes, break down organic matter, recycling nutrients back into the soil and preventing waste accumulation.^[19] Faunal diversity significantly contributes to global biodiversity metrics, including species richness—the total number of species in a given area—and endemism, where species are unique to specific regions, enhancing ecosystem resilience against disturbances. Current estimates indicate approximately 7.8 million animal species worldwide, with insects comprising over 70% of this total, underscoring their dominance in driving biodiversity patterns.^[20] High faunal endemism in isolated areas, such as islands, amplifies local richness and supports unique evolutionary adaptations, as seen in regions like Madagascar where over 90% of certain animal groups are endemic.^[21] This diversity underpins ecosystem services, including pest control by birds that consume agricultural insects, reducing crop losses, and seed dispersal by mammals like rodents and primates, which promote forest regeneration and plant distribution across landscapes.^[18]^[22] The interconnectedness of faunal biodiversity extends to human well-being, providing essential services and resources derived from animal diversity. Biodiversity hotspots like coral reefs, home to over 4,000 fish species and countless invertebrates, support fisheries that feed nearly a billion people globally and protect coastlines from erosion, while also fostering tourism economies.^[23] Faunal diversity has spurred medical advancements, particularly through venoms; for example, captopril, a blockbuster drug for hypertension derived from the Brazilian pit viper's venom, has treated millions since its approval in 1981, exemplifying how animal biochemistry informs pharmaceutical innovation.^[24] These contributions highlight fauna's indispensable role in sustaining both natural and human systems.

Etymology and Historical Development

Etymology

The term "fauna" originates from Latin, referring to Fauna, a Roman goddess of fertility and the earth, who was mythologically portrayed as the sister, wife, or daughter of Faunus, the deity presiding over forests, fields, and wildlife.^[25] This mythological figure embodied the nurturing aspects of nature, particularly in relation to animals and rural life, with her name evoking the woodland spirits known as fauns. The scientific adoption of "fauna" to describe animal assemblages occurred in the mid-18th century, popularized by Swedish naturalist Carl Linnaeus in his 1746 work Fauna Suecica, a systematic catalog of Swedish animal species that paralleled his botanical counterpart Flora Suecica (1745). Linnaeus's use of the term transformed its ancient connotation from a divine entity to a precise descriptor for the totality of animals in a given geographic area, establishing a linguistic symmetry with "flora" for plants.^[26] Early applications in European natural history followed this model, with "fauna" appearing in regional surveys to denote local animal populations, such as in descriptions of continental European wildlife assemblages during the Enlightenment era.^[26] This evolution underscored a broader trend in 18th-century science toward systematic nomenclature, bridging classical mythology with empirical observation.

Evolution of the Concept

The concept of fauna, encompassing the collective animal life of a given region or period, emerged as a structured scientific idea in the 18th century, heavily influenced by Carl Linnaeus's taxonomic system outlined in Systema Naturae (1735 onward). Linnaeus's binomial nomenclature and hierarchical classification of animals provided a standardized framework for cataloging species, shifting from descriptive natural histories to systematic inventories that laid the groundwork for regional faunal studies.^[27] This approach facilitated the documentation of local animal assemblages, marking a transition from anecdotal observations to empirical zoological surveys. By the late 18th and early 19th centuries, naturalists began applying Linnaean principles to specific geographies, as seen in John James Audubon's The Birds of America (1827–1838), which illustrated and described over 400 North American bird species, representing one of the first comprehensive regional faunal works and highlighting geographic variation in animal distributions.^[28] The 19th century saw further evolution through the integration of evolutionary theory and biogeography, expanding fauna beyond mere lists to dynamic assemblages shaped by historical and environmental factors. Charles Darwin's On the Origin of Species (1859) introduced mechanisms like natural selection that influenced understandings of faunal diversity and distribution, prompting studies on how animals adapt to regions. Building on this, Alfred Russel Wallace's The Geographical Distribution of Animals (1876) delineated six major faunal regions (Palearctic, Ethiopian, Indian, Australian, Nearctic, and Neotropical) based on species endemism and barriers like oceans and mountains, establishing biogeography as a key lens for fauna analysis and emphasizing dispersal and evolution over static classification.^[29] These developments in the 20th century intertwined fauna with ecology, viewing animal communities as interdependent systems rather than isolated taxa, influenced by Wallace's and Darwin's legacies in explaining faunal boundaries through geological and climatic history. In the modern era, post-1960s advancements have refined the fauna concept by incorporating conservation, genetics, and environmental dynamics. The International Union for Conservation of Nature (IUCN) established the Red List in 1964, initiating systematic faunal assessments through inventories of threatened species, which evolved into global biodiversity benchmarks tracking faunal declines and informing policy.^[30] Since the 2000s, molecular phylogenetics has revolutionized faunal understanding by revealing evolutionary relationships and cryptic species diversity, as demonstrated in comprehensive animal phylogenies that reshape regional assemblages and challenge traditional boundaries.^[31] Concurrently, studies on climate change impacts, such as those documenting poleward shifts in northeastern North American forest fauna due to warming temperatures, have highlighted fauna as responsive to anthropogenic pressures, integrating predictive modeling into the concept.^[32]

Classification Systems

Taxonomic versus Ecological Classifications

Taxonomic classification of fauna organizes animals into hierarchical groups based on shared evolutionary relationships and ancestry, primarily through methods like cladistics, which constructs phylogenetic trees using shared derived characteristics.^[33] This approach groups species into major phyla such as Chordata (including vertebrates like mammals and birds) and Arthropoda (encompassing insects, crustaceans, and arachnids), reflecting their common descent. Advancements in DNA sequencing have refined these classifications by analyzing genetic data to resolve evolutionary divergences, enabling more precise identification of monophyletic clades—groups comprising an ancestor and all its descendants.^[34] The system employs binomial nomenclature, established by Carl Linnaeus in the 18th century, where each species receives a unique two-part Latin name (genus and species, e.g., Homo sapiens for humans), providing a stable, universal framework for naming across the animal kingdom. In contrast, ecological classification categorizes fauna according to their functional roles, adaptations, and interactions within ecosystems, rather than genetic lineage. Animals are often grouped into guilds—assemblages of species that exploit similar resources or perform comparable functions, such as carnivores that prey on other animals, herbivores that consume vegetation, or detritivores that break down organic matter.^[35] For instance, this approach emphasizes trophic levels and niche partitioning, where species like wolves (Canis lupus) and sharks (Carcharodon carcharias) might both fall into a carnivorous guild despite belonging to distant classes (Mammalia and Chondrichthyes, respectively) within the phylum Chordata.^[36] Such classifications are inherently dynamic, varying by environmental context, and prioritize ecosystem processes over fixed hierarchies. The key differences between these systems lie in their foundations and applications: taxonomic classification is static and lineage-based, aiming for objective, universal hierarchies that do not change with habitat, whereas ecological classification is context-dependent and functional, adapting to specific ecosystems and potentially grouping distantly related species together.^[37] An example of overlap occurs in bird classifications, where taxonomic groupings place all birds in the class Aves based on shared avian ancestry, but ecological guilds might unite disparate orders like raptors (Accipitriformes, carnivorous predators) and passerines (Passeriformes, often insectivorous foragers) under a broader "avian predator" category when assessing food web dynamics.^[38] This convergence highlights how unrelated species can converge evolutionarily to fill similar niches. Modern integrations of these approaches leverage phylogenomics—genome-scale analyses of evolutionary relationships—to inform both taxonomic revisions and ecological assessments, particularly through environmental DNA (eDNA) surveys that detect faunal presence via genetic traces in soil or water. In the 2020s, eDNA metabarcoding has advanced faunal monitoring by combining phylogenetic markers for species identification with ecological data on community composition, outperforming traditional surveys in sensitivity and cost for detecting biodiversity patterns.^[39] For example, phylogenomic studies have resolved deep animal phylogenies, such as those of reef fishes, enabling better integration of evolutionary history into functional diversity metrics for conservation.^[40]

Global and Regional Faunas

The global distribution of fauna is fundamentally shaped by historical biogeographic patterns, as outlined in Alfred Russel Wallace's seminal 1876 work The Geographical Distribution of Animals, which delineates six primary faunal regions based on distinct assemblages of animal species reflecting evolutionary histories and geological barriers. These regions—Palearctic (encompassing Europe and northern Asia), Nearctic (North America), Neotropical (Central and South America), Ethiopian (sub-Saharan Africa), Oriental (Southeast Asia and the Indian subcontinent), and Australasian (Australia, New Guinea, and surrounding islands)—are characterized by unique higher taxa that do not overlap significantly across boundaries, such as the dominance of placental mammals in the Palearctic and Nearctic versus the prevalence of marsupials and monotremes in the Australasian region. Wallace's framework emphasizes how continental drift, isolation, and adaptation have produced these discrete zones, with each region hosting endemic families and genera that underscore faunal uniqueness. Modern analyses have refined this framework, identifying up to 11 biogeographic realms by incorporating phylogenetic relationships and updated species distributions.^[29]^[41]^[42]^[43] Biogeographic patterns within these regions reveal high levels of endemism, particularly in isolated landmasses, where species diversification occurs without external gene flow; for instance, Madagascar, situated within the broader Ethiopian region, exhibits endemism rates of 95–100% for terrestrial vertebrates due to its 88-million-year isolation from other landmasses. Transition zones further highlight these divisions, such as Wallace's Line, an imaginary boundary running through the Indonesian archipelago from the Lombok Strait northward to the Philippines, which sharply separates Oriental faunas (featuring tigers, elephants, and primates) to the west from Australasian assemblages (dominated by marsupials like kangaroos and unique birds like cassowaries) to the east, a demarcation driven by deep ocean barriers limiting dispersal during low sea levels. These patterns not only illustrate faunal homogeneity within regions but also abrupt faunal turnover at boundaries, with endemism serving as a key metric of regional identity.^[44]^[45] Representative regional faunas exemplify these distinctions: the Nearctic region's North American fauna includes iconic large herbivores like the American bison (Bison bison), which once roamed vast grasslands in herds of millions, and raptors such as the bald eagle (Haliaeetus leucocephalus), adapted to coastal and riverine ecosystems across the continent. In contrast, the Ethiopian region's African savanna fauna features apex predators like lions (Panthera leo) and leopards (Panthera pardus) alongside megaherbivores such as the African bush elephant (Loxodonta africana), which shapes landscapes through its foraging and supports diverse trophic interactions in open woodlands and grasslands. These assemblages reflect adaptive radiations tailored to regional climates and vegetation, with the Neotropical region's sloths and New World monkeys or the Oriental region's diverse ungulates providing further parallels in ecological roles.^[46]^[47]^[48]^[49] As of 2025, human-driven globalization is increasingly blurring traditional faunal boundaries through the translocation of species, with invasive alien species contributing to homogenization by outcompeting or hybridizing with native taxa across Wallace's regions; for example, introduced mammals like rats and cats have accelerated extinctions in Australasian island faunas, while global trade facilitates the spread of pathogens and predators that erode endemism rates in isolated hotspots like Madagascar. A 2025 preprint indicates that nearly 40,000 non-native species have become established globally, profoundly reshaping biogeographical boundaries, including across Wallace's Line, with human-mediated dispersal rates orders of magnitude higher than natural and contributing significantly to native species extinctions. These impacts underscore the vulnerability of Wallace's framework to anthropogenic change, as accelerated dispersal rates threaten the evolutionary legacies encoded in global faunal patterns.^[50]

Subdivisions by Habitat and Environment

Terrestrial and Regional Faunas

Terrestrial fauna encompasses the animal assemblages inhabiting land-based environments such as soils, forests, and grasslands, where invertebrates, particularly insects, dominate in both species richness and abundance, comprising at least 80% of all living terrestrial species.^[51] Vertebrates like mammals and birds play key ecological roles, with insects serving as primary decomposers, pollinators, and prey bases that support higher trophic levels. In European temperate forests, for instance, the fauna includes large herbivores such as roe deer (Capreolus capreolus) and red deer (Cervus elaphus), carnivores like the red fox (Vulpes vulpes) and brown bear (Ursus arctos), alongside a vast array of invertebrates including beetles, millipedes, and earthworms that contribute to soil aeration and nutrient cycling.^[52] Adaptations in terrestrial fauna vary markedly across biomes to cope with environmental stressors like aridity or humidity. In desert ecosystems, species exhibit specialized physiological traits for water conservation; the kangaroo rat (Dipodomys spp.), for example, obtains all necessary moisture from metabolic water in seeds and produces highly concentrated urine via efficient kidneys, enabling survival without free-standing water.^[53] In contrast, tropical rainforest biomes support exceptionally high faunal diversity, with these forests harboring over 50% of the world's terrestrial animal species despite covering less than 6% of Earth's land surface; the Amazon Basin, as the largest such biome, contributes significantly to this, hosting millions of insect species, diverse mammals like jaguars (Panthera onca), and birds such as harpy eagles (Harpia harpyja).^[54]^[55] Regional variations in terrestrial fauna often arise from isolation and historical biogeography, leading to high endemism in unique landscapes. Australia's outback, an arid continental interior, exemplifies this with endemic species adapted to sparse vegetation and extreme temperatures, including the emu (Dromaius novaehollandiae), a large flightless bird that forages across grasslands and uses its powerful legs for rapid escape from predators.^[56] Monotremes like the platypus (Ornithorhynchus anatinus), though more associated with eastern waterways, highlight Australia's broader faunal distinctiveness as one of only two extant egg-laying mammals, with distributions influenced by regional aridity gradients.^[57] These patterns underscore how geographic barriers foster specialized assemblages distinct from global norms.

Aquatic Faunas

Aquatic fauna refers to the animal communities inhabiting marine and freshwater environments, shaped profoundly by physical factors such as salinity, depth, and water flow. Marine ecosystems, covering over 70% of Earth's surface, host the majority of aquatic species, with diversity influenced by vertical zonation and substrate type. In contrast, freshwater habitats, comprising less than 1% of global water volume, support a disproportionate share of vertebrate diversity, including about 51% of all known fish species, though total species richness is generally lower than in marine systems due to habitat fragmentation and isolation.^[58]^[59] Marine fauna is broadly divided into pelagic and benthic communities. The pelagic zone encompasses the open water column, away from the seafloor and shores, where highly mobile species dominate; examples include cetaceans like whales and dolphins, large predators such as sharks, and schools of fish like tuna that migrate vast distances for feeding and reproduction.^[60] In the benthic zone, organisms dwell on or within the seafloor sediments, adapting to low oxygen and high pressure; representative species include crustaceans like crabs, echinoderms such as sea stars and sea urchins, and mollusks including clams and polychaete worms that burrow or crawl along the bottom.^[61] A notable phenomenon in deeper marine habitats is deep-sea gigantism, where species grow larger than shallow-water relatives due to cold temperatures slowing metabolism, scarce food resources favoring energy-efficient large bodies, and reduced predation pressure; examples include the colossal squid (Mesonychoteuthis hamiltoni) in pelagic depths and the giant isopod (Bathynomus giganteus) on the benthic abyssopelagic floor.^[62]^[63] Oceanic depth zonation further structures marine fauna, creating distinct communities based on light penetration, temperature, and pressure. The epipelagic zone (0–200 meters) supports abundant photosynthetic-based food chains, hosting diverse planktivores and predators like flying fish and seabirds. The mesopelagic "twilight" zone (200–1,000 meters) features bioluminescent organisms such as lanternfish (Myctophidae) and squid that use light for hunting and camouflage. Deeper bathypelagic (1,000–4,000 meters), abyssal (4,000–6,000 meters), and hadal (>6,000 meters) zones yield sparser populations of resilient species, including anglerfish in the bathypelagic and slow-moving sea cucumbers in the abyssal plains, where chemosynthesis around hydrothermal vents sustains isolated ecosystems.^[64]^[65] Freshwater fauna, often termed limnofauna, thrives in rivers, lakes, and wetlands, facing challenges like seasonal flooding and oxygen variability. In rivers, migratory species such as salmon (Oncorhynchus spp.) undertake long journeys from oceans to spawn in upstream tributaries, while amphibians like frogs and salamanders rely on these waters for breeding and larval development. Lakes host specialized communities, including endemic fish adapted to stable conditions. Overall, freshwater systems exhibit lower total species diversity compared to marine environments—approximately 19,000 freshwater fish species (about 51% of all known fish species) versus over 18,000 marine—but boast high endemism, with nearly 50% of freshwater fish restricted to specific basins; a prime example is the Amazon river dolphin (Inia geoffrensis), a pink-hued cetacean unique to the Amazon and Orinoco river systems, preying on diverse fish like tetras and catfish.^[66] Salinity gradients, particularly in estuarine faunas, create transitional zones where freshwater meets marine waters, fostering euryhaline species tolerant of brackish conditions. These environments support high productivity, with organisms like blue crabs (Callinectes sapidus), oysters (Crassostrea virginica), and juvenile fish such as herring using mangroves and mudflats for nursery habitats. Estuarine biodiversity peaks at intermediate salinities, where consumption rates by predators are maximized, highlighting the gradient's role in structuring food webs.^[67]^[68]

Extreme and Specialized Environments

Faunas inhabiting extreme and specialized environments exhibit remarkable adaptations to conditions such as perpetual cold, total darkness, chemical toxicity, or extreme pressure, enabling survival where typical biodiversity is limited. These ecosystems, often isolated and resource-scarce, foster unique evolutionary pressures that result in specialized physiological, morphological, and behavioral traits. Cryofauna in polar regions and high altitudes, troglofauna and stygofauna in subterranean realms, and chemosynthetic communities at hydrothermal vents represent key examples of such resilience, highlighting the breadth of faunal diversity beyond conventional habitats.^[69] Cryofauna, adapted to frigid polar and high-altitude environments, rely on mechanisms to combat hypothermia and ice formation. In polar seas, notothenioid fish such as Antarctic icefish produce antifreeze glycoproteins that bind to ice crystals in their blood, preventing lethal freezing by lowering the temperature at which blood freezes by several degrees Celsius. These proteins, evolved independently in Arctic gadids like the polar cod, inhibit ice recrystallization and maintain fluid circulation in subzero waters. Terrestrial polar mammals, including the Arctic fox, feature dense insulating fur and compact body shapes to minimize heat loss, while emperor penguins endure Antarctic winters through huddling behaviors and blubber layers up to 5 cm thick for thermal regulation. At high-altitude plateaus like the Tibetan or Andean regions, faunas such as the yak exhibit reduced metabolic rates and efficient oxygen-binding hemoglobins to cope with hypoxia, with hemoglobin variants in species like the gelada monkey enhancing oxygen delivery under low partial pressures. Himalayan marmots, conversely, employ prolonged hibernation—up to seven months annually—to conserve energy in oxygen-poor, cold conditions above 4,000 meters. Snow leopards on these plateaus possess enlarged nasal cavities for efficient air warming and hemoglobin adaptations for better oxygen affinity, allowing predation in thin air.^[70]^[71]^[72]^[73]^[74]^[75] Troglofauna and stygofauna thrive in the aphotic, stable conditions of caves and groundwater aquifers, where energy scarcity drives troglomorphism—extreme adaptations like eye reduction, depigmentation, and elongated appendages for navigation. Troglofauna, air-breathing cave dwellers, include blind cavefish such as the Mexican tetra, which have lost functional eyes and pigmentation through regression, relying instead on lateral line systems for mechanosensory detection of water currents and prey. Other examples encompass troglobitic arthropods like cave millipedes and pseudoscorpions, which exhibit elongated bodies and reduced metabolic rates to navigate narrow fissures and endure nutrient-poor diets, often surviving on detritus washed into caves. In granite talus caves, species like the troglobitic pseudoscorpion Parobisium yttinum display complete eye loss and pale exoskeletons, adaptations honed over millennia in isolated refugia. Stygofauna, aquatic groundwater inhabitants, feature similar traits; the blind cave eel Ophisternon candidum, endemic to Australian aquifers, possesses no eyes or pigment, using chemosensory barbels to forage in dark, oxygen-limited waters. Crustaceans such as stygobitic amphipods (e.g., Niphargus species) and ostracods dominate these systems, with elongated limbs for crawling through gravel interstices and low respiration rates suited to stagnant, low-food environments. These faunas' isolation fosters high endemism but vulnerability to habitat disruption.^[76]^[77]^[78]^[79]^[80]^[81] Hydrothermal vent faunas represent a pinnacle of specialization, harnessing chemosynthesis in lightless, superheated abyssal depths where temperatures exceed 400°C and pressures crush most life. Giant tube worms (Riftia pachyptila) dominate these ecosystems, lacking mouths or digestive systems yet growing up to 2.4 meters long through symbiosis with sulfur-oxidizing bacteria housed in their trophosome—a specialized organ where hydrogen sulfide from vent fluids fuels carbon fixation into organic matter. These bacteria perform chemosynthesis, converting dissolved minerals like hydrogen sulfide into energy, supporting the worm's rapid growth rates of up to 2.5 feet per year. Associated fauna, including vent mussels and shrimp, similarly host endosymbiotic microbes for nutrient processing, while pompeii worms (Alvinella pompejana) tolerate plumes up to 80°C via heat-shock proteins and symbiotic bacteria that detoxify sulfides. These communities, discovered in 1977, form oases of biomass—up to 100 times denser than surrounding seafloor—sustained entirely without sunlight, underscoring chemosynthesis's role in global biogeochemical cycles.^[82]^[83]^[84]^[85]^[86]

Substrate and Hidden Faunas

Substrate and hidden faunas encompass animal communities that inhabit the surfaces, interiors, or concealed microhabitats of soils, sediments, leaf litter, and other substrates, playing crucial roles in nutrient cycling, decomposition, and ecosystem stability. These groups are distinguished by their intimate association with the substrate, which provides protection, food resources, and structural support, often shielding them from surface predators and environmental fluctuations. In both terrestrial and aquatic environments, such faunas contribute to bioturbation— the physical mixing of substrates— which enhances soil aeration, water infiltration, and organic matter breakdown.^[87]^[88] Epifauna consist of animals that live on the surface of substrates, either attached or mobile, without burrowing deeply. Common examples include barnacles affixed to rocky substrates in marine intertidal zones, where they filter plankton from the water column, and various arthropods or mites associated with lichens on tree bark or rocks, utilizing the lichen thallus for camouflage and moisture retention. These organisms often form dense assemblages that modify substrate microenvironments, such as by trapping sediments or providing habitat for smaller associates, thereby influencing local biodiversity and primary productivity. In terrestrial settings, epifaunal communities on leaf litter surfaces, like certain springtails, facilitate initial stages of decomposition by grazing on fungal hyphae.^[88]^[89] Infauna, in contrast, are burrowers that reside within the substrate matrix, exploiting interstitial spaces in sediments or soils for feeding and refuge. Marine infauna, such as clams and polychaete worms in soft seabed sediments, burrow to depths of several centimeters, where they deposit-feed on organic particles or suspension-feed on passing detritus, thereby recycling nutrients and maintaining sediment oxygenation through their activities. Terrestrial counterparts include earthworms in soil profiles, which ingest organic-rich material and excrete nutrient-enriched casts, improving soil fertility and structure; a single earthworm can process its own body weight in soil daily.^[87] These infaunal activities promote vertical mixing, which is essential for root penetration and microbial activity in ecosystems.^[90] Cryptofauna, also termed cryptozoic fauna, refer to animals concealed within crevices, under bark, or in the depths of leaf litter and soil litter layers, where darkness and humidity offer protection from desiccation and predation. In terrestrial environments, this includes soil arthropods like collembolans, mites, and millipedes hidden in leaf mould, which decompose organic debris and regulate microbial populations, contributing significantly to carbon turnover in forest floors. Marine cryptofauna, such as small crustaceans in coral rubble crevices, similarly exploit these hidden niches for ambush predation or scavenging, supporting higher trophic levels while avoiding exposure. The structural complexity of the substrate directly correlates with cryptofaunal diversity, as microhabitats like litter aggregates can harbor up to 10 times more species than exposed surfaces.^[91]

Subdivisions by Size and Scale

Size-based subdivisions of fauna vary by ecological context, such as terrestrial (soil) versus aquatic (benthic) environments, with differing standard thresholds for micro-, meso-, macro-, and meiofauna.

Macrofauna and Megafauna

Macrofauna are defined as multicellular animals larger than 2 mm, typically retained by sieves of this mesh size in ecological sampling, particularly in soil contexts.^[92] These organisms include earthworms, snails, and certain insects, which inhabit soils, sediments, and other substrates. In terrestrial ecosystems, macrofauna play crucial roles in soil health by burrowing and mixing activities that enhance aeration, drainage, and aggregation, thereby improving water infiltration and root penetration.^[92] For instance, earthworms create channels that facilitate oxygen exchange in compacted soils, while snails contribute to organic matter decomposition, fostering nutrient cycling essential for plant growth.^[93] Megafauna, in contrast, refer to the largest animals exceeding 100 kg in adult body mass, such as elephants, whales, and large ungulates, which exert disproportionate influence on ecosystems despite their relatively low population densities and biomass contributions.^[94] These "ecosystem engineers" modify landscapes through foraging, trampling, and seed dispersal, often shaping habitat structure on a broad scale; for example, American bison grazing maintains open prairies by selectively consuming grasses and preventing woody encroachment, thereby promoting biodiversity in grassland ecosystems.^[95] In marine environments, megafauna like whales influence nutrient distribution by vertical migrations that transport deep-sea nutrients to surface waters, supporting phytoplankton blooms.^[96] The late Pleistocene epoch witnessed widespread megafauna extinctions around 10,000 years ago, including woolly mammoths and other large herbivores, likely driven by a combination of human hunting and climate shifts at the end of the last Ice Age.^[94] These losses, affecting over 100 genera globally, led to cascading ecological changes, such as altered vegetation dynamics and reduced landscape heterogeneity, with lasting impacts on modern biomes like the conversion of mammoth-steppe grasslands into forests or tundra.^[97] Today, surviving megafauna continue to face threats from habitat fragmentation and poaching, underscoring their outsized role in maintaining ecosystem resilience despite comprising a minor fraction of total faunal biomass.^[94]

Mesofauna and Meiofauna

Mesofauna are small soil invertebrates typically ranging in size from 0.2 mm to 2 mm in body width, bridging the gap between larger macrofauna and smaller microfauna in terrestrial ecosystems.^[98] These organisms include prominent groups such as mites (Acari), springtails (Collembola), and enchytraeid worms, which inhabit the upper soil layers and litter.^[99] Mesofauna play crucial roles in soil processes, particularly by facilitating organic matter decomposition through fragmentation and grazing on microbes and detritus, thereby enhancing nutrient availability and cycling.^[100] For instance, springtails and mites contribute to litter breakdown by increasing surface area for microbial action, supporting carbon and nitrogen turnover in forest and agricultural soils.^[101] Extraction of mesofauna from soil samples commonly involves wet or dry sieving, where materials are passed through a 2 mm mesh to exclude macrofauna and retained on a 0.5 mm mesh to capture mesofaunal organisms, followed by techniques like Tullgren funnels for live extraction.^[102] This size-based separation allows researchers to quantify mesofaunal abundance and diversity, revealing their sensitivity to environmental changes such as land use or pollution.^[103] Their activities promote soil structure and fertility, underscoring their importance in maintaining ecosystem health. Meiofauna encompass even smaller metazoans, operationally defined as those passing through a 0.5 mm sieve but retained on a 0.063 mm (63 μm) mesh, corresponding to body sizes of approximately 0.045 mm to 0.5 mm, primarily in aquatic sediments but also in interstitial soil spaces. Key representatives include nematodes and harpacticoid copepods, which dominate communities in marine and freshwater environments due to their high reproductive rates and adaptability.^[104] In marine sediments, meiofaunal densities can reach up to 10^6 individuals per square meter, particularly in coastal and deep-sea habitats where nematodes often comprise over 90% of the assemblage.^[105] Meiofauna are vital for benthic nutrient cycling, as their burrowing and feeding stimulate microbial processes like nitrification and denitrification, facilitating nitrogen removal and organic matter remineralization in sediments.^[106] For example, nematode grazing on bacteria enhances sediment oxygenation and carbon flux, while copepods contribute to secondary production as prey for larger organisms.^[107] Extraction methods mirror their size definition, using serial sieving with 0.5 mm and 0.063 mm meshes after sample decantation, often supplemented by density gradient centrifugation for clean separation. These groups thus sustain foundational ecosystem functions, linking microbial activity to higher trophic levels in aquatic systems.

Microfauna

Microfauna encompass microscopic animals typically smaller than 0.1 mm in body size, inhabiting diverse environments such as soil, freshwater sediments, and aquatic biofilms.^[108] These organisms exhibit animal-like characteristics and include groups like protozoans (often classified within microfauna despite their protist status in soil ecology), nematodes, rotifers, and tardigrades.^[108] For instance, protozoans such as amoebae and ciliates, rotifers with their wheel-like cilia for locomotion, and microscopic nematodes represent key components, contributing to the hidden biodiversity of microbial ecosystems.^[109] The diversity of microfauna is immense, with over 50,000 described species of protozoans alone, alongside approximately 30,000 nematode species and more than 2,000 rotifer species.^[109]^[110]^[111] However, current estimates as of 2025 suggest millions of undescribed species, particularly among nematodes, where only a fraction of the projected 0.5 to 10 million total species have been cataloged.^[110] This vast, largely untapped diversity underscores microfauna's role in global biodiversity, far exceeding that of many larger faunal groups in sheer numerical abundance within microscopic scales. Microfauna play critical roles in microbial ecosystems, primarily through predation on bacteria and fungi, which regulates microbial populations and facilitates nutrient cycling.^[111] In soil environments, for example, protozoans and nematodes graze on bacteria, releasing excess nitrogen and other nutrients in soluble forms that enhance plant-available fertility and support ecosystem productivity.^[112] This process not only accelerates decomposition but also maintains soil health by preventing microbial overgrowth and promoting balanced biogeochemical cycles. Their ecological importance extends to broader biodiversity dynamics, influencing energy transfer in food webs. Studying microfauna presents significant challenges, primarily due to their minute size and often transparent morphology, necessitating advanced microscopy techniques for identification.^[113] Traditional light microscopy requires skilled observation to distinguish species, while high-resolution methods like scanning electron microscopy are essential for detailed taxonomy but are labor-intensive and costly. The high cryptic diversity and estimated millions of undescribed species further complicate comprehensive surveys, limiting our understanding of their full ecological contributions as of 2025.

Other Subdivisions

Introduced and Alien Faunas

Introduced and alien faunas, often referred to as non-native or exotic species assemblages, encompass animal populations transported to new regions primarily through human activities such as trade, agriculture, and intentional releases for pest control or ornamental purposes. These introductions can disrupt local ecosystems by altering food webs, competing with native species, and facilitating disease transmission.^[114] A prominent example of human-mediated introduction is the European starling (Sturnus vulgaris), deliberately released in New York City in 1890 by Eugene Schieffelin as part of an effort to establish all birds mentioned in Shakespeare's works in North America. From this initial population of about 100 individuals, the species rapidly expanded across the continent, reaching a population of over 200 million by the late 20th century and causing an estimated $800 million in annual agricultural damage through crop consumption and competition with native cavity-nesting birds.^[115]^[116] Similarly, the cane toad (Rhinella marina) was introduced to Queensland, Australia, in 1935 to control beetles damaging sugarcane crops, with 101 individuals released from Hawaii and Puerto Rico. Instead of targeting the intended pests, the toads proliferated uncontrollably, spreading across more than 1.5 million square kilometers as of 2025 and poisoning native predators like quolls and goannas through their toxic skin secretions, leading to local population declines of up to 90% in some species without causing outright extinctions.^[117]^[118] Impacts of introduced faunas extend to genetic consequences, including hybridization with native species, which can result in introgression— the transfer of genes from non-natives—potentially eroding local adaptations and reducing genetic diversity. For instance, hybridization between introduced rainbow trout and native cutthroat trout in western North America has led to the genetic swamping of endemic populations, contributing to their vulnerability.^[119]^[120] According to the International Union for Conservation of Nature (IUCN), invasive alien species threaten 25.5% of all assessed threatened species on the Red List, underscoring their role as a leading driver of biodiversity loss alongside habitat destruction. Management strategies focus on prevention through biosecurity protocols and eradication efforts, particularly on islands where isolated ecosystems allow for complete removal; for example, the successful eradication of invasive rats from over 100 South Pacific islands since 2000 has enabled the recovery of native seabird populations via ongoing monitoring to prevent reinvasion.^[121]^[122]

Fossil and Extinct Faunas

Fossil faunas represent ancient animal assemblages preserved in the geological record, providing insights into the evolutionary history of life on Earth. During the Paleozoic Era, particularly in the Cambrian and early Ordovician periods, marine ecosystems were dominated by trilobites, a diverse group of arthropods that appeared abruptly around 521 million years ago and comprised a significant portion of the skeletonized fossil remains. These trilobites, ranging from small coin-sized forms to larger species, occupied various niches in shallow seas across ancient continents, alongside other invertebrates like brachiopods and crinoids. As the Paleozoic progressed into the Devonian and Carboniferous, faunas shifted toward more complex assemblages, including early fish and amphibians, marking the transition from marine to terrestrial dominance. In the Cenozoic Era, following the Cretaceous-Paleogene mass extinction that eliminated non-avian dinosaurs approximately 66 million years ago, fossil faunas experienced a profound radiation of mammals. This event released ecological constraints, allowing small, nocturnal mammals to diversify rapidly into larger body sizes and varied niches, with placental mammals evolving at accelerated rates compared to earlier Mesozoic periods. By the Eocene, ecosystems featured diverse orders such as primates, ungulates, and carnivores, filling roles previously held by dinosaurs and contributing to the modern mammalian framework. Extinct faunas highlight catastrophic losses that reshaped biodiversity, as seen in major mass extinction events. The Permian-Triassic extinction, known as the Great Dying and dated to about 252 million years ago, caused the demise of approximately 96% of marine species and around 70% of terrestrial vertebrate genera, likely triggered by massive volcanic activity from the Siberian Traps that led to global warming and ocean anoxia. This event wiped out dominant Paleozoic groups like trilobites and many reef-building organisms, paving the way for Mesozoic faunas. More recently, the extinction of the dodo (Raphus cucullatus) in the late 17th century profoundly altered the fauna of Mauritius, where invasive species such as rats, pigs, and macaques preyed on native birds and disrupted seed dispersal, contributing to the loss of several endemic species in the island's isolated ecosystem. Key concepts in studying fossil and extinct faunas include faunal turnover, which describes the replacement of dominant species assemblages over geological time due to environmental changes or extinctions, and faunal succession, the principle that fossil organisms appear in a consistent, predictable order in stratified rock layers, enabling relative dating of strata. These processes underscore the dynamic nature of ecosystems, with turnover rates accelerating during mass extinctions and recoveries, as evidenced by the sequential replacement of top predators in southern African ecosystems across the end-Permian boundary. Contemporary discussions on extinct faunas extend to de-extinction efforts, where genetic technologies aim to revive lost species. In 2025, researchers at Colossal Biosciences advanced CRISPR-based de-extinction of the woolly mammoth by creating gene-edited "woolly mice" that express mammoth-like traits, such as thick fur, as a proxy for editing Asian elephant genomes toward producing hybrid mammoths by 2027 or 2028. These initiatives highlight potential applications in restoring Pleistocene faunas but raise ethical questions about ecological integration.

Study and Conservation

Methods of Faunal Study

The study of fauna relies on a range of methods to inventory, monitor, and analyze animal communities, encompassing both longstanding fieldwork techniques and emerging technologies. Traditional approaches form the foundation of faunal surveys, enabling direct observation and capture of specimens across diverse ecosystems.^[123] Trapping methods, such as pitfall traps for ground-dwelling invertebrates, malaise traps for flying insects, and hoop-net traps for aquatic species, allow researchers to collect and identify animals systematically while minimizing habitat disturbance.^[123]^[124] Netting techniques, including mist nets for birds and butterflies or fyke nets for fish, facilitate capture for morphological examination and marking for population tracking.^[125] Visual surveys involve direct observation by field teams to record species presence, abundance, and behavior, often supplemented by line transects or point counts in accessible habitats.^[124] For elusive megafauna, camera traps—motion-activated devices that capture images or videos—provide non-invasive, continuous monitoring, as demonstrated in large-scale deployments across savannas and forests to assess distribution and activity patterns.^[126]^[127] Advancements in the 21st century have introduced molecular and digital tools to enhance detection efficiency, particularly for cryptic or low-density species. Environmental DNA (eDNA) sampling extracts genetic material shed by animals into water, soil, or air, enabling metabarcoding to identify species composition without direct capture; for instance, eDNA from seawater has revealed diverse marine fish assemblages in coastal and deep-sea environments.^[128]^[129] Remote sensing, utilizing satellite imagery and LiDAR to map habitat structure and vegetation cover, indirectly infers faunal distributions by correlating environmental variables with species occurrences.^[130] Artificial intelligence (AI) and machine learning algorithms process vast datasets from camera traps or acoustic sensors, automating species identification and behavior classification; deep neural networks, for example, have accurately classified over 48 mammal species in millions of images from African ecosystems.^[131] Drone-based surveys, increasingly adopted in the 2020s, enable aerial counts of birds and large herbivores over expansive areas, reducing human bias and effort compared to ground-based methods.^[132] Once collected, faunal data are analyzed using quantitative metrics to quantify diversity and dynamics. The Shannon diversity index, which accounts for species richness and evenness, provides a measure of community complexity, widely applied in studies of invertebrate and vertebrate assemblages to compare habitats.^[133] Faunal turnover metrics, such as beta diversity components that decompose species replacement and nestedness, assess compositional changes across spatial or temporal gradients in biogeography, revealing patterns driven by environmental filtering or dispersal limitations.^[134] Global initiatives aggregate these data for broader insights, with the Global Biodiversity Information Facility (GBIF) serving as a central repository that, as of 2025, hosts over 3 billion occurrence records contributed by thousands of institutions worldwide.^[135]

Conservation Challenges

Fauna worldwide faces profound conservation challenges driven by anthropogenic pressures, with habitat loss emerging as the predominant threat, affecting 82% of imperiled species in regions like the United States through land-use changes such as agriculture and urbanization.^[136] Climate change exacerbates these issues by altering ecosystems, causing mismatches in migratory patterns and food availability for species like birds, while overhunting and overexploitation further deplete populations.^[137] According to the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES), these interconnected crises— including biodiversity loss, food insecurity, and climate impacts—place approximately one million species at risk of extinction, underscoring the urgency for integrated global responses.^[138] The United Nations Environment Programme highlights that agriculture alone drives threats to 24,000 of the 28,000 assessed species, amplifying faunal declines through habitat fragmentation and pollution.^[139] Island faunas are particularly susceptible to invasive alien species, which have contributed to the extinction of over two-thirds of native species in places like Hawaii and the Mascarenes due to predation and competition from introduced mammals such as rats and feral cats.^[140] These ecosystems, hosting a disproportionate share of global biodiversity despite their small land area, suffer catastrophic losses from invasives, with 41% of highly threatened vertebrates on islands identified as vulnerable to such incursions.^[141] Migratory faunas, including birds, encounter additional barriers from habitat degradation, collisions with infrastructure like wind turbines and buildings, and climate-induced shifts in breeding and stopover sites, leading to population declines in over 1,100 protected species under frameworks like the U.S. Migratory Bird Treaty Act.^[142] These faunal-specific vulnerabilities highlight the need for targeted interventions beyond general biodiversity measures. Conservation strategies have centered on establishing protected areas, which prove 33% more effective at curbing habitat loss than unprotected lands, though their success depends on governance and management quality to mitigate external pressures like nearby human activities.^[143] Multiple-use protected areas, balancing biodiversity goals with equitable community involvement, enhance both ecological outcomes and social acceptance, serving as strongholds for imperiled mammals and other fauna.^[144] Rewilding initiatives, such as the reintroduction of European bison (Bison bonasus) in Romania's Southern Carpathians since 2014, have boosted populations by over 20% through releases of more than 110 individuals, restoring ecosystem functions like carbon sequestration equivalent to offsetting emissions from approximately 43,000 vehicles annually.^[145] International agreements like the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), effective since 1975, regulate trade in over 38,000 species to prevent overexploitation, with 184 parties collaborating to enforce sustainable practices.^[146] Amid the escalating 2025 biodiversity crisis, emerging approaches emphasize genetic conservation through biobanks, which store living cells and tissues to preserve genetic diversity for future reintroductions and adaptation to environmental changes.^[147] The San Diego Zoo Wildlife Alliance's Global Biobanking Network, marking significant progress in 2025, facilitates the cryopreservation of samples from thousands of species, enabling assisted reproductive technologies to counter extinction risks.^[148] Similarly, the International Union for Conservation of Nature's 2025 biobanking guidelines promote standardized repositories as vital tools for safeguarding genetic material during rapid biodiversity loss, complementing in-situ efforts like protected areas.^[149]

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[PDF] 1. Ophisternon candidum (Vulnerable)– Blind Cave Eel - EPA WA
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[PDF] Occasional Papers - Texas Tech University Departments
Sep 30, 2010 · A new species of troglobitic Parobisium pseudoscorpion is described from two caves de- veloped in granite talus slopes in the Yosemite ...
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Subterranean Fishes of the World - Graham Proudlove
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Citizen science approach reveals groundwater fauna in Switzerland ...
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[PDF] Introduction: Chemosynthesis at Hydrothermal Vents
Chemosynthesis is making food from inorganic compounds. Bacteria use chemicals from hydrothermal vents to produce food, like sugar, sulfur, and water.
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The Microbes That Keep Hydrothermal Vents Pumping
Microbes convert mineral-laden fluid into energy via chemosynthesis, using chemical compounds from vents, and are primary producers in the food web.
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How Giant Tube Worms Survive at Hydrothermal Vents
Mar 14, 2022 · This short video explores the symbiotic relationship between giant tube worms and species of chemosynthetic bacteria.
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Weird Science: Hydrothermal Vents and Cold Seeps
Other vent animals, like the giant tubeworms in the genus Riftia, have evolved a symbiotic relationship with the chemosynthetic bacteria (SF Fig. 2.3 C). Adult ...
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Chemosynthesis - NOAA/PMEL
Tubeworms flourish in small clumps, waving in the warm fluids. A typical picture of an active hydrothermal vent is therefore one with shimmering warm ...
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Infaunal Benthic Invertebrate Community
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Glossary of Terminology - NOAA's Coral Reef Information System
... in the ecology of coral reef ecosystems. Image of humpback whale (phylum Chordata) ... epifauna. animals that live upon or are associated with substratum ...
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Lichen, Algae, and Moss on Trees | University of Maryland Extension
Feb 27, 2023 · Ruby-throated hummingbirds line their nests with strips of lichen. Painted lichen moth caterpillars can only eat lichen.
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Functional and phylogenetic responses of motile cryptofauna to ...
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Macrofauna - an overview | ScienceDirect Topics
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[PDF] SOIL MACROFAUNA FIELD MANUAL
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Soil Macrofauna: A key Factor for Increasing Soil Fertility and ... - MDPI
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Megafauna and ecosystem function from the Pleistocene to ... - PNAS
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Global distribution of soil fauna functional groups and their ...
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Mesofauna - Soil Ecology Wiki
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[PDF] Diversity and Function of Soil Mesofauna - ResearchGate
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Investigating the role of soil mesofauna abundance and biodiversity ...
Decomposition is one key functional parameter of soil food webs and indicates the soil's ability to recycle and improve bioavailability of nutrients to support ...
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[PDF] Chapter 19. Sampling Methods for Soil and Litter Fauna
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[PDF] Spatial Structure and Composition of the Soil Mesofauna
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Metagenetic tools for the census of marine meiofaunal biodiversity
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Microfauna - SARE - Sustainable Agriculture Research and Education
$$28.00ECOLOGICAL ROLE: As predators, decomposers and consumers of bacteria, protozoans play a significant role in nutrient cycling. DESCRIPTION: These single ...
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[PDF] European Starlings: A Review of an Invasive Species with Far
European starlings, introduced to North America, are considered one of the "World's Worst" invaders, causing $800 million yearly agricultural damage. They are ...
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Introduction of cane toads | National Museum of Australia
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Cane toads mainly impact large predators through toxic ingestion, but some species recover. No extinctions have occurred, and some species benefit.
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[PDF] The IUCN Red List and invasive alien species: an analysis of ...
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[PDF] Priorities for Restoring U.S. Islands Through Invasive Species ...
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[PDF] Standards and Best Practices for Monitoring and Benchmarking ...
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[PDF] Final Report - Texas Comptroller
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An approach to rapid processing of camera trap images with ...
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(PDF) Pairing camera traps and acoustic recorders to monitor the ...
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Optimizing environmental DNA sampling effort for fish inventories in ...
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We investigate the potential of using metabarcoding of environmental DNA (eDNA) obtained directly from seawater samples to account for marine fish biodiversity.
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Deep learning enables satellite-based monitoring of large ... - Nature
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Automatically identifying, counting, and describing wild animals in ...
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MEASURING BIODIVERSITY VALUE FOR CONSERVATION
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As we approach 2025 we're reflecting on a huge 2024 in
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US Imperiled species and the five drivers of biodiversity loss
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IPBES Report : One Million Species at Risk Due to Human Activities
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Facts about the nature crisis | UNEP - UN Environment Programme
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Want to save 41 percent of the planet's highly threatened vertebrates ...
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Many of the 1,106 species of birds protected under the Migratory Bird Treaty Act are experiencing population declines due to increased threats across the ...
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European bison comeback | Rewilding Europe
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What is CITES?
CITES (the Convention on International Trade in Endangered Species of Wild Fauna and Flora) is an international agreement between governments.
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[PDF] Challenges and prospects for conservation genetics at XXI century
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San Diego Zoo Wildlife Alliance Unveils Milestone Progress in Its ...
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[PDF] IUCN Species Survival Commission biobanking guidelines for ...
Oct 6, 2025 · In light of a worsening biodiversity crisis, views towards the value of biorepositories as well as the relevance of ARTs are fortunately ...