182-459: The hellbender ( Cryptobranchus alleganiensis ), also known as the hellbender salamander, is a species of aquatic giant salamander endemic to the eastern and central United States. It is the largest salamander in North America . A member of the family Cryptobranchidae , the hellbender is the only extant member of the genus Cryptobranchus . Other closely related salamanders in
364-577: A lower house based on population; and the Senate , an upper house based on equal representation for each state. Federalism provides substantial autonomy to the 50 states, while the country's political culture promotes liberty , equality , individualism , personal autonomy , and limited government . One of the world's most developed countries , the United States has had the largest nominal GDP since about 1890 and accounted for over 15% of
546-400: A metamorphosis after around a year and a half of life. At this point, when they are roughly 13.5 cm (5.3 in) long, they lose the gills present during their larval stage. Until then, they are easily confused with mudpuppies, and can be differentiated often only through toe number. After this metamorphosis, hellbenders must be able to absorb oxygen through the folds in their skin, which
728-423: A mutation–selection balance . It is predicted that a viral quasispecies at a low but evolutionarily neutral and highly connected (that is, flat) region in the fitness landscape will outcompete a quasispecies located at a higher but narrower fitness peak in which the surrounding mutants are unfit, "the quasispecies effect" or the "survival of the flattest". There is no suggestion that a viral quasispecies resembles
910-515: A taxonomic rank of an organism, as well as a unit of biodiversity . Other ways of defining species include their karyotype , DNA sequence, morphology , behaviour, or ecological niche . In addition, paleontologists use the concept of the chronospecies since fossil reproduction cannot be examined. The most recent rigorous estimate for the total number of species of eukaryotes is between 8 and 8.7 million. About 14% of these had been described by 2011. All species (except viruses ) are given
1092-542: A two-part name , called a "binomial". The first part of a binomial is the genus to which the species belongs. The second part is called the specific name or the specific epithet (in botanical nomenclature , also sometimes in zoological nomenclature ). For example, Boa constrictor is one of the species of the genus Boa , with constrictor being the species' epithet. While the definitions given above may seem adequate at first glance, when looked at more closely they represent problematic species concepts. For example,
1274-580: A " Special Relationship " with the United Kingdom and strong ties with Canada , Australia , New Zealand , the Philippines , Japan , South Korea , Israel , and several European Union countries ( France , Italy , Germany , Spain , and Poland ). The U.S. works closely with its NATO allies on military and national security issues, and with countries in the Americas through
1456-400: A "classical" method of determining species, such as with Linnaeus, early in evolutionary theory. However, different phenotypes are not necessarily different species (e.g. a four-winged Drosophila born to a two-winged mother is not a different species). Species named in this manner are called morphospecies . In the 1970s, Robert R. Sokal , Theodore J. Crovello and Peter Sneath proposed
1638-424: A 'smallest clade' idea" (a phylogenetic species concept). Mishler and Wilkins and others concur with this approach, even though this would raise difficulties in biological nomenclature. Wilkins cited the ichthyologist Charles Tate Regan 's early 20th century remark that "a species is whatever a suitably qualified biologist chooses to call a species". Wilkins noted that the philosopher Philip Kitcher called this
1820-465: A commercial collector in the 1940s, noted a trend of more crayfish predation in the summer during times of higher prey activity, whereas fish made up a larger part of the winter diet, when crayfish are less active. There seems to be a specific temperature range in which hellbenders feed, as well: between 45 and 80 °F (7 and 27 °C). Cannibalism—mainly on eggs—has been known to occur within hellbender populations. One researcher claimed perhaps density
2002-428: A connected series of neighbouring populations, each of which can sexually interbreed with adjacent related populations, but for which there exist at least two "end" populations in the series, which are too distantly related to interbreed, though there is a potential gene flow between each "linked" population. Such non-breeding, though genetically connected, "end" populations may co-exist in the same region thus closing
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#17327650565682184-542: A coordinated, continent-wide military effort. Finally, deteriorating relations between the British military establishment and the colonists, relations that were already less than positive, set the stage for further distrust and dislike of British troops. Following their victory in the French and Indian War, Britain began to assert greater control over local colonial affairs, resulting in colonial political resistance ; one of
2366-534: A decentralized government that operated until 1789. After the British surrender at the siege of Yorktown in 1781 American sovereignty was internationally recognized by the Treaty of Paris (1783), through which the U.S. gained territory stretching west to the Mississippi River, north to present-day Canada, and south to Spanish Florida . The Northwest Ordinance (1787) established the precedent by which
2548-432: A different species from its ancestors. Viruses have enormous populations, are doubtfully living since they consist of little more than a string of DNA or RNA in a protein coat, and mutate rapidly. All of these factors make conventional species concepts largely inapplicable. A viral quasispecies is a group of genotypes related by similar mutations, competing within a highly mutagenic environment, and hence governed by
2730-560: A dispute by House members who argue that Wehrle's salamander should be given the honor. The legislation did not pass in 2018, but was reintroduced in 2019. On April 23, 2019, Pennsylvania Governor Tom Wolf signed legislation making the eastern hellbender Pennsylvania's official state amphibian. Youth members of the Chesapeake Bay Foundation 's Pennsylvania Student Leadership Council were heavily involved writing and advocating on behalf of this legislation. They hope that
2912-463: A female approaches, the male guides or drives her into his burrow and prevents her from leaving until she oviposits . Female hellbenders lay 150–200 eggs over a two- to three-day period; the eggs are 18–20 mm (0.71–0.79 in) in diameter, connected by five to ten cords. As the female lays eggs, the male positions himself alongside or slightly above them, spraying the eggs with sperm while swaying his tail and moving his hind limbs, which disperses
3094-407: A few prominent industrialists , largely by their formation of trusts and monopolies to prevent competition. Tycoons led the nation's expansion in the railroad , petroleum , and steel industries. The United States emerged as a pioneer of the automotive industry . These changes were accompanied by significant increases in economic inequality , slum conditions , and social unrest , creating
3276-638: A founding member of the World Bank , Organization of American States , NATO , and the United Nations , as well as a permanent member of the UN Security Council . The first documented use of the phrase "United States of America" is a letter from January 2, 1776. Stephen Moylan , a Continental Army aide to General George Washington , wrote to Joseph Reed , Washington's aide-de-camp , seeking to go "with full and ample powers from
3458-508: A genetic boundary suitable for defining a species concept is present. DNA barcoding has been proposed as a way to distinguish species suitable even for non-specialists to use. One of the barcodes is a region of mitochondrial DNA within the gene for cytochrome c oxidase . A database, Barcode of Life Data System , contains DNA barcode sequences from over 190,000 species. However, scientists such as Rob DeSalle have expressed concern that classical taxonomy and DNA barcoding, which they consider
3640-411: A gigantic, dorsoventrally flattened body with thick folds travelling down the sides, a single open gill slit on each side, and hind feet with five toes each. Easily distinguished from most other endemic salamander species simply by their size, hellbenders average up to 60 cm or about 2 ft in length; the only species requiring further distinction (due to an overlap in distribution and size range)
3822-465: A lineage should be divided into multiple chronospecies , or when populations have diverged to have enough distinct character states to be described as cladistic species. Species and higher taxa were seen from the time of Aristotle until the 18th century as categories that could be arranged in a hierarchy, the great chain of being . In the 19th century, biologists grasped that species could evolve given sufficient time. Charles Darwin 's 1859 book On
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#17327650565684004-492: A misnomer, need to be reconciled, as they delimit species differently. Genetic introgression mediated by endosymbionts and other vectors can further make barcodes ineffective in the identification of species. A phylogenetic or cladistic species is "the smallest aggregation of populations (sexual) or lineages (asexual) diagnosable by a unique combination of character states in comparable individuals (semaphoronts)". The empirical basis – observed character states – provides
4186-550: A much lower number of eggs in hellbender nests than would be predicted by egg counts. Adult males are more likely to cannibalize their own offspring in degraded sites with limited food availability. After oviposition, the male drives the female away from the nest and guards the eggs. Incubating males rock back and forth and undulate their lateral skin folds, which circulates the water, increasing oxygen supply to both eggs and adult. Incubation lasts from 45 to 75 days, depending on region. Males are known to show solitary parental care for
4368-408: A number of Eastern US states, from southern New York to northern Georgia , including parts of Ohio , Pennsylvania , Maryland , West Virginia , Virginia , Kentucky , Illinois , Indiana , Tennessee , North Carolina , South Carolina , Alabama , Mississippi , Arkansas , Missouri , and extending into Oklahoma and Kansas . The subspecies (or species, depending on the source) C. a. bishopi
4550-449: A particular species, including which genus (and higher taxa) it is placed in, is a hypothesis about the evolutionary relationships and distinguishability of that group of organisms. As further information comes to hand, the hypothesis may be corroborated or refuted. Sometimes, especially in the past when communication was more difficult, taxonomists working in isolation have given two distinct names to individual organisms later identified as
4732-400: A perfect model of life, it is still a useful tool to scientists and conservationists for studying life on Earth, regardless of the theoretical difficulties. If species were fixed and clearly distinct from one another, there would be no problem, but evolutionary processes cause species to change. This obliges taxonomists to decide, for example, when enough change has occurred to declare that
4914-598: A period of heightened, overt racism following Reconstruction, a time often called the nadir of American race relations . A series of Supreme Court decisions, including Plessy v. Ferguson , emptied the Fourteenth and Fifteenth Amendments of their force, allowing Jim Crow laws in the South to remain unchecked, sundown towns in the Midwest, and segregation in communities across the country , which would be reinforced by
5096-488: A prominent leader in the early 1960s. The Great Society plan of President Lyndon Johnson 's administration resulted in groundbreaking and broad-reaching laws, policies and a constitutional amendment to counteract some of the worst effects of lingering institutional racism . The counterculture movement in the U.S. brought significant social changes, including the liberalization of attitudes toward recreational drug use and sexuality . It also encouraged open defiance of
5278-589: A reported average size of 28 to 40 cm (11 to 16 in) in length, which means that hellbender adults will still generally be notably larger than even the biggest mudpuppies. The genus Cryptobranchus has historically been considered to contain only one species, C. alleganiensis , with two subspecies, C. a. alleganiensis and C. a. bishopi . A recent decline in population size of the Ozark subspecies C. a. bishopi has led to further research into populations of this subspecies, including genetic analysis to determine
5460-433: A result). Nocturnal activity has been found to be most prevalent in early summer, perhaps coinciding with highest water depths. Adult hellbenders can live up to 25–30 years. C. alleganiensis feeds primarily on crayfish and small fish , but also insects , worms , molluscs , tadpoles and smaller salamanders. A study conducted in 2017 found that larval hellbenders eat mayfly and caddisfly nymphs. One report, written by
5642-422: A secretion that is important for innate immunity against chytrid activity. Although their eyesight is relatively poor, they have light-sensitive cells all over their bodies. Those on their tails are especially finely tuned and may help them position safely under rocks without their tails poking out to give themselves away. They have a good sense of smell and move upstream in search of food such as dead fish, following
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5824-526: A sense of manifest destiny . The Missouri Compromise attempted to balance the desire of northern states to prevent the expansion of slavery into new territories with that of southern states to extend it, admitting Missouri as a slave state and Maine as a free state. It further prohibited slavery in all other lands of the Louisiana Purchase north of the 36°30′ parallel . As Americans expanded further into land inhabited by Native Americans,
6006-614: A sense of American identity as well. Men who otherwise never left their own colony now traveled across the continent to fight alongside men from decidedly different backgrounds but who were no less "American". British officers trained American officers for battle, most notably George Washington ; these officers would lend their skills and expertise to the colonists' cause during the American Revolutionary War to come. In addition, colonial legislatures and officials found it necessary to cooperate intensively in pursuit of
6188-400: A short way of saying that something applies to many species within a genus, but not to all. If scientists mean that something applies to all species within a genus, they use the genus name without the specific name or epithet. The names of genera and species are usually printed in italics . However, abbreviations such as "sp." should not be italicised. When a species' identity is not clear,
6370-404: A specialist may use "cf." before the epithet to indicate that confirmation is required. The abbreviations "nr." (near) or "aff." (affine) may be used when the identity is unclear but when the species appears to be similar to the species mentioned after. With the rise of online databases, codes have been devised to provide identifiers for species that are already defined, including: The naming of
6552-523: A species as groups of actually or potentially interbreeding natural populations, which are reproductively isolated from other such groups. It has been argued that this definition is a natural consequence of the effect of sexual reproduction on the dynamics of natural selection. Mayr's use of the adjective "potentially" has been a point of debate; some interpretations exclude unusual or artificial matings that occur only in captivity, or that involve animals capable of mating but that do not normally do so in
6734-400: A species as determined by a taxonomist. A typological species is a group of organisms in which individuals conform to certain fixed properties (a type), so that even pre-literate people often recognise the same taxon as do modern taxonomists. The clusters of variations or phenotypes within specimens (such as longer or shorter tails) would differentiate the species. This method was used as
6916-491: A species. All species definitions assume that an organism acquires its genes from one or two parents very like the "daughter" organism, but that is not what happens in HGT. There is strong evidence of HGT between very dissimilar groups of prokaryotes , and at least occasionally between dissimilar groups of eukaryotes , including some crustaceans and echinoderms . The evolutionary biologist James Mallet concludes that there
7098-685: A species. Generally the term includes the unknown element of a distinct act of creation. Many authors have argued that a simple textbook definition, following Mayr's concept, works well for most multi-celled organisms , but breaks down in several situations: Species identification is made difficult by discordance between molecular and morphological investigations; these can be categorised as two types: (i) one morphology, multiple lineages (e.g. morphological convergence , cryptic species ) and (ii) one lineage, multiple morphologies (e.g. phenotypic plasticity , multiple life-cycle stages). In addition, horizontal gene transfer (HGT) makes it difficult to define
7280-632: A status solidified after its involvement in World War I . After Japan 's attack on Pearl Harbor in December 1941, the U.S. entered World War II . Its aftermath left the U.S. and the Soviet Union as the world's two superpowers and led to the Cold War , during which both countries struggled for ideological dominance and international influence . Following the Soviet Union's collapse and
7462-528: A taxonomic decision at the discretion of cognizant specialists, is not governed by the Codes of Zoological or Botanical Nomenclature, in contrast to the PhyloCode , and contrary to what is done in several other fields, in which the definitions of technical terms, like geochronological units and geopolitical entities, are explicitly delimited. The nomenclatural codes that guide the naming of species, including
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7644-517: A traditional biological species. The International Committee on Taxonomy of Viruses has since 1962 developed a universal taxonomic scheme for viruses; this has stabilised viral taxonomy. Most modern textbooks make use of Ernst Mayr 's 1942 definition, known as the Biological Species Concept as a basis for further discussion on the definition of species. It is also called a reproductive or isolation concept. This defines
7826-447: A variation on the morphological species concept, a phenetic species, defined as a set of organisms with a similar phenotype to each other, but a different phenotype from other sets of organisms. It differs from the morphological species concept in including a numerical measure of distance or similarity to cluster entities based on multivariate comparisons of a reasonably large number of phenotypic traits. A mate-recognition species
8008-515: A variety of reasons. Viruses are a special case, driven by a balance of mutation and selection , and can be treated as quasispecies . Biologists and taxonomists have made many attempts to define species, beginning from morphology and moving towards genetics . Early taxonomists such as Linnaeus had no option but to describe what they saw: this was later formalised as the typological or morphological species concept. Ernst Mayr emphasised reproductive isolation, but this, like other species concepts,
8190-401: A year). Juvenile correctional facilities are operated by local or state governments and serve as longer-term placements for any minor adjudicated as delinquent and ordered by a judge to be confined. As of January 2023, the United States has the sixth-highest per capita incarceration rate in the world—531 people per 100,000 inhabitants—and the largest prison and jail population in
8372-438: Is "an entity composed of organisms which maintains its identity from other such entities through time and over space, and which has its own independent evolutionary fate and historical tendencies". This differs from the biological species concept in embodying persistence over time. Wiley and Mayden stated that they see the evolutionary species concept as "identical" to Willi Hennig 's species-as-lineages concept, and asserted that
8554-400: Is a group of sexually reproducing organisms that recognise one another as potential mates. Expanding on this to allow for post-mating isolation, a cohesion species is the most inclusive population of individuals having the potential for phenotypic cohesion through intrinsic cohesion mechanisms; no matter whether populations can hybridise successfully, they are still distinct cohesion species if
8736-432: Is a predator that can consume a hellbender 1–3 years old. Cannibalism of eggs is also considered a common occurrence. One study found that in areas with increased deforestation, the likelihood of filial cannibalism increases. In another study by Kenison & Wilson (2018), researchers found that young, captive hellbenders showed altered behavior in response to predatory fish nearby. Because of their altered behavior, it
8918-458: Is a set of organisms adapted to a particular set of resources, called a niche, in the environment. According to this concept, populations form the discrete phenetic clusters that we recognise as species because the ecological and evolutionary processes controlling how resources are divided up tend to produce those clusters. A genetic species as defined by Robert Baker and Robert Bradley is a set of genetically isolated interbreeding populations. This
9100-689: Is an indicator that the water is good quality. The hellbender salamander, considered a "habitat specialist", has adapted to fill a specific niche within a very specific environment, and is labeled as such "because its success is dependent on a constancy of dissolved oxygen, temperature and flow found in swift water areas", which in turn limits it to a narrow spectrum of stream/river choices. As a result of this specialization, hellbenders are generally found in areas with large, irregularly shaped, and intermittent rocks and swiftly moving water, while they tend to avoid wider, slow-moving waters with muddy banks and/or slab rock bottoms. This specialization likely contributed to
9282-591: Is believed to be the first widespread culture in the Americas. Over time, indigenous North American cultures grew increasingly sophisticated, and some, such as the Mississippian culture , developed agriculture , architecture , and complex societies . In the post-archaic period , the Mississippian cultures were located in the midwestern , eastern , and southern regions, and the Algonquian in
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#17327650565689464-413: Is blotchy brown or red-brown in color, with a paler underbelly. Hellbenders can also be described as having a gray, or yellowish-brown, to even black coloration. Both males and females grow to an adult length of 24 to 40 cm (9.4 to 15.7 in) from snout to vent , with a total length of 30 to 74 cm (12 to 29 in), making them the fourth-largest aquatic salamander species in the world (after
9646-414: Is called speciation . Charles Darwin was the first to describe the role of natural selection in speciation in his 1859 book The Origin of Species . Speciation depends on a measure of reproductive isolation , a reduced gene flow. This occurs most easily in allopatric speciation, where populations are separated geographically and can diverge gradually as mutations accumulate. Reproductive isolation
9828-528: Is confined to the Ozarks of northern Arkansas and southern Missouri, while C. a. alleganiensis is found in the rest of these states. Some hellbender populations—namely a few in Missouri, Pennsylvania, and Tennessee—have historically been noted to be quite abundant, but several man-made threats have converged on the species such that it has seen a serious population decline throughout its range. In Missouri, it
10010-527: Is derived from the Ancient Greek kryptos (hidden) and branchion (gill). The subspecies name bishopi honors the American herpetologist Sherman C. Bishop . C. alleganiensis has a flat body and head, with beady dorsal eyes and slimy skin. Like most salamanders, it has short legs with four toes on the front legs and five on its back limbs, and its tail is keeled for propulsion. Its tail
10192-403: Is described formally, in a publication that assigns it a unique scientific name. The description typically provides means for identifying the new species, which may not be based solely on morphology (see cryptic species ), differentiating it from other previously described and related or confusable species and provides a validly published name (in botany) or an available name (in zoology) when
10374-630: Is done on both subspecies, as preserving extant genetic diversity is of crucial ecological importance. The Ozark hellbender has been listed as an endangered species under the Endangered Species Act by the US Fish and Wildlife Service since October 5, 2011. This hellbender subspecies inhabits the White River and Spring River systems in southern Missouri and northern Arkansas, and its population has declined an estimated 75% since
10556-513: Is estimated that the populations have declined by 77% since the 1980s. Hellbender populations were listed in 1981 as already extirpated or endangered in Illinois, Indiana, Iowa, and Maryland, decreasing in Arkansas and Kentucky, and generally threatened as a species throughout their range by various human activities and developments. Hellbenders are found in clear, clean water and their presence
10738-671: Is further weakened by the existence of microspecies , groups of organisms, including many plants, with very little genetic variability, usually forming species aggregates . For example, the dandelion Taraxacum officinale and the blackberry Rubus fruticosus are aggregates with many microspecies—perhaps 400 in the case of the blackberry and over 200 in the dandelion, complicated by hybridisation , apomixis and polyploidy , making gene flow between populations difficult to determine, and their taxonomy debatable. Species complexes occur in insects such as Heliconius butterflies, vertebrates such as Hypsiboas treefrogs, and fungi such as
10920-726: Is hard or even impossible to test. Later biologists have tried to refine Mayr's definition with the recognition and cohesion concepts, among others. Many of the concepts are quite similar or overlap, so they are not easy to count: the biologist R. L. Mayden recorded about 24 concepts, and the philosopher of science John Wilkins counted 26. Wilkins further grouped the species concepts into seven basic kinds of concepts: (1) agamospecies for asexual organisms (2) biospecies for reproductively isolated sexual organisms (3) ecospecies based on ecological niches (4) evolutionary species based on lineage (5) genetic species based on gene pool (6) morphospecies based on form or phenotype and (7) taxonomic species,
11102-498: Is known as the presidential system , in contrast to the parliamentary system , where the executive is part of the legislative body. Many countries around the world imitated this aspect of the 1789 Constitution of the United States , especially in the Americas. The Constitution is silent on political parties. However, they developed independently in the 18th century with the Federalist and Anti-Federalist parties. Since then,
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#173276505656811284-485: Is largely behind the need for fast-moving, oxygenated water. If a hellbender ends up in an area of slow-moving water, not enough of it will pass over its skin in a given time, making it difficult to garner enough oxygen to support necessary respiratory functions. A below-favorable oxygen content can make life equally difficult. Hellbenders are preyed upon by diverse predators, including various fish and reptiles (including both snakes and turtles). Particularly, largemouth bass
11466-472: Is listed as Vulnerable on the IUCN Red List of Threatened Species due to the impacts of disease and widespread habitat loss and degradation throughout much of its range. The origin of the name "hellbender" is unclear. The Missouri Department of Conservation says: The name 'hellbender' probably comes from the animal's odd look. One theory claims the hellbender was named by settlers who thought "it
11648-405: Is maintained, and density dependence in turn created, in part by intraspecific predation. When feeding on large prey items relative to themselves, it has been found that they use suction feeding. The hellbenders' breeding season begins in late August or early- to mid-September and can continue as late as the end of November, depending on region. They exhibit no sexual dimorphism , except during
11830-403: Is no easy way to tell whether related geographic or temporal forms belong to the same or different species. Species gaps can be verified only locally and at a point of time. One is forced to admit that Darwin's insight is correct: any local reality or integrity of species is greatly reduced over large geographic ranges and time periods. The botanist Brent Mishler argued that the species concept
12012-478: Is not valid, notably because gene flux decreases gradually rather than in discrete steps, which hampers objective delimitation of species. Indeed, complex and unstable patterns of gene flux have been observed in cichlid teleosts of the East African Great Lakes . Wilkins argued that "if we were being true to evolution and the consequent phylogenetic approach to taxa, we should replace it with
12194-438: Is now the United States experienced conflicts with Native Americans, they also engaged in trade, exchanging European tools for food and animal pelts. Relations ranged from close cooperation to warfare and massacres. The colonial authorities often pursued policies that forced Native Americans to adopt European lifestyles, including conversion to Christianity. Along the eastern seaboard, settlers trafficked African slaves through
12376-421: Is of paramount importance to ensure its continued existence in the wild. The hellbender faces a significant threat due to habitat degradation, primarily caused by activities like dam construction, which disrupts water flow and submerges vital riffle habitats. Logging, mining, and road construction contribute to sedimentation, covering essential nesting and shelter sites. Chemical pollutants and misconceptions about
12558-416: Is shaped like a rudder, but it is rarely used for swimming; these salamanders use pads on their toes instead to grip rocks and walk up and down streams instead of swimming. The hellbender has working lungs, but gill slits are often retained, although only immature specimens have true gills; the hellbender absorbs oxygen from the water through capillaries of its side frills. The frills run from their neck down to
12740-586: Is similar to Mayr's Biological Species Concept, but stresses genetic rather than reproductive isolation. In the 21st century, a genetic species could be established by comparing DNA sequences. Earlier, other methods were available, such as comparing karyotypes (sets of chromosomes ) and allozymes ( enzyme variants). An evolutionarily significant unit (ESU) or "wildlife species" is a population of organisms considered distinct for purposes of conservation. In palaeontology , with only comparative anatomy (morphology) and histology from fossils as evidence,
12922-434: Is sometimes an important source of genetic variation. Viruses can transfer genes between species. Bacteria can exchange plasmids with bacteria of other species, including some apparently distantly related ones in different phylogenetic domains , making analysis of their relationships difficult, and weakening the concept of a bacterial species. United States The United States of America ( USA ), commonly known as
13104-466: Is the common mudpuppy ( Necturus maculosus ). This demarcation can be made by noting the presence of external gills in the mudpuppy, which are lacking in the hellbender, as well as the presence of four toes on each hind foot of the mudpuppy (in contrast with the hellbender's five). Furthermore, the average size of C. a. alleganiensis has been reported to be 45–60 cm (with some reported as reaching up to 74 cm or 30 in), while N. m. maculosus has
13286-552: Is the federal agency charged with addressing most environmental-related issues . The idea of wilderness has shaped the management of public lands since 1964, with the Wilderness Act . The Endangered Species Act of 1973 provides a way to protect threatened and endangered species and their habitats. The United States Fish and Wildlife Service implements and enforces the Act. In 2024, the U.S. ranked 34th among 180 countries in
13468-472: Is the salamander chytrid fungus ( Batrachochytrium salamandrivorans , or "Bsal"), which has caused severe declines in other salamander species. Although not confirmed in the Americas, Bsal's potential introduction poses a substantial risk. If introduced, the impacts on hellbender populations could be swift and severe, necessitating immediate mitigation measures. Karel Čapek War with the Newts (Válka s Mloky in
13650-594: Is threatened by hybridisation, but this can be selected against once a pair of populations have incompatible alleles of the same gene, as described in the Bateson–Dobzhansky–Muller model . A different mechanism, phyletic speciation, involves one lineage gradually changing over time into a new and distinct form (a chronospecies ), without increasing the number of resultant species. Horizontal gene transfer between organisms of different species, either through hybridisation , antigenic shift , or reassortment ,
13832-603: The Allies of World War I , helping to turn the tide against the Central Powers . In 1920, a constitutional amendment granted nationwide women's suffrage . During the 1920s and '30s, radio for mass communication and the invention of early television transformed communications nationwide. The Wall Street Crash of 1929 triggered the Great Depression , which President Franklin D. Roosevelt responded to with
14014-553: The Atlantic slave trade . The original Thirteen Colonies that would later found the United States were administered as possessions of Great Britain , and had local governments with elections open to most white male property owners . The colonial population grew rapidly, eclipsing Native American populations; by the 1770s, the natural increase of the population was such that only a small minority of Americans had been born overseas. The colonies' distance from Britain allowed for
14196-549: The Environmental Performance Index . The country joined the Paris Agreement on climate change in 2016 and has many other environmental commitments. The United States is a federal republic of 50 states and a separate federal capital district, Washington, D.C. It also asserts sovereignty over five unincorporated territories and several uninhabited island possessions . The U.S. is
14378-500: The Federal Bureau of Investigation (FBI) and the U.S. Marshals Service have national jurisdiction and specialized duties, such as protecting civil rights , national security and enforcing U.S. federal courts ' rulings and federal laws. State courts conduct most civil and criminal trials, and federal courts handle designated crimes and appeals of state court decisions . There is no unified "criminal justice system" in
14560-664: The Great Lakes , Mississippi River and Gulf of Mexico . British colonization of the East Coast began with the Virginia Colony (1607) and Plymouth Colony (1620). The Mayflower Compact and the Fundamental Orders of Connecticut established precedents for representative self-governance and constitutionalism that would develop throughout the American colonies. While European settlers in what
14742-719: The Great Lakes region and along the Eastern Seaboard , while the Hohokam culture and Ancestral Puebloans inhabited the southwest . Native population estimates of what is now the United States before the arrival of European immigrants range from around 500,000 to nearly 10 million. Christopher Columbus began exploring the Caribbean for Spain in 1492, leading to Spanish-speaking settlements and missions from Puerto Rico and Florida to New Mexico and California . France established its own settlements along
14924-530: The ICZN for animals and the ICN for plants, do not make rules for defining the boundaries of the species. Research can change the boundaries, also known as circumscription, based on new evidence. Species may then need to be distinguished by the boundary definitions used, and in such cases the names may be qualified with sensu stricto ("in the narrow sense") to denote usage in the exact meaning given by an author such as
15106-596: The Midwest . At the same time, about one million French Canadians migrated from Quebec to New England . During the Great Migration , millions of African Americans left the rural South for urban areas in the North. Alaska was purchased from Russia in 1867. The Compromise of 1877 effectively ended Reconstruction and white supremacists took local control of Southern politics . African Americans endured
15288-688: The National Park Service and other agencies. About 28% of the country's land is publicly owned and federally managed, primarily in the Western States . Most of this land is protected , though some is leased for commercial use, and less than one percent is used for military purposes. Environmental issues in the United States include debates on non-renewable resources and nuclear energy , air and water pollution , biodiversity , logging and deforestation , and climate change . The U.S. Environmental Protection Agency (EPA)
15470-596: The New Deal , a series of sweeping programs and public works projects combined with financial reforms and regulations . All were intended to protect against future economic depressions. Initially neutral during World War II , the U.S. began supplying war materiel to the Allies of World War II in March 1941 and entered the war in December after the Empire of Japan 's attack on Pearl Harbor . The U.S. developed
15652-778: The Organization of American States and the United States–Mexico–Canada Free Trade Agreement . In South America, Colombia is traditionally considered to be the closest ally of the United States. The U.S. exercises full international defense authority and responsibility for Micronesia , the Marshall Islands , and Palau through the Compact of Free Association . It has increasingly conducted strategic cooperation with India , but its ties with China have steadily deteriorated. Since 2014,
15834-648: The Pentium microprocessor in accordance with Moore's law , rechargeable lithium-ion batteries , the first gene therapy trial, and cloning either emerged in the U.S. or were improved upon there. The Human Genome Project was formally launched in 1990, while Nasdaq became the first stock market in the United States to trade online in 1998. In the Gulf War of 1991, an American-led international coalition of states expelled an Iraqi invasion force that had occupied neighboring Kuwait . The September 11 attacks on
16016-708: The South China giant salamander , the Chinese giant salamander and the Japanese giant salamander , respectively) and the largest amphibian in North America, although this length is rivaled by the reticulated siren of the southeastern United States (although the siren is much leaner in build). While males and females grow at similar rates, the females tend to live longer and therefore grow larger. An adult weighs 1.5 to 2.5 kg (3.3 to 5.5 lb), making them
16198-591: The Spanish–American War . (The Philippines was granted full independence from the U.S. on July 4, 1946, following World War II. Puerto Rico and Guam have remained U.S. territories.) American Samoa was acquired by the United States in 1900 after the Second Samoan Civil War . The U.S. Virgin Islands were purchased from Denmark in 1917. The United States entered World War I alongside
16380-623: The United States ( U.S. ) or America , is a country primarily located in North America . It is a federal union of 50 states and a federal capital district, Washington, D.C. The 48 contiguous states border Canada to the north and Mexico to the south, with the states of Alaska to the northwest and the archipelagic Hawaii in the Pacific Ocean . The United States also asserts sovereignty over five major island territories and various uninhabited islands . The country has
16562-700: The cotton gin made the institution increasingly profitable for Southern elites . This sectional conflict regarding slavery culminated in the American Civil War (1861–1865). Eleven slave states seceded and formed the Confederate States of America , while the other states remained in the Union . War broke out in April 1861 after the Confederates bombarded Fort Sumter . After
16744-523: The end of the Cold War in 1991, the U.S. emerged as the world's sole superpower , wielding significant geopolitical influence globally . The U.S. national government is a presidential constitutional federal republic and liberal democracy with three separate branches : legislative , executive , and judicial . It has a bicameral national legislature composed of the House of Representatives ,
16926-537: The fall of communism and the collapse of the Soviet Union , which marked the end of the Cold War and left the United States as the world's sole superpower . The 1990s saw the longest recorded economic expansion in American history , a dramatic decline in U.S. crime rates , and advances in technology . Throughout this decade, technological innovations such as the World Wide Web , the evolution of
17108-399: The fly agaric . Natural hybridisation presents a challenge to the concept of a reproductively isolated species, as fertile hybrids permit gene flow between two populations. For example, the carrion crow Corvus corone and the hooded crow Corvus cornix appear and are classified as separate species, yet they can hybridise where their geographical ranges overlap. A ring species is
17290-400: The global economy in 2023. It possesses by far the largest amount of wealth of any country and has the highest disposable household income per capita among OECD countries. The U.S. ranks among the world's highest in economic competitiveness , productivity , innovation , human rights , and higher education . Its hard power and cultural influence have a global reach. The U.S. is
17472-507: The jaguar ( Panthera onca ) of Latin America or the leopard ( Panthera pardus ) of Africa and Asia. In contrast, the scientific names of species are chosen to be unique and universal (except for some inter-code homonyms ); they are in two parts used together : the genus as in Puma , and the specific epithet as in concolor . A species is given a taxonomic name when a type specimen
17654-436: The siltation and sedimentation, blocking of dispersal/migration routes, and destruction of riverine habitats created by dams and other development, as well as pollution, disease and overharvesting for commercial and scientific purposes. As many of these detrimental effects have irreversibly damaged hellbender populations, it is important to conserve the remaining populations through protecting habitats and—perhaps in places where
17836-406: The "cynical species concept", and arguing that far from being cynical, it usefully leads to an empirical taxonomy for any given group, based on taxonomists' experience. Other biologists have gone further and argued that we should abandon species entirely, and refer to the "Least Inclusive Taxonomic Units" (LITUs), a view that would be coherent with current evolutionary theory. The species concept
18018-813: The 1775 Battles of Lexington and Concord , igniting the American Revolutionary War . At the Second Continental Congress , the colonies appointed George Washington commander-in-chief of the Continental Army , and created a committee that named Thomas Jefferson to draft the Declaration of Independence . Two days after passing the Lee Resolution to create an independent nation the Declaration
18200-559: The 1980s, with only about 590 individuals remaining in the wild. Degraded water quality, habitat loss resulting from impoundments, ore and gravel mining, sedimentation, and collection for the pet trade are thought to be the main factors resulting in the amphibian's decline. When chytridiomycosis killed 75% of the St. Louis Zoo's captive hellbender population between March 2006 and April 2007, tests began to be conducted on wild populations. The disease has been detected in all Missouri populations of
18382-520: The 2010s, political polarization in the country increased between liberal and conservative factions. This polarization was capitalized upon in the January 2021 Capitol attack , when a mob of insurrectionists entered the U.S. Capitol and sought to prevent the peaceful transfer of power in an attempted self-coup d'état . The United States is the world's third-largest country by total area behind Russia and Canada. The 48 contiguous states and
18564-596: The American West have an alpine climate . The climate is arid in the Southwest, Mediterranean in coastal California , and oceanic in coastal Oregon , Washington , and southern Alaska . Most of Alaska is subarctic or polar . Hawaii , the southern tip of Florida and U.S. territories in the Caribbean and Pacific are tropical . States bordering the Gulf of Mexico are prone to hurricanes, and most of
18746-475: The Declaration of Independence on July 4, 1776. The term "United States" and the initialism "U.S.", used as nouns or as adjectives in English, are common short names for the country. The initialism "USA", a noun, is also common. "United States" and "U.S." are the established terms throughout the U.S. federal government , with prescribed rules. In English, the term "America" rarely refers to topics unrelated to
18928-573: The District of Columbia occupy a combined area of 3,119,885 square miles (8,080,470 km ). The coastal plain of the Atlantic seaboard gives way to inland forests and rolling hills in the Piedmont plateau region. The Appalachian Mountains and the Adirondack massif separate the East Coast from the Great Lakes and the grasslands of the Midwest . The Mississippi River System ,
19110-573: The Elder concluded that major military resources needed to be devoted to North America to win the war against France. For the first time, the continent became one of the main theaters of what could be termed a " world war ". The British colonies' position as an integral part of the British Empire became more apparent during the war, with British military and civilian officials becoming a more significant presence in American life. The war increased
19292-683: The January 1863 Emancipation Proclamation , many freed slaves joined the Union army . The war began to turn in the Union's favor following the 1863 Siege of Vicksburg and Battle of Gettysburg , and the Confederacy surrendered in 1865 after the Union's victory in the Battle of Appomattox Court House . The Reconstruction era followed the war. After the assassination of President Abraham Lincoln , Reconstruction Amendments were passed to protect
19474-626: The Origin of Species explained how species could arise by natural selection . That understanding was greatly extended in the 20th century through genetics and population ecology . Genetic variability arises from mutations and recombination , while organisms themselves are mobile, leading to geographical isolation and genetic drift with varying selection pressures . Genes can sometimes be exchanged between species by horizontal gene transfer ; new species can arise rapidly through hybridisation and polyploidy ; and species may become extinct for
19656-574: The Ozark efforts, head-starting programs, in which eggs are collected from the wild and raised in captivity for re-release at a less vulnerable stage, have been initiated in Indiana, New York, and Ohio. Members of the Pennsylvania State Senate have voted to approve the eastern hellbender as the official state amphibian in an effort to raise awareness about its endangered status. The legislation has been mired in controversy due to
19838-522: The Ozark hellbender. NatureServe treats C. a. alleganiensis as an Imperiled Subspecies , C. a. bishopi as a Critically Imperiled Subspecies , and the species as a whole as Vulnerable . The Ozark hellbender was successfully bred in captivity for the first time at the St. Louis Zoo , in a joint project with the Missouri Department of Conservation, hatching on November 15, 2011. Apart from
20020-611: The Rocky Mountains, the Yellowstone Caldera , is the continent's largest volcanic feature. In 2021, the United States had 8% of global permanent meadows and pastures and 10% of cropland. With its large size and geographic variety, the United States includes most climate types. East of the 100th meridian , the climate ranges from humid continental in the north to humid subtropical in the south. The western Great Plains are semi-arid . Many mountainous areas of
20202-647: The U.S. has become a key ally of Ukraine ; it has also provided the country with significant military equipment and other support in response to Russia's 2022 invasion . The president is the commander-in-chief of the United States Armed Forces and appoints its leaders, the secretary of defense and the Joint Chiefs of Staff . The Department of Defense , which is headquartered at the Pentagon near Washington, D.C., administers five of
20384-661: The U.S. territories." Despite disparate systems of confinement, four main institutions dominate: federal prisons , state prisons , local jails, and juvenile correctional facilities . Federal prisons are run by the Federal Bureau of Prisons and hold people who have been convicted of federal crimes, including pretrial detainees. State prisons, run by the official department of correction of each state, hold sentenced people serving prison time (usually longer than one year) for felony offenses. Local jails are county or municipal facilities that incarcerate defendants prior to trial; they also hold those serving short sentences (typically under
20566-464: The United States and the peaceful transfer of power , respectively. The Louisiana Purchase of 1803 from France nearly doubled the territory of the United States. Lingering issues with Britain remained , leading to the War of 1812 , which was fought to a draw. Spain ceded Florida and its Gulf Coast territory in 1819. In the late 18th century, American settlers began to expand westward , many with
20748-714: The United States entered the Cold War, where geopolitical tensions between the U.S. and the Soviet Union led the two countries to dominate world affairs . The U.S. utilized the policy of containment to limit the USSR's sphere of influence, and prevailed in the Space Race , which culminated with the first crewed Moon landing in 1969. Domestically, the U.S. experienced economic growth , urbanization , and population growth following World War II . The civil rights movement emerged, with Martin Luther King Jr. becoming
20930-666: The United States has operated as a de facto two-party system , though the parties in that system have been different at different times. The two main national parties are presently the Democratic and the Republican . The former is perceived as relatively liberal in its political platform while the latter is perceived as relatively conservative . In the American federal system , sovereign powers are shared between two levels of elected government: national and state. People in
21112-593: The United States in 2001 by the pan-Islamist militant organization al-Qaeda led to the war on terror , and subsequent military interventions in Afghanistan and Iraq . The cultural impact of the attacks was profound and long-lasting. The U.S. housing bubble culminated in 2007 with the Great Recession , the largest economic contraction since the Great Depression. Coming to a head in
21294-597: The United States of America to Spain" to seek assistance in the Revolutionary War effort. The first known public usage is an anonymous essay published in the Williamsburg newspaper, The Virginia Gazette , on April 6, 1776. By June 1776, the "United States of America" appeared in the Articles of Confederation and the Declaration of Independence . The Second Continental Congress adopted
21476-593: The United States, despite the usage of "the Americas " as the totality of North and South America. "The States" is an established colloquial shortening of the name, used particularly from abroad; "stateside" is the corresponding adjective or adverb. The first inhabitants of North America migrated from Siberia across the Bering land bridge about 12,000 years ago; the Clovis culture , which appeared around 11,000 BC,
21658-478: The United States, except Iran , North Korea , and Bhutan . Though Taiwan does not have formal diplomatic relations with the U.S., it maintains close unofficial relations. The United States regularly supplies Taiwan with military equipment to deter potential Chinese aggression. Its geopolitical attention also turned to the Indo-Pacific when the United States joined the Quadrilateral Security Dialogue with Australia, India, and Japan. The United States has
21840-509: The United States. The American prison system is largely heterogenous, with thousands of relatively independent systems operating across federal, state, local, and tribal levels. In 2023, "these systems [held] almost 2 million people in 1,566 state prisons, 98 federal prisons, 3,116 local jails, 1,323 juvenile correctional facilities, 181 immigration detention facilities, and 80 Indian country jails, as well as in military prisons , civil commitment centers, state psychiatric hospitals, and prisons in
22022-405: The abbreviation "sp." in the singular or "spp." (standing for species pluralis , Latin for "multiple species") in the plural in place of the specific name or epithet (e.g. Canis sp.). This commonly occurs when authors are confident that some individuals belong to a particular genus but are not sure to which exact species they belong, as is common in paleontology . Authors may also use "spp." as
22204-570: The amount of hybridisation is insufficient to completely mix their respective gene pools . A further development of the recognition concept is provided by the biosemiotic concept of species. In microbiology , genes can move freely even between distantly related bacteria, possibly extending to the whole bacterial domain. As a rule of thumb, microbiologists have assumed that members of Bacteria or Archaea with 16S ribosomal RNA gene sequences more similar than 97% to each other need to be checked by DNA–DNA hybridisation to decide if they belong to
22386-400: The base of their tail on each side of their body. The frills’ function is to increase the surface area of the hellbender and to help the hellbender breathe. Only occasionally leaving the water, the hellbender makes little use of these lungs and the juveniles lose their external gills after around 18 months or about 125 mm in length. Hellbenders use their lungs for buoyancy more than breathing. It
22568-563: The best method for conservation. Crowhurst et al., for instance, found that the "Ozark subspecies" denomination is insufficient for describing genetic (and therefore evolutionary) divergence within the genus Cryptobranchus in the Ozark region. They found three equally divergent genetic units within the genus: C. a. alleganiensis , and two distinct eastern and western populations of C. a. bishopi . These three groups were shown to be isolated, and are considered to most likely be "diverging on different evolutionary paths". Hellbenders are present in
22750-474: The biological species concept, "the several versions" of the phylogenetic species concept, and the idea that species are of the same kind as higher taxa are not suitable for biodiversity studies (with the intention of estimating the number of species accurately). They further suggested that the concept works for both asexual and sexually-reproducing species. A version of the concept is Kevin de Queiroz 's "General Lineage Concept of Species". An ecological species
22932-505: The biological species concept, a cladistic species does not rely on reproductive isolation – its criteria are independent of processes that are integral in other concepts. Therefore, it applies to asexual lineages. However, it does not always provide clear cut and intuitively satisfying boundaries between taxa, and may require multiple sources of evidence, such as more than one polymorphic locus, to give plausible results. An evolutionary species, suggested by George Gaylord Simpson in 1951,
23114-428: The boundaries between closely related species become unclear with hybridisation , in a species complex of hundreds of similar microspecies , and in a ring species . Also, among organisms that reproduce only asexually , the concept of a reproductive species breaks down, and each clone is potentially a microspecies. Although none of these are entirely satisfactory definitions, and while the concept of species may not be
23296-401: The breeding season. While the range of two hellbenders may overlap, they are noted as rarely being present in the overlapping area when the other salamander is in the area. The species is at least somewhat nocturnal, with peak activity being reported by one source as occurring around "two hours after dark" and again at dawn (although the dawn peak was recorded in the lab and could be misleading as
23478-433: The concept of a chronospecies can be applied. During anagenesis (evolution, not necessarily involving branching), some palaeontologists seek to identify a sequence of species, each one derived from the phyletically extinct one before through continuous, slow and more or less uniform change. In such a time sequence, some palaeontologists assess how much change is required for a morphologically distinct form to be considered
23660-405: The contiguous United States are in the State of California, about 84 miles (135 km) apart. At an elevation of 20,310 feet (6,190.5 m), Alaska's Denali is the highest peak in the country and continent. Active volcanoes are common throughout Alaska's Alexander and Aleutian Islands , and Hawaii consists of volcanic islands. The supervolcano underlying Yellowstone National Park in
23842-493: The country's GDP. The U.S. has 42% of the world's nuclear weapons —the second-largest share after Russia. The United States has the third-largest combined armed forces in the world, behind the Chinese People's Liberation Army and Indian Armed Forces . The military operates about 800 bases and facilities abroad, and maintains deployments greater than 100 active duty personnel in 25 foreign countries. State defense forces (SDFs) are military units that operate under
24024-413: The country's territory would expand with the admission of new states , rather than the expansion of existing states. The U.S. Constitution was drafted at the 1787 Constitutional Convention to overcome the limitations of the Articles. It went into effect in 1789, creating a federal republic governed by three separate branches that together ensured a system of checks and balances . George Washington
24206-475: The decline in their populations, as collectors could easily identify their specific habitats. One collector noted, at one time, "one could find a specimen under almost every suitable rock", but after years of collecting, the population had declined significantly. The same collector noted, he "never found two specimens under the same rock", corroborating the account given by other researchers that hellbenders are generally solitary; they are thought to gather only during
24388-584: The development of self-governance, and the First Great Awakening , a series of Christian revivals , fueled colonial interest in religious liberty . For a century, the American colonists had been providing their own troops and materiel in conflicts with indigenous peoples allied with Britain's colonial rivals , especially France, and the Americans had begun to develop a sense of self-defense and self-reliance separate from Britain. The French and Indian War (1754–1763) took on new significance for all North American colonists after Parliament under William Pitt
24570-414: The dispossession of native inhabitants . As more states were admitted , a North–South division over slavery led to the secession of the Confederate States of America , which fought states remaining in the Union in the 1861–1865 American Civil War . With the victory and preservation of the United States, slavery was abolished nationally . By 1900, the country had established itself as a great power ,
24752-591: The eggs and larvae for at least 7–8 months. Hatchling hellbenders are 25–33 mm (0.98–1.30 in) long, have a yolk sac as a source of energy for the first few months of life, and lack functional limbs. Hellbenders are superbly adapted to the shallow, fast-flowing, rocky streams in which they live. Their flattened shape offers little resistance to the flowing water, allowing them to work their way upstream and also to crawl into narrow spaces under rocks. The wrinkles and folds along their skin are used to expand surface area for cutaneous respiration. Their skin also has
24934-407: The environment for labor unions to begin to flourish . This period eventually ended with the advent of the Progressive Era , which was characterized by significant reforms. Pro-American elements in Hawaii overthrew the Hawaiian monarchy ; the islands were annexed in 1898. That same year, Puerto Rico , the Philippines , and Guam were ceded to the U.S. by Spain after the latter's defeat in
25116-435: The evidence to support hypotheses about evolutionarily divergent lineages that have maintained their hereditary integrity through time and space. Molecular markers may be used to determine diagnostic genetic differences in the nuclear or mitochondrial DNA of various species. For example, in a study done on fungi , studying the nucleotide characters using cladistic species produced the most accurate results in recognising
25298-401: The fall mating season, when males have a bulging ring around their cloacal glands. Unlike most salamanders, the hellbender performs external fertilization . Before mating, each male excavates a brood site, a saucer-shaped depression under a rock or log, with its entrance positioned out of the direct current, usually pointing downstream. The male remains in the brood site awaiting a female. When
25480-414: The federal government often applied policies of Indian removal or assimilation . The Trail of Tears (1830–1850) was a U.S. government policy that forcibly removed and displaced most Native Americans living east of the Mississippi River to lands far to the west. These and earlier organized displacements prompted a long series of American Indian Wars west of the Mississippi. The Republic of Texas
25662-408: The fifth heaviest living amphibian in the world after their South China, Chinese and Japanese cousins and the goliath frog , while the largest cane toads may also weigh as much as a hellbender. Hellbenders reach sexual maturity at about five years of age, and may live 30 years in captivity. The hellbender has a few characteristics that make it distinguishable from other native salamanders, including
25844-434: The first nuclear weapons and used them against the Japanese cities of Hiroshima and Nagasaki in August 1945, ending the war. The United States was one of the " Four Policemen " who met to plan the post-war world , alongside the United Kingdom , Soviet Union , and China . The U.S. emerged relatively unscathed from the war, with even greater economic power and international political influence . After World War II,
26026-411: The first settlement of the Thirteen Colonies in Virginia in 1607. Clashes with the British Crown over taxation and political representation sparked the American Revolution , with the Second Continental Congress formally declaring independence on July 4, 1776. Following its victory in the 1775–1783 Revolutionary War , the country continued to expand westward across North America , resulting in
26208-463: The integration of Army National Guard units and personnel into the U.S. Army and (since 1947) the U.S. Air Force. There are about 18,000 U.S. police agencies from local to national level in the United States. Law in the United States is mainly enforced by local police departments and sheriff departments in their municipal or county jurisdictions. The state police departments have authority in their respective state , and federal agencies such as
26390-653: The mating season. If rocks are lacking, hellbenders have been known to use holes in stream banks as habitat. On average, their home range is estimated to be 198 square meters as of 2005. The ideal habitat for a hellbender has a large amount of gravel, low pH, cool water temperatures, and low specific conductivity. The large amounts of gravel enable the hellbender to hide, cool water temperatures allow for more efficient cutaneous gas exchange, and low specific conductivity may indicate an undisturbed stream. Hellbender communities may be more concentrated in undisturbed areas. Both subspecies, C. a. alleganiensis and C. a. bishopi undergo
26572-427: The military draft (leading to the end of conscription in 1973) and wide opposition to U.S. intervention in Vietnam (with the U.S. totally withdrawing in 1975). A societal shift in the roles of women was significantly responsible for the large increase in female paid labor participation during the 1970s, and by 1985 the majority of American women aged 16 and older were employed. The late 1980s and early 1990s saw
26754-612: The most attractive to the population are the most vulnerable. The U.S. is one of 17 megadiverse countries containing large numbers of endemic species : about 17,000 species of vascular plants occur in the contiguous United States and Alaska, and over 1,800 species of flowering plants are found in Hawaii, few of which occur on the mainland. The United States is home to 428 mammal species, 784 birds, 311 reptiles, 295 amphibians , and around 91,000 insect species. There are 63 national parks , and hundreds of other federally managed parks, forests, and wilderness areas , managed by
26936-478: The most violent, the California genocide of thousands of Native inhabitants, lasted into the early 1870s, just as additional western territories and states were created. During the colonial period, slavery had been legal in the American colonies , though the practice began to be significantly questioned during the American Revolution. States in the North enacted abolition laws , though support for slavery strengthened in Southern states , as inventions such as
27118-429: The numerous fungi species of all the concepts studied. Versions of the phylogenetic species concept that emphasise monophyly or diagnosability may lead to splitting of existing species, for example in Bovidae , by recognising old subspecies as species, despite the fact that there are no reproductive barriers, and populations may intergrade morphologically. Others have called this approach taxonomic inflation , diluting
27300-401: The original Czech), also translated as Salamander Wars, is a 1936 satirical science fiction novel by Czech author Karel Čapek Species A species ( pl. : species) is a population of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring , typically by sexual reproduction . It is the basic unit of classification and
27482-525: The paper is accepted for publication. The type material is usually held in a permanent repository, often the research collection of a major museum or university, that allows independent verification and the means to compare specimens. Describers of new species are asked to choose names that, in the words of the International Code of Zoological Nomenclature , are "appropriate, compact, euphonious, memorable, and do not cause offence". Books and articles sometimes intentionally do not identify species fully, using
27664-674: The person who named the species, while the antonym sensu lato ("in the broad sense") denotes a wider usage, for instance including other subspecies. Other abbreviations such as "auct." ("author"), and qualifiers such as "non" ("not") may be used to further clarify the sense in which the specified authors delineated or described the species. Species are subject to change, whether by evolving into new species, exchanging genes with other species, merging with other species or by becoming extinct. The evolutionary process by which biological populations of sexually-reproducing organisms evolve to become distinct or reproductively isolated as species
27846-408: The policy of redlining later adopted by the federal Home Owners' Loan Corporation . An explosion of technological advancement accompanied by the exploitation of cheap immigrant labor led to rapid economic expansion during the late 19th and early 20th centuries , allowing the United States to outpace the economies of England, France, and Germany combined. This fostered the amassing of power by
28028-420: The primary colonial grievances was a denial of their rights as Englishmen , particularly the right to representation in the British government that taxed them . To demonstrate their dissatisfaction and resolve, the First Continental Congress met in 1774 and passed the Continental Association , a colonial boycott of British goods that proved effective. The British attempt to then disarm the colonists resulted in
28210-487: The result of misclassification leading to questions on whether there really are any ring species. The commonly used names for kinds of organisms are often ambiguous: "cat" could mean the domestic cat, Felis catus , or the cat family, Felidae . Another problem with common names is that they often vary from place to place, so that puma, cougar, catamount, panther, painter and mountain lion all mean Puma concolor in various parts of America, while "panther" may also mean
28392-515: The rights of African Americans . National infrastructure, including transcontinental telegraph and railroads , spurred growth in the American frontier . From 1865 through 1917 an unprecedented stream of immigrants arrived in the United States, including 24.4 million from Europe. Most came through the port of New York City , and New York City and other large cities on the East Coast became home to large Jewish , Irish , and Italian populations, while many Germans and Central Europeans moved to
28574-429: The ring. Ring species thus present a difficulty for any species concept that relies on reproductive isolation. However, ring species are at best rare. Proposed examples include the herring gull – lesser black-backed gull complex around the North pole, the Ensatina eschscholtzii group of 19 populations of salamanders in America, and the greenish warbler in Asia, but many so-called ring species have turned out to be
28756-437: The rocky Great Basin and Chihuahua , Sonoran , and Mojave deserts. In the northwest corner of Arizona , carved by the Colorado River over millions of years, is the Grand Canyon , a steep-sided canyon and popular tourist destination known for its overwhelming visual size and intricate, colorful landscape. The Sierra Nevada and Cascade mountain ranges run close to the Pacific coast . The lowest and highest points in
28938-502: The same family are in the genus Andrias , which contains the Japanese and Chinese giant salamanders . The hellbender is much larger than any other salamander in its geographic range, and employs an unusual adaption for respiration through cutaneous gas exchange via capillaries found in its lateral skin folds. It fills a particular niche — both as a predator and prey — in its ecosystem , which either it or its ancestors have occupied for around 65 million years. The species
29120-508: The same species. This concept was narrowed in 2006 to a similarity of 98.7%. The average nucleotide identity (ANI) method quantifies genetic distance between entire genomes , using regions of about 10,000 base pairs . With enough data from genomes of one genus, algorithms can be used to categorize species, as for Pseudomonas avellanae in 2013, and for all sequenced bacteria and archaea since 2020. Observed ANI values among sequences appear to have an "ANI gap" at 85–95%, suggesting that
29302-529: The same species. When two species names are discovered to apply to the same species, the older species name is given priority and usually retained, and the newer name considered as a junior synonym, a process called synonymy . Dividing a taxon into multiple, often new, taxa is called splitting . Taxonomists are often referred to as "lumpers" or "splitters" by their colleagues, depending on their personal approach to recognising differences or commonalities between organisms. The circumscription of taxa, considered
29484-464: The six service branches, which are made up of the U.S. Army , Marine Corps , Navy , Air Force , and Space Force . The Coast Guard is administered by the Department of Homeland Security in peacetime and can be transferred to the Department of the Navy in wartime. The United States spent $ 916 billion on its military in 2023, which is by far the largest amount of any country , making up 37% of global military spending and accounting for 3.4% of
29666-407: The sole authority of a state government. SDFs are authorized by state and federal law but are under the command of the state's governor . They are distinct from the state's National Guard units in that they cannot become federalized entities. A state's National Guard personnel, however, may be federalized under the National Defense Act Amendments of 1933 , which created the Guard and provides for
29848-456: The species concept and making taxonomy unstable. Yet others defend this approach, considering "taxonomic inflation" pejorative and labelling the opposing view as "taxonomic conservatism"; claiming it is politically expedient to split species and recognise smaller populations at the species level, because this means they can more easily be included as endangered in the IUCN red list and can attract conservation legislation and funding. Unlike
30030-556: The species have led to declines. Over-collection for sale and deliberate eradication efforts have also been detrimental. The salamander's habitat is further jeopardized by habitat modifications stemming from industrialization and urbanization, including increased stream channelization and pollution from agricultural runoff, mining, and thermal pollution. Diseases, like Batrachochytrium dendrobatidis (Bd) and Ranavirus infections, have been detected in hellbender populations, contributing to population declines. An emerging disease threat
30212-476: The species was once endemic and has been extirpated—by augmenting numbers through reintroduction . Due to sharp decreases seen in the Ozark subspecies, researchers have been trying to differentiate C. a. alleganiensis and C. a. bishopi into two management units. Indeed, researchers found significant genetic divergence between the two groups, as well as between them and another isolated population of C. a. alleganiensis . This could be reason enough to ensure work
30394-420: The sperm uniformly. The male often tempts other females to lay eggs in his nest, and as many as 1,946 eggs have been counted in a single nest. Males also exhibit mate and shelter guarding. Mortality rate is high for hellbender eggs. Studies have found that until the female successfully reproduces, the male hellbender will guard her in his territory until the reproduction is complete. Cannibalism , however, leads to
30576-543: The states are also represented by local elected governments , which are administrative divisions of the states. States are subdivided into counties or county equivalents , and further divided into municipalities . The District of Columbia is a federal district containing the U.S. capital, Washington, D.C. . The federal district is an administrative division of the federal government. Federally recognized tribes govern 326 Indian reservations . The United States has an established structure of foreign relations, and it has
30758-451: The structure and responsibilities of the national federal government and its relationship with the individual states. The U.S. Constitution is the world's oldest national constitution still in effect (from March 4, 1789). Composed of three branches, all headquartered in Washington, D.C., the federal government is the national government of the United States. It is regulated by a strong system of checks and balances . The three-branch system
30940-420: The success of the hellbender bill in the Pennsylvania Senate will contribute to clean water efforts and raise awareness for the hellbender's struggling population. The hellbender faces an array of challenges that jeopardize its habitat and overall well-being. These challenges include habitat degradation, habitat modifications, pollution, and the looming threat of emerging diseases. The conservation of this species
31122-472: The trail of scent molecules. Smell is possibly their most important sense when hunting. They also have a lateral line similar to those of fish, with which they can detect vibrations in the water. Research throughout the range of the hellbender has shown a dramatic decline in populations in the majority of locations. As of 2022, the species is classified as Vulnerable by the IUCN. Many different anthropogenic sources have contributed to this decline, including
31304-494: The wild. It is difficult to define a species in a way that applies to all organisms. The debate about species concepts is called the species problem. The problem was recognised even in 1859, when Darwin wrote in On the Origin of Species : I was much struck how entirely vague and arbitrary is the distinction between species and varieties. He went on to write: No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of
31486-406: The world's fourth-longest river system , runs predominantly north–south through the heart of the country. The flat and fertile prairie of the Great Plains stretches to the west, interrupted by a highland region in the southeast. The Rocky Mountains , west of the Great Plains, extend north to south across the country, peaking at over 14,000 feet (4,300 m) in Colorado . Farther west are
31668-449: The world's second-largest diplomatic corps as of 2024 . It is a permanent member of the United Nations Security Council , and home to the United Nations headquarters . The United States is a member of the G7 , G20 , and OECD intergovernmental organizations. Almost all countries have embassies and many have consulates (official representatives) in the country. Likewise, nearly all countries host formal diplomatic missions with
31850-461: The world's third-largest land area , largest exclusive economic zone , and third-largest population , exceeding 334 million. Its three largest metropolitan areas are New York , Los Angeles , and Chicago , and its three most populous states are California , Texas , and Florida . Paleo-Indians migrated across the Bering land bridge more than 12,000 years ago, and formed various civilizations and societies . British colonization led to
32032-444: The world's oldest surviving federation, and its presidential system of national government has been adopted, in whole or in part, by many newly independent states worldwide following their decolonization . It is a liberal representative democracy "in which majority rule is tempered by minority rights protected by law ". The Constitution of the United States serves as the country's supreme legal document , also establishing
32214-423: The world's tornadoes occur in the country , mainly in Tornado Alley . Overall, the United States receives more high-impact extreme weather incidents than any other country. Extreme weather became more frequent in the U.S. in the 21st century, with three times the number of reported heat waves as in the 1960s. In the American Southwest , droughts became more persistent and more severe. The regions considered as
32396-458: Was annexed in 1845, and the 1846 Oregon Treaty led to U.S. control of the present-day American Northwest . Victory in the Mexican–American War resulted in the 1848 Mexican Cession of California, Nevada, Utah, and much of present-day Colorado and the American Southwest . The California gold rush of 1848–1849 spurred a huge migration of white settlers to the Pacific coast, leading to even more confrontations with Native populations. One of
32578-613: Was a creature from hell where it's bent on returning." Another rendition says the undulating skin of a hellbender reminded observers of "horrible tortures of the infernal regions." In reality, it's a harmless aquatic salamander. In a study conducted in Indiana, informing the public about the rarity and locality of the hellbender resulted in more positive attitudes toward this species than were previously held. Other vernacular names include snot otter, lasagna lizard, devil dog, mud-devil, mud dog, water dog, grampus, Allegheny alligator, and leverian water newt. The generic name , Cryptobranchus ,
32760-703: Was adopted on July 4, 1776. The political values of the American Revolution included liberty , inalienable individual rights ; and the sovereignty of the people ; supporting republicanism and rejecting monarchy , aristocracy , and all hereditary political power; civic virtue ; and vilification of political corruption . The Founding Fathers of the United States , who included Washington, Jefferson, John Adams , Benjamin Franklin , Alexander Hamilton , John Jay , James Madison , Thomas Paine , and many others, were inspired by Greco-Roman , Renaissance , and Enlightenment philosophies and ideas. The Articles of Confederation were ratified in 1781 and established
32942-404: Was elected the country's first president under the Constitution, and the Bill of Rights was adopted in 1791 to allay skeptics' concerns about the power of the more centralized government. His resignation as commander-in-chief after the Revolutionary War and his later refusal to run for a third term as the country's first president established a precedent for the supremacy of civil authority in
33124-446: Was observed and concluded that hellbenders are capable of detecting kairomones , which are chemical cues emitted by predatory species. This suggests that hellbenders can recognize kairomones as stressful stimuli and identify potential predators. Once a hellbender finds a favorable location, it generally does not stray too far from it—except occasionally for breeding and hunting—and will protect it from other hellbenders both in and out of
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