HOME

TheInfoList



OR:

The American coot (''Fulica americana''), also known as a mud hen or pouldeau, is a bird of the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Rallidae The rails, or Rallidae, are a large cosmopolitan family of small- to medium-sized, ground-living birds. The family exhibits considerable diversity and includes the crakes, coots, and gallinules. Many species are associated with wetlands, althou ...
. Though commonly mistaken for ducks, American coots are only distantly related to ducks, belonging to a separate order. Unlike the webbed feet of ducks, coots have broad, lobed scales on their lower legs and toes that fold back with each step which facilitates walking on dry land. Coots live near water, typically inhabiting wetlands and open water bodies in North America. Groups of coots are called covers or rafts. The oldest known coot lived to be 22 years old. The American coot is a migratory bird that occupies most of North America. It lives in the Pacific and southwestern United States and Mexico year-round and occupies more northeastern regions during the summer breeding season. In the winter they can be found as far south as Panama. Coots generally build floating nests and lay 8–12 eggs per clutch. Females and males have similar appearances, but they can be distinguished during aggressive displays by the larger ruff (head plumage) on the male. American coots eat primarily algae and other aquatic plants but also animals (both vertebrates and invertebrates) when available. The American coot is listed as “Least Concern” under the IUCN conservation ratings. Hunters generally avoid killing American coots because their meat is not as sought after as that of ducks. Much research has been done on the breeding habits of American coots. Studies have found that mothers will preferentially feed offspring with the brightest plume feathers, a characteristic known as chick ornaments. American coots are also susceptible to conspecific brood parasitism and have evolved mechanisms to identify which offspring are theirs and which are from parasitic females.


Taxonomy and systematics

The American coot was formally described in 1789 by the German naturalist Johann Friedrich Gmelin in his revised and expanded edition of
Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...
's '' Systema Naturae''. He placed it with all the other coots in the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
'' Fulica'' and coined the binomial name ''Fulica americana''. Gmelin based his description on the "Cinereous coot" from North America that had been described in 1785 by the English ornithologist John Latham in his book ''A General Synopsis of Birds''. Two subspecies are recognised: * ''F. a. americana'' Gmelin, JF, 1789 – southeast Alaska and Canada to Costa Rica and the West Indies * ''F. a. columbiana'' Chapman, 1914 – Colombia and north Ecuador The Caribbean coot, a color morph of the American coot is now included with the nominate subspecies. Coot
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s from the Middle
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...
of
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
have been described as ''Fulica hesterna'' but cannot be separated from the present-day American coot. However, the Pleistocene coot ''Fulica shufeldti'' (formerly ''F. minor''), famously known as part of the
Fossil Lake Fossil Butte National Monument is a United States National Monument managed by the National Park Service, located west of Kemmerer, Wyoming, United States. It centers on an assemblage of Eocene Epoch (56 to 34 million years ago) animal and pl ...
fauna, quite possibly was a
paleosubspecies A chronospecies is a species derived from a sequential development pattern that involves continual and uniform changes from an extinct ancestral form on an evolutionary scale. The sequence of alterations eventually produces a population that is p ...
of the American coot (as ''Fulica americana shufeldti'') as they only differed marginally in size and proportions from living birds. Thus, it seems that the modern-type American coots
evolved Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
during the mid-late
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...
, a few hundred thousand years ago. The American coot's genus name, ''Fulica'', is a direct borrowing of the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
word for "coot". The specific epithet ''americana'' means "America".


Description

The American coot measures in length with a
wingspan The wingspan (or just span) of a bird or an airplane is the distance from one wingtip to the other wingtip. For example, the Boeing 777–200 has a wingspan of , and a wandering albatross (''Diomedea exulans'') caught in 1965 had a wingspan o ...
of . Adults have a short, thick, white bill and white
frontal shield A frontal shield, also known as a facial shield or frontal plate, is a feature of the anatomy of several bird species. Located just above the upper mandible, and protruding along the forehead, it is composed of two main parts: a hard, proteinaceo ...
, which usually has a reddish-brown spot near the top of the bill between the eyes. Males and females look alike, but females are smaller. Body mass in females ranges from and in males from .''CRC Handbook of Avian Body Masses'' by John B. Dunning Jr. (Editor). CRC Press (1992), . Juvenile birds have olive-brown crowns and a gray body. They become adult-colored around 4 months of age.


Vocalizations

The American coot has a variety of repeated calls and sounds. Male and female coots make different types of calls to similar situations. Male alarm calls are ' while female alarm calls are '. Also, stressed males go ' or ' while females call '.


Distribution and habitat

American coots are found near water reed-ringed lakes and ponds, open marshes, and sluggish rivers. They prefer freshwater environments but may temporarily live in saltwater environments during the winter months. The American coot's breeding habitat extends from
marsh A marsh is a wetland that is dominated by herbaceous rather than woody plant species.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p Marshes can often be found a ...
es in southern
Quebec Quebec ( ; )According to the Canadian government, ''Québec'' (with the acute accent) is the official name in Canadian French and ''Quebec'' (without the accent) is the province's official name in Canadian English is one of the thirtee ...
to the Pacific coast of North America and as far south as northern South America. Birds from temperate North America east of the Rocky Mountains migrate to the southern United States and southern
British Columbia British Columbia (commonly abbreviated as BC) is the westernmost province of Canada, situated between the Pacific Ocean and the Rocky Mountains. It has a diverse geography, with rugged landscapes that include rocky coastlines, sandy beaches, ...
. It is often a year-round resident where water remains open in winter. The number of birds that stay year-round near the northern limit of the species' range seems to be increasing. Autumn migration occurs from August to December, with males and non-breeders moving south before the females and juveniles. Spring migration to breeding ranges occurs from late February to mid-May, with males and older birds moving North first. There has been evidence of birds travelling as far north as Greenland and Iceland.


Caribbean coot

Coots resident in the Caribbean and Greater and Lesser Antilles lack the red portion of the frontal shield, and were previously believed to be a distinct species, the Caribbean coot (''Fulica caribaea''). In 2016, due to research showing that the only distinguishing characteristic between American and Caribbean coots, the presence or absence of red in the frontal shield, was not distinct to Caribbean coots as some American coots, in locations where vagrancy from Caribbean populations was highly unlikely, had fully white shields and, therefore, there was no way to reliably distinguish the species, and there was no evidence of Caribbean and American coots engaging in
assortative mating Assortative mating (also referred to as positive assortative mating or homogamy) is a mating pattern and a form of sexual selection in which individuals with similar phenotypes or genotypes mate with one another more frequently than would be ex ...
, the
American Ornithological Society The American Ornithological Society (AOS) is an ornithological organization based in the United States. The society was formed in October 2016 by the merger of the American Ornithologists' Union (AOU) and the Cooper Ornithological Society. Its m ...
lumped the Caribbean coot as a regional variation of the American coot.


Behavior and ecology

The American coot is a highly
gregarious Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...
species, particularly in the winter, when its flocks can number in the thousands. When swimming on the water surface, American coots exhibit a variety of interesting collective formations, including single-file lines, high density synchronized swimming and rotational dynamics, broad arcing formations, and sequential take-off dynamics.


Feeding

The American coot can dive for food but can also forage and scavenge on land. Their principal source of food is aquatic vegetation, especially algae. Yet they are
omnivorous An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
, also eating
arthropod Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chiti ...
s,
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
, and other aquatic animals. During breeding season, coots are more likely to eat aquatic insects and
mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is e ...
s—which constitute the majority of a chick's diet.


Breeding

The coot mating season occurs during May and June. Coot mate pairings are monogamous throughout their life, given they have a suitable territory. A typical reproductive cycle involves multiple stages:
pairing In mathematics, a pairing is an ''R''-bilinear map from the Cartesian product of two ''R''-modules, where the underlying ring ''R'' is commutative. Definition Let ''R'' be a commutative ring with unit, and let ''M'', ''N'' and ''L'' be ''R''-mod ...
,
nest A nest is a structure built for certain animals to hold eggs or young. Although nests are most closely associated with birds, members of all classes of vertebrates and some invertebrates construct nests. They may be composed of organic materi ...
ing,
copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetra ...
, egg deposition, incubation, and
hatching Hatching (french: hachure) is an artistic technique used to create tonal or shading Shading refers to the depiction of depth perception in 3D models (within the field of 3D computer graphics) or illustrations (in visual art) by varying ...
. The American coot typically has long
courtship Courtship is the period wherein some couples get to know each other prior to a possible marriage. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marriage. A courtship may be an informal and private m ...
periods. This courtship period is characterized by billing, bowing, and nibbling. Males generally initiate billing, which is the touching of bills between individuals. As the pair bond becomes more evident, both males and females will initiate billing only with each other and not other males or females. After a pair bond is cemented, the mating pair looks for a territory to build a nest in. A pair bond becomes permanent when a nesting territory is secured.
Copulation Sexual intercourse (or coitus or copulation) is a sexual activity typically involving the insertion and thrusting of the penis into the vagina for sexual pleasure or reproduction.Sexual intercourse most commonly means penile–vaginal penetra ...
behavior among coot pairs always falls under the same general pattern. First the male chases the female. Then, the female moves to the display platform and squats with her head under the water. The male then mounts the female, using his claws and wings to balance on the female's back while she brings her head above the water. Sexual intercourse usually takes no longer than two seconds.


Nests

The American coot is a prolific builder and will create multiple structures during a single breeding season. It nests in well-concealed locations in tall reeds. There are three general types of structures: display platforms, egg nests and brood nests. * Display platforms are used as roosting sites and are left to decompose after copulation. * Egg nests are typically in diameter with a ramp that allows the parents to enter and exit without tearing the sides of the nests. Coots will often build multiple egg nests before selecting one to lay their eggs in. * Brood nests are nests that are either newly constructed or have been converted from old egg nests after the eggs hatch, becoming larger egg nests. Since American coots build on the water, their structures disintegrate easily and have short life spans. Egg and brood nests are actually elaborate rafts, and must be constantly added to in order to stay afloat. Females typically do the most work while building.


Egg-laying and clutch size

Females deposit one egg a day until the clutch is complete. Eggs are usually deposited between sunset and midnight. Typically, early season and first clutches average two more eggs than second nestings and late season clutches. Early season nests see an average of 9.0 eggs per clutch while late clutches see an average of 6.4 eggs per clutch. There is an inverse relationship between egg weights and laying sequence, wherein earlier eggs are larger than eggs laid later in the sequence. It is possible to induce a female coot to lay more eggs than normal by either removing all or part of her clutch. Sometimes, a female may abandon the clutch if enough eggs are removed. Coots, however, do not respond to experimental addition of eggs by laying fewer eggs. The American coot is a persistent re-nester, and will replace lost clutches with new ones within two days of clutch-loss during deposition. One study showed that 68% of destroyed clutches are eventually replaced. Re-nested clutches are typically smaller than original clutches by one or two eggs, but this could be attributed to differences in time and habitat quality instead of food or nutrient reserves and availability. Younger females reproduce later in the season and produce smaller eggs than older females. Their offspring are also smaller. However, there is no difference in
clutch size __NOTOC__ A clutch of egg (biology), eggs is the group of eggs produced by birds, amphibians, or reptiles, often at a single time, particularly those laid in a nest. In birds, destruction of a clutch by predators (or removal by humans, for exam ...
between older and younger females as there is in other avian species.


Incubation and hatching

Incubation start time in the American coot is variable, and can begin anywhere from the deposition of the first egg to after the clutch is fully deposited. Starting incubation before the entire clutch has been laid is an uncommon practice among birds. Once incubation starts it continues without interruption. Male and female coots share incubation responsibility, but males do most of the work during the 21-day incubation period. Females will begin to re-nest clutches in an average of six days if clutches are destroyed during incubation. Hatch order usually follows the same sequence as laying order. Regardless of clutch size, eight is the typical maximum size of a
brood Brood may refer to: Nature * Brood, a collective term for offspring * Brooding, the incubation of bird eggs by their parents * Bee brood, the young of a beehive * Individual broods of North American Periodical Cicadas: ** Brood X, the largest b ...
. Egg desertion is a frequent occurrence among coots because females will often deposit more than eight eggs. Brood size limits incubation time, and when a certain number of chicks have hatched the remaining eggs are abandoned. The mechanism for egg abandonment has not yet been discovered. Food resource constraints may limit the number of eggs parents let hatch, or the remaining eggs may not provide enough visual or tactile stimulation to elicit incubation behavior. An American coot can be forced to hatch more eggs than are normally laid. These additional offspring, however, suffer higher
mortality rate Mortality rate, or death rate, is a measure of the number of deaths (in general, or due to a specific cause) in a particular population, scaled to the size of that population, per unit of time. Mortality rate is typically expressed in units of d ...
s due to inadequacy in brooding or feeding ability.


Maternal effects

Hormones that are passed down from the mother into the egg affect offspring growth, behavior, and social interactions. These nongenetic contributions by the mother are known as
maternal effect A maternal effect is a situation where the phenotype of an organism is determined not only by the environment it experiences and its genotype, but also by the environment and genotype of its mother. In genetics, maternal effects occur when an org ...
s. In the American coot, two levels of
androgen An androgen (from Greek ''andr-'', the stem of the word meaning "man") is any natural or synthetic steroid hormone that regulates the development and maintenance of male characteristics in vertebrates by binding to androgen receptors. This in ...
and
testosterone Testosterone is the primary sex hormone and anabolic steroid in males. In humans, testosterone plays a key role in the development of male reproductive tissues such as testes and prostate, as well as promoting secondary sexual characteristi ...
variation have been discovered—within-clutch and among-clutch variation. Within the same clutch, eggs laid earlier in the sequence have higher testosterone levels than eggs laid later in the sequence. Females that lay three clutches deposit more androgens into their yolks than females who lay only one or two clutches.


Brood parasitism

The American coot has a mixed reproductive strategy, and
conspecific Biological specificity is the tendency of a characteristic such as a behavior or a biochemical variation to occur in a particular species. Biochemist Linus Pauling stated that "Biological specificity is the set of characteristics of living organis ...
brood parasitism Brood parasites are animals that rely on others to raise their young. The strategy appears among birds, insects and fish. The brood parasite manipulates a host, either of the same or of another species, to raise its young as if it were its ow ...
is a common alternative reproductive method. In one 4-year study, researchers found that 40% of nests were parasitized, and that 13% of all eggs were laid by females in nests that were not their own. Increasing reproductive success under social and ecological constraints is the primary reason for brood parasitism. Floater females without territories or nests use brood parasitism as their primary method of reproduction, if they breed at all. Other females may engage in brood parasitism if their partially complete clutches are destroyed. Conspecific brood parasitic behavior is most common among females trying to increase their total number of offspring. Food supply is the limiting factor to chick survival and starvation is the most common cause of chick morbidity. Parasitic females bypass the parental care constraint of feeding by laying additional parasitic eggs in addition to their normal nest. When a parasitic female lays her egg in a host female's nest, the host female experiences a deposition rate of two eggs per day. Host females may recognize parasitic eggs when the egg deposition pattern deviates from the traditional one egg per day pattern. The occurrence of brood parasitism may be influenced by the body size of the potential parasitic female relative to the potential host female. Parasitic females are generally larger than their host counterparts, but on average, there is no size difference between the parasite and the host. The American coot, unlike other parasitized species, has the ability to recognize and reject conspecific parasitic chicks from their brood. Parents aggressively reject parasite chicks by pecking them vigorously, drowning them, preventing them from entering the nest, etc. They learn to recognize their own chicks by imprinting on cues from the first chick that hatches. The first-hatched chick is a reference to which parents discriminate between later-hatched chicks. Chicks that do not match the imprinted cues are then recognized as parasite chicks and are rejected. Chick recognition reduces the costs associated with parasitism, and coots are one of only three bird species in which this behavior has evolved. This is because hatching order is predictable in parasitized coots—host eggs will reliably hatch before parasite eggs. In other species where hatching order is not as reliable, there is a risk of misimprinting on a parasite chick first and then rejecting their own chicks. In these species, the cost of accidentally misimprinting is greater than the benefits of rejecting parasite chicks.


Chick ornaments

The first evidence for parental selection of exaggerated, ornamental traits in offspring was found in American coots. Black American coot chicks have conspicuously orange-tipped ornamental plumes covering the front half of their body that are known as “chick ornaments” that eventually get bleached out after six days. This brightly colored, exaggerated trait makes coot chicks more susceptible to predation and does not aid in thermoregulation, but remains selected for by parental choice. These plumes are not necessary for chick viability, but increased chick ornamentation increases the likelihood that a chick will be chosen as a favorite by the parents. Experimental manipulation of chick ornamentation by clipping the bright plumes has shown that parents prefer ornamented chicks over non-ornamented ones.


Predation

The American coot is fairly aggressive in defense of its eggs and, in combination with their protected nesting habitat, undoubtedly helps reduce losses of eggs and young to all but the most determined and effective predators.
American crow The American crow (''Corvus brachyrhynchos'') is a large passerine bird species of the family Corvidae. It is a common bird found throughout much of North America. American crows are the New World counterpart to the carrion crow of Europe an ...
s, black-billed magpies and
Forster's tern Forster's tern (''Sterna forsteri'') is a tern in the family Laridae. The genus name ''Sterna'' is derived from Old English "stearn", "tern", and ''forsteri'' commemorates the naturalist Johann Reinhold Forster. It breeds inland in North America ...
can sometimes take eggs. Mammalian predators (including red foxes, coyotes, skunks and
raccoon The raccoon ( or , ''Procyon lotor''), sometimes called the common raccoon to distinguish it from other species, is a mammal native to North America. It is the largest of the procyonid family, having a body length of , and a body weight of ...
s) are even less likely to predate coot nests, though nests are regularly destroyed in usurpation by muskrats. Conversely, the bold behavior of immature and adult coots leads to them falling prey with relative regularity once out of the breeding season. Regular, non-nesting-season predators include
great horned owl The great horned owl (''Bubo virginianus''), also known as the tiger owl (originally derived from early naturalists' description as the "winged tiger" or "tiger of the air"), or the hoot owl, is a large owl native to the Americas. It is an extre ...
s,
northern harrier The northern harrier (''Circus hudsonius''), or ring-tailed hawk, is a bird of prey. It breeds throughout the northern parts of the northern hemisphere in Canada and the northernmost USA. The northern harrier migrates to more southerly areas ...
s, bald eagles,
golden eagle The golden eagle (''Aquila chrysaetos'') is a bird of prey living in the Northern Hemisphere. It is the most widely distributed species of eagle. Like all eagles, it belongs to the family Accipitridae. They are one of the best-known birds of ...
s,
American alligator The American alligator (''Alligator mississippiensis''), sometimes referred to colloquially as a gator or common alligator, is a large crocodilian reptile native to the Southeastern United States. It is one of the two extant species in the gen ...
s, bobcats, great black-backed and
California gull The California gull (''Larus californicus'') is a medium-sized gull, smaller on average than the herring gull but larger on average than the ring-billed gull, though it may overlap in size greatly with both. Description Adults are similar in ap ...
s. In fact, coots may locally comprise more than 80% of the bald eagle diet.


In culture

On the Louisiana coast, the
Cajun The Cajuns (; French: ''les Cadjins'' or ''les Cadiens'' ), also known as Louisiana ''Acadians'' (French: ''les Acadiens''), are a Louisiana French ethnicity mainly found in the U.S. state of Louisiana. While Cajuns are usually described as ...
word for coot is ''pouldeau'', from French for "coot", ''poule d'eau'' – literally "water hen". Coot can be used for cooking; it is somewhat popular in
Cajun cuisine Cajun cuisine (french: cuisine cadienne , es, cocina acadiense) is a style of cooking developed by the Cajun–Acadians who were deported from Acadia to Louisiana during the 18th century and who incorporated West African, French and Spanish c ...
, for instance as an ingredient for
gumbo Gumbo (Louisiana Creole: Gombo) is a soup popular in the U.S. state of Louisiana, and is the official state cuisine. Gumbo consists primarily of a strongly-flavored stock, meat or shellfish (or sometimes both), a thickener, and the Creole "h ...
s cooked at home by duck hunters. The bird is the mascot of the
Toledo Mud Hens The Toledo Mud Hens are a Minor League Baseball team of the International League and the Triple-A affiliate of the Detroit Tigers. They are located in Toledo, Ohio, and play their home games at Fifth Third Field. A Mud Hens team has played in ...
Minor League Baseball team.


Conservation and threats

The American coot is listed under "least concern" by the IUCN Red List of Endangered Species. They are common and widespread, and are sometimes even considered a pest. They are rarely the targets of hunters since their meat is not considered to be as good as that of ducks; although some are shot for sport, particularly in the southeastern United States. Because they are found in wetlands, scientists use them to monitor toxin levels and pollution problems in these environments.


References


Cited sources

*


External links


American Coot – ''Fulica americana''
– USGS Patuxent Bird Identification InfoCenter.

– Cornell Lab of Ornithology * * {{Taxonbar, from=Q470016
American coot The American coot (''Fulica americana''), also known as a mud hen or pouldeau, is a bird of the family Rallidae. Though commonly mistaken for ducks, American coots are only distantly related to ducks, belonging to a separate order. Unlike the w ...
American Coot The American coot (''Fulica americana''), also known as a mud hen or pouldeau, is a bird of the family Rallidae. Though commonly mistaken for ducks, American coots are only distantly related to ducks, belonging to a separate order. Unlike the w ...
Birds of North America Birds of the United States Birds of the Caribbean Birds of Hispaniola Birds of the Dominican Republic Birds of Haiti Birds of Central America
American coot The American coot (''Fulica americana''), also known as a mud hen or pouldeau, is a bird of the family Rallidae. Though commonly mistaken for ducks, American coots are only distantly related to ducks, belonging to a separate order. Unlike the w ...
Taxa named by Johann Friedrich Gmelin