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The tricolored bat (''Perimyotis subflavus'') is a species of
microbat Microbats constitute the suborder Microchiroptera within the order Chiroptera (bats). Bats have long been differentiated into Megachiroptera (megabats) and Microchiroptera, based on their size, the use of echolocation by the Microchiroptera a ...
native to eastern North America. Formerly known as the eastern pipistrelle, based on the incorrect belief that it was closely related to European ''
Pipistrellus ''Pipistrellus'' is a genus of bats in the family Vespertilionidae and subfamily Vespertilioninae. The name of the genus is derived from the Italian language, Italian word , meaning "bat" (from Latin "bird of evening, bat"). The size of the ge ...
'' species, the closest known relative of the tricolored bat is now recognized as the
canyon bat The canyon bat (''Parastrellus hesperus''), also known as the western pipistrelle, is a species of vesper bat. It is found in Mexico and in the western United States. The species has historically been placed in the genus ''Pipistrellus ''Pip ...
. Its common name "tricolored bat" derives from the coloration of the hairs on its back, which have three distinct color bands. It is the smallest bat species in the eastern and midwestern US, with individuals weighing only . This species mates in the fall before hibernation, though due to sperm storage, females do not become pregnant until the spring. Young are born helpless, though rapidly develop, flying and foraging for themselves by four weeks old. It has a relatively long lifespan, and can live nearly fifteen years. In the summer, females
roost Roost may refer to: Animal resting * Roosting, resting behavior of birds * Communal roosting, a behavior of some birds and other animals * Monarch butterfly roosts, communal resting sites in monarch butterfly migration * Bat roost, a list of pla ...
in small groups and males roost solitarily in tree foliage or beard lichen. It eats a diverse array of insects, foraging with a slow, erratic flight and navigating via echolocation. Though once considered one of the most common bat species in its range, its populations have declined rapidly since 2006 with the introduction of the fungal disease
white-nose syndrome White-nose syndrome (WNS) is a fungal disease in North American bats which has resulted in the dramatic decrease of the bat population in the United States and Canada, reportedly killing millions as of 2018. The condition is named for a distincti ...
. It was listed as an endangered species in 2012 in Canada, and has been petitioned for inclusion on the US endangered species list. Along with the
silver-haired bat The silver-haired bat (''Lasionycteris noctivagans'') is a solitary migratory species of vesper bat in the family Vespertilionidae and the only member of the genus ''Lasionycteris''. Etymology The species name translates as night-wandering, ref ...
, the tricolored bat is one of two bat species whose
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. Early symptoms can include fever and tingling at the site of exposure. These symptoms are followed by one or more of the following symptoms: nausea, vomiting, vi ...
variants have most frequently been implicated in human rabies deaths in the US, with sixteen deaths from 1958–2000.


Taxonomy

The tricolored bat was described as a new species in 1832 by French zoologist
Frédéric Cuvier Georges-Frédéric Cuvier (28 June 1773 – 24 July 1838) was a French zoologist and paleontologist. He was the younger brother of noted naturalist and zoologist Georges Cuvier. Career Frederic was the head keeper of the menagerie at the Muséu ...
, who placed it in the genus ''
Vespertilio ''Vespertilio'' is a genus of bats in the family Vespertilionidae. The common name for this family is vesper bats, which is a better-known classification than ''Vespertilio''. They are also known as frosted bats. Species within the genus ''Vespe ...
'' with a scientific name of ''Vespertilio subflavus''. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
had been collected in the US state of
Georgia Georgia most commonly refers to: * Georgia (country), a country in the Caucasus region of Eurasia * Georgia (U.S. state), a state in the Southeast United States Georgia may also refer to: Places Historical states and entities * Related to the ...
by American naturalist
John Eatton Le Conte John Eatton Le Conte, Jr. (sometimes John Eatton LeConte or John Eaton Leconte) (February 22, 1784 – November 21, 1860) was an American naturalist. He was born near Shrewsbury, New Jersey, the son of John Eatton Le Conte and Jane Sloane ...
. In 1897, American zoologist Gerrit Smith Miller Jr. placed it in the genus ''
Pipistrellus ''Pipistrellus'' is a genus of bats in the family Vespertilionidae and subfamily Vespertilioninae. The name of the genus is derived from the Italian language, Italian word , meaning "bat" (from Latin "bird of evening, bat"). The size of the ge ...
'' for the first time as a result of its physical similarities to other members of the genus. Its common name was thus the "eastern pipistrelle". In 1984 its status was reviewed, with H. Menu concluding that it was more similar in appearance to species in the genus ''
Myotis The mouse-eared bats or myotises are a diverse and widespread genus (''Myotis'') of bats within the family Vespertilionidae. The noun "''myotis''" itself is a New Latin construction, from the Greek "''muós'' (meaning "mouse") and "''oûs''" (mea ...
'' rather than ''Pipistrellus'', erecting a new genus for the
taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
: ''Perimyotis''. The name "''Perimyotis''" means "around ''Myotis''", alluding to its similarities with the genus. This classification was not immediately accepted, however, with a 1985 study instead grouping it and the
canyon bat The canyon bat (''Parastrellus hesperus''), also known as the western pipistrelle, is a species of vesper bat. It is found in Mexico and in the western United States. The species has historically been placed in the genus ''Pipistrellus ''Pip ...
into the genus ''
Parastrellus The canyon bat (''Parastrellus hesperus''), also known as the western pipistrelle, is a species of vesper bat. It is found in Mexico and in the western United States. The species has historically been placed in the genus ''Pipistrellus ''Pip ...
'' and a 1987 publication maintaining that the tricolored bat was part of the ''Pipistrellus'' genus, though they listed ''Perimyotis'' as one of seven
subgenera In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the International Code of Zoological Nomenclature, a subgeneric name can be used independently or included in a species name, in parentheses, placed between t ...
. However, in 2003, a genetic study concluded that the tricolored bat was distinct from ''Pipistrellus'' species (and the canyon bat), confirming the validity of the genus ''Perimyotis''. Usage of the scientific name ''Perimyotis subflavus'' is widely accepted as of 2019. Its common name was changed from "eastern pipistrelle" to "tricolored bat" to reflect its revised classification. A 2010 study found that the
sister taxon In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
of the tricolored bat is the canyon bat, with the authors suggesting that these two genera form a
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...
, though declining to name and formally describe the tribe. Instead, they referred to it as the "perimyotine group", which they gave as the most basal member of a clade that also included the following tribes: Nycticeiini,
Eptesicini Eptescini is a tribe of bats in the family Vespertilionidae. This tribe has a cosmopolitan distribution. Species Species in the tribe include: * Genus ''Arielulus'' ** Bronze sprite, ''Arielulus circumdatus'' ** Coppery sprite, ''Arielulus cu ...
, Vespertilioni, and another unnamed tribe referred to as the "hysugine group" (including ''
Chalinolobus ''Chalinolobus'' is a genus of bats, commonly known as pied, wattled, or long-tailed bats. They have fleshy lobes at the bottom edge of their ears and on their lower lips. The bats otherwise classified in the genus ''Glauconycteris'' are included ...
'', ''
Hypsugo The genus ''Hypsugo'' contains many bats referred to as pipistrelles or pipistrelle bats (not to be confused with the genus ''Pipistrellus''). They belong to the family Vespertilionidae or vesper bats. They are primarily found throughout Asia, th ...
'', ''
Laephotis ''Laephotis'' (known as the African long-eared bat) is a genus of bats in the family Vespertilionidae. Species within this genus are: * Angolan long-eared bat (''Laephotis angolensis'') * Botswanan long-eared bat (''Laephotis botswanae'') * Cape ...
'', ''
Neoromicia ''Neoromicia'' is a genus of vesper bat in the family Vespertilionidae. It contains the following species: * Anchieta's serotine (''Neoromicia anchietae'') * Kirindy serotine (''Neoromicia bemainty'') * Yellow serotine (''Neoromicia flavescen ...
'', '' Nycticeinops'', ''
Tylonycteris The bamboo bats are genus of vesper bats Vespertilionidae is a family of microbats, of the order Chiroptera, flying, insect-eating mammals variously described as the common, vesper, or simple nosed bats. The vespertilionid family is the most d ...
'', and ''
Vespadelus ''Vespadelus'' is a genus of Australian bats in the family Vespertilionidae. The genus name was first published by Ellis Le Geyt Troughton and Tom Iredale in 1934, but without an adequate description; the type was cited as ''Scotophilus pumilus' ...
''). The tribe was later described as
Perimyotini Perimyotini is a tribe of bats in the family Vespertilionidae. It contains two species of bats found in North America, each in their own monotypic genus. Although this name is already in use by taxonomic authorities, such as the ''Handbook of the ...
. Its
species name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
"''subflavus''" means "a little less than yellow", meaning yellowish or pale yellow. Four
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
are recognized: *''P. s. clarus'' (Baker, 1954) *''P. s. floridanus'' (Davis, 1957) *''P. s. subflavus'' (Cuvier, 1832) *''P. s. veraecrucis'' (Ward, 1891)


Description

The tricolored bat has blond fur that is distinctly tricolored on its back. Individual hairs tricolored: dark gray at the base, buffy (yellowish brown) in the middle, and brown or reddish brown at the tip. It is the smallest bat species found in the eastern and midwestern US. Individuals have a forearm length of and weigh . Its forearms are reddish, contrasting sharply with the black membranes of its wings. Its wingspan is . It has a
dental formula Dentition pertains to the development of teeth and their arrangement in the mouth. In particular, it is the characteristic arrangement, kind, and number of teeth in a given species at a given age. That is, the number, type, and morpho-physiolo ...
of for a total of 34 teeth.


Biology and ecology


Reproduction and life cycle

The tricolored bat is a
seasonal breeder Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and cha ...
, with
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 ...
(mating) occurring in the fall before hibernation.
Ovulation Ovulation is the release of eggs from the ovaries. In women, this event occurs when the ovarian follicles rupture and release the secondary oocyte ovarian cells. After ovulation, during the luteal phase, the egg will be available to be fertilized ...
does not occur until the spring, however, and females store the males'
sperm Sperm is the male reproductive cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm with a tail known as a flagellum, whi ...
in their
uterus The uterus (from Latin ''uterus'', plural ''uteri'') or womb () is the organ in the reproductive system of most female mammals, including humans that accommodates the embryonic and fetal development of one or more embryos until birth. The uter ...
es through the winter.
Gestation Gestation is the period of development during the carrying of an embryo, and later fetus, inside viviparous animals (the embryo develops within the parent). It is typical for mammals, but also occurs for some non-mammals. Mammals during pregna ...
(pregnancy) length is about forty-four days, with females giving birth in June or July. The litter size is typically two individuals. As newborns, the combined mass of twin pups can be as great as 58% of the mother's postpartum mass. At birth, the young lack fur and their eyes are closed. The mother leaves the offspring behind at the roost while she forages at night. Offspring develop rapidly, beginning to fly at three weeks old. By four weeks old, they are foraging for themselves. Young do not reach sexual maturity in their first fall; they do not breed until their second fall. Individuals can live at least 14.8 years in the wild; juveniles have higher mortality rates than adults, and females have higher mortality rates than males.


Behavior

During the summer, the tricolored bat will roost in tree foliage or buildings, with females alone or in maternity colonies of up to thirty individuals. Trees used for this purpose include
oak An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably ''L ...
,
maple ''Acer'' () is a genus of trees and shrubs commonly known as maples. The genus is placed in the family Sapindaceae.Stevens, P. F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, June 2008 nd more or less continuously updated since http ...
, the
eastern cottonwood ''Populus deltoides'', the eastern cottonwood or necklace poplar, is a cottonwood poplar native to North America, growing throughout the eastern, central, and southwestern United States as well as the southern Canadian prairies, the southernm ...
, and
American tulip tree ''Liriodendron tulipifera''—known as the tulip tree, American tulip tree, tulipwood, tuliptree, tulip poplar, whitewood, fiddletree, and yellow-poplar—is the North American representative of the two-species genus ''Liriodendron'' (the other ...
. Males are solitary and do not form colonies. In
Nova Scotia Nova Scotia ( ; ; ) is one of the thirteen provinces and territories of Canada. It is one of the three Maritime provinces and one of the four Atlantic provinces. Nova Scotia is Latin for "New Scotland". Most of the population are native Eng ...
, researchers discovered nearly one hundred roosts of this species, finding that all sampled individuals were roosting not in tree foliage, but rather in a species of beard lichen, '' Usnea trichodea''. This was thought to be the first documentation of a bat using beard lichen as a roosting substrate. It was speculated that the tricolored bat may use the lichen to ward off parasites.
Usnic acid Usnic acid is a naturally occurring dibenzofuran derivative found in several lichen species with the formula C18H16O7. It was first isolated by German scientist W. Knop in 1844 and first synthesized between 1933-1937 by Curd and Robertson. Usnic a ...
, which has anti-insect and anti-bacterial properties, naturally occurs in beard lichens, and no
ectoparasites Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
(external parasites) have been documented on the tricolored bat in Nova Scotia. In the winter, the tricolored bat
hibernate Hibernation is a state of minimal activity and metabolic depression undergone by some animal species. Hibernation is a seasonal heterothermy characterized by low body-temperature, slow breathing and heart-rate, and low metabolic rate. It most ...
s in mines, caves, or other human structures. A years-long study of a mine in the US state of Nebraska found that in the summer, the mine was mostly occupied by males. Total number of tricolored bats using the mine at one time ranged from zero to forty-three. Bats were in a state of
torpor Torpor is a state of decreased physiological activity in an animal, usually marked by a reduced body temperature and metabolic rate. Torpor enables animals to survive periods of reduced food availability. The term "torpor" can refer to the time ...
from November to April. More bats used the mine as a roost in the winter than in the summer. In another abandoned mine in the US state of Indiana, researchers found that the tricolored bat roosts solitarily during hibernation for the majority of the time (96.8% of observations were singletons). During hibernation, males lose an average of , while females lose an average of .


Diet and foraging

The tricolored bat is insectivorous, consuming small prey of in length. Prey taxa include
mosquito Mosquitoes (or mosquitos) are members of a group of almost 3,600 species of small flies within the family Culicidae (from the Latin ''culex'' meaning " gnat"). The word "mosquito" (formed by ''mosca'' and diminutive ''-ito'') is Spanish for "li ...
es,
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s,
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s,
moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
s, and
cicada The cicadas () are a superfamily, the Cicadoidea, of insects in the order Hemiptera (true bugs). They are in the suborder Auchenorrhyncha, along with smaller jumping bugs such as leafhoppers and froghoppers. The superfamily is divided into two ...
s. It forages with slow, erratic flight in areas near water or forest edges. It is one of the first bat species to begin foraging each night. It navigates and searches for prey via echolocation (sonar). Its echolocation calls are at a high frequency; visual renderings of the calls show a distinctive hook-shaped call profile, with the lowest frequencies (bottom of the hook shape) at 42 k Hz.


Predators, parasites, and disease

The tricolored bat's natural predators include the
northern leopard frog ''Lithobates pipiens''Integrated Taxonomic Information System nternet2012''Lithobates pipiens'' pdated 2012 Sept; cited 2012 Dec 26Available from: www.itis.gov/ or ''Rana pipiens'', commonly known as the northern leopard frog, is a species of le ...
, birds of prey,
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, snakes,
skunk Skunks are mammals in the family Mephitidae. They are known for their ability to spray a liquid with a strong, unpleasant scent from their anal glands. Different species of skunk vary in appearance from black-and-white to brown, cream or ginge ...
s, and
prairie vole The prairie vole (''Microtus ochrogaster'') is a small vole found in central North America. The vole has long, coarse grayish-brown fur on the upper portion of the body and yellowish fur on the lower portion of the body. It has short ears and a ...
s. Feral cats will also catch and kill them as they leave their hibernacula (cave or mine used for hibernation). There are also records of it being attacked by the
hoary bat The hoary bat (''Lasiurus cinereus'') is a species of bat in the vesper bat family, Vespertilionidae. It lives throughout most of North America (possibly including Hawaii, although this is disputed). Taxonomy The hoary bat was described as a ...
. The tricolored bat is the host to several species of endoparasites (internal parasites) and ectoparasites. Oocysts (spores) of an ''
Eimeria ''Eimeria'' is a genus of apicomplexan parasites that includes various species capable of causing the disease coccidiosis in animals such as cattle, poultry and smaller ruminants including sheep and goats. ''Eimeria'' species are considered to be ...
'' species ('' Eimeria heidti'' ) have been recovered from its
guano Guano (Spanish from qu, wanu) is the accumulated excrement of seabirds or bats. As a manure, guano is a highly effective fertilizer due to the high content of nitrogen, phosphate, and potassium, all key nutrients essential for plant growth. G ...
(feces). The tricolored bat has experienced severe population decline as a result of the fungal disease
white-nose syndrome White-nose syndrome (WNS) is a fungal disease in North American bats which has resulted in the dramatic decrease of the bat population in the United States and Canada, reportedly killing millions as of 2018. The condition is named for a distincti ...
, which arrived in the US in 2006, with losses of 70% and greater detected in multiple US states. The disease kills bats by colonizing their skin during the winter, causing them to arouse from torpor and burn through their limited fat reserves. Though its population experienced dramatic reduction, subsequent studies have found that their numbers may be stabilizing, though hibernacula where many individuals once roosted may only host fewer than five bats, or even one solitarily.


Range and habitat

The tricolored bat is common throughout eastern North America, with the southern extent of its range in Central America and the northern extent in southern Canada. Its range has expanded since the 1980s, with westward expansion reaching the US states of Colorado, New Mexico, South Dakota, and Wyoming. Additionally, its range started to include the
Great Lakes Basin The Great Lakes Basin consists of the Great Lakes and the surrounding lands of the states of Illinois, Indiana, Michigan, Minnesota, New York, Ohio, Pennsylvania, and Wisconsin in the United States, and the province of Ontario in Canada, whose dir ...
.


Conservation

As of 2018, it is listed as a
vulnerable species A vulnerable species is a species which has been Conservation status, categorized by the International Union for Conservation of Nature as being threatened species, threatened with extinction unless the circumstances that are threatened species, ...
by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
. In 2008, however, it was considered a
least-concern species A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
, which is the IUCN's lowest conservation priority. In the time between the two assessments, the fungal disease white-nose syndrome became widespread in eastern North America, severely impacting the tricolored bat. Historically, it was one of the most common bat species of eastern North America. After an emergency assessment, the tricolored bat is considered an endangered species under the
Committee on the Status of Endangered Wildlife in Canada The Committee on the Status of Endangered Wildlife in Canada (COSEWIC, French: Comité sur la situation des espèces en péril au Canada, COSEPAC) is an independent committee of wildlife experts and scientists whose "raison d'être is to identify s ...
as of 2012. In 2016, the
Center for Biological Diversity The Center for Biological Diversity is a nonprofit membership organization known for its work protecting endangered species through legal action, scientific petitions, creative media and grassroots activism. It was founded in 1989 by Kieran Suckl ...
and
Defenders of Wildlife Defenders of Wildlife is a 501(c)(3) non-profit conservation organization based in the United States. It works to protect all native animals and plants throughout North America in their natural communities. Background Defenders of Wildlife is a n ...
submitted a petition to the
United States Fish and Wildlife Service The United States Fish and Wildlife Service (USFWS or FWS) is an agency within the United States Department of the Interior dedicated to the management of fish, wildlife, and natural habitats. The mission of the agency is "working with othe ...
(USFWS) to list the tricolored bat under the
Endangered Species Act of 1973 The Endangered Species Act of 1973 (ESA or "The Act"; 16 U.S.C. § 1531 et seq.) is the primary law in the United States for protecting imperiled species. Designed to protect critically imperiled species from extinction as a "consequence of ec ...
. The USFWS determined that the petition demonstrated that listing the species as "threatened" or "endangered" may be warranted. In September 2022, the USFWS announced it would protect the bat as endangered under the Endangered Species Act, following a lawsuit from the Center for Biological Diversity.


Relationship to people

Along with the
silver-haired bat The silver-haired bat (''Lasionycteris noctivagans'') is a solitary migratory species of vesper bat in the family Vespertilionidae and the only member of the genus ''Lasionycteris''. Etymology The species name translates as night-wandering, ref ...
, the tricolored bat is one of two species of bat whose rabies variants are responsible for 70% of human rabies cases from bats in the US. Additionally, the rabies virus variants associated with these two species caused 75% of
cryptic rabies Cryptic rabies refers to infection from unrecognized exposure to rabies virus. It is often phylogenetically traced to bats. It is most often seen in the southern United States. Silver-haired bats (''Lasionycteris noctivagans'') and tricolored ba ...
deaths in the US (rabies cases where the exposure route is unclear, and no bite history exists). From 1958–2000, sixteen people in the US died from the rabies variant associated with tricolored bats, or 46% of all nationwide indigenous (not acquired elsewhere) rabies fatalities. The tricolored bat is infrequently encountered by humans and submitted for rabies testing, with only thirty-one individuals tested for rabies in the US in 2017. Of those, one (3.4%) tested positive for the virus.


See also

*
Bats of Canada There are eighteen indigenous species of bats in Canada, which are found in many parts of the country. They are insectivores, and are prey to falcons, hawks, owls, snakes, cats, and raccoons.Health Canada Species The little brown bat is the mos ...
*
Bats of the United States Most of the many bat species found in the United States are insectivorous except for three flower eating species that migrate from Mexico and one that inhabits the Florida Keys. __TOC__ Species Bats belong to the biological order of Chiropte ...


References


External links

* * {{Authority control Vesper bats Bats of Canada Bats of the United States Mammals of Mexico Bats of Central America Fauna of the Eastern United States Fauna of the Great Lakes region (North America) Mammals described in 1832 Taxa named by Frédéric Cuvier