Tennessee Naturalist Program

Tennessee Mammals Creatures of Habitat

Enhanced Study Guide

12/2015 Tennessee Naturalist Program www.tnnaturalist.org

Inspiring the desire to learn and share Tennessee’s nature

These study guides are designed to reflect and reinforce the Tennessee Naturalist Program’s course curriculum outline, developed and approved by the TNP Board of Directors, for use by TNP instructors to plan and organize classroom discussion and fieldwork components and by students as a meaningful resource to review and enhance class instruction.

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2 Tennessee Mammals Creatures of Habitat

Objectives Present an overview of mammals including characteristics particular to this class of animals and the different groups of mammals found in Tennessee. Explore their behavior, physiology, and ecology, relating these to habitat needs, environmental adaptations, and ecosystem roles, including human interactions.

Time Minimum 4 hours – 2 in class, 2 in field

Suggested Materials ( * recommended but not required, ** TNP flash drive) • Mammals of North America, Fourth Edition (Peterson Field Guides), Fiona Reid * • Mammals of North America, Second Edition, (Princeton Field Guides), Roland W. Keys and Don E. Wilson • Tennessee Mammals Enhanced Study Guide, TNP ** • TWRA Bone Box

Expected Outcomes Students will gain a basic understanding of 1. the diversity and distribution of mammals in Tennessee, including rare species 2. the major groups of mammals and their systematic relationships 3. the distinguishing characteristics of all mammals and general characteristics of each major group 4. mammal behavior - breeding, feeding, shelter, hibernation, and range/territory 5. mammal physiology and morphology - environmental adaptations, functions, species identification 6. mammal ecology - habitat needs, ecosystem roles 7. animal tracks and signs 8. animal and human interactions

3 Mammals Curriculum Outline

I. Mammals A. Class characteristics

II. Mammals of Tennessee A. Diversity and distribution B. Common mammal Orders, their relationships and differences 1. opossums 2. moles and shrews 3. bats 4. rabbits 5. rodents 6. carnivores 7. hoofed mammals C. Nonnative species and their impacts D. Rare species and their conservation concerns

III. Mammal Behavior A. Feeding B. Breeding C. Nesting, territory, home range D. Hibernation and winter survival E. Defensive strategies

IV. Mammal Physiology and Morphology A. Environmental adaptations B. Functions C. Identification 1. skins, pelts 2. skulls 3. teeth

V. Mammal Ecology A. Habitat variety and ‘home’ needs B. Ecological role 1. food web 2. predators 3. keystone species C. Diurnal, crepuscular, nocturnal

4 VI. Observations in the Wild A. Tracks B. Signs C. Scat

VII. Wild Mammals and Human Interactions A. Habitat loss, fragmentation, and encroachment B. Management concerns and strategies 1. population monitoring and measuring 2. trapping and tagging C. Hurt, captured or orphaned animals

VIII. Resources A. Publications B. Organizations C. Internet

5 I. Mammals Introduction

There are over 5,400 species of mammals worldwide. Of these, nearly 25 percent are threatened and at risk of extinction. The United States has over 410 species and 80 are on the endangered species list. Tennessee’s mammals (current and historical) number 86 (Kennedy et al. 2012), representing a diversity from northern species in the mountains to western species.

Common mammal characteristics As a class (Mammalia), mammals exhibit several common characteristics that separate them from other vertebrate animal groups possessing an internal skeleton with a backbone. All mammals share the three asterisked (*) characteristics; exceptions may be found for the others.

• hair or fur on the body at some point during life * (most have hair their entire lives) • generally endothermic, producing body heat internally, and homeothermic, maintaining constant temperature • give birth to live young (egg-laying exceptions not found in Tennessee) • milk producing mammary glands to feed young * • three middle ear bones (hammer, anvil, stirrup) * In addition, most mammals have a single bone lower jaw, four-chambered heart, highly developed brain, secondary palate separating air and food passages in the mouth, muscular diaphragm between the thoracic and abdominal cavities, internal fertilization, and separate sexes with embryo sex determined by the presence of a Y or two X chromosomes.

Differences Differences among mammal species in physical morphology, physiology, behavior, ecology, etc., form the basis for identification.

• anatomy -- hard (skeleton, particularly skulls) and soft tissue • teeth -- differentiated tooth types; also give clues to diet and habitat • embryonic development -- placental, marsupial, and egg laying (not in Tennessee) • morphological extremes -- diversity of forms, bodies designed for flying, running, hopping, burrowing, climbing, gliding, or swimming and vast size differences • active time of day -- nocturnal (night), diurnal (day), or crepuscular (dawn and dusk)

6 Habitat Tennessee has a great diversity of land types. Therefore, habitats also vary widely. Within these habitats, different mammal species find homes suited to their life history strategies. There are four main home types with local examples of each.

• arboreal -- squirrels, golden mouse • aquatic -- beaver, muskrat • home seekers (look for shelter in caves, logs or manmade structures) -- bats, bobcat • homeless (bed down in a different place each night) -- deer, elk Each species is adapted to specific habitats. Problems arise when habitats are altered, whether by loss, fragmentation, disease, or the introduction of exotic species.

Non-native species in Tennessee • roof rat • brown rat (Norway rat) • house mouse • nutria or coypu • domestic dog and domestic cat • wild hog (European boar)

Extirpated native species in Tennessee • snowshoe hare • North American porcupine • gray wolf • red wolf • cougar

Extirpated and reintroduced native species in Tennessee • fisher • elk • bison (captive herds, not free roaming)

7 Tennessee Mammals: Quick Facts

Virginia Opossum (1 sp.) Order Didelphimorphia only marsupial in TN (related to the kangaroo of Australia) pouch runs front to back, bears up to 13 young that crawl from vagina to pouch coarse, grizzled grayish fur, naked tail and ears, pink toes and nose opposable thumbs on hind feet relatively disease free 50 teeth, more than any other mammal in TN sometimes ‘play possum’ (feign death) when threatened young can hang by tail, adults are a bit too heavy ears and tail can get frostbitten range moving north

Shrews (9 spp.) and Moles (3 spp.) Order Soricomorpha Shrews have musk glands, stink voracious predator with insatiable appetite, must eat 2-3 times weight daily insectivores, also eat smaller mammals, aggressive attackers brown-tipped teeth and tiny eyes shrews are hard to identify to species a couple of species echolocate food one is poisonous with neurotoxin toward prey some species prefer grassland habitats, some woodland pygmy shrew is one of the smallest mammals in the world East Tennessee has more species of shrews

Moles: eastern mole statewide, star-nosed and hairy-tailed moles limited and uncommon hair stands on end, unidirectional to repel dirt regardless of animals direction source of Victorian powder puffs tunnels using broad forepaws, white teeth voracious eaters, insect and worm fossorial (spend most time underground) incisors small, tiny eyes, no external ears skulls distinctive among three species of moles predators include coyotes, hawks, owls, foxes, and house cats

8 Bats (15 spp.) Order Chiroptera (hand-wing) evening bats -- tragus (small pointed structure, part of external ear) ID character TN has small bats, larger fruit-eating bats are not in TN eastern red bat, perhaps most common species in TN eastern pipistrelle, smallest, often in houses, can carry rabies less than 1% (0.5 %) of the bat population have rabies echolocate insects, shut down ears during cry and open again to hear echo 8-15 cries per second normally, sudden movements 150-200 cries/sec nest in caves, trees (under bark), buildings most go into torpor in winter, Little Brown Bat is a true hibernator two listed endangered species bats are sensitive to their environment White Nose Syndrome, a fungal disease of cave bats, results in high mortality

Armadillo (1 sp.) Order Cingulata Dasypus (hairy foot) novemcinctus (nine banded) non-native, but relatively benign, moving north and east of historic range back is covered in skin plates born with plates, harden as they mature peg teeth, eat insects, ants, and termites (including fire ants) claws for digging, reduced teeth in reproduction, the egg splits to form four genetically identical young move fast and jump when scared, which is why cars kill so many called ‘possum on the half shell’ coyotes are predators carry leprosy and are used in research

Rabbit (4 spp.) Order Lagomorpha eastern cottontail, swamp rabbit (W TN), Appalachian cottontail (Blue Ridge) snowshoe hare extirpated four incisors are clustered in two pairs, one behind the other skull differences separate species in identification young are altricial, born blind and hairless [Hares are precocial] swamp rabbits poop on logs in wetlands eat poop to glean all nutrition from food (coprophagia) relies on early successional habitats, weedy, meadows “rabbitat” population declines can occur from habitat loss due to land use changes

9 Rodents (31 spp.) Order Rodentia rodents have long incisors that never stop growing, must chew to wear them down Squirrels (7 spp.) tree squirrels (5 spp.) -- gray squirrel (most common), fox squirrel (largest, dark color), red squirrel (“boomer” Blue Ridge), flying squirrels (southern and northern species, latter only in spruce-fir forests) population of albino squirrels lives in Kenton, TN (Gibson County in West TN) chipmunk or ground squirrel (western prairie dogs are ground squirrels too) woodchuck or groundhog, also called whistle pig or land beaver (a marmot) burrows in the ground woodchucks are true hibernators

Beaver aquatic with waterproof fur, nictating eye membrane, and ear, nose and throat valves chestnut brown fur, flat scaly tail, and webbed hind feet build lodges and modify landscape for own purpose live in family units eat cambium layer of trees, use sticks and small branches for lodge eat poop to get all nutrition, coprophagia castor sacs fluid (castoreum) used in expensive perfumes for leathery scent

Native mice, rats, voles, lemming, muskrat, jumping mice (18 spp.) have lighter bellies, seed eaters wood rat (largest) to harvest mouse (smallest) woodland vole has a short tail, is a pest in yards deer, cotton, and white-footed mice have big eyes and big ears hispid cotton rat is common in old fields, coyotes love them golden mouse builds its nest up high, little condo with roof on it muskrat is aquatic and along with beaver among the largest of our rodents jumping mice have long tails and long hind feet, and broad dark stripe on back one of the few mammals in TN that truly hibernate

Non-native rodents (4 spp.) roof rat (black rat) and house mouse (beady eyes, small ears, scaly tail) are all gray with uniform color Norway rat (brown rat) is found all over world nutria (coypu) is aquatic, looks like a beaver or giant muskrat, prolific and invasive; harms wetland marshes affecting waterfowl and muskrat habitat; native to South America

Porcupines have been extirpated from Tennessee. There are historical reports of the species from early settlement before statehood.

10 Carnivores (18 spp.) Order Carnivora Canine (6 spp.) tails -- dogs’ are often held up; coyote tails held down; fox tails held straight out canine claws are not retractable, paw prints will have claw marks

red and gray wolves have been extirpated from TN attempts to reintroduce red wolves to the Smokies failed gray fox is size of bobcat (15 lbs), grayish body with reddish sides and legs black patches on sides of muzzle, black tip and upper stripe on tail can climb trees, likes deeper woods very susceptible to canine distemper red fox is an edge species reddish fur with black legs, feet, ears, white tip on tail shed fur and become mangy in summer populations tend to dip in the presence of coyotes susceptible to canine distemper coyotes moved into TN by 1960s, good populations established by 1980s population density-dependent reproductive pattern coyote is an edge species but adaptable to various habitats need skull to ID, especially upper jaw solitary, not communal with alpha males like wolves mate in pairs, raise young who disperse following year 30 pounds is typical various fur coloration blonde, off-white, gray, rusty brown primarily carnivores but also eat vegetable matter, basically omnivorous dog skulls have a pronounced dip to the snout and wider palate coyotes skulls follow a straighter path to the snout with a skinny palate coyotes don’t like to mate with dogs, TN doesn’t have many “coydogs” coyotes can get parvovirus

Black Bear omnivorous -- fruits, nuts, insects, carrion tolerant of different environments historic range was statewide, current range is expanding from mountains, with individuals found around state Cumberland Plateau is next site for stable populations mate in July, give birth in mid winter cubs often born in pairs, remain with mother 18 months don’t truly hibernate, just long periods of sleep may leave den briefly during warm spells in winter or if disturbed

11 Raccoon related to bears ringed tails used historically for food, fur, other items now more of a problem species around human habitation rabies in raccoons is a concern, inoculation efforts create an immunity barrier

Mustelids (5 spp.) -- river otter, weasels, mink, fisher not often seen, very secretive odorous glands river otters are aquatic, minks are somewhat aquatic weasels and fishers are terrestrial otters may pose problems around fish ponds or hatcheries fishers reintroduced on Cumberland Plateau, success is not certain weasels are ferocious, mean

Skunks (2 spp.) spotted skunks (employ handstand pose as threat) striped skunks (spray from raised tail) striped skunk genus Mephitic means bad odor striped skunks may not always have distinctive stripes, might just have white spot on head and black body spotted has distinct behavior differences, fast and nervous disposition can get canine distemper and rabies and therefore cannot be translocated

Cats (3 spp.) cats have retractable claws, paw prints will not have claw marks cougars extirpated western species range expanding into the Midwest with recent sightings in KY/TN bobcats only native wild cat in TN actually a lynx, often with a spotted coat very secretive but common formidable predator, can kill deer

12 Ungulates (4 spp.) Order Artiodactyla -- even-toed, cloven hooves Wild hog (European boar) introduced, invasive, destructive dark fur, long nose, leaner animal with tusks damage soil (rototiller) and plants tusk marks on trees, dirt wallows very smart, prolific reproduction, impossible to eradicate produce a baseball-sized knot of poop illegal to transport black/white hybrid pigs, they revert to wild species generationally hybrids introducing diseases TWRA used to promote them as big-game species but doesn’t anymore used for hunting, people move them and worsen the situation TWRA needs backing of agriculture community to address problem

Elk reestablished in the Smokies and northwest of Knoxville bugle in October bulls shed antlers in early spring

White-tailed deer bucks typically shed antlers in winter antlers are fastest growing tissue known an antler consists of one main beam with points coming off it rodents chew on shed antlers popular for hunting

Bison historically occurred in pre-settlement prairie habitats in TN reintroductions are captive populations, not free-roaming

Fallow deer, Eurasian exotic on hunting preserves sometimes escape are smaller sized, spotted as adults and juveniles, with straighter back

13 II. Annotated Checklist of the Mammals of Tennessee

Kennedy, Michael L.; Kennedy, Phyllis K.; Warr, Edward L.; Wyatt, Robert L. "Pete" June 1, 2012 Journal of the Tennessee Academy of Science [Reprinted here with permission of the authors.]

Abstract This investigation presents an annotated checklist of the living mammals reported to occur in Tennessee as well as recently extirpated species and reintroduced taxa. Data were obtained from field collections, literature records, and selected mammal collections in North America. Species included in the checklist represent 8 orders, 23 families, and 86 species. The work provides information relating to distribution, habitat, and conservation status of each species and should be useful in planning management and conservation programs in Tennessee.

Introduction As pointed out by Kennedy and Harvey (1980), information relating to mammals in Tennessee is scattered. However, several works have contributed to our understanding of the mammalian fauna of the state. For examples, see Rhoads (1896), Komarek and Komarek (1938), Kellogg (1939), Calhoun (1941), Goodpaster and Hoffmeister (1952), Howell and Conway (1952), Conway and Howell (1953), Beasley and Severinghaus (1973), Severinghaus and Beasley (1973), Graves and Harvey (1974), Smith et al. (1974), Hall (1981), Kennedy et al. (1984), Kennedy (1991), and Linzey (1995). Overall, the species richness of mammals in Tennessee is high, yet it has not been summarized in many years. Therefore, the purpose of this investigation was to review available information concerning mammals in Tennessee and provide an annotated checklist of the species occurring within the state. Our checklist includes species currently residing within the state's boundaries that were present when Europeans first arrived, as well as introduced, extirpated (including extirpated and reintroduced), and exotic species. Additionally, we provide information relating to distribution, habitat, and conservation status for each taxon; such information should be useful in planning management and conservation programs.

Materials and Methods Scientific and common names employed in the checklist are those used by Wilson and Reeder (2005). In each account, an attempt has been made to provide the reader with a brief understanding of the taxon's distribution in Tennessee, general habitat, and conservation status. Where there is an official state [Tennessee Department of Environment and Conservation (TDEC), http://www.tennessee.gov/environment/na/ or Tennessee Wildlife Resources Agency

14 (TWRA), http://www.tn.gov/twra/pdfs/endangered.pdf, http://www.tn.gov/twra/pdfs/ wildlifeinneed.pdf] or federal (United States Fish and Wildlife Service, http://www.fesgov/) listing of a taxon for protection or concern, the state listing is given first in the accounts (Conservation status) followed by federal listing. The state ranking system employed was as follows: S1--extremely rare and critically imperiled in the state (often with five or fewer occurrences) or very few remaining individuals, or because of some special condition where the species is particularly vulnerable to extirpation; S2--very rare and imperiled within the state, six to 20 occurrences, or few remaining individuals, or because of some factor(s) making it vulnerable to extirpation; S3--vulnerable, rare, and uncommon in the nation or state due to a restricted range, relatively few populations (often 80 or fewer), recent and widespread declines, or other factor(s) making it vulnerable to extirpation; S4--uncommon but not rare, and apparently secure within the state, but with cause for long-term concern due to declines or other factors; S5--demonstrably common, widespread, and secure in the state. The federal (global) ranking system used was a five-tier system ranking (G1-G5) to describe rarity, from G1 (extremely rare) to G5 (widespread). A rank range (e.g., S2-S3; G3-G4) denotes uncertainty in rarity. T# refers to a subspecific taxon rank at the global level. Endangered (state and federal levels) refers to any species or subspecies of wildlife whose prospects of survival or recruitment is in jeopardy or is likely to become so within the foreseeable future. "Deemed in Need of Management" refers to any species or subspecies of nongame wildlife which TDEC or TWRA believe should be investigated in order to develop information relating to populations, distribution, habitat needs, limiting factors, and other biological and ecological data to determine management measures necessary for their continued ability to sustain themselves successfully (analogous to Special Concern utilized in some states). References to abundance or common/uncommon in the accounts reflect our assessment of the species' status based on the number of published records and number of museum specimens available in collections and general observations made by the authors. A conservation status of none reflects that no conservation measure is in place at this time to protect the species.

The list of species in the present report is based partly on voucher specimens in the following museum collections: Austin Peay State University; Carnegie Museum of Natural History; Field Museum of Natural History; Louisiana State University, Museum of Natural Science; National Museum of Natural History; University of California, Berkeley, Museum of Vertebrate Zoology; University of Illinois, Museum of Natural History; University of Kansas; The University of Memphis Museum of Zoology; University of Michigan, Museum of Zoology; University of Oklahoma Sam Noble Museum of Natural History; University of Tennessee at Martin, University of Tennessee at Knoxville, Department of Biological Sciences. Following Beasley and Severinghaus (1973) and Severinghaus and Beasley (1973), western Tennessee is defined as that part of Tennessee west of the Tennessee River. Other regions of Tennessee (middle and eastern) follow Corgan (1976). In the list of species, references to the status of taxa as common or uncommon and to abundance of individual species are based on the number of published records and number of museum specimens available in collections and on field collections and general observations made by the authors.

15 Discussion We present an annotated checklist of mammals for Tennessee that includes native species currently residing within the state's boundaries or that resided within the state's boundaries when European man first arrived. Additionally, we have noted introduced, extirpated (including extirpated and reintroduced) and exotic species. In total, these taxa represent 8 orders, 23 families, and 86 species. This is comparable to 95 species reported for seven south-central states by Choate et al. (1994). The richness of species in Tennessee is likely associated with its diversity in physiographic regions that range from the Mississippi River Valley in western Tennessee to the Unaka Mountains in the eastern part of the state (see Miller 1974).

Kennedy (1991) noted that many species of mammals in western Tennessee had not been studied in detail, especially many of the nongame species. This also is the case for the state as a whole. Few distributional records exist for many taxa. Generally, the mammalian fauna of Tennessee needs additional investigation. Future study may add a few additional species to the list of mammals known for the state. The following species have been reported in proximity to Tennessee: Geomys bursarius (plains pocket gopher), Reithrodontomys megalotis (western harvest mouse), R. fulvescens (fulvous harvest mouse), and Taxidea taxus (badger). Specimens of T. taxus are known from western Tennessee but are thought to represent captive animals that escaped or were released; breeding populations are unknown in the state. Until statewide inventories are completed, the status and distribution of a number of taxa will remain uncertain However, the annotated checklist list presented within this report should provide managers of natural resources useful information for planning processes.

16 Species Accounts

Order Didelphimorphia -- Pouched Mammals Family Didelphidae (Opossums) Didelphis virginiana (Virginia opossum). Statewide in distribution; utilizes a variety of habitats but typically found in wetter areas associated with streams and rivers. Conservation status: none; common in most habitats.

Order Soricomorpha -- Insectivores Family Soricidae (Shrews) Sorex cinereus (Cinereus shrew). Found throughout eastern Tennessee with peripheral populations in middle Tennessee; habitat varies from moist woodlands (especially, woodlands with decaying stumps and logs associated with thick leaf litter) to open fields and marshy areas. Conservation status: Deemed in Need of Management (TDEC; TWRA). S4, G3; can be locally abundant. Sorex Iongirostris (Southeastern shrew). Statewide in distribution; occupies a number of habitat types including river floodplains, old fields, sandy river bluffs, and forest edges. Conservation status: Deemed in Need of Management (TDEC; TWRA). S4, G5; widespread in distribution but never found in large numbers. Sorex palustris (American water shrew). Known only from the mountains of extreme eastern Tennessee, most often found in association with permanent water (e.g., streams, rivers, lakes, and bogs) and habitats that include cold, fast-flowing mountain streams with overhanging banks, rocks, and roots. Conservation status: Deemed in Need of Management (TDEC; TWRA). S2, G5; uncommon throughout its distribution in the state. Sorex fumeus (Smoky shrew). Occurs in eastern and middle Tennessee; habitat is moist woodlands with decaying logs and thick leaf litter associated with moss-covered rocks in both coniferous and deciduous forest. Conservation status: Deemed in Need of Management (TDEC; TWRA). S4, G5; can be locally abundant. Sorex dispar (Long-tailed shrew). Known only from a few sites in the mountains of eastern and middle Tennessee; habitat includes cool and moist talus slopes in forested areas; some captures have been associated with rocky areas covered with mosses and lichens. Conservation status: Deemed in Need of Management (TDEC; TWRA). S2, G4; uncommon throughout its distribution in the state. Sorex hoyi (American pygmy shrew). Reported in eastern and middle Tennessee westward to the Tennessee River, occurs in a variety of habitats that include both wet and dry areas; an area with fallen trees and tree stumps in a forest clearing associated largely with beech, maple, buckeye, and hawthorn is the only reported description of habitat for the species in Tennessee.

17 Conservation status: Deemed in Need of Management (TDEC). S2, G5; uncommon throughout its distribution in the state. Blarina brevicauda (Northern short-tailed shrew). Occurs in middle and eastern Tennessee; occurs in a variety of habitats that include forest, forest and field edges, grasslands, and marshy areas. Conservation status: none; may be common in preferred habitat. Blarina carolinensis (Southern short-tailed shrew). Statewide in distribution; may occur in a variety of habitats that includes woodlands, grasslands, marshy areas, and relatively dry areas. Conservation status: none, may be locally abundant. Cryptotis parva (North American least shrew). Statewide in distribution; habitat primarily early successional areas (e.g., fields, meadows, weedy fencerows, and grassy road sides). Conservation status: none; widespread in distribution but never found in large numbers. Family Talpidae (Moles) Parascalops breweri (Hairy-tailed mole). Occurs in the mountains of eastern and middle Tennessee; habitat includes forests, pastures, gardens, lawns, and other areas with loose, well- drained soils. Conservation status: Deemed in Need of Management (TDEC; TWRA). 33, G5; uncommon throughout its distribution in the state. Scalopus aquaticus (Eastern mole). Statewide in distribution; inhabits a variety of habitats (usually associated with relatively light, well-drained soils). Conservation status: none; a common fossorial species. Condylura cristata (Star-nosed mole). Known only from eastern Tennessee; prefers damp and muddy soils associated with forests, bogs, swamps, and meadows. Conservation status: Deemed in Need of Management (TDEC; TWRA). S2, G5; uncommon throughout its distribution in the state.

Order Chiroptera -- Bats Family Vespertilionidae (Vespertilionid Bats) Myotis lucifugus (Little brown myotis). Occurs in eastern and middle Tennessee (east of the Tennessee River Valley); primarily inhabits caves and buildings. Conservation status: none; an uncommon species. Myotis austroriparius (Southeastern myotis). Distribution in the state is uncertain; appears to be primarily west of the Tennessee River Valley; habitat includes trees, caves, and buildings. Conservation status: Deemed in Need of Management (TDEC). S3, G3-G4; relatively common west of the Tennessee River. Myotis grisescens (Gray myotis). Known in eastern and middle Tennessee (east of the Tennessee River Valley); known in caves or cave-like habitats. Conservation status: Endangered Species. S2, G3.

18 Myotis septentrionalis (Northern myotis). Occurs in eastern and middle Tennessee (east of the Tennessee River Valley); habitat primarily includes caves, mines, or similar shelters. Conservation status: none; an uncommon species. Myotis sodalis (Indiana myotis). Reported in eastern and middle Tennessee (east of the Tennessee River Valley); only one specimen is known from west of the Tennessee River (Shelby County; Graves and Harvey 1974); occurs in caves, mines, or similar sites; in summer, maternity colonies are formed in hollow trees and behind loose slabs of bark in forested areas. Conservation status: Endangered Species. S1, G2. Myotis leibli (Eastern small-footed myotis). Occurs in eastern and middle Tennessee (east of the Tennessee River Valley); habitat includes caves, rock fissures, abandoned mines, bridges, and barns. Conservation status: Deemed in Need of Management (TDEC; TWRA). S2-S3, G3, relatively rare and has a spotty occurrence throughout its distribution. Lasionycteris noctivagans (Silver-haired bat). Statewide in distribution; primarily inhabits forest associated with watercourses. Conservation status: none; large numbers of this species are unknown for the state; an uncommon migratory species. Perimyotis subflavus (Tri-colored bat). Reported statewide; occurs in trees, caves, and buildings. Conservation status: none; common in preferred habitat. Eptesicus fuscus (Big brown bat). Known statewide; found in a number of habitats (e.g., old buildings, caves, hollow trees, under bridges, and deep crevices in rocky cliffs). Conservation status: none; common in suitable habitat. Lasiurus borealis (Eastern red bat). Occurs statewide; a forest species. Conservation status: none; common bat that roosts in trees. Lasiurus seminolus (Seminole bat). Statewide in distribution; habitat is primarily forest associated with watercourses. Conservation status: none; an uncommon species. Lasiurus cinereus (Hoary bat). Statewide in distribution; a tree-dwelling species. Conservation status: none; an uncommon species. Nycticeius humeralis (Evening bat). Reported statewide; habitat includes forest and old buildings. Conservation status: none; a common bat. Corynorhinus rafinesquii (Rafinesque's bigeared bat). Occurs statewide; habitat includes trees, caves, and buildings. Conservation status: Deemed in Need of Management (TDEC; TWRA). S3, G3-G4; an uncommon species. Family Molossidae (Free-tailed Bats) Tadarida brasiliensis (Brazilian free-tailed bat). In Tennessee, the species is known from only two records in middle Tennessee (a building in Montgomery Co and the Austin Peay Dunn Center); habitat for the species in the southern United States is primarily buildings but it has been recorded in hollow trees and similar habitats. Conservation status: unknown; further study needed to verify the status of the species in the state.

19 Order Cingulata -- Strange-jointed Mammals Family Dasypodidae (Armadillos) Dasypus novemcinctus (Nine-banded armadillo). Known statewide except in the most eastern part of Tennessee; occupies a variety of habitats that includes forested bottomlands and upland woodlands. Conservation status: none; a common species in suitable habitat.

Order Lagomorpha -- Hare-shaped Mammals Family Leporidae (Hares and Rabbits) Sylvilagus floridanus (Eastern cottontail). Statewide in distribution; occurs in a variety of habitats that includes old fields, brushy edges, and other habitats with a mixture of herbaceous and shrubby plants. Conservation status: none; may be a common species in suitable habitat. Sylvilagus obscurus (Appalachian cottontail). Occurs in mountainous regions of eastern and middle Tennessee; habitat most often associated with dense forest and boreal environments. Conservation status: none; an uncommon species throughout most of its distribution. Sylvilagus aquaticus (Swamp rabbit). Known throughout western Tennessee and may occur in parts of middle Tennessee; habitat includes lowland swamps and other wetlands. Conservation status: none; may be common in preferred habitat. Lepus americanus (Snowshoe hare; Extirpated). Found within historic times in the mountains of extreme eastern Tennessee; primary habitat included forest associated with brushy areas and areas with secondary growth. Conservation status: none.

Order Rodentia -- Gnawing Mammals Family Sciuridae (Squirrels) Tamias striatus (Eastern chipmunk). Occurs statewide; found in a variety of habitats in both rural and urban areas (e.g., forest edge, log piles, wooded banks, piles of stones, around stone walls, under sidewalks, and old outbuildings). Conservation status: none; a common species. Marmota monax (Woodchuck). Statewide in distribution; found in woodlands, roadside habitats, and fields. Conservation status: none; a common species in many parts of the state. Sciurus carolinensis (Eastern gray squirrel). Statewide in distribution; inhabits forests of rural and urban areas. Conservation status: none; a common species. Sciurus niger (Eastern fox squirrel). Statewide in distribution; found primarily in hardwood forests. Conservation status: none; a common species in many parts of the state. Tamiasciurus hudsonicus (Red squirrel). Occurs in eastern Tennessee; habitat is primarily forest (at higher elevations). Conservation status: none, common species in eastern part of the state. Glaucomys volans (Southern flying squirrel). Statewide in distribution; habitat is primarily forest. Conservation status: none; the species may be locally abundant in a variety of forested habitats.

20 Glaucomys sabrinus (Northern flying squirrel). Known only from the mountains of eastern Tennessee; habitat includes both coniferous and deciduous forest at higher elevations; subspecies occurring in Tennessee is G. s. coloratus; a federally listed endangered subspecies. Conservation status: Endangered Subspecies. S1-S2, G5T1. Family Castoridae (Beavers) Castor canadensis (American beaver). Reported statewide in distribution; inhabits a variety of aquatic sites. Conservation status: none; a common mammal in wetland habitats; locally abundant in many areas. Family Cricetidae (Cricetids) Oryzomys palustris (Marsh oryzomys). Statewide in distribution; found in a variety of habitats (especially marshes and wet meadows). Conservation status: none; a common species in suitable habitat. Reithrodontomys humulis (Eastern harvest mouse). Statewide in distribution, habitat includes oldfields, meadows, and thickets. Conservation status: none; an uncommon species rarely found in any large numbers at individual sites. Peromyscus maniculatus (North American deermouse). Statewide in distribution; a short-tailed form (P. m. bairdii) is found in the western part of the state and inhabits primarily open habitats (e.g., meadows, pastures, cultivated fields, and along field borders and fencerows), a long-tailed form (P. m. nubiterrae) inhabits forests of eastern Tennessee. Conservation status: none; P. m. bairdii, uncommon; P. m. nubiterrae, common. Peromyscus leucopus (White-footed deer-mouse). Statewide in distribution, found in a variety of habitats that includes wooded areas and forest edge associated with brush piles, grassy areas, old fields, and fencerows. Conservation status: none; common in suitable habitat. Peromyscus gossypinus (Cotton deermouse). Statewide in distribution; usually found in moist, timbered areas (e.g., swamps and river bottoms) and in caves and crevices around rocky bluffs. Conservation status: none; locally abundant in some areas. Ochrotomys nuttalli (Golden mouse). Statewide in distribution; habitat includes woodlands, forest edges, moist thickets (with vines and dense understory), and mid-successional fields. Conservation status: none; an uncommon species; widespread in distribution but never found in large numbers at individual sites. Sigmodon hispidus (Hispid cotton rat). Statewide in distribution; occurs in grassy fields, borders of cultivated fields, along roadsides in dense, grassy overgrowth, and other early successional habitats. Conservation status: none; a common species in suitable habitat. Neotoma floridana (Eastern woodrat). Distribution includes western and portions of eastern Tennessee (absent in most of middle Tennessee); in the western part of the state, N. f. illinoensis occurs in timbered areas but most common in lowland hardwood forest; may occupy old buildings; in the eastern portion of Tennessee, N. f. haematoreia occupies brush piles in forest or at forest edge, caves in limestone bluffs or crevices of rocky outcroppings. Conservation

21 status: Deemed in Need of Management (TDEC; TWRA). N. f. illinoensis, S3, G5T5; N. f. haematoreia, S2, G5T4Q; both subspecies are uncommon throughout most of their distribution. Neotoma magister (Allegheny woodrat). Known from the central part of Tennessee; occurs in a variety of disjunct forested habitats that include talus slopes, cliff faces, rock outcrops, boulder fields, and caves. Conservation status: Deemed in Need of Management (TDEC). S3, G3-G4; the species is uncommon but may be locally abundant. Myodes gapperi (Southern red-backed vole). Occurs in eastern and middle Tennessee; habitat includes primarily mesic areas of woodlands and moss-covered boulders as well as wet meadows. Conservation status: none; common in preferred habitat. Microtus pennsylvanicus (Meadow vole). Found in the eastern part of the state; habitat includes moist fields with thick vegetation mainly of grasses and sedges, species may be locally abundant. Conservation status: none; common in preferred habitat. Microtus chrotorrhinus (Rock vole). Known only from extreme eastern Tennessee; primary habitat includes forest clearings associated with early successional vegetation and moss- covered boulders in mountainous areas. Conservation status: Deemed in Need of Management (TDEC; TWRA). S2, G4T3; uncommon. Microtus ochrogaster (Prairie vole). Ranges throughout Tennessee except in the extreme eastern portion (mountains) of the state; primary habitat is grassy fields. Conservation status: none; may be locally abundant. Microtus pinetorum (Woodland vole). Statewide in distribution; common in a variety of habitats (e.g., fields adjacent to wooded areas, in orchards, along fencerows, and in thickets along forest edges). Conservation status: none; may be locally abundant. Ondatra zibethicus (Common muskrat). Statewide in distribution; inhabits many aquatic habitats (e.g., lakes, ponds, swamps, streams, and rivers). Conservation status: none; may be locally abundant. Synaptomys cooperi (Southern bog lemming). Statewide in distribution; habitat is mainly marshy meadows and low, wet areas. Conservation status: Deemed in Need of Management (TDEC; TWRA). S4, G5; uncommon. Family Muridae (Murids) Rattus rattus (Roof rat; Introduced). Statewide in distribution; associated with human habitation; usually upper stories of buildings. Conservation status: none; may be locally abundant. Rattus norvegicus (Brown rat; introduced). Statewide in distribution; found in many habitats but usually most common near human habitation. Conservation status: none; common in suitable habitat. Mus musculus (House mouse; Introduced). Statewide in distribution; inhabits a variety of habitats, which are usually associated with humans. Conservation status: none; common in suitable habitat.

22 Family Dipodidae (Jumping Mice) Zapus hudsonius (Meadow jumping mouse). Statewide in distribution; habitat includes open grassy field and other early successional vegetation. Conservation status: Deemed in Need of Management (TDEC; TWRA). S4, G5; widespread in distribution but an uncommon species. Napaeozapus insignis (Woodland jumping mouse). Known from mountainous areas of eastern and middle Tennessee; habitat includes moist areas in the forest or near the forest edge. Conservation status: Deemed in Need of Management (TDEC; TWRA). S4, G5; widespread in suitable habitat but never found in large numbers at individual sites. Family Erethizontidae (Porcupine) Erethizon dorsatum (North American porcupine; Extirpated). In historic times, the species was known in eastern Tennessee; primary habitat was likely forested areas. Conservation status: none. Family Myocastoridae (Coypu or Nutria) Myocastor coypus (Coypu or Nutria; Introduced). Reported only in western Tennessee; habitat is primarily freshwater marshes, streams, and rivers. Conservation status: none; increasing in numbers and distribution spreading eastward.

Order Carnivora -- Flesh-eating Mammals Family Canidae (Dogs, Foxes, and Allies) Canis familiaris (Domestic dog; Introduced). Feral or free-ranging dogs are known statewide; found in most rural and urban habitats. Conservation status: none; may be locally common. Canis latrans (Coyote). Statewide in distribution; found in a variety of rural and urban habitats but most common in brushy areas, forest edge, and open farmland. Conservation status: none; may be locally common. Canis lupus lupus (Gray wolf; Extirpated). In historic times, the species probably occupied much of eastern and middle Tennessee; found in a variety of habitats that included upland and bottomland forest. Conservation status: none. Canis lupus rufus (Red wolf; Extirpated). In historic times, the taxon probably ranged over much of western Tennessee; habitat included upland and bottomland forest as well as coastal prairies and marshes. Conservation status: none. Vulpes vulpes (Red fox). Statewide in distribution; inhabits borders of forested areas and adjacent open lands. Conservation status: none; may be locally common. Urocyon cinereoargenteus (Gray fox). Statewide in distribution; inhabits numerous forested areas and open brushland. Conservation status: none; may be locally common.

23 Family Ursidae (Bears) Ursus americanus (American black bear). Known in the eastern and middle parts of the state; inhabits wooded and swampy bottomlands. Conservation status: none; thought to be increasing in numbers across the state. Family Procyonidae (Raccoons) Procyon lotor (Raccoon). Statewide in distribution; found in a wide variety of rural and urban habitats; most abundant in bottomland hardwood forest associated with water. Conservation status: none; may be locally abundant. Family Mustelidae (Weasels and Allies) Lontra canadensis (North American river otter). Statewide in distribution; inhabits rivers, streams, lakes, swamps, and ponds often bordered by forest. Conservation status: none; may be locally abundant. Martes pennanti (Fisher; Extirpated, Reintroduced). In historic times, occurred in northeastern Tennessee; reintroduced in Cumberland Co. in 2002; habitat is primary mixed forest. Conservation status: none. Mustela nivalis (Least weasel). Occurs in eastern and middle Tennessee; primary habitat includes forested areas with talus slopes but may occur in grasslands and edge habitats. Conservation status: Deemed in Need of Management (TDEC). S2, G5; uncommon. Mustela frenata (Long-tailed weasel). Statewide in distribution; found in a variety of habitats (e.g., forest edges, stream banks, brushland, and fencerows). Conservation status: none; an uncommon species in many areas. Neovison visors (American mink). Statewide in distribution; habitat includes lakes, ponds, rivers, streams, swamps, and other areas with permanent water; additionally, forest stands adjacent to water are utilized by the species. Conservation status: none; may be common in suitable habitat. Family Mephitidae (Skunks) Spilogale putorius (Eastern spotted skunk). Occurs in eastern and middle Tennessee; habitat included rugged terrain (rocks and cliffs) as well as fencerows, brushy fields, and farmlands. Conservation status: none. S3, G5; uncommon. Mephitis mephitis (Striped skunk). Statewide in distribution; occurs in a variety of habitats (e g., forests, fields, forest/field edges, rocky outcrops, fencerows, pastures, and cultivated sites) in rural and urban areas. Conservation status: none; may be locally abundant.

24 Family Felidae (Cats) Felis catus (Domestic cat; Introduced). Feral or free-ranging cats are known statewide; they occupy most rural and urban habitats. Conservation status: none; may be locally abundant. Puma concolor (Cougar; Extirpated). In historic times, the species occurred statewide; occupied a wide range of habitats that included bottomlands and rough mountainous areas. Conservation status: none. Lynx rufus (Bobcat). Statewide in distribution; occurs in a variety of rural and urban habitats that include river bottoms, rocky outcrops, and forested areas with thick underbrush. Conservation status: none; may be locally abundant.

Order Artiodactyla -- Even-toed Hoofed Mammals Family Suidae (Pigs) Sus scrofa (Wild pig; Introduced). Statewide in distribution; inhabits a variety of habitats, does well in upland and bottomland hardwood forest. Conservation status: none; may be locally abundant. Family Cervidae (Deer) Cervus elaphus (Red deer or elk; Extirpated, Reintroduced). Found only in eastern Tennessee; reintroduced population; habitat mainly a mixed open and woodland landscape. Conservation status: none; population sustaining itself. Odocoileus virginianus (White-tailed deer). Statewide in distribution; common in a variety of rural and urban habitats that includes forest and fields. Conservation status: none, may be locally abundant. Family Bovidae (Bovids) Bison bison (American bison; Extirpated; Reintroduced in captivity). Captive herds only; a grassland species. Conservation status: none; no free-roaming populations.

25 Acknowledgments We thank the many students, personnel of the Tennessee Wildlife Resources Agency, Tennessee Department of Environment and Conservation, Tennessee Valley Authority, US Fish and Wildlife Service, US Forest Service, National Park Service, US Department of Defense, Ames Plantation, Milan Army Ammunition Plant, and numerous individuals within the community who contributed to building the collection of mammals housed at The University of Memphis and other sites. Appreciation is extended to the curators of the respective institutions listed under Materials and Methods for access to their collections.

Literature Cited Beasley, L. E., and W. D. Severinghaus. 1973. A survey of the cricetine rodents of West Tennessee. Journal of the Tennessee Academy of Science 49: 106-112. Calhoun, J. B. 1941. Distribution and food habits of mammals in the vicinity of the Reelfoot Lake Biological Station. Journal of the Tennessee Academy of Science 6: 177-225. Choate, J. R., J. K. Jones Jr., and C. Jones. 1994. Handbook of Mammals of the South-central States. Louisiana State University Press, Baton Rouge, Louisiana. Conway, C. H., and J. C. Howell. 1953. Observations on the mammals of Johnson and Carter counties, Tennessee, and Avery County, North Carolina. Journal of the Tennessee Academy of Science 28: 53-61. Corgan, J. X. 1976. Vertebrate fossils of Tennessee. Bulletin 77. Tennessee Department of Conservation. Nashville. Tennessee. Goodpaster, W. W., and D. F. Hoffmeister. 1952. Notes on the mammals of western Tennessee. Journal of Mammalogy 33: 362-371. Graves, F. F., and M. J. Harvey. 1974. Distribution of Chiroptera in western Tennessee. Journal of the Tennessee Academy of Science 49: 106-109. Hall, E. R. 1981. The Mammals of North America. Ronald Press Company, New York. Howell. J. C., and C. H. Conway. 1952. Observations on the mammals of the Cumberland Mountains of Tennessee. Journal of the Tennessee Academy of Science 27: 153-158. Kellogg, R. 1939. Annotated list of Tennessee mammals. Proceedings of the US National Museum 86: 245-303. Kennedy, M. L. 1991. Annotated checklist of the mammals of western Tennessee. Journal of the Tennessee Academy of Science 66: 183-185. Kennedy, M. L., and M. J. Harvey. 1980. Mammals. Pp. C1-050 in Tennessee's Rare Wildlife. Vol. 1: The Vertebrates (D. C. Eagar and R. M. Hatcher. eds.). Tennessee Department of Environment and Conservation. Nashville. Tennessee.

26 Kennedy, M. L., P. K. Kennedy. and G. D. Baumgardner. 1984. First record of the Seminole bat (Lasiurus seminolus) in Tennessee. Journal of the Tennessee Academy of Science 59: 89-90. Komarek, E. V., and R. Komarek. 1938. Mammals of the Great Smoky Mountains. Bulletin of the Chicago Academy of Science 5: 137-162. Linzey, D. W. 1995. Mammals of Great Smoky Mountains National Park. The McDonald and Woodward Publishing Company, Blacksburg, Virginia. Miller, R. A. 1974. The geologic history of Tennessee. Tennessee Division of Geology Bulletin 74: 1-100. Rhoads, S. N. 1896. Contributions to the zoology of Tennessee. No. 3. Mammals. Proceedings of the Academy of Natural Sciences of Philadelphia 48: 175-205. Severinghaus, W. D., and L. E. Beasley. 1973. A survey of the microtine and zapodid rodents of west Tennessee. Journal of the Tennessee Academy of Science 48: 129-133. Smith, C. K., J. Giles, M. E. Richmond, J. Nagel, and D. W. Yambert. 1974. The mammals of northeastern Tennessee. Journal of the Tennessee Academy of Science 49: 88-94. Wilson, D. E., and D. M. Reeder, 2005. eds. 2005. Mammal Species of the World: A Taxonomic and Geographic Reference. 316 ed. Johns Hopkins University Press, Baltimore, Maryland. Michael L. Kennedy, Phyllis K. Kennedy, Edward L. Warr, and Robert (Pete) L. Wyatt Ecological Research Center, The University of Memphis. Memphis, TN 38152 (MLK, PKK) Tennessee Wildlife Resources Agency, Wildlife Management Division, PO Box 40747, Nashville, TN 37204 (ELW) Tennessee Wildlife Resources Agency, Region IV Office, 330 Wildlife Way, Morristown, TN 37814 (RLW) Present address: 500 Nilson Court, Nashville, TN 37211 (ELW)

Copyright 2012 Tennessee Academy of Science Copyright 2012 Gale, Cengage Learning. All rights reserved.

27 III. Mammal Skulls

Tooth Types and Diets

Teeth stab, tear, slice, grind, crush, and chop food in hundreds of different ways, and more can be revealed about animals’ lifestyles by examining their teeth than by any other single physical characteristic. This is because tooth size, shape, and arrangement in the mouth are important determinants of the type of food an animal can obtain and consume. Its diet and method of feeding are, in fact, closely related to its dentition (as in number, kinds, and arrangement of teeth). In fact, dentition can be used to identify an animal to type.

Mammalian Tooth Types Incisors are the flat, chisel-shaped teeth at the front of the mouth (behind the lips) that are used in biting, cutting, nibbling, and stripping. [3 pairs] Canines are behind the incisors, and are located on the cheek-side of the mouth. They are cone- shaped, and may even be dagger-like. They are used for seizing, piercing and tearing. [1 pair] Premolars are transitional teeth located between the canines and the molars at the rear. They have 1 or more cusps (points) and are generally used for grinding and crushing, but they may also be used in slicing food. [4 pairs] Molars have many major cusps and are located in the back of the mouth on the cheek side. Molars function in grinding and crushing. [2 pairs]

28 Diet Types

Animals can generally be assigned to one of four major groups with respect to diet. Carnivore (meat eater) Herbivore (plant eater) Insectivore (insect eater) Omnivore (an animal that eats a variety of foods including meat and plants)

Carnivores share special adaptations for life as predators. Most notably, they have large grasping (holding) prey and slicing canines. All of their teeth have long roots. The fourth upper premolar and first lower molar carnivores are called carnassial teeth. These are the longest and sharpest teeth in the carnivore’s mouth and are specialized for cutting flesh. The cusps on a carnivore’s molars are also high and pointed. Even these teeth in the back of the mouth are used in tearing flesh. Carnivores also have front facing eyes, heavy skulls, and jaws that are strong but that can only open and shut, not move side to side.

Herbivores have tall molar teeth that are very broad. These molars have flat upper surfaces sometimes with ridges on them to help grind plant material. Associated with the grinding tooth structure, many herbivores are able to move their jaws from side to side. This increases the ability of the herbivore to grind plant material between the surfaces of the molars. Many herbivores lack canines entirely as these teeth are specially adapted to tearing flesh, and herbivores feed only on plants. The incisor in the herbivore is not dagger-like, but rather may be clipper-like, for use in cutting off plant stems. In many herbivores, the incisors grow continuously. Cutting hard plant material wears an herbivore’s incisors down, so continuous tooth growth keeps them at an optimal length for this work.

Insectivores have a mouthful of sharp little teeth that are similar in size and shape. These are used in seizing and crushing hard-shelled insects and other small animals like worms. They use incisors for slicing and premolars.

Omnivores are generalists and have all four tooth types. They have sharp, long canines for puncturing and grabbing onto animal prey. Wide molar teeth with low bumpy crowns are often present to handle both the chewing of meat and grinding of plant material. Greater variability in the shape of different teeth is the key to this type of animal given its broad diet.

29 Dental Formulas and Skull Descriptions

Dental formulas tell the number of each type of tooth of a species. This helps biologists identify species using their skulls. The following dental formulas are for one side of the mouth, top row referencing the upper jaw, bottom row the lower jaw. Multiply by two to get the total number of teeth. Key: I incisor, C canine, P premolar, M molar.

The dental formula for Homo sapiens is:

I = 2 C= 1 P= 0 M= 2 = 20 Deciduous I = 2 C= 1 P= 2 M= 3 = 32 Permanent 2 1 0 2 2 1 2 3

Humans get all of their permanent teeth by the age of 21. All but the third molars (the wisdom teeth) have erupted by the end of adolescence.

Dental formulas and skull descriptions of some common Tennessee wild mammals

Opossum (Didelphis virginiana) The opossum has more teeth than any other North American mammal. Adults have 50 teeth. Skulls have a large saggital crest and a small braincase.

I= 5 C= 1 P= 3 M=4 = 50 4 1 3 4

American Beaver (Castor canadensis) The largest rodent in N. America, the American beaver has a large, heavy skull. The auditory meatus is elevated to accommodate an aquatic lifestyle. The front surface of the incisors is orange.

I= 1 C= 0 P= 1 M= 3 = 20 1 0 1 3

Raccoon (Procyon lotor) The raccoon skull is triangular in outline with a short, broad rostrum, long spreading zygomatic arches and a large braincase. Temporal ridges form a V.

I= 3 C= 1 P= 4 M= 2 = 40 3 1 4 2

30 Domestic cat (Felis catus) Auditory bullae are large, oval and inflated.

I= 3 C= 1 P= 3 M= 1 = 30 3 1 2 1

Mountain Lion or Cougar (Felis concolor) Mountain lions have a large, rounded skull with a short, broad rostrum. Carnassials are long and blade-like. The eye orbits of cougars are large and forward facing to provide excellent binocular vision.

I= 3 C= 1 P= 3 M= 1 = 30 3 1 2 1

White-tailed Deer (Odocoileus virginanus) Deer skulls have a long, slender rostrum. Deer also lack upper incisors requiring them to rip the vegetation when they browse.

I= 0 C= 0 P= 3 M= 3 = 32 3 0 3 3

Eastern Cottontail (Sylvilagus floridanus) The Eastern cottontail is in the order of animals called Lagomorphs. Lagomorphs have 4 upper incisors, one pair behind the other. The forward pair of incisors is grooved and rodent-like. The second pair is reduced and peg-like.

I= 2 C= 0 P= 3 M= 3 = 28 1 0 2 3

Big Brown Bat (Eptesicus fuscus) All bats in Tennessee are insectivores. They have different types of teeth, but all of the teeth are similar looking.

I= 2 C= 1 P= 1 M= 3 = 32 3 1 2 3

31 Skull Glossary

Auditory bullae -- The hollow bulbous structures at the back of the skull that houses the inner and middle ear Auditory meatus -- Ear holes Carnassials -- Large teeth found in many carnivorous mammals used for shearing flesh and bone. In the carnivores, the carnassials are the modified last upper premolar and the first lower molar. Lagomorphs -- The order of mammals consisting of hares, rabbits and pikas. Rostrum -- Snout Saggital crest -- Raised ridge that runs along the center of the back portion of the skull. Divides the back of the skull into right and left sides Temporal ridges -- Raised ridge of skull that runs along the sides of the back portion of the skull. Zygomatic arches -- Cheek bones

32 Skull Key to Certain Orders of Mammals

1. a. Homodont – or teeth absent ……………………………………………………………………………….. 10 1. b. Heterodont – different teeth present ……………...... ……………………... 2

2. a. Upper incisors present ………………………………………………………………………………………….. 3 2. b. Upper incisors absent, or if present (pigs and peccaries) canines almost triangular in cross section ………………………………..Artiodactyla (even-toed ungulates, i.e. deer)

3. a. Diastema (space) present between incisors and cheek teeth .……………………………… 4 3. b. Diastema absent between incisors and cheek teeth ……………………………………………. 6

4. a. Upper incisors 6; no incisors enlarged ……………………………………………….Perissodactyla (odd-toed ungulates, i.e. horses) 4. b. Upper incisors 2 or 4; two upper incisors much enlarged ……………………………………. 5

5. a. Upper incisors in two pairs one behind the other ………………...Lagomorpha (rabbits) 5. b. Upper incisors one pair only ……………………………………………………..Rodentia (rodents)

6. a. Diastema (space) present between upper incisors ..……………………..Chiroptera (bats) 6. b. Diastema not present between upper incisors ..………………………………………………….. 7

7. a. Enlarged canine teeth present, zygomatic arch well developed ..…………………………. 8 7. b. Canine teeth not enlarged, zygomatic arch slender or absent ……………………………….. ………………………………………………………………………………...Soricomorpha (insectivores)

8. a. Angular process of mandible bent in toward midline, skull less than 6 cm. long …….. ………………………………………………………………….Didelphimorphia (pouched mammals) 8. b. Angular process of mandible not bent in toward midline (except in some Pinnipedia, then skull over 6 cm. long) ……………………………………………………………………………….. 9

9. a. All molars uniformly conical, some with prominent accessory cusps in symmetrical pattern of threes ..……………………………………………..….. Order Carnivora (carnivores) Suborder Pinnipedia (fin-footed, i.e. seals) 9. b. Molars not uniformly conical or not as above ..……………………………...Order Carnivora Former Suborder Fissipeda (most carnivores)

10. a. Lacks incisors and canines, teeth peg-like, nostrils open on front end of skull ..…….. ……………………..…………..Order Cingulata (strange-jointed mammals, i.e armadillos) 10. b. Nostril opening (1 or 2) located on top of skull ……..……Cetacea (marine mammals)

From University of Tennessee, Knoxville 1982 Mammalogy Class – Forestry, Wildlife and Fisheries Department, Dr. Michael Pelton. Edited by Pandy English, 2015

33 IV. Mammal Tracks and Signs

Mammal Tracking Station

Tracks can be used to detect the types of animals that use an area. A tracking or scent station can be build to record these tracks.

1) Find an area that appears well traveled by animals, possibly with existing paths or footprints. These “travel lanes” are often located at the wood’s edge or where there is cover for escape. Also check drainage areas, rock walls and wood piles, especially for smaller mammals. 2) Clear an approximate 2-foot radius circle, removing all vegetation. This area will need to remain clear, requiring occasional weeding. 3) Cover the area with fine sand such as masonry or play sand. Wet the sand with a mist bottle. 4) Take a tongue depressor or a popsicle stick and split the end. In the split, place a small cotton ball soaked in drops of sardine oil or cat food. Place this lure in the circle’s center. 5) Set the lure in late afternoon or evening. Return the following morning to investigate any clues revealing the night’s activities. Look up tracks in a field guide to identify overnight visitors.

Scent Animals use odors from urine or scent gland oils to mark territory, and often attract other animals with these smells. If one animal marks a cache or food supply, others can follow their noses to a free lunch. Some animals are just curious to know who is in the neighborhood. These smells provide important avenues of communication within the biological community, indicating an animal’s health, dominance, and fertility. Wildlife lures, such as sardine oil, tuna, and cat food, may be used to mimic these strong odors and attract animals to a tracking station.

Other Commonly Seen Signs Besides tracks, other signs indicate the presence of mammals: scat, rubbings, claw marks, burrows, dens, nests, and tunnels. Look carefully at scat. Signs of the animal’s diet may be found, such as seeds from paw paw or persimmon. If it belongs to a carnivore, it will often contain fur from a recent meal. Male deer may often leave rubbings from their antlers on small trees as well as urine. A bobcat may leave claw markings on the trunks of trees. However, cats retract their claws when walking, and paw prints with claw impressions indicate a canine not a cat.

34 V. Resources

Field Guides will help distinguish one mammal genus from another, but to distinguish at the species level requires more in-depth examination of technical reference materials and literature.

Booth, Ernest S. How to Know the Mammals. Literary Licensing, 2012. Elbroch, Mark. Animals Skulls: A Guide to North American Species. Stackpole Books, 2006. Elbroch, Mark. Mammal Tracks and Sign. Stackpole Books, 2003. Kays, Roland W. and Don E. Wilson. Mammals of North America, 2nd Edition. Princeton Field Guides. Princeton University, 2009. Levine, Lynn. Mammal Tracks and Scat: Life-size Tracking Guide. Heartwood Press, 2008. Reed, Fiona A. A Field Guide to Mammals of North America, 4th Edition. Peterson Field Guides. Houghton Mifflin Co., 2006 Schwartz, Charles W. and Elizabeth R. Schwartz. The Wild Mammals of Missouri, 2nd Edition. University of Missouri, 2002. Wilson, Don E. and DeeAnn M. Reeder. Mammal Species of The World, 3rd Edition. John Hopkins University Press, 2005. (primary authority for taxonomy and names)

Additional Reading

Blackman, Haden W. Field Guide to North American Monsters: Everything You Need to Know about Encountering over 100 Terrifying Creatures in the Wild. Three Rivers Press, 1998. (mythology) Dunn, Rob. Every Living Thing: Man’s Obsessive Quest to Catalog Life. Harper Perennial, 2010.

Online

Animal Diversity Web, University of Michigan Museum of Zoology http://animaldiversity.org/accounts/Mammalia/

35 VI. Review Questions

1. Which of the following statements is TRUE about mammals? a. Mammals are ectothermic animals. b. Mammals have milk-producing glands and hair. c. Mammals that approach humans are usually friendly. d. There are no true hibernators among Tennessee’s mammals.

2. Marsupials comprise one group of mammals. Tennessee has a. no marsupial species b. one marsupial species (opossum) c. two marsupial species (opossum and raccoon) d. several marsupial species

3. In Tennessee, the current established range of black bears a. is shrinking b. remains restricted to Southern Appalachian counties along the North Carolina border c. is expanding westward onto the Cumberland Plateau d. extends statewide

4. Deer and elk are considered “homeless” mammals because a. they bed down in different places at night b. their natural habitats have disappeared c. they prefer urban environments d. they depend on humans for food

5. The beaver is distinguished by which of the following features: a. ear and nose valves b. polygamous lifestyle c. light brown, coarse hair d. a long, narrow and scaly tail

6. Which statement is TRUE about Tennessee’s bats? a. All Tennessee bat species are true hibernators. b. White Nose Syndrome is not affecting bats in this area. c. Tennessee has mainly larger, fruit-eating bats. d. Approximately half of 1% of the bat population have rabies.

7. Crepuscular mammals a. have opposable thumbs b. are mostly active at dawn and dusk c. have powerful hind legs for running and jumping d. are active during the day

36 8. This mammal utters a high pitched whistle when alarmed which earned it the common name ‘whistle pig.’ a. wild hog b. muskrat c. chipmunk d. woodchuck

9. Which statement is FALSE? a. Wildlife scents only serve to mark territory. b. A scent marking from one species may attract individuals of different species. c. Strong scents like sardine oil can be used as lures to attract wildlife. d. Scent, scat, and urine can communicate an animal’s health and sex.

10. Match the description to the correct animal. a. rich, dark chestnut brown fur b. predominantly grey or rusty brown or off-white with a black-tipped tail c. coarse, grizzled grayish fur d. rusty-red or reddish yellow coat; black legs, feet and ears _____ red fox _____ beaver _____ opossum _____ coyote

11. Two extirpated native species that have been successfully reintroduced to the wild in Tennessee are a. fisher and elk b. bison and snowshoe hare c. porcupine and elk d. red wolf and elk

12. Rabbits and beavers eat their excrement to extract additional nutrients. This behavior is termed a. cephalagia b. narcolepsy c. dysphagia d. coprophagia

13. Rodents’ ______keep growing and must be worn down to avoid problems. a. front claws b. incisor teeth c. canine teeth d. molar teeth

37 14. To distinguish canine tracks from cat tracks, note a. the presence or absence of a ‘fifth’ toe b. the shape of the center foot pad c. the presence or absence of claw marks d. the width of the toe pads

15. Which statement regarding elk and deer antlers is FALSE? a. bucks shed their antlers in winter b. bulls shed their antlers in spring c. antler tissue is the fastest growing mammal tissue d. shed antlers remain untouched for years

16. Much can be learned about a mammal’s lifestyle by examining its a. teeth b. feet c. skull d. tail

17. Herbivores often lack this type of teeth. a. incisors b. canines c. premolars d. molars

18. Rabbit teeth a. are composed entirely of similar sized incisors b. are characterized by two pairs of canines c. feature two pairs of upper incisors, one set behind the other d. are replaced annually

19. This mammal’s skull has a large saggital crest and small braincase. a. American beaver b. raccoon c. opossum d. white-tailed deer

20. Indicate whether the following traits are true (T) or false (F) regarding coyotes. ____ solitary animals that move in family units during summer to raise young ____ feed only on very specific foods ____ able to adapt to and exploit almost any habitat ____ hold their tails up

38 Answer Key

1. b 2. b 3. c 4. a 5. a 6. d 7. b 8. d 9. a 10. d, a, c, b

11. a 12. d 13. b 14. c 15. d 16. a 17. b 18. c 19. c 20. T, F, T, F

39 Appendix A: Mammals of Tennessee Species List

Common Name Scientific Name Virginia Opossum Didelphis virginiana Cinereus shrew Sorex cinereus Southeastern shrew Sorex Iongirostris American water shrew Sorex palustris Smoky shrew Sorex fumeus Long-tailed shrew Sorex dispar American pygmy shrew Sorex hoyi Northern short-tailed shrew Blarina brevicauda Southern short-tailed shrew Blarina carolinensis North American least shrew Cryptotis parva Hairy-tailed mole Parascalops breweri Eastern mole Scalopus aquaticus Star-nosed mole Condylura cristata Little brown myotis Myotis lucifugus Southeastern myotis Myotis austroriparius Gray myotis Myotis grisescens Northern myotis Myotis septentrionalis Indiana myotis Myotis sodalis Eastern small-footed myotis Myotis leibli Silver-haired bat Lasionycteris noctivagans Tri-colored bat Perimyotis subflavus Big brown bat Eptesicus fuscus Eastern red bat Lasiurus borealis Seminole bat Lasiurus seminolus Hoary bat Lasiurus cinereus Evening bat Nycticeius humeralis Rafinesque's bigeared bat Corynorhinus rafinesquii Brazilian free-tailed bat Tadarida brasiliensis Nine-banded armadillo Dasypus novemcinctus Eastern cottontail Sylvilagus floridanus Appalachian cottontail Sylvilagus obscurus Swamp rabbit Sylvilagus aquaticus

40 Snowshoe hare; Extirpated Lepus americanus Eastern chipmunk Tamias striatus Woodchuck Marmota monax Eastern gray squirrel Sciurus carolinesis Eastern fox squirrel Sciurus niger Red squirrel Tamiasciurus hudsonicus Southern flying squirrel Glaucomys volans Northern flying squirrel Glaucomys sabrinus American beaver Castor canadensis Marsh oryzomys Oryzomys palustris Eastern harvest mouse Reithrodontomys humulis North American deer-mouse Peromyscus maniculatus White-footed deer-mouse Peromyscus leucopus Cotton deer-mouse Peromyscus gossypinus Golden mouse Ochrotomys nuttalli Hispid cotton rat Sigmodon hispidus Eastern woodrat Neotoma floridana Allegheny woodrat Neotoma magister Southern red-backed vole Myodes gapperi Meadow vole Microtus pennsylvanicus Rock vole Microtus chrotorrhinus Prairie vole Microtus ochrogaster Woodland vole Microtus pinetorum Common muskrat Ondatra zibethicus Southern bog lemming Synaptomys cooperi Roof rat; Introduced Rattus rattus Brown rat; Introduced Rattus norvegicus House mouse; Introduced Mus musculus Meadow jumping mouse Zapus hudsonius Woodland jumping mouse Napaeozapus insignis North American porcupine; Extirpated Erethizon dorsatum Coypu or Nutria; Introduced Myocastor coypus Domestic dog; Introduced Canis familiaris Coyote Canis latrans Gray wolf; Extirpated Canis lupus lupus

41 Red wolf; Extirpated Canis lupus rufus Red fox Vulpes vulpes Gray fox Urocyon cinereoargenteus American black bear Ursus americanus Raccoon Procyon lotor North American river otter Lontra canadensis Fisher; Extirpated, Reintroduced Martes pennanti Least weasel Mustela nivalis Long-tailed weasel Mustela frenata American mink Neovison visors Eastern spotted skunk Spilogale putorius Striped skunk Mephitis mephitis Domestic cat; Introduced Felis catus Cougar; Extirpated Puma concolor Bobcat Lynx rufus Wild pig; Introduced Sus scrofa Red deer or elk; Extirpated, Reintroduced Cervus elaphus White-tailed deer Odocoileus virginianus American bison; Extirpated, Reintroduced Bison bison in captivity

Kennedy, Michael L.; Kennedy, Phyllis K.; Warr, Edward L.; Wyatt, Robert L. "Pete" Journal of the Tennessee Academy of Science June 1, 2012

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