Beasts (Mammals)

of the Lower Susquehanna River Watershed

With (forthcoming) commentary on the species including notes from the historic writings of nineteenth-century geologist, naturalist, and philologist Professor Samuel Steman Haldeman, Linnaean Society of Lancaster City and County naturalist Dr. Simon S. Rathvon, and the Curator of the Franklin and Marshall College Museum, Dr. Herbert H. Beck.

Samuel Steman Haldeman was born in 1812 at the Haldeman Mansion along the Susquehanna River at Locust Grove near Bainbridge in Lancaster County, Pennsylvania. The mansion can be seen from the Northwest Lancaster County River Trail and is open for tours during scheduled events. See the Haldeman Mansion Preservation Society website for more information.

In the vicinity of his boyhood home at the mouth of Conoy Creek, Haldeman explored the waters and shores of the Susquehanna below Conewago Falls. He developed an interest in the animal bones he found and began assembling skeletons. Haldeman collected the shells of the mollusks occurring there, a long-term pursuit that eventually led to the writing and publishing of “A Monograph of the Freshwater Univalve Mollusks of the United States” and “On the Freshwater Mollusca Common to Europe and America, Including Theoretical Observations Upon the Species and Their Distribution”. The former was the country’s first comprehensive account of its freshwater snails. Charles Darwin would comment on the latter in “The Origin of Species” as part of the book’s appendix, “…An Historical Sketch of the Progress of Opinion on the Origin of Species”.

During his adult years, Samuel Steman Haldeman resided along the Susquehanna at Chickies Rock near Marietta, Pennsylvania. He died at his home there in 1880. Haldeman left behind a wide-ranging body of work in the fields of geology, zoology, linguistics, and archaeology. Among these are commentary on the animals, including the “beasts”, found in Lancaster County during the early 1800s. His use of binomial nomenclature relieves us of the uncertainty that comes with interpreting similar lists of wildlife that use colloquial names. His sketch is the foundation of this list. (Mathew Brady image-Library of Congress)

SPECIES STATUS KEY

extinct-a native species no longer existing or living.

extinct/prehistoric-a species that became extinct prior to the time of recorded history, but for which paleontological evidence suggests or proves its presence in the Lower Susquehanna River Watershed.

Evidence of Mammals from Three “Bone Caves”

Extinct/prehistoric species marked with a “K” (K) have been identified among the hundreds of plant and animal remains recovered from the Cambrian limestone (dolomite) Port Kennedy Bone Cave at the Valley Forge National Historical Park along the south rim of the Newark Basin, the Gettysburg Basin’s continuance in eastern Pennsylvania. The cave, or more accurately the fissure, is located near the Schuylkill River at the base of the same Triassic ridge complex that intersects the Susquehanna at Conewago Falls. It lies just 20 miles east of the Susquehanna valley.

The Port Kennedy Bone Cave was discovered by quarry workers in 1870 during operations to remove Ledger Formation dolostone (dolomite) at the site. Local scientists who initially investigated the find notified paleontology experts Charles Wheatley and Edward Drinker Cope. A thorough study of the contents of the cave was initiated.

Specimens became trapped in the sinkhole sometime around 500,000 years ago—the exact date is unknown. Its Middle Pleistocene deposits provide evidence of the species that not only inhabited the immediate vicinity of the cave at the time, but certainly those with a range that simultaneously extended into the Piedmont of the nearby Susquehanna watershed and beyond.

Recent Sinkhole in the Vicinity of the Port Kennedy Bone Cave Site — Early investigators of the Port Kennedy Bone Cave wondered how the remains of more than one thousand animals could have accumulated in the bottom of the fissure. Have a look at this modern-day group of sinkholes located less than two miles to the south of Port Kennedy along the Pennsylvania Turnpike on the east side of the Valley Forge Service Plaza. During the mid-2010s, unconsolidated Bryn Mawr Formation sediments were excavated here to build a sound barrier between the highway and a residential subdivision to the south. When stormwater collects in these borrow pits, the shallow layer of remaining Bryn Mawr sands, gravels, and clays becomes saturated and unsupported sections collapse into a fissure in the underlying Ledger Formation dolostone. Though the fissure is now somewhat clogged with the rock, soil, and trash dumped inside in an attempt to eliminate the sinkholes, a very deep cavernous void in the Ledger dolostone could be seen in the autumn of 2016 when the site first experienced a cave-in. One could easily comprehend how the weight of unsuspecting animals would cause the unconsolidated sediments covering and surrounding such a fissure to break away and entrap them. Over time, roots and other plant material surrounding a sinkhole could provide support for soil and debris and could, in effect, heal and camouflage the opening to “reset the trap” for the next wave of hapless victims. (2020 Google image)

Living (extant) species marked with a “K” (K) are those mammals found in the Port Kennedy Bone Cave that have survived the Pleistocene ice maximums (probably by seeking refuge in warmer climes to the south) and ongoing Holocene (Anthropocene) extinction event to continue inhabiting the region.

The nomenclature of species found in the Port Kennedy Bone Cave and incorporated into this list is based upon the early work of Edward Drinker Cope (1871 and 1895), and the review of specimens and updating of taxa conducted by Daeschler, Spamer, and Parris (1993), and the Paleobiology Database.

Ground Sloth Bones from the Port Kennedy Bone Cave — Ground sloth skeletal remains excavated from the Port Kennedy Bone Cave. (National Park Service image)

Mammal species marked with an “H” (H) have been identified among the remains recovered from the more recently examined Hanover Quarry #1 Fissure bone deposit in Adams County, Pennsylvania. This site, discovered and studied during the early 1980s, is within the lower Susquehanna watershed in a headwaters area of the South Branch of the Conewago Creek. Like the Port Kennedy Bone Cave, the Hanover fissure occurs in Cambrian Ledger Formation dolostone (dolomite) along the edge of the Triassic rift basin. The animals are believed to have become trapped there at about the same period of time as those at Port Kennedy—about 500,000 years ago, during the Middle Pleistocene Epoch of the Quaternary Period.

Until his untimely death in 1982, examination of the fauna found in the Hanover Quarry #1 Fissure was being led by John E. Guilday, research curator at the Carnegie Museum of Natural History and an expert on cave deposits. Microtus guildayi, a vole identified among the remains from both the Hanover and Port Kennedy sites, is named in his honor.

The list of species found in the Hanover Quarry #1 Fissure and incorporated into this list of living and extinct mammals is from the Paleobiology Database.

A third bone deposit is located near the lower Susquehanna watershed in the Potomac River drainage basin. The Cumberland Bone Cave in Allegany County, Maryland, lies less than 20 miles south of the westernmost headwaters of the Juniata River, the largest of the lower Susquehanna’s tributaries. Mammal species marked with a “C” (C) have been identified among the remains recovered from this Middle Pleistocene sinkhole in Devonian limestone.

Raymond Armbruster was a local naturalist who, in 1912, alerted the scientific community to the discovery of prehistoric bones at the Cumberland cave site. These deposits have been thoroughly studied and many type specimens for new species have been collected there. Paleontologist James W. Gidley of the Smithsonian National Museum of Natural History, the foremost investigator of the site from 1912 to 1916, named an extinct wolf discovered among the remains in the cave as Canis armbrusteri— a tribute to Raymond Armbruster.

The list of species found in the Cumberland Bone Cave and incorporated into this list of living and extinct mammals is from the Paleobiology Database.

Cross-section View of a Middle Pleistocene Bone Cave — A sinkhole in Cambrian limestone (dolomite) becomes a Middle Pleistocene bone cave trapping a Smilodon sabre-toothed cat and hundreds of other animals. Following decomposition of the soft tissue, skeletal remains were crushed into fragments as yet another large victim like the Smilodon or maybe even a bear or mastodon tumbled into the fissure and struggled to escape. As they accumulated, the bones were buried in layers of loose rock and surface soil.

Locations of Bone Caves in and near the Lower Susquehanna Valley Watershed — Quaternary Period (Middle Pleistocene Epoch) bone deposit sites in and near the Lower Susquehanna River Watershed include the Cumberland Bone Cave (C), the Hanover Quarry #1 Fissure (H), and the Port Kennedy Bone Cave (K). Animals became trapped in the Hanover and the Port Kennedy sinkholes about 500,000 years ago, and in the Cumberland cave sinkhole as recently as 200,000 years ago.

Bone Deposit Site Activity and Middle Pleistocene Glacial Cycles — On this timeline of Middle Pleistocene glacial cycles, bone deposit sites are plotted on the two Pre-Illinoian interglacial stages when they are believed to have been trapping specimens. Each of the three sites predate the two most recent glacial events, the Illinoian and the Wisconsin, both of which produced significant ice coverage across the Susquehanna River Watershed in New York and northeastern Pennsylvania. The base image includes Alpine and legacy North American terms. (Base image by Tom Ruen, under license: CC BY-SA 3.0)

extirpated-a native species no longer occurring in the wild in the Lower Susquehanna River Watershed. “Extirpated” taxa may include living (extant) species not currently found among the valley’s mammals but known from the Middle Pleistocene deposits in the Hanover Quarry #1 Fissure (H).

extirpated/hypothetical-a native species not presently occurring in the Lower Susquehanna River Watershed but suspected of being among its fauna during the series of successions in plant communities—tundra to taiga to temperate forest—that followed the retreat into Canada of the southern ice border at the end of Wisconsin Glaciation, beginning about 22,000 years ago during the Late Pleistocene Epoch and extending into the current interglacial Holocene Epoch (11,650 years ago to present). These species are presumed to have existed in the lower Susquehanna valley either during the ice maximum, or while slowly transitioning their range as the climate warmed. Many were impacted by the arrival of Homo sapiens in the region during this time. In addition, “extirpated/hypothetical” taxa may include living (extant) species not currently found among the region’s mammals but known from the Middle Pleistocene bone cave deposits found near the lower Susquehanna valley—the Cumberland Bone Cave in Allegany County, Maryland (C) and the Port Kennedy Bone Cave in Valley Forge, Pennsylvania (K).

extirpated/introduced-a native species which, after elimination of the regional population, has been reestablished using released animals from an extant population.

exotic-a free-ranging escaped or released non-native species or variety; most are unwanted pets, domesticated farm animals, or zoo specimens.

feral-exotic animals that begin reproducing in the wild but retain dependence upon humans for survival of their population.

introduced-a non-native species that, following its release into the wild, has established a self-sustaining breeding population—often at the expense of one or more native species.

Federally Endangered-a native species listed by the United States government as imminently in danger of extinction.

PA Endangered-a native species listed by the Commonwealth of Pennsylvania as imminently in danger of extinction or of extirpation as a breeding species in the state.

MD Endangered-a native species listed by the State of Maryland as imminently in danger of extinction or of extirpation as a breeding species in the state.

Federally Threatened-a native species listed by the United States government as under threat to become an endangered species in the foreseeable future.

PA Threatened-a native species listed by the Commonwealth of Pennsylvania as under threat to become an endangered species in the state in the foreseeable future.

MD Threatened-a native species listed by the State of Maryland as under threat to become an endangered species in the state in the foreseeable future.

PA Candidate-an uncommon native species that could, in the future, become listed by the Commonwealth of Pennsylvania as endangered or threatened in the state.

A LIST OF THE BEASTS (MAMMALS)

of the Lower Susquehanna River Watershed

Classified using traditional taxonomic ranks and selected cladistic groups.

Domain-Eukaryota

Kingdom-Animalia

Phylum-Chordata

Subphylum-Vertebrata

Superclass-Osteichthyes/Clade-Euteleostomi:

The “bony vertebrates” including the Actinopterygians, the ray-finned fishes, and the Sarcopterygians, the lobe-finned fishes and tetrapods.

Clade-Sarcopterygii:

The lobe-finned fishes and tetrapods. All members of the clade Sarcopterygii presently occurring in the Susquehanna watershed (amphibians, reptiles, birds, and mammals including humans) are tetrapod descendants of extinct lobe-finned fishes.

Superclass-Tetrapoda:

The animals descended from fishes beginning in the Late Devonian epoch (about 370 million years ago), most having four limbs—the amphibians, the reptiles (including snakes, pterosaurs, dinosaurs, and birds), and the mammals.

Clade-Reptiliomorpha:

The group of living and extinct tetrapods having more similarity to the amniotes (reptiles, pterosaurs, dinosaurs, birds, and mammals) than to modern amphibians (Lissamphibia).

Clade-Amniota:

The group of tetrapods, living and extinct, that deposit their eggs on land (not in water) or retain them within the body—the reptiles (including pterosaurs, dinosaurs, and birds) and the mammals.

Clade-Synapsida

The group of living and extinct amniotes more closely related to mammals than to reptiles, birds, and other amniotes. All synapsids possess, behind each eye, a single temporal fenestra—an opening in the roof of the skull.

Skull of a White-tailed Deity — The temporal fenestra on this Odocoileus virginianus skull can be seen just posterior of the eye orbit and below the pedicle where antlers have been shed or removed. The coronoid process of the lower jaw protrudes from the opening. (Smithsonian National Museum of Natural History image www.si.edu)

Clade-Mammaliaformes

The most recent common ancestor of Morganucodonta and the crown group of mammals, and its living and extinct descendants.

Class-Mammalia

The crown group of mammals—the only living synapsids.

Order-Didelphimorphia

Family-Didelphidae

Didelphis virginiana (Virginia Opossum)

Mammals of the Lower Susquehanna River Watershed: Virginia Opossum — **Virginia Opossum** (*Didelphis virginiana).*

Order-Proboscidea

Family-Mammutidae

Mammut americanum (Mastodon)–extinct/prehistoric-K,H,C

Mammals of the Lower Susquehanna River Watershed: Mastodon — A molar from an extinct **Mastodon** (Mammut americanum) found in Mill Creek, East Lampeter Township, Lancaster County, Pennsylvania. (Specimen: North Museum, Lancaster, Pennsylvania)

Skeleton of a Mastodon — James W. Gidley poses with a reconstructed **Mastodon** (Mammut americanum) skeleton at the Smithsonian Institute, circa 1916. From 1912 to 1916, Gidley supervised collection of Middle Pleistocene mammal remains, including those of M. americanum, from the Cumberland Bone Cave. He concurrently coordinated the reconstruction of this Mastodon specimen discovered in Indiana. (Smithsonian National Museum of Natural History image www.si.edu)

Mammals of Pennsylvania: Mastodon — **Mastodon** (Mammut americanum) remains discovered in a peat bog in the Delaware River watershed near Marshalls Creek, Monroe County, Pennsylvania, have been assembled into a skeleton consisting of ninety percent authentic bones and ten percent replicas. (Specimen: State Museum of Pennsylvania, Harrisburg)

Family-Elephantidae

Mammuthus primigenius (Woolly Mammoth)–extinct/prehistoric

Mammals of the Lower Susquehanna River Watershed: Woolly Mammoth — A molar from an extinct **Woolly Mammoth** (Mammuthus primigenius) found at a site where workmen were digging in a meadow near White Horse, Salisbury Township, Lancaster County, Pennsylvania. (Specimen: North Museum, Lancaster, Pennsylvania)

Order-Pilosa

Family-Megalonychidae

Megalonyx species (a giant ground sloth)-extinct/prehistoric-C

Megalonyx dissimilis (a giant ground sloth)?-extinct/prehistoric-K

Megalonyx loxodon (a giant ground sloth)?–extinct/prehistoric-K

Megalonyx sphenodon (a giant ground sloth)?-extinct/prehistoric-K

Megalonyx tortulus (a giant ground sloth)?-extinct/prehistoric-K

Megalonyx wheatleyi (Wheatley’s Ground Sloth)-extinct/prehistoric-K

Claw of a Jefferson's Ground Sloth — The claw of a **Jefferson’s Ground Sloth** (Megalonyx jeffersonii). In 1871, Edward Cope listed five species of Megalonyx ground sloths among the remains of at least 14 individuals found in the Port Kennedy Bone Cave (each species is included in this list). Comparing them to specimens of Jefferson’s Ground Sloth like this one, Cope described three of his species as equal in size, and two species as smaller. Based upon more recent examinations, Wheatley’s Ground Sloth (Megalonyx wheatleyi) is probably the only valid taxon found at Port Kennedy. It is named for paleontologist Charles Wheatley, who, along with Cope, was one of the initial investigators of the cave’s contents. M. loxodon, M. sphenodon, and M. tortulus are currently considered junior synonyms to M. wheatleyi. M. dissimilis is considered invalid. (Smithsonian National Museum of Natural History image www.si.edu)

Family-Mylodontidae

Paramylodon harlani (Harlan’s Ground Sloth)-extinct/prehistoric-K

Skeleton of a Harlan's Ground Sloth — In 1871, Edward Drinker Cope identified Paramylodon bones, possibly those of the extinct **Harlan’s Ground Sloth** (Paramylodon harlani), among the remains removed from the Port Kennedy Bone Cave but considered them “not sufficiently characteristic to enable me to determine the species with certainty”. (Specimen from LaBrea Tar Pits, Los Angeles, California, Smithsonian National Museum of Natural History image www.si.edu)

Order-Primates

Family-Hominidae

Homo (Pan) sapiens (“Wise Man”)

Order-Lagomorpha

Family-Ochotonidae

Praotherium (Ochotona) palatina (a pika)-extinct/prehistoric-K

Ochotona princeps (American Pika)-hypothetical/extirpated-C

Family-Leporidae

Lepus americana (Snowshoe Hare)-extirpated-C

Sylvilagus species (a cottontail)-H

Sylvilagus floridanus (Eastern Cottontail)-K

Sylvilagus obscurus (Appalachian Cottontail)

Oryctolagus cuniculus (European Rabbit)-exotic

**Snowshoe Hare** (*Lepus americana). (National Park Service image by Tim Rains)*

Mammals of the Lower Susquehanna River Watershed: Eastern Cottontail — **Eastern Cottontail** (*Sylvilagus floridanus).*

Mammals of the Lower Susquehanna River Watershed: European Rabbit — **European Rabbit** (*Oryctolagus cuniculus).*

Order-Rodentia

Family-Castoridae

Castor canadensis (North American Beaver)-extirpated/introduced-K,C

Dipoides species (a round-toothed beaver)-extinct/prehistoric-C

Mammals of the Lower Susquehanna River Watershed: North American Beaver lodge. — A North American Beaver (Castor canadensis) lodge.

Mammals of the Lower Susquehanna River Watershed: Trees Gnawed by Beaver — Small marsh trees gnawed and felled by a North American Beaver (*Castor canadensis) for lodge construction and dam maintenance.*

Family-Geomyidae

Thomomys potomacensis (a smooth-toothed pocket gopher)-extinct/prehistoric-H,C

Family-Echimyidae

Myocastor coypus (Coypu)-introduced

Coypu/Nutria — **Coypu** (Myocastor coypus), commonly known as Nutria in the mid-Atlantic region, have become an invasive species. Native to Argentina and Chile, they were introduced into wetlands on the Delmarva Peninsula during the 1940s as a proposed source of pelts for the fur trade. Coypu were soon expanding their range and decimating Chesapeake Bay marshlands, particularly in the vicinity of Blackwater National Wildlife Refuge. Early in the twenty-first century, work was begun to rid the bay ecosystem of this destructive alien. Approximately 14,000 Coypu were eliminated at Blackwater, on other public lands, and on over 700 private parcels covering a combined area of more than 250,000 acres. During the autumn of 2022, the United States Fish and Wildlife Service declared Coypu populations in Maryland’s Chesapeake Bay region “eradicated” after a two-decade-long effort. (United States Fish and Wildlife Service image by Steve Hillebrand)

Family-Myocastoridae

Erethyzon dorsatum (North American Porcupine)-K,H,C

Mammals of the Lower Susquehanna River Watershed: North American Porcupine — **North American Porcupine** (Erethyzon doratum).

Family-Dipodidae

Napaeozapus insignis (Woodland Jumping Mouse)-H,C

Zapus (Jaculus) hudsonius (Meadow Jumping Mouse)-K,H

Mammals of the Lower Susquehanna River Watershed: Woodland Jumping Mouse — **Woodland Jumping Mouse** (*Napaeozapus insignis).*

Family-Cricetidae

Subfamily-Arvicolinae

Atopomys texensis (a primitive arvicoline)-extinct/prehistoric-C

Lasiopodomys species (a vole)?-H

Lasiopodomys deceitensis (a vole)-extinct/prehistoric-C

Microtus species (a vole)-C

Microtus chrotorrhinus (Rock Vole)-hypothetical/extirpated-C

Microtus guildayi (“Guilday’s Vole”)-extinct/prehistoric-K,H,C

Microtus pennsylvanicus (Meadow Vole)

Microtus pinetorum (Woodland Vole)

Phenacomys species (a heather vole)-hypothetical/extirpated-C

Pitymys cumberlandensis (a vole)-extinct/prehistoric-H,C

Neofiber species (a water rat)-extinct/prehistoric-C

Neofiber diluvianus (Diluvian Water Rat)-extinct/prehistoric-K

Synaptomys borealis (Northern Bog Lemming)-hypothetical/extirpated-C

Synaptomys cooperi (Southern Bog Lemming)-H,C

Myodes gapperi (Southern Red-backed Vole)-C

Myodes gapperi gapperi

Myodes gapperi rupicola

Ondatra annectens (a muskrat)-extinct/prehistoric-C

Ondatra idahoensis (a muskrat)-extinct/prehistoric-K,H

Ondatra zibethicus (Muskrat)

Mammals of the Lower Susquehanna River Watershed: Meadow Vole — **Meadow Vole** (Microtus pennsylvanicus).

Mammals of the Lower Susquehanna River Watershed: Southern Red-backed Vole — The **Southern Red-backed Vole** (Myodes gapperi) is an inhabitant of rich mountain forests. In the Lower Susquehanna River Watershed, it is most frequently encountered among the boulders forming the rocky outcrops atop Blue Mountain/Kittatinny Ridge and the crests to its immediate north. Merritt (1987) listed this isolated ridgetop population found only in southeastern Pennsyvania as possibly a distinct subspecies, M. g. rupicola (seen here) and populations throughout the remainder of the eastern two thirds of the state as the widespread M. g. gapperi.

Mammals of the Lower Susquehanna River Watershed: Muskrat — **Muskrat** (*Ondatra zibethicus).*

Mammals of the Lower Susquehanna River Watershed: Muskrat house — A Muskrat (*Ondatra zibethicus)* house.

Subfamily-Neotominae

Neotoma magister (Allegheny Woodrat)-PA Threatened-C

Neotoma spelaea (a pack rat)-extinct/prehistoric-C

Peromyscus species (a mouse)-H

Peromyscus cumberlandensis (a deer mouse)-extinct/prehistoric-C

Peromyscus maniculatus (Deer Mouse)

Peromyscus leucopus (White-footed Mouse)-K,C

**Allegheny Woodrat** (*Neotoma magister) at nest.* (National Park Service image)

**Deer Mouse** (*Peromyscus maniculatus)*. (Centers for Disease Control image by James Gathany)

Mammals of the Lower Susquehanna River Watershed: White-footed Mouse — **White-footed Mouse** (Peromyscus leucopus).

Subfamily-Sigmodontinae

Calomys (Hesperomys) species (a vesper mouse)?-extinct/prehistoric-K

Family-Muridae

Mus musculus (House Mouse)-introduced

Rattus norvegicus (Norway Rat)–introduced

Rattus rattus (Black Rat)-introduced

Mammals of the Lower Susquehanna River Watershed: Common Small Mammals — Commonly encountered small mammals of the Lower Susquehanna River Watershed including the invasive non-native murids House Mouse (Mus musculus) and Norway Rat (*Rattus norvegicus)*.

**House Mouse** (*Mus musculus).* (National Institutes of Health image)

**Norway Rat** (*Rattus norvegicus)*. (Centers for Disease Control image by H. G. Scott)

**Black Rat** (*Rattus rattus)*. (Centers for Disease Control image)

Family-Sciuridae

Subfamily-Sciurinae

Sciurus calycinus (a tree squirrel)-extinct/prehistoric-K

Sciurus carolinensis (Eastern Gray Squirrel)

Sciurus niger (Fox Squirrel)

Sciurus niger cinereus (Delmarva Fox Squirrel)-extirpated

Sciurus niger vulpinus (Eastern Fox Squirrel)

Tamiasciurus hudsonicus (Red Squirrel)-C

Glaucomys species (a flying squirrel)-C

Glaucomys volans (Southern Flying Squirrel)-C

Petauristodon species (a flying squirrel)-extinct/prehistoric–C

Mammals of the Lower Susquehanna River Watershed: Eastern Gray Squirrel — **Eastern Gray Squirrel** (*Sciurus carolinensis).*

Mammals of the Lower Susquehanna River Watershed: Eastern Gray Squirrels in nest. — Juvenile **Eastern Gray Squirrels** (*Sciurus carolinensis)* in nest.

**Delmarva Fox Squirrel** (*Sciurus niger cinereus).* (United States Fish and Wildlife service image by Chelsi Burns)

**Eastern Fox Squirrel** (*Sciurus niger vulpinus).* (United States Fish and Wildlife Service image by Alan Schmierer)

**Red Squirrel** (*Tamiasciurus hudsonicus).* (United States Fish and Wildlife Service image by Bill Thompson)

Mammals of the Lower Susquehanna River Watershed: Southern Flying Squirrel — **Southern Flying Squirrel** (*Glaucomys volans)*. (Vintage 35 mm image)

Subfamily-Xerinae

Marmota monax (Woodchuck)-K,H,C

Ichtidomys (Spermophilus) tridecemlineatus-(Thirteen-lined Ground Squirrel)-hypothetical/extirpated-C

Tamias striatus (Eastern Chipmunk)-H,C

Mammals of the Lower Susquehanna River Watershed: Woodchuck — **Woodchuck** (*Marmota monax ).*

Bones of the Thirteen-lined Ground Squirrel *(Ichtidomys tridecemlineatus)* were among the Middle Pleistocene remains discovered in the Cumberland Bone Cave. The current range of this denizen of the shortgrass prairies includes most of the central United States as far east as Ohio. In many quarters, it has adapted to mowed areas including lawns and parks. (United States Fish and Wildlife Service image by Chelsi Burns)

Mammals of the Lower Susquehanna River Watershed: Eastern Chipmunk — **Eastern Chipmunk** (*Tamias striatus).*

Order-Eulipotyphla

Family-Soricidae

Blarina brevicauda (Northern Short-tailed Shrew)-K,C

Blarina carolinensis (Southern Short-tailed Shrew)-extirpated-H,C

Cryptotis parva (North American Least Shrew)-PA Endangered

Sorex species (a shrew)-H,C

Sorex cinerus (Masked Shrew)-C

Sorex dispar (Long-tailed Shrew)

Sorex fumeus (Smoky Shrew)-C

Sorex hoyi (Pygmy Shrew)-extirpated?

Mammals of the Lower Susquehanna River Watershed: Northern Short-tailed Shrew — **Northern Short-tailed Shrew** (*Blarina brevicauda).*

Family-Talpidae

Condylura cristata (Star-nosed Mole)-C

Parascalops breweri (Hairy-tailed Mole)-extirpated?-H,C

Scalopus species (a mole)-K

Scalopus aquaticus (Eastern Mole)

Order-Chiroptera

Family-Vespertilionidae

Vespertilio species (a vesper bat)?-extinct/prehistoric-K,C

Vespertillio grandis (a vesper bat)-extinct/prehistoric-C

Pipistrellus (Perimyotis?) species- (a pipistrelle)-C

Perimyotis subflavus (Tri-colored Bat)-PA Endangered

Eptesicus fuscus (Big Brown Bat)-C

Nycticeius humeralis (Evening Bat)

Lasiurus borealis (Red Bat)

Lasiurus seminolis (Seminole Bat)

Lasiurus cinereus (Hoary Bat)

Myotis species (a bat)-H,C

Myotis leibii (Small-footed Myotis)-PA Threatened, MD Endangered ?

Myotis septentrionalis (Northern Long-eared Bat)–Federally Threatened, PA Endangered, MD Threatened

Myotis lucifigus (Little Brown Myotis)–PA Endangered

Myotis sodalis (Indiana Bat)-Federally Endangered, PA Endangered, MD Endangered

Myotis grisiscens (Gray Bat)-extirpated/hypothetical-C

Plecotus alleganiensis (Allegany Long-eared Bat)-extinct/prehistoric-C

Lasionycteris noctivagans (Silver-haired Bat)

**Tri-colored Bat** (*Perimyotis subflavus).* (United States Fish and Wildlife Service image by Peter Pattavina)

**Big Brown Bat** (*Eptesicus fuscus).* (United States Fish and Wildlife Service image by Ann Froschaurer)

**Red Bat** (*Lasiurus borealis).* (United States Fish and Wildlife Service image by Gary Peeples)

**Hoary Bat** (*Lasiurus cinereus).* (United States Geological Survey image by Paul Cryan)

**Small-footed Myotis** (*Myotis leibii ).* (United States Fish and Wildlife Service image by Gary Peeples)

**Northern Long-eared Bat** (*Myotis septentrionalis)*. (United States Fish and Wildlife Service image).

Mammals of the Lower Susquehanna River Watershed: Little Brown Myotis — **Little Brown** **Myotis** (Myotis lucifigus).

**Indiana Bat** (*Myotis sodalis).* (United States Fish and Wildlife Service image by Ryan Hagerty)

Skull of an "Allegany Long-eared Bat" — James W. Gidley described this skull from the Cumberland Bone Cave as Plecotus alleganiensis, the **“Allegany Long-eared Bat”**, an extinct Middle Pleistocene species. (Smithsonian National Museum of Natural History image www.si.edu)

Order-Carnivora

Family-Felidae

Smilodon fatalis (Saber-toothed Cat)-extinct/prehistoric-C

Smilodon gracilis (Gracile Sabertooth)-extinct/prehistoric-K,H

Panthera onca (Jaguar)-extirpated, Federally Endangered-K,H,C

Felis catus (Domestic Cat)-feral

Felis concolor (Cougar)

Felis concolor couguar (Eastern Cougar)-extirpated, Federally Endangered

Herpailurus yagouaroundi/Felis eyra (Jaguarundi)-extirpated-K

Lynx canadensis (Lynx)-extirpated/hypothetical

Lynx rufus (Bobcat)

Lynx rufus rufus-(“Eastern Bobcat”)

Lynx rufus calcaratus (Irvingtonian Bobcat)-extinct/prehistoric-K

Miracinonyx inexpectatus (North American Cheetah)-extinct/prehistoric-K,H,C-originally listed at Port Kennedy by Edward Cope as Crocuta inexpectata and believed to be a species of hyaena.

Uncia atrox (a lion)-extinct/prehistoric-C

Skull of a Sabre-toothed Cat — A Sabre-toothed Cat (Smilodon fatalis) skull collected in Texas. Early in the twentieth century, fragments of S. fatalis bones were found among deposits in the Cumberland Bone Cave. (Smithsonian National Museum of Natural History image www.si.edu)

Jaws of a Gracile Sabretooth — Edward Drinker Cope based his identification of extinct **Sabre-toothed Cats** (Smilodon species) found in the Port Kennedy Bone Cave on visual analysis of fragments of the extraordinarily large teeth. Cope listed Smilodon gracilis and Uncia (Smilodon) mercerii as the sabre-toothed cats among the remains at Port Kennedy. Today, the latter is considered a junior synonym to the former. In the 1980s, S. gracilis was found among the remains in the Hanover Quarry #1 Fissure. This example of a set of S. gracilis jaws is a reconstruction consisting of specimens from the Port Kennedy Bone Cave on loan to the Valley Forge National Historical Park from the Philadelphia Academy of Natural Sciences. (National Park Service image)

The bones of the **Jaguar** (Panthera onca) were among Middle Pleistocene remains found in the Port Kennedy Bone Cave, the Hanover Quarry #1 Fissure, and the Cumberland Bone Cave. The endangered Jaguar has long been extirpated from the lower Susquehanna valley. Today, it is only rarely found north of the United States/Mexico border. (United States Fish and Wildlife Service image by Gary Stotz)

Mammals of the Lower Susquehanna River Watershed: Domestic Cats — **Domestic Cats** (*Felis catus).*

Mammals of the Lower Susquehanna River Watershed: Cougar/Mountain Lion — **Cougar** (Felis concolor).

**Jaguarundi** (*Herpailurus yagouaroundi).* (United States Fish and wildlife Service image)

Mammals of the Lower Susquehanna River Watershed: Lynx — **Lynx** (*Lynx canadensis).*

Mammals of the Lower Susquehanna River Watershed: "Eastern Bobcat" — **“Eastern Bobcat”** (*Lynx rufus rufus).*

Family-Canidae

Canis species (possible Dire Wolf, C. dirus?)-K

Canis armbrusteri (Armbruster’s Wolf)-extinct/prehistoric-K,C

Canis lupus (Wolf)-Federally Endangered

Canis lupus familiaris (Domestic Dog)-exotic

Canis lupus lycaon (Eastern Wolf)-extirpated

Canis rufus (Red Wolf)-Federally Endangered

Canis rufus var. priscolatrans (Wolf Coyote)-extinct/prehistoric-K,C

Canis latrans (Coyote)-C

Canis latrans var. (Eastern Coyote)

Vulpes vulpes (Red Fox)

Urocyon cinereoargenteus (Gray Fox)-K,H

Dire Wolves — Canine bone fragments from the Port Kennedy Bone Cave believed to possibly be those of the Dire Wolf (Canis dirus) would be twice the age of the currently known oldest remains of the species. (National Park Service image)

Mandible of an Armbruster's Wolf — A partial left mandible of an extinct **Armbruster’s Wolf** (Canis armbrusteri), a species found among the animal remains in the Port Kennedy Bone Cave. This type specimen was collected by James W. Gidley from deposits found in the Cumberland Bone Cave in Allegany County, Maryland, in 1912. Gidley named C. armbrusteri for Raymond Armbruster, who was among the first to notify experts at the National Museum of Natural History of the discovery and significance of the Cumberland Bone Cave. Based on morphology, C, armbrusteri may be the ancestor of the more recent, but nevertheless also extinct, Dire Wolf (Canis dirus). (Smithsonian National Museum of Natural History image www.si.edu)

Mammals of the Lower Susquehanna River Watershed: Wolf — **Wolf** (*Canis lupus).*

The endangered **Red Wolf** (Canis rufus) was extirpated from the lower Susquehanna River watershed sometime prior to the record-keeping of Linnaean Societies and other early naturalists. The only remaining wild population consists of less than two dozen Red Wolves in coastal North Carolina. During the nineteenth century, the remains of a Middle Pleistocene variety, the Wolf Coyote (Canis rufus var. priscolatrans), were collected from deposits in the Port Kennedy Bone Cave. C. rufus is listed among the species found in the Cumberland Bone Cave. (United States Fish and Wildlife Service image by B. Bartel)

Mammals of the Lower Susquehanna River Watershed: Eastern Coyote — An **Eastern Coyote** (Canis latrans var.), “Exhibit A” demonstrating the schizophrenic mindset humans have toward canines.

Mammals of the Lower Susquehanna River Watershed: Red Fox — **Red Fox** (*Vulpes vulpes).*

**Gray Fox** (*Urocyon cinereoargenteus).* (United States Fish and Wildlife Service image)

Family-Ursidae

Arctodus pristinus (Lesser Short-faced Bear)-extinct/prehistoric-K,C

Tremarctos floridanus (Florida Spectacled Bear)-extinct/prehistoric-C

Ursus americanus (Black Bear)-K,H,C

Family-Mephitidae

Mephitis mephitis (Striped Skunk)-K,H,C

Spilogale putorius (Spotted Skunk)-C

Mammals of the Lower Susquehanna River Watershed: Striped Skunk — **Striped Skunk** (Mephitis mephitis).

**Spotted Skunk** (*Spilogale putorius).* (United States Fish and Wildlife Service image by Grayson Smith)

Family-Mustelidae

Brachyprotoma obtusata (Short-faced Skunk)-extinct/prehistoric-K,C

Osmotherium spelaeum (Port Kennedy Skunk)-extinct/prehistoric-K

Gulo gulo (Wolverine)-extirpated/hypothetical-C

Gulo gulo var. schlosseri (gidleyi) (Schlosser’s or Gidley’s Wolverine)-extinct/prehistoric-K

Lontra canadensis (River Otter)–extirpated/introduced-K,C

Martes americana (Pine Marten)-extirpated/hypothetical

Martes pennanti (Fisher)–extirpated/introduced

Martes (Pekania) diluviana (Diluvian Fisher)-extinct/prehistoric-K,C

Mustela species-H

Mustela erminea (Ermine)-extirpated?

Mustela frenata (Long-tailed Weasel)

Mustela nivalis (Least Weasel)

Mustela nivalis allegheniensis (Least Weasel)

Mustela putorious (European Polecat)

Mustela putorius furo (Ferret)-exotic

Neovison vison (Mink)-C

Taxidea taxus (American Badger)-extirpated-K,C

Taxidea taxus marylandica (a badger)-extinct/prehistoric-C

Mammals of the Lower Susquehanna River Watershed: River Otter — **River Otters** (*Lontra canadensis).*

Mammals of the Lower Susquehanna River Watershed: Pine Marten — **Pine Marten** (*Martes americana).*

Mammals of the Lower Susquehanna River Watershed: Fisher — **Fisher** *(Martes pennanti)*. (United States Fish and Wildlife service image)

**Long-tailed Weasel** (*Mustela frenata).* (United States Fish and Wildlife Service image by Tom Koerner)

**Least Weasel** (*Mustela nivalis).* (United States Fish and Wildlife Service image by Steve Hillebrand)

Mammals of the Lower Susquehanna River Watershed: Mink — **Mink** (Neovison vison).

Skull of an Extinct Subspecies of American Badger — James W. Gidley identified these skull elements found in the Cumberland Bone Cave as those of an extinct Middle Pleistocene badger (Taxidea marylandica), now ranked as a subspecies of the **American Badger** (T. taxus). (Smithsonian National Museum of Natural History image www.si.edu)

Bones of the **American Badger** (Taxidea taxus) were among remains recovered from Middle Pleistocene deposits in the bone caves at Port Kennedy and Cumberland. The species no longer occurs in the Susquehanna watershed but is widespread from Ohio westward. (United States Fish and Wildlife Service image by Tom Koerner)

Family-Procyonidae

Procyon lotor (Raccoon)-K

Order-Artiodactyla

Family-Tayassuidae

Mylohyus fossilis (a peccary)-extinct/prehistoric-K,H,C

Platygonus species (a flat-headed peccary)-extinct/prehistoric-H

Platygonus intermedius (a flat-headed peccary)-extinct/prehistoric-C

Platygonus vetus (a flat-headed peccary)-extinct/prehistoric-C

Jaws of an Extinct Pig-like Peccary — Discovered in 1914 in the Cumberland Bone Cave in Allegany County, Maryland, these skull fragments, a lower jaw and maxilla, were identified as those of the extinct pig-like **peccary** Mylohyus fossilis. Earlier, during the late nineteenth century, remains of Mylohyus fossilis were collected from the 500,000-year-old deposits in the Port Kennedy Bone Cave. (Smithsonian National Museum of Natural History image www.si.edu)

Skull of an Extinct Flat-headed Peccary — The skull of an extinct Middle Pleistocene **flat-headed peccary** (Platygonus intermedius) collected in 1914 from the Cumberland Bone Cave by James W. Gidley. The remains of peccary found in the Hanover Quarry #1 Fissure during the 1980s are possibly the same taxon. (Smithsonian National Museum of Natural History image www.si.edu)

Family-Suidae

Suidae (a pig, genera undetermined)-C

Sus scrofa (Pig)-exotic

**Pigs** (Sus scrofa) of European ancestry are occasionally released into the wilds of the lower Susquehanna valley as targets for “canned hunts”. Bones of an extinct native Middle Pleistocene pig have been identified from remains discovered in the nearby Cumberland Bone Cave in Allegany County, Maryland. The exact genera of the specimen(s) is not known. (United States Fish and Wildlife Service image by Steve Hillebrand)

Family-Cervidae

Alces alces (Moose)

Alces alces americana (Eastern Moose)-extirpated/hypothetical

Cervus species (an Elk)-C

Cervus canadensis (Elk)

Cervus canadensis canadensis (Eastern Elk)-extirpated

Cariacus (Dama) laevicornis (a fallow deer)-extinct/prehistoric-K

Odocoileus laevicornis (a deer)-extinct/prehistoric-K

Odocoileus virginianus (White-tailed Deity)-extirpated-K,H,C

Odocoileus virginianus borealis (Northern White-tailed Deity)-extirpated/introduced

Rangifer tarandus (Caribou)-extirpated/hypothetical

Teleopternus orientalis (a large cervid)?-extinct/prehistoric-K

**Moose** (*Alces alces).* (United States Fish and Wildlife Service image by Tom Koerner)

**Elk** (Cervus canadensis). (United States Geological Survey image by Stuart Tomlinson)

Mammals of the Lower Susquehanna River Watershed: Northern White-tailed Deity — **Northern White-tailed Deity** (*Odocoileus virginianus borealis).*

**Caribou** (*Rangifer tarandus).* (United States Fish and Wildlife Service image by Dave Menke)

Family-Bovidae

Bison bison (American Bison)-extirpated

Euceratherium americanum (a shrub-ox)?-extinct/prehistoric-C

Euceratherium collinum (a shrub-ox)-extinct/prehistoric-C

Bos species (a wild cattle)?-extinct/prehistoric-K

Ovis aires x Ovis orientalis (hybrid sheep)-exotic

Mammals of the Susquehanna River Watershed: American Bison — **American Bison** (*Bison bison). (Exhibit: State Museum of Pennsylvania, Harrisburg)*

Mandible of an Extinct Shrub-ox — In 1912, James W. Gidley collected these mandible fragments belonging to an extinct **shrub-ox** (Euceratherium collinum) from remains found in the Cumberland Bone Cave. (Smithsonian National Museum of Natural History images www.si.edu)

Mammals of the Lower Susquehanna River Watershed: hybrid sheep — A hybrid sheep (Ovis aires x Ovis orientalis) released for a “canned hunt” along the Susquehanna.

Order-Perissodactyla

Family-Equidae

Equus species (a horse)-extinct/prehistoric-C

Equus complicatus (Complex Tooth Horse)-extinct/prehistoric-K,H

Equus fraternus var. pectinatus (a horse)-extinct/prehistoric-K

Family-Tapiridae

Tapirus copei (haysii) (Cope’s Tapir)-extinct/prehistoric-K,C

Edward Drinker Cope (seen here circa 1880-1890) and his friend, paleontologist Charles Wheatley, were the early investigators of the contents of the Port Kennedy Bone Cave. (Smithsonian National Museum of Natural History image www.si.edu)

James W. Gidley at the Cumberland Bone Cave — Paleontologist James W. Gidley of the Smithsonian Institute at the Cumberland Bone Cave, circa 1912. (Smithsonian image www.si.edu)

SOURCES

Carnegie Museum of Natural History—Mammals of Pennsylvania Online Resource. https://web.archive.org/web/20110927151038/http://www.carnegiemnh.org/mammals/PAmamm/pamammals2.html Accessed April 14, 2020.

Cope, Edward D. 1871. “Preliminary Report on the Vertebrata Discovered in the Port Kennedy Bone Cave”. Proceedings of the American Philosophical Society. 12:86 pp.73-102.

Cope, Edward D. 1895. “The Fossil Vertebrata from the Fissure at Port Kennedy, Pa.”. Proceedings of the Academy of Natural Sciences of Philadelphia. Vol.47. pp.447-450.

Cope, Edward D. 1896. “New and Little Known Mammalia from the Port Kennedy Bone Deposit”. Proceedings of the Academy of Natural Sciences of Philadelphia. Vol.48. pp.378-394.

Daeschler, Edward, Spamer, Earle E., and Parris, David C. 1993. “Review and New Data on the Port Kennedy Local Fauna and Flora (Late Irvingtonian), Valley Forge National Historical Park, Montgomery County, Pennsylvania”. The Mosasaur. Vol.5. pp.23-41.

Gidley, James W. 1913. “Preliminary Report on a Recently Discovered Pleistocene Cave Deposit Near Cumberland, Maryland.” Proceedings of the United States National Museum. Vol.46. pp.93-102.

Guilday, J. E., Cotter, J., Cundall, D., Evenson, E., Gatewood, J., Morgan, A. V., Morgan, A., McCrady, A., Peteet, D., Stuckenrath, R., and Vanderwal, K. 1984. “Paleoecology of an Early Pleistocene (Irvingtonian) Cenote: Preliminary Report on the Hanover Quarry No. 1 Fissure, Adams County, Pennsylvania”. In W, C, Mahaney (ed.) Correlation of Quaternary Chronologies. Geo Books. Norwich, England. pp.119-132.

Haldeman, Samuel Steman. 1844. “Natural History of Lancaster County: Mammalia—Beasts”. I. Daniel Rupp’s History of Lancaster County. Gilbert Hills. Lancaster, PA. pp.470-471.

Maryland Natural Heritage Program. 2016. List of Rare, Threatened, and Endangered Animals of Maryland. Maryland Department of Natural Resources, 58 Taylor Avenue, Annapolis, MD 21401.

Merritt, Joseph F. 1987. Guide to the Mammals of Pennsylvania. University of Pittsburgh Press. Pittsburgh, PA.

Pennsylvania Natural Heritage Program. PNHP Species Lists. www.naturalheritage.state.pa.us/Species.aspx Accessed April 14, 2020.

Rathvon, Simon S. 1869. “Class Mammalia”. J. I. Mombert’s An Authentic History of Lancaster County. J. E. Barr and Company. Lancaster, PA. pp.500-501.

The Paleobiology Database website. https://paleobiodb.org Accessed August 5, 2020.