Lath Pleistocene Cave Deposit in Bedford County, Pennsylvania

BULLETIN of THE NATIONAL SPELEOLOGICAL SOCIETY

VOLUME 26, NUMBER 4 OCTOBER 1964

CONTENTS

NEW PARIS No. 4: A PLEISTOCENE CAVE DEPOSIT IN BEDFORD COUNTY, PENNSYLVANIA - - - - - John E. Guilday, Paul S. Martin, Allen D. McCrady 121

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ASSOCIATE EDITORS THOMAS C. BARR, JR. THOMAS L. POULSON JOHN V. THRAILKILL Department of Zoology Department of Zoology Department of Geology University of Kentucky Yale University Princeton University Lexington, Kentucky New Haven, Connecticut Princeton, New Jersey

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LIBRARY National Speleological Society Library Attention: MRS. JULIA L. STANILAND, Librarian 1251 North Negley Avenue Pittsburgh, Pennsylvania 15206 NEW Paris NIL 4: It Lath Pleistocene Cave Deposit In Bedford County, Pennsylvania

By John E. Guilday*, Paul S. Martin**, Allen D. McCrady*

ABSTRACT

Excavation of a 10-meter column of surface-derived matrix from Sink- hole No. 4 at New Paris, Bedford County, Pennsylvania produced a late Pleistocene biota: over 2,700 vertebrates, with an accompanying pollen profile. C-14 date is approximately 11,300 B.P. The indicated environment is cool taiga parkland during the early phase of infill with progressive reforestation during a subsequent warming (but still boreal) period.

CONTENTS

Introduction 122 Reptiles 142 Acknowledgements 122 Birds 143 History of Investigation 125 Mammals 144 Insectivora 144 Geology 126 Chiroptera 151 Geography of Area 126 Rod entia 152 Lagomorpha 168 History of Excavation, Materials Carnivora 169 and Methods 127 Artiodactyla 171 Procedure 129 The Late-Glacial Environment of Unglaciated Pennsylvania 174 Abbreviations 131 Introduction 174 Bone Analysis 131 Faunal Change 175 The Bergmann Response 179 Carbon-14 Dates 132 Fossil Pollen, Small Vertebrates and the Late-Glacial Climate 181 Faunal List 133 The Prairie Peninsula Problem ..... 183 Early Man in the Eastern Pollen Analysis 135 Late-Glacial 185 Invertebrates 141 Conclusions 187 Amphibians 141 Bibliography 189

* Carnegie Museum ** University of Arizona

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 121 INTRODUCTION in Pleistocene paleoecology, that of separating (whenever possible) boreal forest from The excavation of Sinkhole No. 4 near boreal woodland (see Martin, 1959). New Paris, Pennsylvania, provides the first detailed record of late-glacial range adjust- The term open boreal woodland is ap- ments made by many boreal species of birds plied by Hare to the type of tree growth and mammals in the late Pleistocene period seen in the foreground of 0. J. Muries's of eastern North America. Excavation of about photo of Great Whale River (fig. 5) in which "tall and well-developed spruce (more rarely 10 meters of unstratified mud and rubble fill from the sinkhole produced skeletal re- other conifers) stand scattered in a sea of ( mains of at least 2,769 vertebrates associated Cladonia." Hare, 1950, p. 624). This is a with plant pollen and carbon. very different environment from the more familiar boreal forest to the south, where In the modern environmental gradient of more than 90 per cent of the spruce pulp- eastern North America a zoo-geographer may wood of Canada is cut. While the same spe- recognize (from north to south) Arctic- cies of trees may dominate in both the wood- Alpine, Hudsonian, Canadian and Austral land and the forest, they form a closed can- life-zones corresponding respectively to the opy, grow more rapidly, and to a greater tundra, boreal woodland, boreal forest, and height in the latter. deciduous forest of the ecologist. A major In the fossil pollen record criteria for dis- objective in current studies of the Pleisto- tinguishing boreal forest from boreal wood- cene has been to determine the fate of these land are not clearly established; even tundra environments. Were they dissolved, amalga- mated, or otherwise drastically altered? Did may be difficult to recognize from the pollen record alone. Fortunately, certain of the small they endure, albeit compressed and somewhat modified, during glaciation? What can the vertebrate remains found in such abundance in Pleistocene fossil record tell us about the New Paris No. 4 appear to be sensitive indi- nature of plant and animal communities be- cators of tundra, woodland, and forest. The yond the ice sheet? vertebrates make it possible to determine more precisely the nature of the late-glacial environ- Throughout our text we will attempt to ment in Pennsylvania than would be possible make a distinction we believe is important from pollen evidence alone. Morphological and ecological analysis of the mammals represents the contribution of Guil- day; the excavation proper and its description is the contribution of McCrady, while the results of pollen analysis are presented by Martin. A chapter on reptiles and amphibians is provided by Neil D. Richmond. Our concluding state- ment on "Late-glacial environment and life in unglaciated Pennsylvania" represents a joint effort. We recognize that the Pleistocene fossil record from the unglaciated East is highly fragmentary and poorly dated. Our conclusions may wither in the face of more definitive fossil Figure 1. evidence or more perceptive ecological insight. Map of east-central United States. Hachur- Certain contradictions remain to be resolved ed line delimits approximate southern lim- through analysis of yet undiscovered late Pleis- it of Wisconsin glaciation. NP = New Paris tocene fossil deposits. No. 4, Pa. BS = Bootlegger Sink, Pa. M = Marsh, Pa. CC = Cumberland Cave, Md. ACKNOWLEDGEMENTS NC = Natural Chimneys, Va. CG = Cran- berry Glades, W. Va. RC = Robinson Cave, We are indebted to Mr. and Mrs. Oscar Tenn. Miller and their children, the owners of the

122 THE NATIONAL SPELEOLOGICAL SOCIETY Figure 2. Northern Bedford County, Pennsylvania. a. Eastern rim of Appalachian Plateau, Allegheny Mountain. b. Dunning Mountain. c. Bedford, Pa. NP =New Paris, Pa. Base Map = U. S. Army Corps of Engineers NK 17- 1 2. Series V501 P. Grid = 10 km. site, who have been wonderful and gracious Barton, the late Martin S. Bender, Ralph C. to the steady stream of cavers who have de- Bossart (field supervisor), Susan Booth, John scended on them through the years. Devlin, Mr. and Mrs. Gerald Frederick, Bruce We thank the following people who donated Godwin, Daniel Green, Mr. and Mrs. Allan time and varied talents to the excavation of P. Haarr, Roscoe Hall, Mr. (field supervisor) New Paris No. 4. Without them this project and Mrs. Harold W. Hamilton, Mr. and Mrs. would have been impossible. We can only hope George Howard, William Hunter, Mr. and that other institutions will be so blessed with Mrs. Paul Imblum, W. Roswell Jones, Viera such a capable field force: Susan Allardice, Kulamer, John B. Leppla, John A. Leppla Kenneth Acklin, C. Dale Acklin, Charles Bar- (field supervisor), Ann Montgomery, Jerry ton, Hugh Barton, Mr. and Mrs. W. Galen Paulich, Frank Potter, Monica Rectenwald, BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 123 Figure 3. Looking west from New Paris sinkholes toward Allegheny Mountain. Miller house, center foreground. Dunning Creek appears as wooded corridor running from left to right beyond farm.

Figure 4. Looking east across valley of Dunning Creek to Chestnut Ridge. Arrow points to site of New Paris sinkholes.

Figure 5. Coniferous forest with tundra intervals. Northern edge of Hudsonian 0 life-zone. Great Whale River, lat. 55 17 NI., Quebec, Canada. Simi- lar conditions prevailed at New Paris during the Wisconsin glaciation about 12,000 years ago. From Todd, 1963.

124 THE NATIONAL SPELEOLOGICAL SOCIETY John Reiglci MI'. and Mrs. Edmund Taylor, secretary, Carnegie Museum, whose patience 'Flank Vaichulik, Marjorie Shaw, Valerie and abilities were indispensable to this project. whitehead. We wish to thank Dr. J. Kenneth Doutt, We wish to thank William J. Houston Co.; Miss Caroline Heppenstall, Dr. Craig C. Black, Locke Stove Co.; Morris Knowles, Inc.; R. A. Dr. Mary Dawson, Neil D. Richmond, Harry McCiatly, Esq; Richard Stone; the Pennsyl- K. Clench, Dr. M. Graham Netting and Dr. vania Electric Company; the Pennsylvania Game Claude W. Hibbard for reading and criticizing Commission; the Aluminum Company of portions of this manuscript. America; the Phoenix Memorial Laboratory, Art work was by Richard Lang (figs. 13, University of Michigan; Geochronology Lab- 18, 19, 22, 26), Monica Rectenwald (fig. 6), oratories, University of Arizona; Geochrono- and Donald P. Tanner (figs. 1, 7, 12, 14, metric Laboratory, Yale University; Biology 15, 27 through 34). Photography was by Department, University of Minnesota; Depart- W. Galen Barton (figs. 2, 3, 4, 10, 16, ments of Mammals and of Vertebrate Paleon- 17, 20, 21, 24, 25), John A. Leppla (figs. 8, tology, American Museum of Natural History; 11, 23), M. Graham Netting (fig. 9 ) and Olaus Divisions of Birds, of Mammals and of Verte- J. Murie.(fig. 5). The color plates, center leaf brates Paleontology, United States National are by Harry K. Clench and A. D. McCrady. Museum; the Academy of Natural Sciences We wish to thank Clifford J. Morrow, Chief, of Philadelphia; and the Pittsburgh Grotto of Special Exhibits Staff, Carnegie Museum for the National Speleological Society for their technical advice. We wish to thank W. E. generous gifts and/or services. Clyde Todd, Curator Emeritus of Birds, Car- We thank the following specialists for their negie Museum, for permission to publish help in identification or consultation: Dr. Nell fig. 5. B. Causey, University of Arkansas (millipeds); There are two people who must be men- Juan J. Parodiz, Carnegie Museum (snails); tioned with special feeling on the part of the Dr. Alexander Wetmore, United States Na- senior author: Dr. J. LeRoy Kay, Curator tional Museum (birds ); Dr. Paul W. Parmalee, Emeritus of Vertebrate Fossils, Carnegie Mus- Illinois State Museum (birds ); Dr. Kenneth C. eum who saw the project off to a successful Parkes, Carnegie Museum (birds ); Dr. J. Ken- start, and Mrs. Alice M. Guilday who guided neth Doutt, Carnegie Museum (mammals ); Dr. it between the pages of this report. Jane Gray, University of Oregon (pollen); This research was made possible by grants Dr. Joseph Tihen, University of Notre Dame from the A. W. Mellon Educational and Chari- (amphibians ); Neil D Richmond, Carnegie table Trust and National Science Foundation Museum (reptiles and amphibians ); Dr. Ernest grant no. 20868. L. Lundelius, Jr., Texas Memorial Museum ( Mylobyus); Dr. William H. Burt, University of Michigan (mammalian distributions); Dr. HISTORY OFINVESTIGATION Ralph M. Wetzel, University of Connecticut The New Paris sinkholes were initially in- (Synaptomys). vestigated in 1932 by Charles E. Mohr, now We thank Dr. James B. Griffin, University Director of Kalamazoo Nature Center and of Michigan and Dr. Edward S. Deevey, Yale were described by him in Stone, 1932. In April, University for carbon-14 dates; Dr. Louis 1948, members of the Pittsburgh Binford, University of Chicago for thorium Grotto, National Speleological Society analysis; and Dr. Nicholas M. Short, U.S. (N.S.S.), revisited the site and secured a partial elk (Cervus canadensis) skeleton from Bureau of Mines for advice on clay structure. Sinkhole No. 2. Carnegie Museum, under a The roster of museum colleagues that have grant from the A. W. Mellon Educational assisted through the project includes them all. and Charitable Trust, excavated Sinkhole Na, We shall single out only Albert C. Lloyd (the 2 (or Elk Hole) in 1949 and 1950. A Recent "Lloyd's Rockhole" Lloyd) for years of help fauna of 26 species of mammals was recovered and encouragement and Joseph Y. Quil, clerical (Guilday and Bender, 1958). In August of

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 125 1956 the Pittsburgh Grotto (N.S.S.), working is further enlarging the pits. Infilling is ex- in conjunction with Dr. J. LeRoy Kay, Cura- tremely slow at the present time due to the tor Emeritus, Section of Vertebrate Fossils, thin soil mantle which averages 0.5 meters in Carnegie Museum, began the excavation of thickness. Infilling may have been more rapid Sinkhole No. 4, or "Lloyd's Rock Hole." during late Pleistocene times under the in- Excavation continued until March of 1963 fluence of frost-wedging. when sterile cave-derived sediments were en- A skeleton of a Recent timber wolf (Canis countered at a depth of 9.1 m and bedrock lupus) from New Paris No. 2 yielded an age 0.6 m lower. of 1,875 ± 100 years B.P. (1-743). The animal was buried beneath about 2 meters of mud and GEOLOGY rubble. This date provides a crude estimate of Sinkhole No. 4 or Lloyd's Rock Hole (2.4 the rate of infill, about 1 meter per 1000 years. km [1.5 mi.[. northeast of New Paris, West At the same rate the 9.1 meters of fill in New St. Clair Township, Bedford County, Penn- Paris No. 4 would have required some 10,000 sylvania; lat. 40° 5' N., long. 78° 39' W., years. Internal evidence suggests that the rate alt. 465 m [1500 feet]. See map, Guilday of infilling of New Paris No. 4 was more rapid and Bender, 1958), is one of many along the during the initial phase of accumulation when west flank of Chestnut Ridge*, an anticline the local climate was colder, and slower dur- of upper Silurian-lower Devonian marine lime- ing the terminal phase. Accepting Y-727 (11,- stone and sandstone (fig. 4). The ridge itself 300 ±1,000 B.P.) as a reliable date for the is capped by the Lower Devonian Oriskany 5.7 m level; the upper half of the cave fill sandstone. Streams draining the ridge have accumulated at a minimum rate of 0.5 m formed small coves that locally have exposed per thousand years. the underlying Helderberg limestone. Ground water circulation, locally governed by the joint GEOGRAPHY OF AREA system of the limestone, has produced a series of domepits along the joint structure. Some Chestnut Ridge (fig. 4) is the low 488 m of these have intersected the present land sur- anticline upon which the sinkhole is located. face and are filling with surface-derived mat- It lies in an intermontane valley (fig. 2) 30 erial, soils, humus, rocks, plant and animal to 40 km wide and over 160 km long. The remains. Others, as exposed in a local quarry, valley floor is approximately 425 m above sea have not reached the surface. They are in the level. It is bounded on the northwest by form of deep, crack-like caves floored with 885 m Allegheny Mountain (fig. 3 ), the eastern sterile, yellow-brown clay derived from the rim of the Appalachian Plateau, and on the silicious and argillaceous residue of the chem- southeast by a long, narrow 700 m ridge ical breakdown of the parent limestone. Still variously named Bald Eagle, Canoe, Lock, others have had surface connections in the Loop, Dunning, Evitt's, and Will's Mountain past and have been filled to the surface. Some depending upon its local sinuous configura- still show a slight funnel-shaped depression tion and the presence of water and of wind- on the surface of the ground. Others have breaks. Blue Knob, a local outlier of no surface indication. Allegheny Mountain some 19 km north of the Open sinkholes reach a depth of over 26 sinkholes, rises to 956 m — one of the high- meters. Their walls bell out below the surface est points in the state. into typical dome-pit conformation (see Pohl, In addition to its karst drainage, which con- 1955 and Merrill, 1960). Solution fluting tributes to its surface dryness, Chestnut Ridge and relatively insoluble fossil inclusions pro- lies within the rain shadow of Allegheny truding from the moist walls indicate that Mountain. Prevailing winds are from the west solution initiated by humic-rich ground waters and mean annual precipitation may be 25 to 38 cm less east of Allegheny Mountain. ' not to be confused with the more widely known Average annual precipitation at Somerset, 40 Chestnut Ridge 80 km to the west on the Appa- km west of New Paris on the Appalachian lachian Plateau. Plateau (elev. 640 m) is 129.5 cm per year

126 THE NATIONAL SPELEOLOGICAL SOCIETY (Gifford and Whitebread, 1951). A few kilo- determined by the availability of suitable habi- meters east of the Appalachian Pleateau tats and the adaptability of the species." at Hyndman, 40 km south of New Paris in the same intermontane valley, it drops to HISTORY OF EXCAVATIONS, MATERIALS 89.3 cm per year (Kauffman, 1960). A pre- AND METHODS historic rain shadow is also quite likely. Initially, Sinkhole No. 4 resembled at the It is conceivable that the relative aridity of surface a bowl-shaped depression 9 m in diam- the site may have been attractive to jack pine eter, 2 m deep in its center (fig. 8). A 2 x 2 (Pinus Banksiana) during late Pleistocene m shaft (Station 1) was dug in the sinkhole times. This species finds its best development fill directly in the center of the depression at the present time on well-drained sandy (fig. 6). No contact was made with the walls tracts throughout the northern coniferous for- of the sinkhole and square-set shoring was est belt of North America. Contemporaneous- required to a depth of 7.6 m. This shaft pro- ly, the poorly drained rim of the Appalachian duced no bones. At the bottom of the shaft, Plateau 8 km west and 460 m higher may on the west side, a passage (Station 2) led to have supported a tundra/taiga biota. It seems an intersecting dome-pit. Talus from its bottom evident, both from the make-up of the New revealed a rich deposit of late Pleistocene bones Paris No. 4 local fauna and the complicated (Station 3). Excavation of this collapse mater- physiography of the area, that several biotic ial, which had lost its original stratigraphy, zones could live in juxtaposition under a cri- was not conducted by levels. At this point, tical climatic regime. They do so today, but 5.2 m southwest of the original shaft, a sec- only to a limited extent. Snowshoe hares (Le- ond shaft was started from the surface. There pus americanus), for example, formerly oc- was no surface indication of this shaft prior to curred in hemlock bogs along the crest of excavation. Shoring was not required below Allegheny Mountain within 16 km of New ground level because this narrow fissure was Paris (Rhoads, 1903) but not at New Par- separated from the original shaft by a lime- is where the lagomorph fauna during Recent stone wall. The excavation was rich in organic times contained only cottontails ( Sylvila- inclusions (bone, pollen, charcoal) to a depth gus floridanus and S. transitionalis). Ca- of 9.1 in, below which it was abruptly under- nadian life-zone conditions are suggested in laid by sterile yellow cave-derived sediments a few places, such as the Allegheny Moun- which could be traced into the original shaft tain crest, but the biota is generally transi- demonstrating that the two shafts were origi- tional in character. nally part of the same subterranean system (figs. 10 and 11). Gifford and Whiteb read (1951, p. 18) states: "The part of this sector (south-central Penn- The large doline represented by the surface sylvania) that lies south and east of Blue Moun- depression was a massive collapse phenomenor tain (Dauphin County) would properly be apparently the result of the failure of the ceiling considered part of the Carolinean Biotic of a large cave room. Surface fill rapidly slump- Province of Dice ( 1943 ). While the areas along ed in from the large feed-cone at the surface. the Allegheny Front ... are most like the south- This collapse was apparently too large in diam- ern portion of the Canadian Biotic Province of eter and consequently too gentle in slope to Dice. However, the Ridge and Valley section function as an efficient trap. The small dome- generally is difficult to assign to either of pit (Station 3) on the other hand, had a these broad classes. Although it has many of restricted surface opening due to its narrow the features of a transitional zone between width. With a small debris-cone, a slow rate the two biotic provinces, it is also an area of infilling and vertical walls, it served as an where the distribution of plants and animals efficient trap (fig. 6). is strongly influenced by the local and varied There was no apparent internal stratigraphy topography and soils with the result that the to the 9.1 m column of debris. Deposition distribution of many of the plants and animals took the form of a conical talus so that an found in this area appear to be primarily object falling in may have gone no further, BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 127 11111111111111111II

3 4

tripod

II ILION II 11 11 11 a 11 a MI ,E1

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BON 1111,-11 MORI MINI I

uL4 AMMO AMMO 1 41 11 1,11:0 1,1.4•Wonoonarmuum 1 111111,14,, 1[11

Figure 6. Postulated steps in the history of the New Paris No. 4 bone deposit. Black = bedrock. Gray = cave-derived sediments. Hatched = surface-derived materials. B = bones, etc. or may have worked its way down slope considerable slope or slumpage. On the other or deep into a crack. Remains Of the one hand many of the skeletons recovered were peccary (Mylohyus) recovered, although in uncrushed and still in articulation, indicating a semi-articulated condition, were scattered that little if any post-depositional disturbance ovet a vertical distance of 1.5 m indicating had occurred. A few rodent-gnawed snow- 128 THE NATIONAL SPELEOLOGICAL SOCIETY I 33. 303: 3130333(\33 \

.W3 21' 133\\VVRIE 22 22 24 4. 25 a. 26 • a 27 :\N3 - 28 3 6 TI 9 29 30

Figure 7. Figure 9. Percentage of total catalogued bones and Sinkhole No. 4, New Paris, Pa. Washing teeth recovered per level in Station 3, and screening. Coarse screen above, fine Sinkhole Na. 4, New Paris, Pa. White= screen below. Water recirculated by pump. Microtinae. Bla•' = Chiroptera. Hatched= others.

Figure 8. Sinkhole No. 4, New Paris, Pa. Original test shaft, Station 2, view southwest. shoe hare bones and the recovery of a concen- Figure 10. tration of rodent droppings containing a Sinkhole No. 4, New Paris, Pa. Looking modern pollen flora (see p. 137) indicate that from surface to the 7.0 m level. Excava- there must have been some disturbance. De- tion in progress, view southwest. spite the homogeneous nature of the fill, and the uneven deposition, a convincing faunal and floral succession can be demonstrated. Similar PROCEDURE sites are known from the late-Wiirm of eastern Excavation of the original test shaft and the Europe and demonstrate parallel ecological connecting passage (Stations 1 and 2) pro- replacement. Different species are involved ceeded with no stratigraphic controls. As soon of course but the similarity is too close to as the bone deposit was encountered and the be ignored. second surface shaft (Station 3) was begun,

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 129 smaller than a chipmunk were recovered on the washing screens rather than during excavation. Cataloguing was conducted with the objective of establishing a species census for each 0.3 m level. All skulls, all maxillae, all mandibles and all first lower molars (in the cricetids and zapodids) were catalogued by stratigraphic position. Skeletal material of animals that were hare-sized or larger was also catalogued. Isolated bones of forms already known from cranial parts were not catalogued as a rule. No bone has been discarded or destroyed (except a small sample for C-14 Figure 11. and thorium analysis) but is stored with the Sinkhole No. 4, New Paris, Pa. Contact collection boxed by depth level. Catalogue between surface-derived fossiliferous mat- information was placed on punch cards and erial above and sterile, cave-derived sedi- analyzed stratigraphically. Census percentages ments below. Southwest face, 9.1 m level. are based upon minimum number of individuals per level. This was based solely upon skulls strict stratigraphic control of all excavated or skull parts, and may not coincide with the matrix was initiated. The roughly circular minimum number of individuals present in shaft was divided into four quarters. Matrix the whole deposit. The total number of yellow- was removed in 0.3 m (one foot) levels. cheeked voles (Microtus xanthognathus), Labeled lots of matrix were placed on drying for instance, is derived from a count of all racks consisting of an upper screen of 6 mm lower first molars of that form that were mesh and a lower screen of 1.5 mm mesh. recovered. But two other species of the genus, When the matrix was dry enough to break the the rock vole (M. chrotorrhinus) and the colloidal bonds of the mud, it was hosed meadow vole (M. pennsylvanicus), cannot through these two grades of screens (fig. 9) be told apart by the morphology of the lower and any bones or teeth placed in vials with the first molar. The skulls, or the upper denti- proper stratigraphic data. Picking of the finest tions of all three forms are separable, and their grade of matrix was carried out largely without use in establishing percentages by level gives the aid of magnification. It is certain that a a truer relative picture than if lower teeth of small percentage of microfauna was not re- the first and upper teeth of the second and covered that could have been picked up by third form were employed. Skulls or maxillae magnification. The large number ofspecimens of the two bats of the genus Myotis present recovered, and the meaningful percentages are readily separated, but mandibles are diffi- derived from them, together with the excellent cult to identify to species in many cases with preservation and the thoroughness of the any real assurance. Due to the presence of search, all lead us to believe that little of any so many well-preserved bat skulls this was consequence to the project was missed. This not attempted. There were more sets of mandi- would not be true where the objective was bles than there were skull parts recovered, morphological reconstruction of poorly- so that the minimum number of bats of the known or new forms, but the primary objec- genus Myotis (based upon mandibles) is tive here was establishing percentage relation- greater than the sum of the individual species ships in a numerically rich and morphologi- of that genus (based upon skull parts). This cally well-known fauna. A minimum of 125 is true for some other groups as well. The tons of matrix were processed. mammal, bird and reptile remains are cata- It was possible in some instances to recover logued in the Section of Vertebrate Fossils, articulated skeletons even of some ofthe smaller Carnegie Museum: Accession numbers 18786 shrews. These were excavated and preserved and 19109; catalogue numbers 5440-5503, as such. But almost all of the specimens 5505-5515, 5517-5931, 6004-6386, 6647-

130 THE NATIONAL SPELEOLOGICAL SOCIETY

7890, 7948-7953. Gastropods are in the snowshoe hare bones from an animal furnished Section of Recent Invertebrates, Carnegie Mu- by the Pennsylvania Game Commission. The seum. Diplopods were retained by Dr. Nell modern bones were degreased in benzene, the Causey, University of Arkansas. fossil sample cleared of matrix in a water bath. Both samples were leached in a 50 ml. bath of hot HCL for 30 minutes. Samples AB13121.V1ATIONS were then ashed and a qualitative spectro- The following abbreviations are useded scopic analysis run. Results are summarized throughout this paper. below:

ANSP Academy of Natural Sciences, Phila- Fossil sample Modern sample

delphia. Weight of bones 6.8351 gm. 6.5996 gm. Weight after leach 2.2053 2.8923

CM Carnegie Museum (Section of Verte- % of original bone after leach 43.8% 32.3% Weight of ash of bones 1.9444 0.6895 brate Fossils ). % of original bone after ashing 28.5% 10.4% CM mammal no. Carnegie Museum (Section

of Mammals ). Elements detected in ash

UMMP University of Michigan, Museum of Relative concentration Leach of Old bones Leach of New bones PaleOntology. old bones after leach new bones after leach

USNM United States National Museum. 10 to 100% Calcium Calcium Calcium Calcium Silicon TMM Texas Memorial Museu m. Aluminum

BP Before present. Phosphorus Phosphorus Phosphorus Phophorm Canine tooth Iron Potassium d. Deciduous dentition .1 to 1% Aluminum Sodium Magnesium Sodium Incisor tooth Iron Magnesium Sodium Magnesium Sodium Titanium Potassium Molar tooth Strontium Magnesium Premolar tooth Strontium

NAP Non-arboreal pollen .01 to .1% Barium Strontium Barium Iron Sample size POLIS5i11111 Barium Iron Strontium Manganese Rubidium Barium Sample mean Boron Potassium Lithium Zinc o' Standard deviation Zircon' Silicon Lead Coefficient of variation Aluminum O.R. Observed range. In,, than .10% Boron Mangsmise Silicon Boron Lithium lciad Lead Lithium The following cave faunas from the central Copper Ytterbium Aluminum Manganese Nickel Vanadium B Copper Appalachian Region are mentioned repeatedly Chromium Gallium Manganese Chromium Lead Chromium Copper Nickel in the text. The references are listed here to Copper Nickel avoid needless repetition. Beryllium

.eh, 500 C 81.42% 88.17% 95.77% 23.04% Cave Fauna Primary Reference

Port Kennedy Cave, Pennsylvania Cope, 1899

Frankstown Cave, Pennsylvania Peterson, 1926

Hartman's Cave, Pennsylvania Leidy, 1889 The fossil bones are much higher in alumi- New Paris No. 2, Pennsylvania Guilday and Bender, 1958

Bootlegger Sink, Pennsylvania Guilday, Hamilton, McCrady, num and silicon than their modern counter- in press parts, with a percentage rise of from less than Hay, 1920 Cavetown, Maryland Cumberland Cave, Maryland Gidley and Gazin, 1938 one-tenth of one percent to over 10 percent.

Robinson Cave, Tennessee McCrady and Schmidt, 1963 Iron and potassium showed a comparable in- Guilday, 1962 Natural Chimneys, Virginia crease. Traces of the original clay matrix infiltrating the bone may be responsible for some QUALITATIVE CHEMICAL ANALYSIS OF BONE of this increase. Elements present in the fossil Snowshoe hare bone from the sinkhole was bone but not recorded from the modern submitted to the Alcoa Research Laboratories, sample include titanium, rubidium, zirconium, Aluminum Company of America, New Ken- vanadium, ytterbium, gallium, and beryllium. sington, Pa., for qualitative chemical analysis. The fluorine in modern hare bones was 0.25 This was compared with a sample of modern percent, in the fossil sample, 1.69 percent.

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 131 The fossil sample was lower in water and properties for soft, spongy bone as opposed to organic constituents, with a total loss on hard, compact bone. These findings make it ashing of 16.3 percent in weight compared imperative that in further investigations de- with a 35.8 percent weight loss of modern signed for answering questions about deposi- hare bone. tional or hydrological history of deposit speci- RADIOMETRIC ANALYSIS OF BONE mens of the same species and preferablybones Selected samples of snowshoe hare long of the same anatomical part be used for com- bones and vertebrae distributed from the 3.0 parisons." (Binford [1960], p. 7-8). m to 5.8 m levels, and a sample of the clay matrix from the 6.1 m level, were submitted CARBON - 14 DATES, NEW PARIS No. 4 for radiometric analysis to Dr. Lewis H. Carbon-14 dating tests were run by two Binford, then with the Phoenix Memorial laboratories, one on charcoal particles, pre- Laboratories, University of Michigan. The sumably derived from a forest fire, by Yale results will be published elsewhere, but a brief Geochronometric Laboratory through the summary is presented here. Two problems courtesy of Dr. Edward S. Deevey; the other were involved: 1. Did bone samples from var- on bone, primarily Lepus americanus, by ious stratigraphic levels show significant dif- Michigan Memorial Phoenix Project Radio- ferences in terms of beta radiation that would carbon Laboratory, University of Michigan bring any evidence to bear on the rate of the through the courtesy of Dr. James B. Griffin. infilling of the sinkhole? 2. Since samples of

both vertebrae and limb bones were submitted Depth from each level, did these two types of bones, Sample from surface Date Y-727 charcoal 4.6 m 11,300 ± 1,000 yrs. B.P. one cancellous, one dense, absorb radioactive M-1067 bones, small mammals 5.2 m — 6.4 rn 9,540 500 yrs. B.P. thorium salts at the same rate? Binford concluded: Dr. Binford (letter of Dec. 28, 1960) "The radiometric analysis of bone and soil commented on the above dates: specimens from Lloyd's Rock Hole, (Sinkhole No. 4), has revealed that there is no real "The observed difference, 1760 years, can be difference in the beta radiation characteristic expressed in units of the standard error — of bone specimens from levels 15 (5 m) 1760/1118.03 = 1.57 standard errors (a val- through 19 (6.3 m) (this is also probably ue expressed in this standard form is called true as high as level 10 [3.3 m1). The distri- "t"), and from a table of areas of the nor- bution of the specimens of long bone from mal curve or of "t" for infinite degrees of these levels was found to be a normal curve freedom the probability of obtaining so large with a negative skew. Although no conclusive a value can be read off. demonstration was possible, this was interpreted as indicative of a slight shift in the rate "In your case the probability is greater of accumulation of the deposit, accumulation than 10 per cent. That is, you would expect having occurred rather rapidly for the mass of to obtain so large a discrepancy, more than the deposit with a diminished accretion over a ten out of a hundred separate runs on the period of time at least as long as that required identical specimen. This information, plus for the original deposition of the mass of the fact that the Michigan date was a short run, the deposit. suggests that both dates are estimates of a single "true date" and that it is likely that the "In addition to the findings relative to the in- "true date" is closer to the 11,300 estimate terpretation of the deposits in Lloyd's Rock than to the 9,540 estimate." Hole, it was found that long bones and vertebrae, both from Lepus americanus and taken from the same levels, represented distinct pop- CARBON - 14 DATES, NEW PARIS No. 2 ulations with respect to beta count. These A carbon-14 date is now available for Sink- findings were interpreted, at least for the time hole No. 2, 183 m to the east of Sinkhole No. being, as indicative of differential absorptive 4. The fauna has been reported on by Guilday

132 THE NATIONAL SPELEOLOGICAL SOCIETY and Bender, 1958 and is Recent in age. Samples Striatura ferrea Morse of wolf (Canis lupus) limb bones from this Ventridens intertextus Binney 1,875± 100 deposit were dated at yrs. B.P., STROBILOPSIDAE sample no. 1-743 by Isotopes, Inc. Strobilops aenea Pilsbry FAUNAL LIST, NEW PARIS NUMBER 4 ARTHROPODA CHORDATA Diplopoda (identifications by Dr. Nell B. Amphibia (identifications by Neil D. Rich- Causey) mond) Dixidesmus or Pseudopolydesmus sp. CAUDATA Scytonotus sp. AMBYSTOMIDAE Abacyon sp. Ambystoma sp. salamander

AIOLLUSCA PLETHODONTIDAE (identifications by Juan J. Parodiz) Pletbodon cf. glutinosus (Green) GASTROPODA slimy salamander CIONELLIDAE Diemictylus cf. viridescens (Rafinesque) Cionella lubrica (Mueller) red-spotted newt Desmognathus sp. HAPLOTREMATIDAE salamander Haplotrema concava (Say) Eurycea? VALLONIIDAE salamander ValIonia costata (Mueller) SALIENTIA Minimum No. SUCCINEIDAE Bufo cf. americanusamericanus) of Individuals Succinea ova/is Say (Holbrook) Succinea avara Say American toad 74 Bufo cf. americanus copei POLYGYRIDAE (Yarrow and Henshaw) Triodopsis albolabris Say Hudson Bay toad Triodopsis denotata (Ferussac) Hyla cf. crucifer Wied Triodopsis tridentata (Say) spring peeper Mesodon tbyroideus (Say) Rana cf. pipiens Schreber Stenotrema birsutum (Say) leopard frog 66 Stenotrema stenotrema Ferussac Rana sylvatica (LeConte) Stenotrema fraternum cavum (Pilsbry wood frog Vannata) REPTILIA ENDODONTIDAE (identifications by Neil D. Richmond) (Say) Helicodiscus parallelus SERPENTES Discus patalus ( Deshayes) CROTALIDAE Discus cronkbitei (Newcomb) Anguispira alternata Say Crotalus borridus (Linnaeus) Anguispira alternata mordax timber rattlesnake (Shuttleworth) Ancistrodon cf. contartrix ( Daudin) copperhead ZONITIDAE Euconulus fulvus (Mueller) COLUBRINAE Mesompbix sp. COLUBRIDAE Retinella indentata (Say) Elapbe cf. obsoleta (Say) Retinella virgin/ca Morr rat snake Retinella sculpt//is Bland Coluber cf. constrictor (Linnaeus) Hawaiia minuscula (A. Binney) black racer

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 133 Diadopbis cf. punctatus (Linnaeus) Sorex cinereus Kerr 35 ring-necked snake masked shrew Carpbophis amoena (Say) Sorex fumeus Miller 1 worm snake smoky shrew Lampropeltis cf. doliata (Lace-pede) Sorex dispar Batchelder 4 milk snake • big-tailed shrew NATRICINAE Sorex arcticus Kerr 6 arctic shrew Natrix cf. s ipeefon (Linnaeus) Sorex palustris Richardson 1 water snake water shrew Tbamnopbis cf. sirtalus (Linnaeus) Sorex s p. 12 garter snake Blarina brevicauda (Say) 37 species Storer/a, ? short-tailed shrew brown and/or red-bellied snake Natricinae, unidentified. CHIROPTERA AVES VESPERTILIONIDAE (identifications by Alexander Wetmore) Eptesicus cf. grandis (Brown) 1 big-brown bat TETRAONIDAE Minimum N. of Individuals Pipistrellus cf. subflavus (F. Cuvier) 13 Bonasa umbel/us (Linnaeus) 3 pipistrelle ruffed grouse Myotis keenii (Merriam) 302 Pedioecetes pbasianellus (Linnaeus) Keen's bat sharp-tailed grouse Myotis cf. lucifugus (Le Conte) 225 MELEAGRIDAE little brown bat Meleagris gallopavo Linnaeus 1 Myotis sp. probably keenii turkey or lucifugus 694 PICIDAE RoDENTiA Colaptes auratus (Linnaeus) 1 scruRnmE yellow-shafted flicker Marmota monax Linnaeus 1 Dryocopus pilatus (Linnaeus) 1 woodchuck pileated woodpecker Cite/Ins tridecemlineatus (Mitchell) 8 MIMIDAE 13-lined ground squirrel Tam/as striatus (Linnaeus) 27 Toxostoma rufum (Linnaeus) 1 chipmunk brown thrasher Glaucamys sabrinus (Shaw) 15 FRINGILLIDAE northern flying squirrel Passerella iliaca (Merrem) 1 Glaucomys volans (Linnaeus) 2 fox sparrow southern flying squirrel MAMMALIA Tamiasciurus budsonicus (identifications by John E. Guilday) tents/dens (Hay) 30 red squirrel INSECTIVORA TALPIDAE CRICETIDAE Condylura cristata (Linnaeus) 2 Neotoma cf. floridana (Ord) 3 star-nosed mole woodrat Parascalops braveri (Bachman) 1 Peromyscus leucopus (Rafmesque) 11 hairy-tailed mole white-footed mouse Peromyscus man iculatus (Wagner) 128 SORICIDAE deer mouse Microsorex boyi (Baird) 11 Peromyscus sp. pygmy shrew (leucopus or maniculatus) 96

134 THE NATIONAL SPELEOLOGICAL SOCIETY Minimum No. of Ind duals POLLEN ANALYSIS - INTRODUCTION Dicrostonyx budsonius (Pallas)) 3 The interpretation of fossil pollen counts Labrador collared lemming is often burdened with uncertainty. Before Pbenacomys cf. ungava Merriam 46 describing results of pollen analysis at New spruce vole Paris, it seems desirable to make explicit a Synaptomys cooperi Baird 16 few of the difficulties to be expected. southern bog lemming Pollen profiles of full-glacial or late-glacial Synaptomys borealis (Richardson) 71 age from near an ice margin commonly con- northern bog lemming tain more shrub and herb (non arboreal or Cletbrionomys gapperi (Vigors ) 260 NAP) pollen than those from a post-glacial red-backed vole forest. A high frequency of herb pollen would Microtus pennsylvanicus (Ord) 164 be expected either in treeless landscape akin meadow vole to Arctic tundra or else one in which scattered Microtus chrotorrhinus (Miller) 45 trees form a boreal woodland similar to that rock vole shown in fig. 5. But ecologists have recognized Microtus xantbognatbus (Leach) 344 that the late-glacial herb pollen zones could be yellow-cheeked vole explained in other ways and that the presence Microtus sp. 460 of grass and sedge pollen is by no means Pitymys p inetorum (Le Conte) 12 proof of either tundra, woodland, or steppe. pine vole For example, in a glaciated region a certain Ondatra zibetbicus (Linnaeus)) 1 portion of the NAP might be attributed to muskrat primary plant succession, in which an environment otherwise suitable for tree growth was ZAPODIDAE temporarily dominated by herbs during early stages of plant succession. Thus, while Zapus budsonicus (Zimmermann) 4 the 70-90 percent NAP found in Zone I meadow jumping mouse at Madelia, Minnesota could indicate cold cli- Napaeozapus insignis (Miller) 13 mate herbs and open vegetation, Jelgersma woodland jumping mouse (1962) felt it represented the pioneer phase ERETHIZONTIDAE of plant succession following ice retreat. In Eretbizon dorsatum (Erxleben) 3 geologically stable unglaciated areas, such as porcupine New Paris primary plant succession cannot be invoked to explain the record of herb LAGOMORPHA pollen found in late- or full-glacial deposits. LEP 3RIDAE A second problem encountered in many Lepus americanus Erxleben 52 pollen studies and inescapable in the present snowshoe hare case is the question of rebedding. Within the glaciated region the oldest late-glacial silts CARNIVORA may contain large amounts of temperate for- MUSTELIDAE est pollen washed out of adjacent glacial Mustela cf. rixosa ( Bangs) till. Redeposition can also complicate ecolo- least ? weasel gical interpretations of the pollen record south Martes americana (Turton) 1 of the ice margin, as in the Carolina Bays pine marten (Frey, 1953). Martes pennant/ (Erxleben) 1 At New Paris redeposition from till was fisher out of the question, but there was another source of older microfossils - the easily recog- ARTIODACTYLA nized Devonian-age spores in the limestone en- TAYASSUIDAE closing the sinkholes. More vexing was the Mylobyus nasutus (Leidy) 1 matter of intrusion of younger pollen into long-nosed peccary older fill, established with reasonable certainty

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 135 3 3 *** * ° " °° s 3 7171 171 ° ,1117■ ,7.7,1■,f, °

— 212

— LIS

—

203

—

".7.1 Lau.

Figure 12. Pollen analysis, Sinkhole No. 4, New Paris, Pa. in the case of rodent scats and "cave earth" kettles recommended by Iversen (1960) as de- from 5.9 m (table 1), less certain but suspect- sirable in describing the nature of the Atlantic ed in one of two samples from 6.7 m (table 2) climax forest from the Danish pollen record. and admittedly possible in all samples. Intru- Unfortunately, a major disadvantage of sub- sion may also account for the record of the aerial deposits such as sinkhole fill is the ten- pine vole (Pitymys pinetorum) and the white- dency for poor pollen preservation. Further- footed mouse (Peromyscus leucopus) from more, if sufficient light can reach the bottom certain levels. of the sinkhole, plants may grow on the debris cone, to be "over represented" in the fossil A third problem confronting those who record. But this does not seem to be the case would interpret pollen profiles is the matter with the narrow shaft of Sinkhole No. 4. of how much came from the upland "climatic In the absence of primary plant succession, climax." In regions of closed-canopy forest, with little likelihood of unrecognized redeposi- large amounts of non-arboreal pollen may tion of older pollen, and under a topography sometimes appear in sediments. The source that precludes hygric plant communities, New can generally be traced to local hygric habitats Paris Sinkhole No. 4 offers decided advantages near the site under study. In the late- or full- in paleo-ecological interpretations. Weighed glacial of the ice margin the major herb pollen against these must be the matter of poor types — grass, sedge, and much of the pollen preservation, possible over representa- Composites — could be derived from either an tion of pollen from plants growing on the fill, upland climax tundra or from poorly drained and the intrusion of younger pollen. hygric habitats within either tundra, woodland, The major paleo-ecological opportunity pre- or forest. sented by the sinkhole is the association of a pollen-bearing matrix with a fauna of nearly For topographic reasons the New Paris 3,000 identified birds and mammals. Small sinkholes should be ideally suited to trap pollen vertebrates are generally viewed as much more only from a well-drained upland habitat. In dependent on the local environment than are this regard they resemble the steep-walled large vertebrates and in this regard are corn- 136 THE NATIONAL SPELEOLOGICAL SOCIETY

Source 4 Rbamnuf-Vitil

1. Duff 20 1 4 1 72 2 1 3 4 1 2 6 117 4 60 2 2 1 1 13 200 1 PSM

2. "Caw earth- 13 1 11 103 3 1 3 2 3 11 151 15 1 30 4 1 5 22 229 1 '

3. Scat a 3 3 165 2 1 1 3 178 4 12 1 1 4 200 '

4. 6 1 129 1 1 1 1 140 4 9 2 4 159 JG

5. Scat b 18 6 1 75 6 2 3 1 3 1 7 123 18 1 53 1 3 3 20 222 PSM

6. 7 6 1 42 1 2 1 3 63 8 9 2 2 7 91 JG

7. Mixture o15 scats 29 5 105 5 4 1 1 6 156 22 1 21 2 11 213 1 PSM

Total 96 2 36 3 691 20 7 10 4 4 10 5 3 37 928 75 3 194 7 3 9 14 81 1314 3

Table 1. - Miscellaneous modern pollen samples from New Paris No. 4. Although samples 2-7 came from matrix between 5.9 and 6.1 m depth within the sinkhole, their pollen content closely resembles that of Sample 1 from leaf mould outside the cave and probably represent contamination. All samples are considered less than 200 years old.

.11

1.81 7 1 38 232 1 15 36.1

0.27 12 131 106 117 34.5

5.03 4 116 33 33.8

5.49 15 22 215 74 55.2

5.79 1 27 a 212 6 33.1

6.40 17 118 5 17.9

A 6.71 18 1 29 201 1 62 35.5

6.71 88 10 137 29.5

7.32 119 203 21.8

7.92 119 35 202 29.0

8.53 1 24 205 17 23.4

70141 1 502 60 40 26 86 1 02 83 2226 • 71 65

LIME A 72 8 734 2 54 37 2 3 24 4 5 1 1 40 38 94 66 15 4 9 9 2 71 1 295 .13 50

5.6 06 56.8 0.2 4 2 2.9 0.2 0.2 1.9 0_3 02 0 1 0 1 1_1 2.9 7.3 5.1 1.2 0.3 O./ 0.7 0.2 5.5 100.0 34.2 3.9

1.188 B 92 568 6 1 1 2 1 40 / 02 11 36 20 1 II 13 931 28 1 5

9.9 61.0 4.9 11.0 1.3 3.9 2.6 0.1 1.2 0 0 1.4 100.0

Table 2. - Fossil pollen counts from New Paris No. 4

parable to many plants in their sensitivity as the region today. While most of the small ecological indicators. vertebrates recovered near the bottom of the sinkhole presently occupy boreal environments Evidently the vertebrate fauna of the late- of southern and central Canada, two represent glacial was quite different from that known in prairie and one a tundra species.

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 137 During the excavation of New Paris Sink- pecially numerous between 6.7 m and 7.3 m, hole No. 4 samples of both fill and of rodent presumably weathered from the Devonian lime- feces were saved for fossil pollen analysis. stone forming the cave. In addition to their Field collectors sought to avoid surface con- distinctive morphology the spores were easily tamination and to find samples that were inti- recognized by their failure to stain in fuchsin mately associated with the fossil bones. Pollen and by their flattened appearance. sampling did not begin until the excavation Poor preservation of certain fill samples was at the 5.8 m level; remnants of matrix left complicated the problem of pollen identifica- in the walls of the sinkhole above this point tion. Within the Betulaceae well-preserved ex- were collected to represent the upper parts of amples of birch (Betula), hornbeam/ the deposit. Below 5.8 m samples were col- ironwood ( Carp inus- Os trya), and rarely, lected at the time excavation was in progress. hazelnut (Corylus) were encountered, but the The New Paris sinkholes lie on the boun- high frequency of poorly preserved grains in dary of Braun's Oak-Chesnut Forest region the triporate group made it impossible to and Mixed Mesophytic Forest region, immedi- determine relative numbers accurately. Most ately south of the Hemlock-White Pine- of the fern spores were of the monolete, Northern Hardwoods Forest region (Braun, bean-shaped Polypodiaceae. Fungal spores 1950). They are surrounded by red oak-red were common in scat and forest duff samples maple woods; other trees in the immediate but not in samples of fill. vicinity include big tooth aspen, white oak, Dr. Jane Gray kindly analyzed two of the white pine, and formerly chestnut. The woods rodent scat slides and reported sycamore are free of an understory and are relatively dry. (Platanus). Otherwise, her list of pollen The dominant pollen type found in the modern types and her percent estimates were similar pollen rain is oak ( Quercus); large numbers to those of P.S. Martin (see table 1). of ragweed (Ambrosia) pollen grains blow Results: Pollen counts from 11 samples be- into the forest from adjacent cultivated lands. tween 3.8 m and 8.5 m are shown in table 2 and fig. 12. The dominant type through- Extraction and Analysis: A few grams of out is pine pollen of small size. The counts can each sample were subjected to HC1 and HF be divided into two units. The upper, 3.8 m extraction following the schedule of Faegri to 5.8 m (unit A) contains more pollen of and Iversen (1950); the residue was stained oak (Quercus), alder (Alnus), fir (Abies), in fuchsin and mounted in glycerine. In most Betulaceae, temperate trees, Liguliflorae, un- cases at least 200 pollen grains were present knowns, and ferns than the lower (unit B), and counted on each slide. 6.4 m to 8.5 m, which contains more pollen Pollen was well preserved and abundant of sedges and spruce (table 2). Above 5.0 m in rodent scats collected at the 5.9 m level the ratio of arboreal to non-arboreal pollen and in a mixed sample of humus collected in rises. Pine breakage is significantly lower be- May of 1961 from beneath the leaf litter at low 5.8 m than above. The better preservation four points near the sinkhole. In contrast of pine in the deeper levels suggests a more pollen from the sinkhole fill was poorly pre- rapid rate of filling under a climate more served compared with the humus-dung sample favorable for frost action than the present one. and with Pleistocene alluvial clays from Marsh, Means of pine bladder measurements (20 Pa. and Cranberry Glades, West Virginia. grains each) in unit B are 6.4 m 33.5,u, Four samples from New Paris Sinkhole 6.7 m-32.2A, 7.3 m-33.4,u, 7.9 m-33.0,u, No. 8 (0.5 m, 2.6 m, 5.3 m and 7.5 m) and 8.5 m--34.6,u. Those from unit A may proved sterile. At Sinkhole No. 4 sufficient be slightly larger, 5.8 m--37.5 it, 5.5 m- pollen for analysis was obtained in 12 of 15 30.1,u, 5.0 m--34.8,u,4.3 m-38.8,u, 3.8 m-- fill samples, with sterile samples coming from 34.9a. These means resemble those obtained 2.0 m, 7.6 m and 8.7 m. The 8.7 m level from Zones F-2 and F-4 in Chester County was regarded by the excavators as solution alluvium (Martin, 1958) and are much smaller material deposited prior to perforation of the than means of post-glacial pine pollen from sinkhole. Rebedded Paleozoic spores were es- that area.

138 THE NATIONAL SPELEOLOGICAL SOCIETY The changes in fossil pollen content are which is on bone. Accepting Y-727 at face accompanied by changes in the character of value, pollen unit A (above 6.0 m) can be the vertebrate fauna. Above 6.0 m there are correlated with the Alleri5d oscillation of Eu- more temperate elements both in the verte- rope and with pollen zone A, of New Eng- brate record and in the pollen count. Below land. Both pollen content and vertebrate re- 6.0 m in a pollen zone with sedge and vir- mains in unit A indicate that while still relatively tually no temperate decidious tree pollen, the cool the climate in Pennsylvania at that time vertebrate fauna is dominated by cold wood- was decidedly warmer than during deposition land (boreal) species with very few elements of unit B. If unit A is contemporaneous with characteristic of the temperate zone. the pollen zone A, of southern New England, Two counts were made at the 6.7 m level, unit B with its higher frequency of herb one relatively poor in pollen but similar in pollen and much lower frequency of deciduous pollen content to adjacent levels. The other tree pollen should correlate with either pollen count, shown as a dotted line on fig. 12 zones A 2 , A„ or with zone T of southern and as part of unit A on table 2, was matrix New England. As Davis (1961) has shown, associated with the foot of a peccary, Mylobyus the distinction between A1 , A2, and A, is This pollen count resembles levels of unit slight. At Cambridge, Massachusetts Davis A more than those of unit B and may repre- found a pronounced rise in birch (Betula), sent contamination of unit B by matrix from oak ( Quercus), and a small but significant unit A. If so, the peccary foot may also be increase in other tree pollen types on the intrusive. boundary between A, and T. Abovethe T zone Intrusion of much younger (p o s t- there was an accompanying decrease in grasses settlement ) pollen is evident in both matrix and sedges. The change from unit B to unit and rodent scats thought to be contemporane- A at New Paris No. 4 is quite comparable. ous with the fossil fauna from the 5.9 m Unit A at New Paris may represent not only level. A sample of matrix at this depth very zone A, but also A, and A2 of southern closely resembles the count of pollen in de- New England, while unit B at New Paris is cayed forest litter adjacent to the cave (table 1). equivalent to New England zone T. Can the In the case of the rodent scats there is relatively New Paris record be correlated with the pollen high variation between samples, as one com- record farther south? monly expects in animal dung. Nevertheless, Schrock (1944) analyzed cores taken from the counts closely resemble the modern for- Glade Run Bog, Somerset County, Pennsyl- est litter, especially in a relatively large amount vania, 80 km southwest of New Paris at an of oak and ragweed pollen (listed as altitude of 825 m in the Negro Mountains, low-spine Compositae in table 1). For this reason the rodent scats must postdate the clear- approximately 160 km south of the Wisconsin terminal moraine. An undated early-pine- ing, cultivation, and accompanying ragweed maximum containing large amounts of spruce invasion of central Pennsylvania within the and traces of fir, hemlock, Betulaceae and oak last 200 years. In view of the extirpation of preceded the spruce-fir-maximum. She states chestnut (Castanea) 40 years ago it is of (ibid. p. 30) "The pines represented are interest to find chestnut pollen present in doubtless several species, those of the lower two of the rodent scat samples. levels as well as those of the top levels being Correlation: Presence of only two pollen definitely of larger size than the ones present units in 4.7 m of late-glacial fill is likely to in the spruce-fir-maximum." She suggests that reflect rapid deposition. During excavation jack pine (Pinus Banksiana) is represented no evidence of soil formation or other strati- by the small pollen associated with her spruce- graphic break was seen. Two radiocarbon fir maximum. Correlation with the New Paris dates associated with the matrix are shown in No. 4 pollen record is not possible at the fig. 12. The upper, Y-727 (11,3001-1000 yrs. present time. Dr. Schrock believes that the B.P. ), is of charcoal and is considered more early-pine-maximum at Glade Run Bog pre- reliable than M-1067 (9,5401-500 yrs. B.P.), ceded the glacial maximum to which she assigns

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 139 Locality

Cranberry Glade, W. Va. 1.70 11.5 20.5 3.7 7.1 11.5 10.4 6.7 1.1 3.0 75.5 11.9 7.4 3.0 0.7 0.4 1.1 24.5 269 1.95 30.0 24.5 .5 1.0 1.0 57.0 28.5 9.0 3.5 1.0 1.0 43.0 200 " 2.20 23.1 17.6 2.2 2.2 1.1 1.1 47.3 43.4 6.6 0.5 0.5 0.5 0.5 0.5 52.5 182 2.45 21.5 17.5 .5 .5 .5 1.0 41.5 41.0 16.5 0.5 0.5 58.5 200

Marsh, Pa. M, 0.74- 1.15 22.0 7.7 1.2 0.1 1.8 1.1 2.2 0.6 0.1 0.7 37.5 37.7 20.3 0.5 1.2 0.2 05 0.3 2.0 62.7 3668 1.82- 2.07 19.3 1.7 0.0 0.1 1.3 0.4 0.5 0.5 0.4 24.2 34.5 37.4 0.5 1.0 0.2 1.0 0.5 0.6 75.7 2415

Marsh, Pa. M. 0.95- 1.45 17.2 10.1 5.3 0.1 2.3 0.8 3.5 0.5 0.4 40.2 28.3 21.6 0.7 2.7 0.1 0.1 0.6 0.2 5.6 59.9 2176

New Paris No. 4, 6.40- Unit B 8.53 61.0 9.9 0.6 0.2 0.1 0.1 0.1 72.0 11.0 4.9 2.6 3.9 2.3 0.1 0.3 1.2 1.7 28.0 931

Table 3. - Late or full-glacial pollen records f om unglaciated areas the over-lying spruce-fir maximum. If this is cent tree pollen, almost exclusively of pine so it is of extreme importance in reconstruct- and spruce, with 40 percent sedge. There is ing full-glacial conditions in the central Appa- extremely little temperate tree pollen in these lachians. It is probable, however, that the Glade lower samples (table 3). The two upper sam- Run Bog spruce-fir maximum may be a late- ples contain more tree pollen, with oak, birch, glacial readvance of the ice front, such as the hemlock, and alder common in the uppermost Valders. Only arboreal species were con- level. The deeper clay samples from Cranberry sidered. It would be of interest to examine Glades contain somewhat more spruce and less the NAP picture as well. grass than similar clays from the F zone at Martin (1958) reported that the F zones Marsh, but in general the two sets of counts in alluvium of Marsh, eastern Pennsylvania, are quite similar and may be contemporaneous. contained less than 40 percent tree pollen, They are quite different from the post-glacial mainly of small-sized pine grains and of spruce records in eastern North America. (see table 3). The upper part of the F zones Can New Paris pollen unit B be considered yielded radiocarbon dates of about 13,500 contemporaneous with the F zones of Marsh years (Y-478 and Y-479). Are the F zones and of Cranberry Glades? In the latter localities unique to Marsh? One of the more promising non-arboreal pollen is much more abundant localities for a pollen comparison with Marsh and pine pollen much less abundant than in appeared to be Cranberry Glades, West Vir- the New Paris Sinkhole. During accumulation ginia where Darlington (1943) reported clays of extensive flood plain deposits of clay and of unknown age beneath a post-glacial peat silt there were disturbed habitats in which trees profile dated at its base as 9,434±840 years could not grow and in which local herb pollen B.P. (C-336). production was especially heavy. The relatively In November of 1957 Martin visited Cran- high concentration of herb pollen close to the berry Glades briefly and obtained a 1.5 m site of alluvial deposition would profoundly core from the clays beneath peat found in affect the local pollen rain along Marsh Creek Round Glade. Pollen is abundant and well and at Cranberry Glades, as it does in alluvial preserved in this material. The counts of two flood plains in the southwest (Martin, 1963). samples below 2 m in depth (table 3) from On the other hand, at New Paris No. 4 the Round Glade contain approximately 50 per- pollen-collecting surface - a feed cone in the

140 THE NATIONAL SPELEOLOGICAL SOCIETY top of a sinkhole - existed in the middle of mid portion of the stratigraphic column to an upland environment without the flood plain rare in the lower levels. The numbers of the plant communities and flood plain pollen rain. large woodland snail Triodopsis albolabris Unless large numbers of herbs grew on the Say diminished dramatically below the 7.0 m disturbed top of the debris come in the sink- level. The commonest snail in all upper levels, hole (while may indeed be the case of the Lig- it appeared to have been absent from the ulfiflores and ferns in unit A), the sinkhole lower 2.7 m of the deposit. would be likely to trap only the natural upland pollen, free of any riparian influence. AMPHIBIA — AMPHIBIANS The main difference between counts from by Neil D. Richmond New Paris No. 4, unit B, and the F zones of Caudata--s alamander both Marsh and Cranberry Glades is the higher Remarks: Most of the salamander vertebrae frequency of pine pollen and the lower fre- from this deposit have not been studied. Only quency of sedge and grass. This difference those sufficiently large or distinctive in struc- could simply mean that hygric communities ture to be easily recognized have been identi- of grass and sedge, which certainly occurred fied. The five genera are all forms found at at the Marsh and Cranberry Glades, did not the site today and while the red-spotted newt exist around the New Paris sinkholes. Tenta- ( Dtemictylus) remains represent trapped tively we suggest that pollen unit B in the New specimens, the other species are fossorial and Paris sinkhole was contemporaneous with the tend to follow crevices in the limestone and top of pollen zone F-4 from Marsh. Beyond could be expected alive at almost any depth a greater representation of spruce, conditions in this formation. were similar at Cranberry Glades. The T zone of New England may locally contain more Salientia--frogs and toads pollen of Betula or Populus than known Bufo --toads from Cranberry Glades or the Marsh, but it Material: This genus is represented at all levels. is clear that the T zone of southern New A total of 116 ilia, 34 fronto-parietals and 18 England is more like Marsh zone F than any sacral vertebrae were recovered as well as other part of the New England pollen record. large numbers of limb bones, vertebrae- and Although it has more pine pollen, New Paris miscellaneous skull elements. At least 74 indi- unit B seems best correlated with New Eng- viduals are represented in this material. land T zones. Admittedly, stratigraphic control and radiocarbon dating of sites south Remarks: On the basis of the ilia it would of the ice margins is insufficient for a vig- be possible to assign three names to this col- orous defense of this or any other correla- lection; using the more diagnostic fronto- tion scheme. parietals, only two groups appear. Since there are changes in the ilia and fronto-parietals from the top of the deposit to the bottom, INVERTEBRATES and since they are both well represented, I Three species of millipedes and 26 spe- am assuming that only two forms occurred cies of land snails were recovered. All may here, that they were separated by time, and be found in western Pennsylvania and the were allopatric. The form at the top and mid- sinkholes area today. depths is identified as the American toad (B u- It is difficult to separate the land c nails that fo a. americanus] and the form in the lower were contemporaneous with the late Pleistocene levels as the Hudson Bay toad (B. a. copei). fauna of Sinkhole No. 4 from those that kept Admittedly to recognize subspecies on the basis infiltrating during the years of excavation. of skeletal remains is a dubious procedure Snails are very common in this limestone area especially when the subspecies involved is also and preservation very good. a questionable one. For a discussion of the Two general observations could be made current status of copei see Logier and Toner, however: the numbers of all species of snails 1961. Most of the skeletal characters of B. dropped from common in the upper and a. copei are also those of small but sexually

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 141 ous limb bones. At least 66 individuals represented. Remarks: This genus is well represented in the deposit. Most of the identifiable bones appear. to be those of the leopard frog (R. pipiens). Only one illium, CM 8017, has the pronounced knob-like dorsal prominence characteristic of the wood frog (R. sy Iva t ica). The bull frog (Rana catesbeiana), the green frog (Rana clam/tans), and the pickerel frog (Rana pa lustris), all common in the New Paris area today, appear to be absent from the fossil material. REPTILIA — REPTILES Serpentes –snakes Figure 13. Material: Vertebrae, frontals and toothed elements were recovered. Most of the determin- Ilia a, b, c,) and fronto-parietals (d, e), Bufo americanus, Sinkhole No. 4, New Par- ations are based on precaudal vertebrae and is, Pa. in some cases on maxillae. a. CM 7950 - B. a. ame-icanus, level ? Remarks: Several features of this large collec- b. CM 7952 - B. a. americanus, level ? tion are noteworthy. One of the surprising c. CM 7951 - B. a. cope', 8.5 m level features is the relative frequency of the rattle- d. CM 7949 - B. a. americanus, 6.4 m snake (Crotalus horridus) compared with level. that of the copperhead (Ancistrodon con- e. CM 7948 - B. a. copei, 8.5 m level t (atrix). Rattlesnake vertebrae were abundant near the top of the deposit and represented mature B. a. americanus, i.e., open fronto- all sizes from newborn to individuals larger parietal canals and relatively low ilial promi- than most modern specimens. By contrast nences. The assumption that the lower levels only a few copperhead vertebrae were recovered of this deposit received only juveniles while and only 2 maxillae, all in the upper part the upper levels received only large adults of the deposit. Most of the rattlesnakes (12 seems unlikely. Instead, I believe that the maxillae) were in the upper portion but oc- lower, earlier levels received a small form curred at all levels possibly as a contaminant (B. a. copei) while the upper, more recent from above. (See fig. 34). levels, the larger form B. a. americanus. The most numerous snake vertebrae were Most ilia in the upper levels are easily as- natricine, most of which are probably garter signed to Bufo a. americanus. Associated snake ( Tbamnophis sp. ). Only those vertebrae with these bones are others that have a spike- that represented mature individuals and that like dorsal prominence (fig. 13a) resembling compared in detail with Recent material were such forms as B. hibbardi and B. rep entinus. named. Unlike the colubrine genera which (See Tihen, 1962 for figures of these species. ) became scarcer with depth, the natricine genera Since only two types of fronto-parietals were were abundant from top to bottom (fig. 34). recovered, these large ilia are considered to be This is not surprising in view of the wide from very large old specimens of B. a. amer- environmental tolerance of the garter snake icanus. Associated with size and age are ( Thamnoph IS sirtalis) today. Its present changes of shape and hypertrophy of the vari- range, from the Labrador Peninsula to Florida, ous areas of muscular attachment. includes all of the climatic possibilities that might have been in the vicinity of the sinkhole Rana--frogs at any time in its history. Nor is the garter Material: 102 ilia, 29 sacral vertebrae, numer- snake sensitive to changes from forest to grass-

142 THE NATIONAL SPELEOLOGICAL SOCIETY land. It has been recorded as far north as Remarks: This nearly complete skeleton (dis- 0 Churchill, Manitoba, Lat. 58 N. at the juncture cussed in Wetmore, 1959) was found against of the Hudsonian and Arctic life zones. (Shel- the north wall at the 5.2 m level. The skeleton ford and Twomey, 1941). Only two snakes in was uncrushed and in essential articulation. this collection are southern in their affinities, The skull was not recovered and had presum- the worm snake (Carphophis) and the copper- ably been crushed or removed by later rock head (Ancistrodon). Their remains were con- falls. The species has also been recorded fined to the upper levels of the sinkhole. from the Natural Chimneys local fauna of Although both of these range farther north Virginia. along the coast they are near the northern Several limb bones of the ruffed grouse edge of their present range at New Paris. ( Bonasa umbellus) also catalogued under All of the other forms in this deposit range CM 5440 were found in association with the at least into southern Canada today. skeleton. Sauna and Chelonia--lizards and turtles The presence of both sharp-tailed and ruffed Material: None grouse in the upper levels of the sinkhole Remarks: The absence of both lizard and suggests a closing forest canopy. Bonasa turtle remains is probably significant. Today is essentially a forest species with a preference there are three species of lizards in the New for coniferous-deciduous forest-edge situations Paris area. The Recent faunas of Sinkholes (Aldrich and Duvall, 1955; Edminister, 1947). 2 and 3 both contained box turtle ( Terrupene Pedioeceter is a grouse of narrow habitat carolina), but the boreal climate of New requirements. It inhabits mixed prairie-forest- Paris No. 4 times was too severe for terres- edge situations and can tolerate neither open trial turtles to survive. The range of the box prairie where it is replaced by the prairie chicken turtle does not extend much farther north (7'ympanuchu.■ cup/do) in the more southerly than Pennsylvania today. portions of its range, nor forest situations where the canopy is too dense to support AVES— BIRDS an understory of grasses. In the northern In contrast to the birds of the Natural part of its range it is found typically in open Chimneys local fauna (40 species, 78 individ- or brushy muskeg situations interspersed in uals) bird remains were scarce in the New coniferous forest. Ammann (1957) working Paris No. 4 deposit. (7 species, 13 individuals ). with this grouse in Michigan states that 20- 40 percent woody cover is optimum for the The Natural Chimneys deposit was accumu- species if the trees occur in copses. Occasional lated primarily by carnivorous birds. The individuals were found in areas with up to numerous avian remains include both shore 75 percent woody cover. It was found that a and upland game species and a variety of minimum of slightly under one square mile swallows, woodpeckers, jays, and smaller song of near-optimum habitat is required to sustain birds of varied habits that had fallen as prey and were brought into the cave. a population although individual birds may occupy areas as small as 10-15 acres in size. New Paris No. 4, however, was a sinkhole It is possible that trap fed from above by animals that accidentally Pedioecetes and Bonasa fell in. Once a bird blundered into the shaft did not occupy the immediate area contem- (like a bird in a chimney), despite its ability poraneously. Charcoal particles in the matrix indicates at least one forest fire during the for horizontal flight it was trapped in this period of infilling. This would impose vege- vertical passage. All of the species with the exception of the pileated woodpecker are tational changes of shorter duration than those dictated by climatic change. The two prone to search for food on the surface of the grouse may then have inhabited the area ground. suc- cessively--Pedioecetes being replaced by Bon- Pedioecetes phasianellus–(Linnaeus) asa as forest cover returned in the wake of Sharp-tailed grouse fire. Material: CM 5440, partial skeleton. It cannot be assumed that the one speci-

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 143 men of Pedioecetes represents optimum range aquatic and partial to saturated soils and its conditions for that species in the sinkhole distribution is usually governed by this, but area. Its presence, however, in the unit A it is occasionally taken in some numbers in fauna points to areas of open ground still well-drained atypical upland sites (Richmond lingering in the vicinity. Conditions were such and Rosland [sic], 1949, p. 36). that the bird, now largely confined to por- Specimens are more robust than modern tions of the Canadian and Hudsonian life- Pennsylvania material. An additional individual zones south and east to northern Michigan, from Station 2, CM 6952, is believed to be occurred in late Wisconsin time, south at Recent. Its limb bones are less robust and least as far as lat. 38° in what is now the appear peculiarly "greasy." Carolinean life-zone of Virginia's Shenandoah Condylura cristata, at the present time, Valley. shows a slight, positive "Bergmann's re- Meleagris gallopavo Linnaeus – sponse." (Jackson, 1915, p. 90). The New Wild Turkey Paris No. 4 specimens, in common with so Material: CM 5896, 5897, 5900. 1 tarsome- many other species from the deposit, find tatarsus, 1 humerus, 5 vertebrae, 2 phalanges. modern size equivalents in the northern por- Stratigraphy: 5.2 m - 6.7 m level. tion of their present range. Remarks: The wild turkey is considered to be The alveolar length of P. -M3 measured 6.8 a bird of the deciduous forests of thesouthern mm, 6.9 mm and 7.2 mm. portions of the continent north tothe northern Parascalops brewer/ (Bachman)--Hairy- boundary of the Carolinean life-zone. Cleland, tailed mole 1963, found turkey remains at the Juntenen Material: 1 left mandible, no dentition; 1 Site (20 Mk 1), an archaeological site on humerus: 1 ulna; 1 isolated M.. Bois Blanc Island in Lake Huron, 200 km north of its known historic limit and within Stratigraphy: upper levels to 6.0 m (fig. 27). the Canadian life-zone, lat. 45°. Even in the light Remarks: Possibly two individuals are pre- of Recent distributional patterns, it is possible sent. Specimens agree in all particulars with for the turkey to have been a member of the modern Pennsylvania material. This mole, as unit A avifauna, advancing its range as the well as Condy/ura cristata, is common in glacier receded. Turkey remains are also known the area today. Because of a positive "Berg- from Frankstown Cave, Port Kennedy Cave, mann's response," the latter is believed to and Natural Chimneys, Virginia. be a member of the late Pleistocene fauna. Parascalops brewer/ may or may not have MAMMALIA — MAMMALS been although it does range extensively throughout the Canadian life-zone today. It Order: Insectivora has been recorded from Frankstown Cave, Family: Talpidae Pa., Bootlegger Sink, Pa., Natural Chimneys, Condylura cristata (Linnaeus )--Star-nosed Va., and Robinson Cave, Tenn. mole Family: Soricidae Material: 1 left, 3 right humeri; 2 left, 2 right partial mandibles; 2 right scapulae; 1 left, Microsorex boy/ (Baird).-Pygmy shrew 2 right tibiae; 1 left, 1 right femur; 1 radius. Material: 4 partial skulls; 2 left, 2 right max- Stratigraphy: from 5.5 m to 7.0 m level (fig. illae, 11 left, 11 right mandibles. 27). Stratigraphy: evenly distributed from the 5.2 m Remarks: Ranging farther north than any through the 8.5 m level (fig. 31). other North American mole, throughout the Remarks: This was the third commonest shrew Canadian life-zone and locally, into the Hud- in the deposit. The pygmy shrew is rare in sonian life-zone (Edwards, 1963), this is the Recent mammal collections from eastern North only species of mole that occurred beneath America. There is only one modern Pennsyl- the 6.0 m level. Only three individuals were vania record (Roslund, 1951, p. 40). This present. This mole is considered to be semi- shrew was apparently widespread and common 144 THE NATIONAL SPELEOLOGICAL SOCIETY in the East during the late Pleistocene. It is known from Natural Chimneys, Va., Robin- s ALASKA son Cave, Tenn., and Bootlegger Sink, Pa. 8- Measurements of the New Paris No. 4 55° 5-LA8RADOR specimens appear to be average for the species as defined and measured by Jackson (1928); smaller than modern far-northern races, but larger than the southern M. b. winnemuna. 50 Both M. minutus Blown (1908) and M. pratensis Hibbard are reported to be larger than modern M. boy/ (Paulson, 1961, p. 5- MINN ES OTA 134). 45

4-RHODE ISLAND Table 4 - Measurements in mm. Microsorex boy,' (Baird) OHIO VIODE RN PE NN5 -NA. 40 PAR IS#4 New Paris No. 4, Pa. (fossil) Mandible 3-MARYLAN D height, total total 4-NORTH CAROLINA Cat. MrM3 ascending length, length No. ramus including I, dentary 35 6240 2.8 3.0 8.0 6.5 5.5 6.0 6.5 6667 2.8 2.9 8.2 6.6 Size in mrn. 6672 2.9 2.9 7.9 6.5 6676 - 3.0 8.2 6.6 7102 2.8 3.0 7.6(worn) 6.6 7330 2.8 2.9 7.7 6.4 Figure 14. 7330b 2.9 2.9 7.8 6.4 7428 2.9 7.8 6.4 Increase in mean length of palate with 7103 2.8 3.0 8.1 6.4 increase in latitude. Sorex cinereus, 7104 3.1 8.4 6.6 various Recent North American localities mean 2.86 2.94 7.97 650 and Sinkhole No. 4, New Paris, Pa. Late Pleistocene. Data, in part, from Jackson, 1928; Bole and Moulthrop, 1942. Skull (measurements, see Jackson, 1928, p. 13) dons of the continent. It tolerates a wide Cat. Maxillary Palatal Interorbital . variety of habitats and with the exception of No. Width Length 3 Breadth P -M Micros orex is the only insectivore found at 5849 4.0 5.2 2.9 3.4 all levels of the sinkhole that produced an 6239 4.2 5.3 2.9 3.5 adequate sample. 7330 4.3 5.1 2.8 3.4 4755 4.1 5.3 2.9 3.5 Sorex cinereus, in accordance with Berg- mean 4.15 5.2 2.87 3.45 mann's Rule, exhibits a positive correlation between size and latitude except in the tundra area of the western Nearctic where it under- goes a reduction in size. (Jackson, 1928, So rex cinereus Kerr--Masked shrew p. 38). Measurements of the New Paris No. 4 population are larger than those from more Material: 13 partial skulls; 35 left, 35 right southerly areas of the animal's range (fig. mandibles. 14) and agree in superior size with modern Stratigraphy: common at all levels from 4.9 northern populations. Unfortunately, all pub- m to 8.5 m (fig. 31). lished measurements dealing with the neotaxo- Remarks: This animal has the widest range nomy of S. cinereus are confined to either of any North American soricid, including external measurements which are of little use almost all of the central and northern por- when dealing with skeletal material or to a few BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 145 cranial measurements which require skulls in Table 5 - Size of third unicuspid relative to almost perfect condition. There is a need for fourth, Sorex cinereus . published measurements of mandibles and larger equal smaller lower dentitions (those parts most commonly Modern south-central Pa. 10 12 1 recovered in good condition from paleontol- New Paris No. 4 2 ogical sites) of most of our Recent mammals. These could be easily incorporated into modern taxonomic studies and could be utilized by Sorex fumens Miller-Smoky shrew students working with comparable Pleistocene Material: 1 right mandible. CM 6673. material. Stratigraphy: 3.9 m level (fig. 31). Remarks: This shrew is found in a variety Ordinarily the mandible of Sorex cinereus of wooded situations in the Appalachian area. lacks the post-mandibular foramen. However, It occurs in Bedford County at the present five mandibles with this foramen present were time but the immediate sinkhole area would noted in a collection of 24 Recent individuals appear to be too dry to support it. Condi- from south central Pennsylvania. In four spec- tions seem never to have been optimum for imens it was small and occasionally confluent it throughout the period of infill. C -M with the mandibular foramen. In one (CM 1 3 measured 4.7 mm. mammal no. 33716), however, it was extremely well-developed. One example was noted in the Sorex dispar Batchelder-Big-tailed shrew New Paris No. 4 collection (CM 7116). Material: 4 left, 1 right mandible.

Table 6 - Selected Measurements, Sorex cinereus. measurements in millimeters

Locality O.R. V Total length, condyle to most anterior point of dentary New Paris No. 4 7.6 ± .06 7.2-7.8 .27 ± .05 3.55 ± .60 17 Modern Penna. 7.28 ± .02 7.1-7.7 .10 ± .01 1.37 ± .20 22

C1-M3, antero-posterior crown length New Paris No. 4 3.9 ± .04 3.7-4.3 .16 ± .02 4.09 ± .53 29 Modern Penna. 3.69 ± .01 3.6-3.9 .07 ± .01 1.89 ± .29 20 M,, length of crown New Paris No. 4 1.33 ± .01 1.2-1.5 .05 ± .005 3.75 ± .39 44 Modern Penna. 1.28 ± .02 1.2-1.4 .07 ± .01 6.01 ± .90 22 Length of palate* New Paris No. 4 6.4 6.3-6.6 - 5 Modern Penna. 6.09 ± .04 5.5-6.6 .21 ± .03 3.44 ± .50 23 Interorbital breadth* New Paris No. 4 3.05 3.0-3.1 - 2 Modern Penna. 2.99 ± .01 2.9-3.1 .05 ± .007 .67 ±- .24 23 Maxillary breadth* New Paris No. 4 4.2 4.0-4.4 - 4 Modern Penna. 4.23 ± .02 4.0-4.5 .11 ± .01 2.60 ± .39 22 P4-M3, antero-posterior crown length New Paris No. 4 3.7 3.5-4.0 - 5 Modern Penna. 3.52 ± .02 3.4-3.7 .10 ± .01 2.83 -± .41 23

* Defined: Jackson, 1928. 146 THE NATIONAL SPELEOLOGICAL SOCIETY Stratigraphy: 3.6 m to 7.3 m level (fig. 31). (Blarina). This is the fourth known late Remarks: This animal, endemic to the Appa- Pleistocene locality for the Arctic shrew. It lachians, requires cool, moist, rocky mountain has been reported from the Natural Chimneys forest. The presence of surface cover seems local fauna of Virginia, the Robinson Cave less essential than deep passages beneath local fauna of Tennessee, and Bootlegger loosely piled talus. Sorex dispar does not Sink, Pennsylvania. occur at the site today but is not uncommon in suitable areas of moist talus along the flanks of the Appalachian Mountains and at Sorex palustris Richardson --Water shrew no greater altitude (Richmond and Grimm, Material: 1 partial skeleton including anterior 1950). portion of skull with associated mandible. CM 7251. Table 7 - Measurements in millimeters, Sorex dispar, New Paris No. 4, Pa. Stratigraphy: A single individualwas recovered from the 8.5 m level (fig. 31). CM Dentary CI-M3 M, Remarks: This shrew is semi-aquatic and is 6243 1.3 not normally found far from water. This would seem to indicate the near presence of sur- 6241 8.2 4.3 1.3 face water, but the fact that only one indivi- 7116 8.2 1.4 dual was recovered suggests that this may have been an itinerant individual in an atypi- Sorex tactical Kerr --Arctic shrew cal habitat. The accompanying fauna and pol- Material: 4 partial skulls; 6 left, 5 right man- len profile at that level suggests a relatively dibles. dry environment. Stratigraphy: 6.1 m to 8.5 m level (fig. 31). Remarks: The Arctic shrew inhabits northern Table 9 - Measurements, Sorex palustris, North America south to Wisconsin and New CM 7251, New Paris No. 4, Pa. Brunswick. It is most often found in conifer- ouse forest boglands. Although it is a minor Total length of dentary component of the fauna, it did persist to the (to and including condyle but deeper levels and is a distinct boreal element not angular process) 9.5 mm. replacing more southerly forms such as the Length C1-1143 5.2 mm. smoky shrew (S. fa eu s), thebig-tailed shrew Length M, ...... 1.6 mm. (S. and the short-tailed shrew dispar), Length of palate 8.3 mm. 4 3 P .41 ...... 5.0 mm. Table 8 - Measurements in millimeters, Sorex arcticus, New Paris No. 4, Pa. Sorex, species ? CM Dentary C1-M3 M, Material: 4 partial skulls; 1 right maxilla; 6643 8.7 4.2 1.4 8 left, 12 right partial mandibles. 6662 8.7 4.5 1.5 Remarks: All of this material is exceedingly 6941 9.2 4.4 1.4 fragmentary. It is doubtful if any unrecog- 6942 8.7 4.5 1.5 nized species is present. 7101 9.3 4.5 1.5 Blariaa brevicauda (Say )--Short-tailed shrew 7109 - - 1.5 Material: 17 partial skulls; 6 left, 7 right 7111 - 4.4 1.3 maxillae; 37 left, 25 right mandibles. Stratigraphy: Mean 8.9 4.4 1.4 Commonest insectivore in upper and middle levels of sinkhole from 2.1 m 7738 Length of palate 7.8 mm. to 7.0 m. Absent in lower levels (fig. 31). BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 147 Remarks: The Blarina material from Sink- hare bones indicate that infilling was relatively hole No. 4 presents a mixture of two genetic rapid and was completed during the boreal stocks. One, presumably B. b. kirtlandi, episode. The sinkhole filled to the top during is indistinguishable from the living form in late Pleistocene-early Recent times. The prob- the area. The other is a large, late Pleisto- able explanation for the correlation is an in- cene form, typical examples of which are creasing contamination of the deposit by mod- readily distinguishable from modern B. b. ern shrews as one approaches the surface. kirtlandi but not from the modern Minne- The late Pleistocene B. b. cf. brevicauda sota specimens of B. b. brevicauda used extended throughout the stratigraphic column as comparative material. but became increasingly contaminated by the remains of the smaller modern B. b. kirtlandi burrowing, or pursuing cracks from

m. mm6.0 .1 2 .3 4 5 6 .7 8 .9 7.0 n. the surface.

5' 4 -2- That the large sinkhole Blarina finds its modern population equivalent in the north- -3- 4310' 0 2 western sector of its present range, and is indeed of B. b. brevicauda stock is quite 15' 0 -5- probable, and in harmony with the range changes of the yellow-cheeked vole ( Microtus -6- 20 xanthognatbus), the thirteen-lined grotmd ( C -M ) -7- 1 3 squirrel (Cite//us tridecemlineatus), and Blarina brevicauda 25' 1 the sharp-tailed grouse (Pedioecetes phasi- 0 anellus), which have shown similar shifts. The late Pleistocene occurrence of a living subspecies south of its present range in what Figure 15. is now the territory of a morphologically dis- Increase in mean length C,-M, with depth. tinct subspecies has been elsewhere demon- Blarina brevicauda, Sinkhole No. 4, New strated by finds of the Scandinavian red fox Paris, Pa. ( Vu/pes v. vu/pes ) associated with the arctic fox (Alopex lagopus) in cave deposits in Figure 15 illustrates the increase in an aver- Wiirttenberg, Germany (Lehmann, 1954), age size of specimens of Blarina correlated Willendorf, Austria (Thenius, 1959) and with depth. At each station, representatives of Karnten, Austria (Thenius, 1960). Vu lpes both "races" could be picked out as well as v. crucigera, the present middle European unassignable intermediates. The high coeffi- fox, is a smaller animal with pronounced den- cients of variation of the entire sample of tal differences (see Thenius, 1960, p. 37). Blarina from the sinkhole and the bimo- Hibbard (1963) discusses a large form of dality of certain measurements (condylewidth, Blarina brevicauda, from the late Illinoian C1-1113 ) suggests that it was made up of more of Kansas which is indistinguishable from than one population. B. b. Brevicauda the large modern northern race. This form is apparently succeeded in The increase in size with depth shown by Sangamon deposits from the same area by a figure 15, although seemingly correlated with small B. b. cf. carolinensis. time, is apparently not. There are two populations present at all levels from which a A skull of Blarina b. cf. brevicauda large enough sample was available to decide. complete and uncrushed (fig. 16a) was re- This increase is due rather to a shift in the covered from the 5.2 in level of the sinkhole. racial composition of the samples from each Its measurements (table 10) are greater than level. The presence of boreal forms in the the maximum observed in over 1,100 crania upper levels, the pollen picture, and the results of Blarina b. kirtlandi from Pennsylvania, of the thorium analysis of the snowshoe but can be matched by Minnesota speci- 148 THE NATIONAL SPELEOLOGICAL SOCIETY Figure 16. Skulls of Blarina brevicauda, dorsal view. Note positive "Bergmann's response." Bar = 1 cm. CM 5838 - B. b. cf brevicauda, 5.8-6.4 m level, Sinkhole No. 4, New Paris, Pa. Late Pleistocene. CM Mammal 4355 - B. b. brevicauda, Minnesota. Recent. Remainder - B. b. kirtlandi, West- ern Pa., Recent.

Its ex- mens of Blarina b. brevicauda. .4111 , treme ruggedness and the accentuated crests -1"9 " and fossae for muscular attachments seem to indicate its advanced physiological age and can be approached by large, old skulls of B. b. 4r2.A47 kb. Zell brevicauda. f") Blarina remains have been recovered from at least five other Appalachian cave deposits. '4 1. Frankstown Cave, Pennsylvania. Three stist 1* mandibles indistinguishable from modern Pennsylvania stock. Due to the circumstances of recovery of the Frankstown local fauna Figure 17. there is no guarantee that the specimens are, Mandibles of Blarina brevicauda from Sink- in fact, Pleistocene. hole No. 4, New Paris, Pa. Bar = 1 cm. 2. Port Kennedy Cave, Pennsylvania. A sin- B. b. cf. kirtlandi, CM 5848a, b, 2.4 - 3.3 gle mandible, the type specimen of Blarina m level. B. b. cf. brevicauda, CM 5843, simplicidens Cope. Re-examined by Hibbard 5847, 6027, 6087, 3.6 - 6.1 m level. (1957) the species B. simplicidens was pears comparable to New Paris No. 4 in that found to be invalid, a misinterpretation of a there is a mixture of smaller (presumably broken specimen of B. brevicauda. kirtlandi) and large (B. b. cf. brevicauda) 3. Cumberland Cave, Maryland. ".. . rostral specimens. If the value for length of C1-M3 portion of a skull. .. two maxillary fragments is disregarded due to the small sample size, . . . eight lower jaws." (Gidley and Gazin, the coefficients of variation appear quite large 1938). These specimens were described as when compared with modern samples, sug- of a "size somewhat greater than the average gesting that the collection consists of more in Recent specimens." than one population. 4. Wytheville, Virginia (Cope, 1869). One The late Pleistocene occurence of a large mandible in breccia. Lost before any obser- form of Blarina in lowland Virginia is also vations could be made on it. suggested by the weekly differentiated Blari- 5. Natural Chimneys, Virginia. 100 man- na telmalestes Merriam, a relatively large dibles, 4 partial skulls, 11 maxillae, Measure- torm confined to Dismal Swamp and sur- ments presented below. This collection ap- rounded by the small race of the southern

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 149 lowlands B. b. carolinensis. It appears to incisor, from dorsal to ventral border at have been isolated at some late Wisconsin/ junction of incisor and first unicuspid. Mea- early Recent period, a situation reminiscent of surements with ocular micrometer at 10X the isolation of Clethrionomys gapperi in mm. rhoadsi in the cedar swamps and sphag- SPECIMENS EXAMINED num bogs of southern New Jersey. Blarina Carnegie Museum: Section of Mammals. telmalestes, according to Bole and Moul- throp (1942) is more closely related to the Moorhead, Minnesota 13; Detroit, Minnesota northeastern race B. b. talpo ides than to 1; 1 mi. NE Osterburg, Bedford Co., Pa. 9; 2 mi. E. Osterburg, Bedford Co., Pa. B. b. carolinensis, its immediate neighbor. 3; 1 mi. NE Spruce Creek, Huntingdon Co., Skull measurements as defined by Guilday Pa. 5; 2 mi. SW Pennsylvania Furnace, Hunt- (1957, p. 43). Mandibular measurements: ingdon Co., Pa. 1; 1-1/2 mi. W Immler, 1. Condyle - greatest length of condyle, mea- Bedford Co., Pa. 2; 7 mi. NW Immler, Bed- sured as jaw lies on buccal surface. 2. C-M3 - ford Co., Pa. 1; 3-1/2 mi. E Tyrone, Blair Co., greatest length, crowns. 3. Depth of lower Pa. 2.

Table10- Summary, cranial measurements in millimeters, Blarina brevicauda Locality O.R. Total length of skull Minnesota 14 24.15 23.5-25.7 .70 2.90 E. Springfield, Pa. 73 23.44 21.8-25.0 .57 2.43 Sink No. 4, CM 5838 1 25.6 Cranial Breadth Minnesota 15 12.94 12.3-13.9 .41 3.16 E. Springfield, Pa. 72 12.51 11.7-13.2 .33 2.63 Rostral breadth Minnesota 21 3.55 3.2-3.9 .19 5.46 E. Springfield, Pa. 77 3.14 2.9-3.4 .13 4.14 Sink No. 4 2 3.55 3.4-3.7 Natural Chimneys 3 3.40 3.2-3.7 Maxillary breadth Minnesota 21 8.43 8.0-9.2 .35 4.13 E. Springfield, Pa. 77 7.84 7.0-8.3 .22 2.80 Sink No. 4 2 8.50 8.1-8.9 Natural Chimneys 4 8.31 8.0-8.7 Upper toothrow Minnesota 21 6.42 5.6-7.2 .41 6.33 E. Springfield, Pa. 77 6.25 5.8-6.6 .19 3.04 Sink No. 4 2 6.45 6.2-6.7 Natural Chimneys 8 6.59 6.4-6.8 .18 2.73 Interorbital breadth Minnesota 17 6.24 5.7-7.0 4.65 E. Springfield, Pa. 77 5.95 5.7-6.6 .19 3.19 Sink No. 4 1 6.30 Natural Chimneys 2 6.15 6.0-6.3

150 THE NATIONAL SPELEOLOGICAL SOCIETY Table 11 - Summary of mandibular measurements in millimeters, Blarina brevicauda Locality X O.R. Total length Minnesota 19 16.20 14.9-17.6 .67 4.13 Penna., modern 17 15.15 14.6-16.0 .32 2.09 Sink No. 4 29 15.33 13.8-17.2 .88 5.74 Toothrow, C,- Minnesota 20 6.29 ± .04 5.8-6.6 .19 3.02 Penna., modern 24 6.06 ± .03 5.7-6.3 .19 3.13 Sink No. 4 36 6.14 ± .06 5.5-6.8 .39 6.48 Natural Chimneys 9 6.43 6.2-6.6 .13 2.02 Width of lower incisor Minnesota 19 1.49 1.3-1.6 .07 4.69 Penna., modern 22 1.31 1.2-1.5 .08 5.92 Sink No. 4 41 1.42 1.3-1.6 .09 6.84 Frankstown Cave 1 1.3 -- Natural Chimneys 16 1.39 1.3-1.45 .07 5.03 Width of mandibular condyle Minnesota 22 4.16 3.9-4.6 .19 4.56 Penna., modern 24 3.85 3.7-4.1 .13 3.37 Sink No. 4 37 3.91 3.3-4.5 .30 7.67 Frankstown Cave 3 3.80 3.7-3.9

Carnegie Museum: Section of Vertebrate 225 M. cf. lucifugus, 13 Pipistrellus, 1 Fossils. Natural Chimneys local fauna, Au- Eptesicus). It was also the commonest gusta Co., Va., entire collection; Frankstown Myotis at Natural Chimneys, Va. ( 39 M. Cave, Blair Co, Pa., entire collection; Sinkhole keenii, 5 Myotis sp.). Keen's bat is not the No. 4, New Paris, Bedford Co., Pa., entire commonest Myotis in the area today. The collection. Pennsylvania mammal survey during the years Minnesota Museum of Natural History. 1946 to 1951 collected an average of 6 M. 1 mi. NE Valentine Lake, Ramsey Co., 3; lucifugus for every Myotis keen ii (see bibli- Corlos Avery Game Refuge, Anoko Co., 1; ography, Roberts and Early, 1952). 2 mi. S of Reno, Houston Co., 1; La Cres- ,11yotis cf. lucifugus (LeConte)--Little brown cent, Houston Co., 2; Washington Co., 1. bat Order: Chiroptera Material: 194 partial skulls; 29 left, 31 right Family: Vespertilionidae maxillae. .11y otis keenii (Merriam )--Keen's bat Stratigraphy: From 4.3 m to 8.5 m level; Material: 261 partial skulls; 41 left, 40 right diminishing in numbers with depth. At the maxillae. 5.2 level, 14 percent of all individual animals were of this species; at the 8.5 m level, only Stratigraphy: From 4.3 m to 8.5 m level; dimin- 2 percent (fig. 28). ishing in numbers with depth. At 4.3 m, 24 percent of all individual animals were of Remarks: This is the commonest species of this species. At 8.5 m, only 2 percent (fig. Myotis in the Appalachians today. The iden- 28). tification remains provisional because of the Remarks: Myotis k een i was the commonest close similarity of skeletal remains of M. bat at all levels of the deposit. (302M. keen ii, lucilugus, .11. soda/is and M. austroripar-

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 151 his. Both M. keenii and M. luctfugus range Burrowing rodents, such as the woodchuck, north into the Canadian life-zone today and are a constant source of contamination of are the only eastern species of Myotis to do older by younger material in archaeological so. sites throughout the East. The New Paris .11y otis, species ? --Little brown bat No. 4 specimen may or may not have been Material: 1,221 left, 1,216 right mandibles; intrusive although Marmota is common in 106 left, 98 right maxillae; 34 partial skulls. most late Pleistocene deposits in the area and Stratigraphy: From the 2.3 m to the 9.7 m level. ranges north to 65° N lat. Alveolar length (see fig. 28). P4 to M, measured 19.6 mm (CM 5746), a small adult. Remarks: No attempt was made to identify mandibles or skull fragments beyond genus. Cite//us tridecemlineatus (Mitchell)-- No specimens were observed that fell into the Thirteen-lined ground squirrel large M. griscescens size range nor the small Material: 2 partial skulls; 3 left, 5 right maxillae; M. s. leibii class. It is felt that most if not 8 left, 6 right partial mandibles. all of these specimens are M. keenii and Stratigraphy: From 5.2 m level to the 8.2 m M. cf. /ucifugus as represented by the cranial level (fig. 29). material in the two previous species accounts. Remarks: A minimum of eight animals were Pipistrellus cf. subflavus (F. Cuvier )-- represented. Seven were immature with de- Pipistrelle ciduous premolars. One, CM 6041, a right Material: 1 skull; 4 left, 1 right maxilla; 8 maxilla with M2 -M3 was a young adult. left, 13 right mandibles. The specimens appear to be slightly larger Stratigraphy: From the 4.3 m to the 6.4 m than the average for the modern northern level. Absent from lower levels (fig. 28) race C. t. tridecemlineatus (Mitchell) but Remarks: Pipistrellus occurs only locally in well within its range of variation (Howell, the southern portions of the Canadian life 1938, p. 107). zone today. Though never common in any one eastern late Pleistocene deposit, these ground squirrels Eptesicus cf. grandis (Brown)—Big brown bat are known from four other sites in what is now the forested East: Cumberland Cave, Mary- Material: 1 mandible; 1 maxilla with P4 -M3 , land; Bootlegger Sink, Pennsylvania; Natural 1 humerus. Chimneys, Virginia; and Robinson Cave, Ten- Stratigraphy: From 5.2 m and 6.4 m level nessee. (fig. 28). The animal is presently confined to areas Remarks: Referred to E. grandis because of well-drained upland prairie north to central of size. Measurements agree with the type Alberta and Saskatchewan. Its presence series and are larger than modern Pennsyl- throughout a large area of the East in early vania specimens. Whether E. grandis is a post-glacial time is compelling evidence for the subspecies (Brown, 1908), a species (Gidley former presence of grasslands in that area. and Gazin, 1938) or is invalid as a taxon must await further studies on both living and Tam ias st riatus (Linnaeus )--Chipmunk fossil brown bat populations. Material: 1 partial skull; 5 left, 9 right maxillae; 24 left, 27 right mandibles. Order: Rodentia Stratigraphy: From the 3.3 m levelto the 7.3 m Family: Sciuridae level. They were not present in the bottom 2 m of the deposit (fig. 29). Marmota monax Linnaeus—Woodchuck Remarks: Chipmunks constituted 32 percent Material: 1 left mandible with M, to 1V13 , 3 of the sciurids from New Paris No. 4 and are isolated molars, 2 partial incisors. represented by at least 27 individuals. In the Stratigraphy: From the 6.0 m level. Recent New Paris No. 2 collection they account Remarks: At least one animal was represented. for 83 percent (114 individuals ) of all sciurids.

152 THE NATIONAL SPELEOLOGICAL SOCIETY No correlation was found between stratigraphic position in the sinkhole and size. However, Sinkhole No. 4 specimens were larger than Recent Pennsylvania material, as represented by the Sinkhole No. 2 collection. Alveolar length of the lower cheek teeth was compared and the results are presented below. The populations differ significantly. The probability of their means being drawn from the same population is less than .1 percent.

4 Table 12 — Tam/as striatus (Linnaeus), alveolar length, P —M', measurements in millimeters.

Locality Approx. Age X O.R. v N New Paris No. 4 11,300 B.P. 6.74 6.3-7.1 .2 6.67 30 New Paris No. 2 1,875 B.P. 6.28 5.8-6.8 .18 2.86 114 One left maxilla, CM 6044, with full dentition, had a well developed 133 (see fig. 18c). This is a character of the genus Eutamias (see Jones, 1960 for exception) but virtually unknown in Tam ias. One unilateral example of P3 was found in an examination of 360 modern skulls (an adult male, Erie County, Pa., CM Mammal No. 25145, fig. 18b). De- spite the presence of other midwestern forms Figure 18. in the deposit (Pedioecetes, CiteIlus), CM Presence or absence of P3 in chipmunks. 6044 is more probably an aberrant Tam/as a. CM Mammal 12679- Eutamias sp., right rather than a Eutamias. Its size agrees with P3-M3. Note normal, well developed P3 at the rest of the Sinkhole No. 4 chipmunk right. b. CM Mammal 25145 - Tamias stria- material, and is much larger than Eutamias tug, right P 3-M3. Note abnormal presence minimus the only "eastern" species of the of P3 at right. c. CM 6044 - d. Tamias stra- genus occurring as far east as western Quebec tus. Sinkhole No. 4, New Paris, Pa. 5.2 about 800 km north of New Paris. Unless m level. Left upper toothrow, note P3 at additional material can be found to substan- left. tiate the presence of Eutamias in the late Pleistocene of the Appalachians, this specimen, from the Natural Chimneys local fauna are despite the presence of 33 1 , is considered a of equal size and both are larger than modern Tam /as. Unfortunately, the maxilla shows no material from the area. Measurements for length other diagnostic characters. of lower toothrow (n=17) were larger than the observed maximum of 37 G. s. macro/us, Glaucomys sabrinus (Shaw )--Northern fuscus, and co/oratus from the eastern Appa- flying squirrel lachians. The fossil material averaged 13 per- Material: 13 left, 15 right maxillae; 13 left, cent larger than modern Pennsylvania G. s. 14 right mandibles. macro/us, 7 percent larger than central Cana- a Stratigraphy: From 5.2 m to 6.7 m levels dian G. s. sabrinus (4 specimens), and can (fig. 29). be matched by Alaskan specimens of G. s. Remarks: Glaucomys sabrinus is another yukonensis and G. s. zapbaeus. They are species that follows "Bergmann's Rule." exceeded in size by specimens of G. s. Specimens from eastern United States are the tus from Idaho. Comparative measurements smallest. Size increases to the north and to the are given below. Measurements were taken from west. Specimens from New Paris No. 4 and Howell, 1918; Handley, 1953; and Guilday,

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 153 1962. Fifteen C. s. macrotus from Pennsyl- ments, to average 16 percent larger than the vania (CM Mammal no. 31612-17; 36390-9) smaller form in those deposits. There was no were used for direct comparison. Alveolar overlap in measurements. This late Pleistocene length of upper toothrow, when found in the G. sabrinus is 13 percent larger than modern literature, was converted into alveolar length Appalachian races of that species, again with of lower toothrow by multiplying by .94, a no overlap in measurements. Comparable sized constant found to work among modern Penn- individuals are found in northern and western sylvania specimens. races today (see table 13, p. 154). The smaller late Pleistocene species has been identified as Table 13 — Alveolar length, lower toothrow, P4 —M3, G. voIans. Unfortunately, their measurements measurements in millimeters, Glaucomys sabrinus average larger than modern G. volans by some 8 percent. This makes them as large N X O.R. as many modern G. sabrinus macrotus. The Late Pleistocene size difference is not statistically significant Natural Chimneys, Va. 14 7.8 7.3-8.4 and the ranges are almost identical. New Paris No. 4, Pa. 3 7.7 7.6-8.1 The smaller late Pleistocene form is iden- Modern fled as G. volans, not on morphological Eastern U. S. grounds — in this respect they are signifi- G. s. coloratus 7 6.9 6.7-7.1 cantly smaller than the G. sabrinus with G. J. fuscus 5 6.7 6.5-6.9 which they occur. In other words, the slightly G. s. macrotus 23 6.6 6.0-7.0 larger G. volans from the late Pleistocene Canada cannot be confidently separated from mod- G. .r. sabrinus 4 7.2 6.8-7.6 ern G. s. macrotus of the same area, but G. c. makkovikensis 2 7.1 6.9-7.4 can be differentiated from the extremely large Alaska G. sabrinus from these same late Pleisto- G. s. yukonensis cene deposits. G. v. zaphaeus 5 7.6 7.3-7.9 An alternate treatment assigning the small- Idaho er group to G. s. macrotus and assuming a G. s. hullatus 3 8.7 8.6-8.7 chronocline would require the presence of intermediate individuals. These are not pre- Glaucomys volans (Linnaeus )—Southern sent. Likewise, the assumption that the small- flying squirrel er form is G. sabrinus while the larger form is an undescribed taxon is equally as Material: 2 left maxillae; 1 left, 1 right partial unrealistic. The best explanation appears to mandible, fragmentary. be that both G. volans and G. sabrinus Stratigraphy: From the 2.7 m to 6.1 m levels were of larger size during the late Pleistocene (fig. 29). in the central Appalachian region. Remarks: Glaucomys volans can usually be distinguished from Glaucomys sabrinus by its inferior size. Occasional individuals of G. sabrinus from the southeastern portion of their modern range are as small as some Table 14 — Alveolar length, lower toothrow, G. no/ans. If the dental measurements of a measurements in millimeters, GIaucomys volans particular individual fall within this area of overlap, specific identification by those charac- N X O.R. ters is not possible. Modern, Pennsylvania* 38 6.0 5.6-6.4 There are two sharply differentiated species Late Pleistocene of flying squirrels in both the New Paris Natural Chimneys 17 6.5 6.3-6.9 No. 4 and the Natural Chimneys local faunas. New Paris No. 4 2 6.4 The larger has been identified as G. sabrinus. Data from Howell, 1918; Guilday, 1962; Doutt, It was found, on the basis of denial measure- unpubl.*

154 THE NATIONAL SPELEOLOGICAL SOCIETY Tumiasciurus hudsonieus tenuidens ( Hay) as well. The ratio in 36 T h. loquax from --Red squirrel Pennsylvania averaged 38 percent; in 30 7'. Sciurus tenuidens Hay, 1920 h. tenuidens from New Paris No. 4, 37 Sciurus (Tamiasciurus)tenuidens Hay, percent. T. h. tenuidens had a relatively massive jaw apparatus averaging 10 percent larg- Gidley and Gazin, 1938 er than the modern Appalachian populations Material: 18 partial skulls; 10 left, 12 right of the species. The lower incisors are, in maxillae; 17 left, 21 right mandibles, plus fact, of greater antero-posterior diameter than associated skeletal material. in modern comparative material, but they are Stratigraphy: From 3.6 m to 8.2 m level not relatively narrower. (fig. 29). Remarks: The extinct Tamiasciurns tenui- Gidley and Gazin's observations on the Cumberland Cave skull (USNM 8164) are dens Hay was based upon a lower right jaw confirmed by the New Paris No. 4 collection. fragment and incisor, mistaken by its des- While T. h. tenuidens is distinctive, none criber for an upper left incisor and premax- of its morphological features are unique. They illary fragment. The characters given in the can be duplicated, singly or in combination, type description (Hay, 1920, p. 105): "Up- throughout the range of the species. Most per incisors broad and unusually thin; front of Gidley and Gazin's characters are a func- border rounded." merely describe differences tion of size — broad frontal region, deep muz- between upper and lower incisors of tree zle, deeper jugal, etc, and are thus redundant squirrels, rather than any interspecific differ- (although these differences are obvious when ence. Gidley and Gazin (1938, pp. 54-56, making comparisons between T. h. tenuidens fig. 30) redescribed the form using three and most modern subspecies ). T. h. tenni- partial skulls and four mandibles from the dens is equaled in size by T. h. fremonti Cumberland Cave, Maryland. Seven partial and exceeds all eastern forms in dental mea- mandibles, two maxillae and one humerus surements. Measurements of body elements, from Natural Chimneys, Virginia are referred if anything, are a little smaller than modern to this form as well. Pennsylvania material, however(table 15). The additional well preserved material from Viewed from the side, the dorsal profile of New Paris No. 4, representing the remains the skull is stated to be flatter than in the of at least 30 animals, allows the reappraisal modern "species". This is true if the com- of the taxonomic status of the form tenui- parison is made between T. h. tenuidens dens. It is here considered an extinct sub- and T h. loquax, but a series of T h. species of the modern Tam iasciurus hud- preblei from Aklavik, Northwest Territories sonicus (Erxleben). agreed with T. b. tenuidens, and the char- In their redescription of T. h. tenuidens, acter appears to be of subspecific value at Gidley and Gazin (1938, p. 54) stated that most. the type incisor "is relatively narrow trans- Inflation of the auditory bullae, noticeable versely and of greater antero-posterior diam- in T. h. tenuidens, appears to be correlated eter than in S. !Judson/ens" and that the with latitude in modern populations. It is lower incisors of the Cumberland Cave ma- more pronounced in specimens of T. h. terial are "of greater antero-posterior diam- preblei and T. h. ungavensis, less so in eter" than in modern hudsonicus. Neither T. h. minnesota and T h. loquax. Hay's nor Gidley and Gazin's measurements The most striking character complex seen bear this out, however. The ratios of trans- in T. h. tenuidens is that associated with verse diameter to antero-posterior diameter, the jaw musculature. The zygomatic plate of based upon their measurements, are 33-1/3 the maxilla is flaring and deeply excavated percent and 42.8 percent respectively. Modern above the infra-orbital foramen. Ridges and ratios vary from 32 percent to 44 percent, fossae for attachment of masticatory muscula- a range which includes all of the fossil material ture, both on the skull and mandible, are BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 155 sharp and prominent. Some individuals of UTAH T. b. fremonti agree quite well with the Daggett County - 4, Duchesne County - 2, fossil material although the norm for the Summit County - 15, Uintah County - 9. 7'. h. fremonti appeared less rugged. WEST VIRGINIA Compared with T. b. loquax, the T. b. Marion County - 1, Preston County - 2. tenuidens mandible is quite distinctive, short BRITISH COLUMBIA and deep, with a heavy, deep symphysis and McLennan and Fraser Rivers - Caribou a distinct upsweep to the incisor that con- District - 1, Snowshoe - 3. trasts with the longer, shallower, more pro- LABRADOR cumbent anterior portion of the mandible as Hamilton River - 12. seen in T. b. loquax. The loquax condition is the common one throughout the range MANITOBA of the species. Short, heavy mandibles with cor- Churchill - 4. responding deep masseteric ridge profiles ap- NEW BRUNSWICK pear to occur sporadically in some modern Tabusintac - 1. populations. Examples were noted in series of T. b. fremonti and T. b. ungavensis. NORTHWEST TERRITORIES Aklavik - 8, Charleton Island, James Bay - The massive jaw in T. b. tenuidens appears to be constant in T. h. tenuidens, 6, Great Slave Lake - 8. however, and is quite distinctive when com- ONTARIO parison is made with modern 7'. b. loquax Cochrane - 1, Crooked Lake - 1, Kapus- now inhabiting the area. kasing River - 3, Louisville, Kent County - 2, Missanobi - 1, Moose Factory - 12, Twin Fifty-five modern skulls from all over the Lake, T. & N. 0., Ry. - 1. modern range of the squirrel were checked QUEBEC for the presence of P3. It was absent in 18 cases, present in 37. In 14 examples of T. 3 East Main - 5, Fort Chimo -2, Fort George - b. tenuidens P was absent in 2. 4, Great Whale River - 3, Keromska - 1, Little Mecatina River -1, Natishquan River-11, Richmond Gulf- 1, Rupert House - 1,2 miles SPECIMENS EXAMINED - 182 AS FOLLOWS: below 5th Camp, St. Margaret River - 1, Camp 15, St. Margaret River - 1, 9th Camp, St. Carnegie Museum Mammal Collection Margaret River - 1. Upper Forks, St. Mar- Tamiasciurus hudsonicus garet River - 1. ALASKA SASKATCHEWAN Seward - 1. Emma River - 2. ARIZONA 8200 ft., Coconino County -4, Kaibab, Coconino County - 1. SPECIMENS EXAMINED 14 AS FOLLOWS: MARYLAND Garrett County - 6. Carnegie Museum Mammal Collection MINNESOTA Tam iasc in rus donglasii Detroit - 4, Moorhead - 9. CALIFORNIA PENNSYLVANIA Humboldt County - 2, Plumas County - 2, Allegheny County - 1, Butler County - 2, Tuolumne County - 3. Crawford County - 14, Erie County - 1, Forest OREGON County - 2, Fayette County - 3, Lycoming Bly, Klamath County - 1, Klamath Falls, County - 1, Somerset County - 1, Venango Klamath County - 2, Portland - 1, Joseph - 1. County - 2, Warren County - 1, Potter County WASHINGTON - 2. Olympic Mts. -2.

156 THE NATIONAL SPELEOLOGICAL SOCIETY Maps A and B, Center Leaf

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VOLUME 26, NO. 4, OCTOBER, 1964

Table 15- Skeletal measurements: Tam ias- Locality Y( O.R. N dunes hudson icus tenuidens (Hay), from Sinkhole No. 4, New Paris, Pa. and Recent Occlusial length, P 4-M 3 Pennsylvania s p ec im ens of Tam iasciurus hudsonicus loquax, Carnegie Museum Col- 1 7.37 6.7-8.1 30 lection. Measurements in millimeters. 2 7.58 7.2-8.4 17 Size of T. h. tenuidens 3 7.47 7.3-7.7 4 relative to that of g O.R. N T. h. loquax - percent 4 7.36 7.0-7.6 6 5 7.52 7.1-8.0 19 Total length, humerus 6 7.75 7.5-8.1 16 tenuidens 31.07 30.4-33.1 7 7 7.45 7.3-7.6 2 loquax 31.76 28.9-33.6 31 97.8 8 8.00 7.8-8.2 4 Width, distal end of ulna 7.6-8.3 4 tenuidens 7.93 7.4-8.4 10 9 7.95 loquax 7.98 7.3-8.8 32 99.4 10 8.16 8.1-8.2 3 Ulna, total length 11 8.25 8.2-8.3 2 tenuidens 35.33 33.4-37.5 6 12 8.09 7.7-8.4 9 loquax 100.2 35.25 33.6-37.8 20 13 7.85 7.2-8.4 10 Radius, total length 14 ------tenu idens - - - - loquax 29.32 27.5-31.3 22 Femur, total length tenuidens 39.18 38.5-40.3 11 Length, P4 -IA , loquax 39.69 35.6-41.9 32 98.7 1 6.8-7.8 Femur, width of distal end 7.39 33 ten u Edens 6.83 6.6-6.9 12 2 7.66 7.3-8.0 20 loquax 6.72 6.0-7.5 31 101.64 3 7.65 7.3-8.0 4 Femur, shaft diameter 4 7.31 6.9-7.5 6 tenuidens 2.79 2.7-3.2 16 5 7.58 7.1-7.8 19 loquax 2.94 2.7-3.3 94.9 32 6 7.84 7.6-8.2 15 Tibia, total length tenuidens 44.32 44.0-44.6 5 7 7.60 7.5-7.7 2 loquax 44.40 41.2-47.6 17 99.8 8 7.95 7.8-8.1 4 9 8.05 7.7-8.5 4 10 8.23 8.0-8.4 3 Table 16 - Cranial measurements, in mm, 11 8.30 8.3-8.3 2 Tumiusciarus budsonicus (Erxleben), Car- 12 8.20 8.0-8.4 9 negie Museum Collection. 13 7.92 7.2-8.5 11 Localities: 1. Pennsylvania 14 8.20 8.0-8.3 4 2. Natishquan R., Quebec; Ham- ilton R., Labrador 3. St. Margaret R., Quebec Antero-posterior diameter, 4. Charleton Is., James Bay, N. lower incisor W. T. 1 3.00 2.5-3.3 36 5. Ontario 2 2.80 2.4-3.1 23 6. Hudson Bay, Rupert Houseto 3 2.90 2.8-3.0 4 Richmond Gulf, Quebec 4 2.76 2.4-3.0 6 7. Fort Chimo, Quebec 5 2.95 2.7-3.1 19 8. Churchill, Manitoba 6 2.72 2.6-3.1 15 9. Snowshoe, British Columbia 7 2.55 2.5-2.6 2 10. Great Slave Lake, N.W.T. 8 3.00 2.7-3.3 4 11. Emma Lake, Saskatchewan 9 3.07 2.9-3.4 4 12. Aklavik, N. W. T.; Seward, 10 3.00 2.9-3.2 3 Alaska 11 2.60 2.6-2.6 2 13. Moorhead, Detroit, Minn. 12 2.79 2.6-3.0 9 14. T. b. tenuidens, New Paris 13 2.94 2.3-3.2 13 No. 4, Penna., late Pleistocene 14 3.30 3.0-3.6 29

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 157 Table 16 (Continued) of Georgia. Despite the fact that the well- marked race N. f magister ranges only as far Transverse diameter, lower as lat. 42°, it does not act like a "southern" incisor form. It may occur on mountain summits in 1 1.00 .9-1.2 36 excess of 900 m throughout its range, and 2 .95 .8-1.1 23 appears restricted to the Appalachian mountain 3 1.02 1.0-1.1 4 and plateau region. Neotoma has been re- 4 .95 .9-1.0 6 corded from Cumberland Cave, Maryland, 5 1.03 .9-1.2 19 Natural Chimneys, Virginia and Robinson 6 .97 .8-1.1 15 Cave, Tennessee-all late Pleistocene in age. 7 .85 .8- .9 2 It also inhabits most of the caves in the region 8 1.05 1.0-1.1 4 today. 9 1.05 1.0-1.1 4 Schwartz and Odum (1957, pp. 197-199) 10 1.10 1.1-1.1 3 suggest that Neotoma floridana invaded the 11 .90 .9- .9 2 southern United States from the southwest; 12 1.04 1.0-1.1 9 one branch, the "magister" stock, ascending 13 1.19 1.0-1.5 13 the Appalachians; the other, more southerly 14 1.20 1.0-1.6 29 "floridana" stock, invading the southern Inter-orbital width lowlands and ascending the Atlantic coastal area. This invasion presumably was post- 1 13.68 13.0-14.5 36 Wisconsin. 2 13.17 12.0-14.1 21 3 13.65 13.1-13.9 4 It would appear probable, in the light of 4 13.3 12.8-14.3 6 the animal's presence as far north as New 12.2-15.1 18 Paris during the late Pleistocene, that N. f. 5 13.3 magisier, the largest, and, morphologically 6 13.8 12.8-14.4 14 and ecologically, the most well-marked sub- 7 12.2 12.1-12.4 2 species, survived the Wisconsin glaciation in 8 13.2 13.0-13.5 4 the southern Appalachians. It then repossessed 9 14.2 13.5-15.0 4 the northern sector of its present range very 10 13.7 13.0-14.5 3 rapidly after deglaciation had commenced. Its 11 12.8 12.7-13.0 2 northern advance may have been halted more by 12 14.2 13.6-14.8 9 the lack of cliff-talus-cave habitat to which it 13 13.6 12.0-14.5 13 appears strongly attracted than by climatic 14 13.9 13.3-14.7 10 factors. Peromyscus cf. man iculatas (Wagner )--Deer Family: Cricetidae mouse Subfamily: Cricetinae Material: 4 partial skulls; 72 left, 93 right Neotoma cf. floridana (Ord)--Wood rat maxillae; 128 left, 117 right mandible.l.. Material: 1 partial left mandible, M1 -M, ; 1 partial left mandible, M2 -1V13. j 1 right IA' ; Pero myscus cf. leacopus (Rafinesque)-- 2 upper, 1 lower incisor; 1 left, 1 right M i . White-footed mouse Stratigraphy: 5.5 m to 6.4 m level. Material: 2 left, 1 right maxillae; 7 left, 11 Remarks: Two animals were represented. The right mandibles. presence of on one mandible, CM flowstone Peromyscus, species ? 5751, argues for some age. Neotoma , despite right the fact that it ranges only two degrees further Material: 3 partial skulls; 72 left, 93 north today, may have been a member of the maxillae; 64 left, 70 right mandibles. Pleistocene fauna. Its modern distribution in Remarks: Mice of the genus Peromyscus the East is enigmatic - present in the Appalach- were common throughout the deposit. P. ian mountains and in the Atlantic lowlands lencofius (the only Peromyscus living at the of the South as far north as Virginia, but site today) occurred uncommonly at all levels absent from the intervening Piedmont north with little change of numbers while P. Mani-

158 THE NATIONAL SPELEOLOGICAL SOCIETY u/atus increased markedly in the middle and P. /rat-opus is probably not a member lower levels. of the late Pleistocene fauna. The presence Table 17— Stratigraphic distribution of two species of modern rodent droppings at the 5.9 m of Peromyscres, New Paris No. 4, Pa. level plus the ever present possibility of con- Depth in Meters tamination of the lower levels by modern P. in Species 0.25 m 2.6-5.5 m 5.6-8.5 m 8.6-10.0 m leacopus (and Pitymys p etorum) falling in during the four year excavation period P. leucopur 100% 10% 7% 4% P. maniculatus 0 90% 93% 96% leaves the presence of this temperate species individuals 3 10 86 24 in the primary boreal fauna open to question.

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Figure 19. Crown patterns, upper and lower molars, Microtinae, Sinkhole No. 4, New Paris, Pa. I 3 1. CM left left 1 3 5633, 6267 - Synaptomys cooperi, M -M , 3 M -M I 2. CM 6911, 6724 - Synaptomys borealis, right M'-M 2, left M -M, 3. CM 6723, 6258 - Dicrostonyx hudsonius, left M`-M , left M,M; I 3 4. CM 6725, 5869- Phenacomys cf. ungava, left M -1%A1 , left M1-M3 5. CM 5647, 6701 - Clethrionomys gapperi, left MLM , left M I-M, 1 t 1 3 6. CM 6249, 5541 - Microtus xanthognatus, left M -M3 , left M M 7. CM 6188- Microtus pennsylvanicus, left M'-M , left3 M1-M, 8. CM 5595, 6186 - Microtus chrotorrhinus, leftt 3M'-M , right M1-M 3 9. CM 5683, 6015- Pitymys pinetorum, right M -M , right M,M3

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 159 P. led/cop/is ranges throughout central and Presence of this lemming is strong evidence eastern North America north to lat. 46° N. for the near presence of tundra. It may oc- It is usually a deciduous forest form and does casionally range into the fringes of the Hud- not penetrate far into the coniferous forests of sonian life-zone (Harper, 1961) but is a true the Canadian life-zone. P. man icalatus ranges barren-ground form. The fact that only three over most of North America except the low- animals were recovered suggests that typical lands of the southeastern United States and tundra per se did not persist near the mouth the far northern tundra. It ranges locally into of the sinkhole during the period of infill. the Arctic life zone (Harper, 1961), and its The source could have been a relict colony presence is quite compatible with boreal con- on Allegheny Mountain 10 km to the west or ditions. perhaps on the crest of Chestnut Ridge itself. Unlike the brown lemming (L emmus) which Table 18 — Peromyscus, length of first lower molar, is notorious for its cyclic migrations the measurements in millimeters, New Paris No. 4, Pa. collared lemming is relatively sedentary. This Species O.R. makes it likely that the source of the New P. leucopus 18 1.58 1.4-1.7 Paris No. 4 specimens was nearby. P. maniculatus 48 1.52 1.4-1.7 Subfamily: Microtinae Table 19 — Measurements, in mm, Dicrostonyx D icros I onyx buds on ins ( Pallas )--Labrador budsonius (Pallas) New Paris No. 4, Pa. collared lemming Occlusal length Upper Lower Material: CM 6258. 1 partial right mandible, 1st molar 2.4 mm 3.4 mm full dentition (FIG. 20). 1 partial left mandible, 2.5 3.1 1 2 full dentition; 1 partial palate, left M -M . 3.4 CM 6695. 1 left M . CM 7639. 1 upper 1 2nd molar 1.9 1.7 left incisor. CM 6723. 1 partial palate, left 2 t 2 2 1.9 1.7 M' -M , 1 right M , 2 right M , 1 left M . 1.9 Stratigraphy: One at the 6.1 m, two at the 1.9

6.4 m level (fig. 30) 3rd molar 1.5 Remarks: A minimum of three animals were 1.5 recovered (erroneously reported as four in Alveolar length, M1—M3 Guilday, 1963). Dicrostonyx is found throughout the Arctic regions of the world 6.8 mm wherever there is tundra. The seasonal hyper- 7.3 trophy of the front claws and the fact that it turns white in the winter make it unique among rodents. New Paris No. 4 is the only SOMESTME,„. North American fossil record ofDicrostonyx, 1 outside of Alaska although it is common in the ..0-0000"t"' Pleistocene of Europe. The species D. budson- ins is isolated at the present time on the Ungava Peninsula of northern Quebec and Labrador. It is differentiated from D. groenlandicus of the western Nearctic, the Canadian Arch- ipelago and Greenland, by characters in both upper and lower dentitions (Anderson and Rand, 1945; Hall and Kelson, 1962). On the basis of these, the New Paris No. 4 specimens are D. !Eidson ius . (See fig. 19 for drawing of dental pattern.) The possible Figure 20. zoogeographical history of Dicrostonyx in CM 6258- Dicrostonyx hudsonius. S in k- North America during the late Pleistocene is hole No. 4, New Paris, Pa. Crown and discussed in Guilday, 1963. buccal view, right mandible, mm grid.

160 THE NATIONAL SPELEOLOGICAL SOCIETY Pbenacomys cf. angava Merriam--Spruce of its present range, becoming smaller with vole increasing latitude. Smallest modern popula- Material: 13 partial skulls; 2 left, 4 right max- tions are from eastern Canada. illae; 46 left, 37 right mandibles, or M, 's. Measurements of one specimen from the Stratigraphy: General from the 4.9 m through Recent Sinkhole No. 2 fauna lie at the upper the 8.5 m level, becoming scarcer with depth limit of the modern Pennsylvania material (fig. 30). and are significantly larger than those of the Remarks: The spruce vole lives in pine and late Pleistocene Sinkhole No. 4 sample. spruce forests of the Canadian and Hudsonian Wetzel, 1955, postulated a post-glacial life-zones of North America (Foster, 1961). spread of Synaptomys north and east from It is instructive to note that both the spruce a southwestern Appalachian "refuge" into vole and the red-backed vole (Cletbrionomys), those areas of central and eastern North Amer- another woodland microtine, become com- ica freed from glacial ice or its effects. The moner in the higher level's of Sinkhole No. 4 small late Pleistocene Sinkhole No. 4 popu- while other microtines with the exception of lation, finding its closest modern counterparts the pine vole (Pitymys) and the southern bog in population samples from the extreme lemming (S. cooperi) decrease in number northern limits of its present range, would in the upper levels. Presumably, this is in seem to indicate that the modern clinal pattern response to an ecological change from more open conditions to a closed forest. in the Appalachian area was established as early as the Wisconsin and reflects adaptations to full-glacial conditions. Late Pleistocene S. coop- Table 20- Measurements in millimeters, Pbrearo cf. Itngava. New Paris No. 4, Pa. eri from the boreal Natural Chimneys local Measurement I O.R. ti V fauna are equally as small, while those from length, M, 2.67 .01 1.8-3.3 .08 .1 .007 3.10 7 .28 59 length, M,-M, 5.50 I .09 4.9-6.1 .29 1.06 5.27 1.17 10 Robinson Cave, Tennessee, south of the ani- length, M4-M' 4.5 4.3-5.0 4 mal's present range, and of similar age, are incisive foramen 3.5 3.3-3.7 4 inter-orbital breadth 2.9 2.9-3.0 3 larger than modern Pennsylvania S. C. cooperi and presumably represent the S. c. stonei Synaptomys cooperi Baird - Southern bog stock of the southern Appalachians. If sub- lemming specific patterns were established by the rigor- Material: 2 partial skulls; 15 left, 16 right man- ous selection of full-glacial times, then sub- dibles or M, 's. sequent reinvasion by the animal into more Stratigraphy: From the 4.9 m to the 6.7 m level (fig. 30). Table 21 - Synaptomys cooperi, occlusal length M,-M 5 Remarks: The southern bog lemming was one Measurements in millimeters. of only four microtines from Sinkhole No. Localities: 1. Recent south-central Pennsyl- 4 (Cletbrionomys, Pitymys, Phenacomys, vania, CM 2. Recent, West Virginia. From S. cooperi) that did not increase in numbers McKeever, 1954. with depth. It was concentrated in the upper 3. Recent, Sinkhole No. 2, New Paris, Pennsylvania. CM levels of the deposit in contrast to the northern 4. Recent, Quebec, Canada. CM bog lemming S. borealis which replaced 5. Late Pleistocene, Sinkhole No. it in the lower levels of the sinkhole. The 4, New Paris, Pennsylvania. CM 6. Late Pleistocene, Natural Chim- Sinkhole No. 4 population of S. cooperi neys, Virginia. CM differs significantly in size from a modern 7. Late Pleistocene, Robinson Cave, Tennessee. CM sample of Synaptomys c. cooper/ Baird from the same area (south-central Pennsyl- Locality O.R. V vania). It is smaller in all dimensions. It agrees 1. 5.92 5.07 5.3-6.6 .36 5.05 6.08 5 .85 25 2. 6.35 5.9-6.8 17 most closely with specimens from eastern 3. 6.6 1 Quebec and Canada. Synaptomys cooperi 4. 5.77 ± 09 5.6-6.1 .21 2 .06 3.63.1 1.15 5 exhibits a negative Bergmann's response. It 5. 5.62 5..04 5.4-5.9 .13 5.03 2.305 .57 8 6. 5.5 5.4-5.6 2 is largest in the southern and western portions 7. 6.25 6.1-6.4 2

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 161 Table 22- Synaptomy! cooperi, occlusal length M,, Cietbrionomys gapperi (Vigors)-- measurements in millimeters. Red-backed vole Localities as in Table 21 (Locality 2 and 4 ommitted). • Material: 15 partial skulls; 254 left, 260 right Locality O.R. d V mandibles or M, 's; 54 left, 51 right maxillae. 1. 2.48 1.03 2.1-2.7 .19 .1- .02 7.66 I 1.08 25 3. 2.80 1 Stratigraphy: Present at all levels below 4.0 5. 2.41 1.02 .L.5 -2.5 .09 5.01 3.73 5 .60 20 m. At 4.9 m Cletbrionomys was the com- 6. 2.39 .01 2.2-2.5 .05 5.01 2.92 5 .55 14 7. 2.57 1- .04 2.3-2.9 .20 5.03 7.78 ± 1.26 19 monest small rodent in the deposit. Below 5.8 m it became progressively scarcer (although northerly areas has kept pace with climatic most microtines increased in numbers) (fig. retreat. Modern clinal variation in this form 30). represents the Pleistocene clinal pattern, dis- placed to the north and undoubtedly modified Remarks: Clethrionomys gapperi and C. by local conditions, but still discernible in rut,/us of the western Nearctic are identical broad outline. in dental pattern but can be differentiated by the degree of ossification of the hard palate. Synaptomys borealis (Richardson)-- Using this as a criterion, where this character Northern bog lemming could be observed, our specimens are C. Material: 31 partial skulls; 71 left, 66 right gapperi. The post-palatal bridge was incom- mandibles or M, 's. plete on one immature specimen, complete Stratigraphy: From the 4.9 m through the on each of five adults. Bee (Bee and Hall, 8.5 level. Becomes commoner in lower levels 1956, p. 116-117) suggests that the degree (fig. 30). of closure of the post-palatal bridge may be a function of the length of growing season, Remarks: At the 5.2 m level, southern bog and that this character may have no real tax- lemming remains exceeded those of the north- onomic value. ern bog lemming (8 S. cooperi, 3 S. boreal- Cletbrionomys does not now occur at is), but below this level S. borealis became New Paris*. Extensive trapping by the Penn- the commoner of the two. From the 7.0 m sylvania Mammal Survey failed to produce it to the 8.8 m level only the northern bog lem- at the site, although it is common on the ming was found. crest of Allegheny Mountain 10 km to the These two species co-exist today only in a west. Cletbrionomys occurs throughout narrow belt of southeastern Canada at the the northern and mountainous areas in the juncture of the Hudsonian and Canadian life- state today in cool, moist, rocky forest with zones. All lines of evidence from Sinkhole good ground cover. It was not present in the No. 4 indicate that similar ecological conditions occurred during the deposition of the Recent Sinkhole No. 2 microtine component. matrix. Changing proportions of the two At present the site appears to be too dry to species with depth is taken as an indication support this vole. Its presence in Sinkhole No. 4 fauna is presumed to be indicative of of climatic change during deposition. moister, cooler conditions. Its reduction in Slightly over four S. borealis were re- numbers with depth, associated with an increase covered for every one S. cooper/. At Natural of grassland microtines and a decrease in other Chimneys, Virginia lat. 38° 22' N, the situation woodland forms, is an indication of less was reversed with four S. cooperi for every heavily forested conditions during the initial one S. borealis. phases of infilling. Table 23 - Cranial measurements in millimeters, Svnuplomt , 0 or, a h■ (Richardson), New Paris No. 4, Pa. Meosumment II OR. V M,, occlusal length 2.91 .1 .01 2.3-3.2 17 ,..01 5.84 .46 HO * Although both Dr. Doutt and the senior author M,-M„ occlusal watched what they took to be a Clethrionomys length 6.61 1 .03 6.0-7.0 24 1 .03 3.63 L .54 22 for a full ten minutes about p.m., April M-M, occlusal 2:00 length 6.94 .04 6.6-7.3 .14 .02 2.01 L .41 12 29, 1962, making repeated short forays for dead incisive foramen leaves from the log shoring of the entrance to length 5.03 4.4-5.4 6 inter-orbital width 3.45 3.1-3.8 4 Sinkhole No. 4.

162 THE NATIONAL SPELEOLOGICAL SOCIETY Comparison of Sinkhole No. 4 Cletbrio- Table 25 - Length in millimeters 11V-34 t, Micron, pennsylvanicus nomys material with modern Pennsylvania A. Modern Pennsylvania (data from Goin, 1943) specimens from Pike County revealed no sig- B. Late Pleistocene, New Paris No. 4, Pa. nificant differences. C. Modern, New Paris No. 2, Pa. Table 24 - Measurements in millimeters, Loc. Sex O.R. V Clethrionomys gappert, New Paris No. 4, Pa. A. male 6.46 5, .05 .51 5- .03 7.88 5 .52 56 A. female 6.42 5.04 .45 5.03 6.99 5 .48 48 O.R. V B. 5.72 5..03 5.0-6.4 .22 ± .02 3.89 5 .34 65 Occlusal length M L-M3 C. 6.16 6.0-6.4 3 5.07 I- .07 4.3-5.6 .31 6.11 5 1.03 17 Occlusal length M1-M3 The one intact skull from New Paris No. - 5.04: .03 4.5-5.4 .22 1 .02 4.36 5 .50 37 4, CM 6945, measured 27.7 mm in total Occlusal length M, length. Judging from Snyder's data (Ibid., 2.23 ± .004 1.8-2.7 .07 J.: .003 3.13 .13 254 p. 211), this is not a small skull. Goin (lb 1(1., Microtus pennsylvanicus (Ord)-- Meadow p. 216) gives 27.2 mm as the average total vole length of skull of population A (above). The Material: 156 partial skulls; 8 left, 8 right fossil skull, CM 6945 (assuming standard maxillae; 1 left, 1 right mandible. mensuration techniques) is larger than the average of that Recent population but the Stratigraphy: Found in increasing numbers length of its upper molar teeth, 5.9 mm is from the 4.9 m level to the bottom of the sinkhole (fig. 30). only slightly larger than the mean of the sample front which it was drawn, and significantly Remarks: The present taxonomic picture of smaller than Modern Pennsylvania material. Microtus pennsylvanicus lacks clarity at the Observations such as this, coupled with the subspecific level. Bailey (1900) presents a pic- lack of knowledge, on a continental scale, ture of decreasing size with increasing latitude. of cranial variation in this vole, make it im- Rand (1943) is in agreement but states (p. 115) possible to assess the taxonomic standing of that while, in general, northern races (drum- the Sinkhole No. 4 population. It is hoped mondi) are smaller than southern races (penn- this collection will be restudied by a stu- sylvanicus, modestus), some northern pop- dent versed in the Recent geographical vari- ulations of drummondi are larger than others ation of the species. further south. Edwards (1963) records specimens (cf. labradorius) from Richmond Gulf, Micro ties chrotorrh inns (Miller )--Rock vole Quebec as large or larger than M. p. penn- Material: 39 partial skulls; 3 left, 6 right max- sylvanicus from central New York (Hamil- illae; 1 right mandible. ton, 1943). Snyder (1954) demonstrated that Stratigraphy: From the 2.7 m to the 8.8 m cranial differences between local populations level (fig. 30). of this vole from Pennsylvania, even when Remarks: The rock vole occurs in the Canadian rigorously selected for age and sex, could equal life-zone 'east of 95°W. long. In Pennsylvania or exceed what had formerly been considered it is known ony from Wayne, Sullivan, and subspecific thresholds for some currently rec- Luzerne counties approximately 320 km north- ognized forms (fontigenus, enixus, labra- east of New Paris. It is rare and local in cool, dorius). rocky forest situations. The New Paris No. 4 Microtus pennsyl- There was no correlation between strati- vanicus sample, unaged and unsexed, but with graphic position and gross size or dental a low coefficient of variation, is large enough pattern. The alveolar length of the upper molars to assess the population from which it was was compared with that of 52 modern indi- drawn. Compared with modern Pennsylvania viduals from six localities in the Appalachian material these were small animals with short Mountain area from Labrador south to North upper molar rows, deep pterygoid pits, small Carolina. A size dine is indicated with the New auditory bullae, and diverging upper tooth Paris No. 4 material agreeing most closely rows. with Labrador-Quebec specimens.

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 163 Table 26 — Alveolar length in millimeters, M Buccal and lingual salient angles varied Microtus cbrotorrhinur from four to six as follows: Locality Lat . O.R. tY V N Table 28 — Microtus cbrotorrhinus, *Labrador 53 5.9 5.4-6.5 — — 7 M3 dental pattern, salient angles Quebec 50 6.2 5.5-6.7 .3 4.86 20 *New Hampshire 44 6.5 6.2-6.9 — — 7 Buccal Lingual Penna. & W. Va. 40 6.4 5.6-7.2 .5 7.62 10 Locality N Aver. Aver. **West Virginia 38 6.6 6.4-7.0 7 *North Carolina 35 7.1 1 Penna./W. Va. 9 4.3 9 4.4 Quebec 22 4.3 20 4.8 New Paris No. 4 6.0 5.4-6.7 .4 6.18 27 (fossil) New Paris No. 28 4.4 29 4.8 *Data from Komarek, 1932. **Data from McKeever, 1954. The fourth lingual re-entrant varies from fully developed, i.e., deep, cement-filled, cur- There appear to be significant differences in ving caudally, to barely suggested by a slight, the dental pattern of the upper third molar shallow concavity in all three samples. It was between modern Quebec and Pennsylvania/ more highly developed in the Quebec material West Virginia specimens as presented below. (60 percent) than in either the modern Penna./ The Sinkhole No. 4 material agrees with W. Va. material (25 percent) or New Paris modern Pennsylvania West Virginia specimens No. 4(10 percent). in this respect. (fig. 19) The degree of isolation of the individual M' and M 2 are as in M. xantbognathus, triangles, presence or absence of posterior buc- except for their much smaller size. Mandibles cal and lingual salient angles, and the degree and lower dentitions of M. cbrotorrbinus of development of the fourth lingual re- are indistinguishable from those of M. penn- entrant valley, all appear to vary significantly sylvanicas. The one listed above was found in the 114 3 's of the samples studies (20 skulls in association with a M. chrotorrbinus skull. from southern Quebec, 10 skulls from Penn- In summary, the New Paris No. 4 cranial sylvania and West Virginia, and 29 fossil material agrees most closely in gross size with specimens from New Paris No. 4, all Car- modern specimens from Labrador and Quebec negie Museum specimens). and more like Penna./VV. Va. material in the Microtas cbrotorrbinus is usually stated seemingly more primitive dental pattern of 3 to have an M 3 pattern of an anterior crescent M (triangles not as tightly isolated; shallow followed by five closed alternating triangles or non-existent fourth lingual re-entrant). (Hall and Kelson, p. 741). This appears to Microtus xunthognatbus (Leach )--Yellow- be the exception rather than the rule. It was so cheeked vole in only 30 percent of the Quebec specimens, Material: 134 partial skulls; 8 left, 6 right 10 percent of the Pennsylvania and West Vir- maxillae; 344 left, 317 right mandibles or ginia material, and not at all in the New Paris M I 's (fig. 21). No. 4 material. "Normal" pattern in all three Stratigraphy: The commonest terrestrial mam- population samples was: an anterior crescent, mal in the deposit. Remains were found in triangles 1 and 2 confluent, triangle 3 isolated, steadily increasing numbers from the 2.1 m triangle 4 and 5 confluent. Triangle closure through the 8.8 m level where it accounted for w as more advanced in the Quebec material over 40 percent of all mammals recovered than in any other. (fig. 30). ' Table 27 —it chro torrhlert, , M dental patterns Remarks: This is a rare mammal in modern Penna./ N ew Paris museum collections but it is widespread in Triangles isolated or confluent Quebec W. Va. No. 4 the western Nearctic throughout the Hudson- a. 1, 2, 3,4, 5 isolated ...... 6 1 0 b. 1, 2,3 isolated; 4-5 confluent ...... 2 6 ian life-zone. It is extremely local and subject c. 1-2 confluent; 3 isolated; 4-5 confluent 111 7 23 to extreme fluctuations in population size. d. 1-2 confluent; 3, 4. 5. isolated ...... 1 0 0 e. 1-2-3-4-5 confluent ...... 1 0 0 Due to these factors it is often missed by tra- 164 THE NATIONAL SPELEOLOGICAL SOCIETY versing collecting parties. That it may become common at times is evident by Lensink 's state- ment that ... "Trappers in the vicinity have remarked upon its abundance and consider it the primary food of martens." (Lensink, 1954, p. 259) and Preble's observation of a colony on the Athabaska River that ..."must have comprised many thousands of individuals, and occupied a heavily wooded area, at least a half a mile (0.8 km) square." (Preble, 1908). Primary sources dealing with the habitat of ill. vaathoga(ftbac are few...... Preble gives: 1. "a strip of young, mixed woods bordering a swamp ... burrows ... were in dry ground in the woods or shrubbery Figure 21. and evidently were quite deep, as I saw nearly Microstus xanthognatus skull. Dorsal, ven- a bushel of dirt at the entrance to a single tral, lateral view. CM 6249. Sinkhole No. burrow." 2. "...CONTRARY to their usual habit 4, New Paris, Pa. 7.0 m level, mm grid. this colony had extended their runways into a wet, sphagnum swamp." 3. "Deep, almost non-existent and the life history of the mixed woods on the summit of the hills animal is unknown. bordering the valley of the Athabaska." 4. Both authors mention this vole's fondness "at the base of a limestone cliff." 5. "a willow for horse tail (Equisetum) and Lensink men- covered island." 6. "poplar woods." 7. tions lichens in stomach contents. "heavily-wooded area.., on the gently sloping Sinkhole No. 4 Micro/us xanthognathur sides of a valley." appears to have been the dominant microtine Lensink describes the habitat at Castle Rock, below the 7.3 m level. Alaska, "thin, boreal forest, black spruce, The presence of M. xantbognathas in larch, white spruce, birch and aspen, ground ever increasing numbers with depth indicates cover sphagnum moss and lichen with scat- that full tundra conditions were either never tered clumps of sedge, cotton grass and horse- attained at the site or occurred prior to the tail ... runways and diggings were found filling of the sinkhole. Remains of the collared throughout the entire area although they were lemming would appear to indicate the near most common in lowlands and swampy areas. presence of tundra. It is conceivable that Runways frequently crossed small puddles." during the periods of cyclic abundance, wan- Preble mentions trapping meadow vole (ill- dering individuals might encroach into adjoin- cm ti'speallrylvanicusi"drunimondi"1) and ing habitats but as far as we know the two least weasel ( Mustela rixosa) in M. xan- species do not normally occupy common tbognathas runways. ground today although both may inhabit the All of the accounts mention boreal wood- same general area. The large number of M. land, ranging from "heavily-wooded" to xanthognatbus from the lower levels of Sink- "thin", elevations ranging from hilltop to hole No. 4 would seem to indicate a suitable island. Ground conditions range from dry habitat at the mouth of the sinkhole rather and well-drained to swampy. than the trapping of itinerant animals such Despite its seeming rarity, M. xanthogna- as may have been the case with the lemming thus appears to be adaptable to a variety of ( Ditro.itoay.x-). habitats within the boreal forest. Grasslands Yellow-cheeked vole remains have also been as such are not mentioned and it does not reported from late Pleistocene deposits at Nat- occur on the barren ground. Full habitat ural Chimneys, Augusta County, Virginia, and descriptions, however, such as Lensink's, are Bootlegger Sink, York County, Pennsylvania.

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 165 There is no apparent change in cranial or dental dimensions with depth. The Sinkhole No. 4 population cannot be adequately distinguished from modern samples (Guilday and Bender, 1960).

Table 29 — Measurements I,, millitmaters. sta swaths, b ( Leach 3 Nor i■ No 4. P. Measurement X length M, 3.4 a .02 2.7-4.0 .23 7.03 6.687 .48 100 lenyah 7.33 7 .04 5.6-8.2 .447 .03 6.007 .44 91 1.01 M'—M' 7.38 7.04 (5.0-8.3 .39 7 .03 5,287 .42 7. For additional measurements, see Guilday and Bender. 1 960.

,A1 icro /as , species? Figure 22. Material: 21 partial skulls; 460 left, 411 right First lower molars of Microtus (presumably chrotorrhinusl mandibles or M, 's. M. pennsylvanicus or from Sinkhole Na. 4, New Paris, Pa., showing Remarks: Most of the material, lower jaws variation of trefoil from "miurus" to pennsil ran icas or and teeth, are either Al. "pennsylvanicus" pattern. CM 6272, 5615, Al. c brot err!, in . M. xanthognat bus man- 5618, 5884, 5616, 6213. CM 6271, 6211, dibles and M, 's are identifiable by large size 5607, 6225, 6225b, 5618b. and usually distinctive pattern of M, and M3 . Young AI. xan Magna thus cannot always be identified, however, and approximately 10 per- pattern occurs in M, 's of the Recent European cent of the mandibles (based upon M, pat- Al. arval is as an uncommon variant and is tern) may be juvenile Al. xan t bognet t bus . there referred to as the "gregal older" type. Although three species of Micro/us are Both Al. grega is Pallas and Al. in in refs believed to be represented in this collection Osgood are in the subgenus Stenocranius the following notes on M, variation may be Kastschenko. (Janossy and Schmidt, 1960). profitable in the analyses of other such deposits. Such variants were given names in the past-- All three species normally possess five alter- "form ass/ill iI is" (Rorig and Borner, 1905 nating triangles. Five alternating triangles were and Schaefer, 1935), but the practice was dis- present in 93 percent of this collection; continued as the true nature of such varia- six alternating triangles in 6.7 percent and tions became clear. seven in .3 percent. One rather consistent The M, of a microtine is of diagnostic variant (5.5 percent) is the suppression of value at the generic level and occasionally to the fourth buccal re-entrant valley (fig. 22, subgenus, but it is of dubious value at the CM 6272, 5615, 6271) producing a pattern specific level (unless we are dealing with Recent identical to the Alaskan M. ill/urns (see Bee monotypic forms ). and Hall, 1956, p. 58). Initially, it was sus- Much of the material above undoubtedly pected in the light of the presence of other belongs to the cranial material listed under the high latitude forms in the deposit that M. appropriate species. In/urns might well have been present. There appears to be no stratigraphic pattern to the Pit.vmys pinet (»-zi (LeConte)--Pine vole distribution of the "miurus" variant in the Material: 1 partial skull; 10 left, 12 right sinkhole. No cranial material of this form was mandibles or M,' s. discovered and a skull of M. penns y/ van icus Stratigraphy: Sporadic from 2.1 m to 8.5 m with an articulated mandible of the "m inrus" level (fig. 30). pattern was unearthed. Every possiblevariation Remarks: One of the rarest microtines in the between a typical "In in rus" (fig. 22, CM 6272) deposit, the pine mouse is the only microtine and a typical ''pen n s y Ivan /cif s " pattern (fig. represented that does not occur north of the 22, CM 6213, 6225b) were present in the southern fringes of the Canadian life-zone collection. It is interesting to notethat the same which it reaches only sporadically (Hamilton,

166 THE NATIONAL SPELEOLOGICAL SOCIETY 1938). In contrast the pine mouse was the area of temperate and boreal North America. commonest rnicrotine in the Recent Sinkhole A single individual was also found in the No. 2 fauna, and is the common microtine Recent Sinkhole No. 2 fauna. at the site today. Family: Zapodidae In contrast to the chipmunk and southern Zap us hudsonicus (Zimmermann )--Meadow bog lemming populations, no difference could jumping mouse. be found between the Pitymys studied from Material: 4 fragmentary right mandibles, 2 Sinkhole No. 2 and those from Sinkhole No. with M, -M3 , 2 with M1 . 4. There appears to be evidence for contamin- Stratigraphy: From the 5.1 m to the 7.2 m ation of the upper levels of Sinkhole No.4 by level (fig. 27) modern short-tailed shrews. Rodent droppings recovered, apparently in situ, at the 5.9 m Remarks: M, -M3 measured 3.6 and 3.7 mm. level contain an unmistakably modern pollen Mi measured 1.3, 1.4 and 1.4 mm in total spectrum. One dead Pitymys, demonstrably length. recent, was found embedded in the mud and NapaeOZUP/IS insignis (Miller )--Woodland was discarded during the excavation. Rabbits jumping mouse (Sy/ri/agus ), woodchuck (Marinota), and Material: 1 partial skull; 7 left, 4 right maxillae; opossum ( Didelphis ) were also recovered 13 left, 12 right mandibles. between excavating trips. These were easily Stratigraphy: From 4.9 m to 8.5 m levels recognized as Recent, but the small size of (fig. 27). Pitymys (and of Peromyscus leucopus) might allow them to escape detection until Remarks: The New Paris No. 4 and the Na- their teeth or jaws showed up in the washing tural Chimneys Napaeozapus, both late Pleis- trays. We suspect that all Pitymys are intru- tocene, differ significantly in size from modern sive. central Appalachian material. They average 10 percent larger in all dimensions and have Table 30 — Occlusal length, NI, in millimeters, Prtron, pro rrrrrror. correspondingly heavier, more pronounced New Pads, Pa. muscle attachments. The two Pleistocene col- Site X OR. Ci V N lections appear closely related if size is taken Sinkhole No. 2 2.65 .01 2.4-2.9 12 :1.008 4.52 , .30 110 Sinkhole No. 4 2.62 .1.05 2.0-2.9 22 , .03 8.39 1, 1.39 18 as a criterion of relationship. This late Pleis- tocene form may be worthy of formal sub- The highly variable fourth buccal re-entrant specific recognition. Although the cave popu- of M, behaved in the same fashion in both lation samples are significantly different from sinkhole samples. (fig. 19). Pennsylvania material, their true relationships with modern populations, especially far north- Table 31 - Fourth buccal re-entrant M1, ern ones, must await a more detailed study of Pitymys pinetoram geographic variation within the species. The New Paris New Paris woodland jumping mouse is also recorded No.2, Pa. No. 4, Pa. from Cumberland Cave, Maryland; Natural absent 14 12.5% 3 26.3% shallow, no cement present 42 Chimneys, Virginia, Bootlegger Sink, Penn- shallow, cement present 49 43.7 6 sylvania, and Robinson Cave, Tennessee. medium, cement present 6 43.7 6 31.5 deep, cement present 1 .8 Sample size 112 SPECIMENS EXAMINED

Ondatra zibethicus (Linnaeus )--Muskrat Modern specimens. CM Mammal No. 1818, Material: 1 right humerus, 1 M3 , 1 partial 3035, 4057, 37013-14, 370116-26, incisor. 370128-35. Stratigraphy: depth unknown. Natural Chimneys, Augusta County, Virginia. CM 7529. Remarks: although muskrats are semi-aquatic marsh dwellers they are also extensive wan- Family: Erethizontidae derers at times. They are found over a wide Erethizon dorsaturn (Erxleben)--Porcupine

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 167 Table 32- Measurements in millimeters, Napaeozapus ins ignis, Carnegie Museum collection Locality O.R. 0' V Occlusal length, M, Pennsylvania, modern 1.6 ± .008 1.5-1.8 .04 ± .006 2.50± .39 20 Quebec, modern 1.6 2 Ontario, modern 1.6 1 New Paris No. 4, Pleistocene 1.8 ±.027 1.7-2.1 .09 ± .019 5.00 21 1.06 11 Natural Chimneys, Pleistocene 1.7 1.6-1.7 6 Occlusal length, M1-M3 Pennsylvania, modern 3.17 ± .023 2.9-3.4 .11 ± .016 3.47 .52 22 Quebec, modern 3.15 2 Ontario, modern 3.21 1 New Paris No. 4, Pleistocene 3.43 3.2-3.6 4 Natural Chimneys, Pleistocene 3.45 3.3-3.6 6 Occlusal length, lower molar row, M1-M3 Pennsylvania, modern 4.06 ±.03 3.8-4.3 .13 ± .02 3.20 .53 18 Quebec, modern 4.15 4.0-4.2 2 Ontario, modern 4.22 1 New Paris No. 4, Pleistocene 4.31 4.2-4.4 2 Natural Chimneys, Pleistocene 4.41 4.2-4.6 2

Material: 2 immature skulls with associated Remarks: These hares deserve more detail- mandibles and partial skeletons; 1 left man- ed study. The effect of physiological age upon dible, adult. the skull and dentition is considerable and Stratigraphy: From 5.5 m to 6.4 m level. may influence the sample parameters. Most Remarks: Porcupine remains are known from specimens of snowshoe hare in museum collections are adults collected by open hunt- the Recent fauna of New Paris No. 2. It is still found in the northern part of the state ing. The sinkhole population has a much today and occurs throughout the Canadian wider age span from obvious nestlings on. and Hudsonian life-zones of North America. Measurements below were taken from what was considered to be adults but no detailed Order: Lagomorpha selection was made. Family: Leporidae Remarks: Modern snowshoe hares, contrary L ep u s americanus (Erxleben) -- to "Bergmann's Rule", attain their smallest Snowshoe hare average size in the northern portion of their Material: 32 partial skulls; 20 left, 13 right range. The largest eastern individuals are maxillae; 49 left, 48 right madibles. Skele- found in the central Appalachians. Smallest tal material representing at least 49 animals. average size occurs from the James Bay area One mounted composite skeleton (fig. 24). north (fig. 23). Stratigraphy: Commonest at 5.8 m level The snowshoe hare sample from Sinkhole where it composed 7.4 percent of the recov- No. 4 is composed of small individuals. They ered fauna. Below the 6.4 m level it virtually are significantly smaller in some measure- disappeared from the fauna (fig. 27). ments than L. a. virgin/anus now in Penn- 168 THE NATIONAL SPELEOLOGICAL SOCIETY eral decrease of woodland forms (red squirrel, flying squirrel, pine vole, spruce vole, short-tailed shrew, red-backed vole, woodland jumping mouse) and the increase in grassland forms (most microtines, thirteen- lined ground squirrel) at the lower levels. Rabbits of the genus Sylvilagus, which did not occur in the Sinkhole No. 4 fauna, occupy the area today to the exclusion of Lepus, and apparently did so during the past several thousand years at least. Sylvi- lagus occurred in the Sinkhole No. 2 local fauna (C-14 date of 1,875 ± 100 yrs. B.P.) Lepus did not.

Figure 23. During the excavation of Sinkhole No. 4, five or six cottontail rabbits ( Sylvilagus ) Skulls of Lepus americanus. Negativ e "Berg- fell into the hole and expired. mann's response." CM 5480. Sinkhole No. 4, New Paris, Pa. 5.8- 6.4 m level, late Pleistocene. CM Mamm al 3273- James Bay, Order: Carnivora Quebec. Recent. CM Mammal 2212 - Penn- Family: Mustelidae sylvania. Recent. Martes americana (Turton) — Pine marten Material: Partial rostrum of skull containing left I', C', P2, P4, right 132 — P4. CM 5737. Stratigraphy: Recovered from the 5.2 m level. Remarks: In some portions of its modern range, Microtus xanthognathus is the primary prey of the marten (Lensink, 1954, p. 259), a relationship which probably held during the late Pleistocene as well. Pine martens occurred in the northern and mountain- Figure 24. ous sectors of the state during early settle- Lepus am ericanus composite skeleton, Sink- ment days, but probably not in the immed- hole No. 4, New Paris, Pa. CM 5944. Hind iate sinkhole area. foot --,. 115 mm. Marten remains have been recovered from Natural Chimneys, Virginia and Robinson sylvania,* but not distinguished from mod- Cave, Tennessee. ern L. a. americanus from the James Bay area of Ontario and Quebec. Martes pennanti (Erxleben) — Fisher The significance of the disappearance of Material: 1 M'. the snowshoe hare from the lowest levels of Stratigraphy: 6.1 m level. Sinkhole No. 4 may be related to the gen- Mustela rzxoa (Bangs) — Least weasel 3 Material: 1 right P,, 14. Stratigraphy: 5.5 m level. * Stocking programs have altered the situation in many areas and make it difficult to obtain racially Remarks: Specimens are referred to M. rixo- pure comparative material. sa solely on their small size.

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 169 Table 33 - Lepus americanus, modern and Pleistocene localities. Measurements in millimeters.

Locality 7 O.R. CI V N Lower incisor width Penna., Mich., -- modern 2.5 L', .03 2.3-2.6 .10 1 .03 4.00 1. 1.00 8 Moose Factory, Ont., - modern 2.35 1 .04 2.2-2.6 .14 1 .03 5.95 .._ 1.48 8 New Paris No. 4, late Pleistocene 2.32 .02 2.0-2.5 .13 1 .01 5.60 1 .79 25 Lower toothrow length Penna., modern 16.0 14.3-17.4 6 Michigan, modern 15.9 15.7-16.2 2 Moose Factory, Ont., modern 15.8 14.5-19.9 8 New Paris No. 4, - late Pleistocene 14.4 L .18 12.7-15.9 .89 L .12 3.70 1 .53 24

Upper toothrow length Penna., modern 15.6 14.2-16.6 6 Michigan, modern 15.25 15.2-15.3 Moose factory, Ont., modern 15.25 14.5-16.8 8 New Paris No. 4 late Pleistocene 14.53 ±. .15 13.6-15.8 .63 1 .10 4.33 .1.74 17 Maxillary width Penna., modern 37.81 34.2-40.1 6 Michigan, modern 36.60 35.8-37.4 2 Moose Factory, Ont., modern 36.82 35.0-39.1 8 New Paris No. 4 late Pleistocene 35.5 33.0-38.1 1.55 .29 4.36 _t .82 14

Incisive foramen Penna., modern 19.90 19.4-20.8 6 Michigan, modern 20.10 19.2-21.0 2 Moose Factory, Ont., modern 19.88 19.2-20.6 8 New Paris No. 4, late Pleistocene 18.53 ± .29 16.6-20.5 1.05 1 .20 8.07 1 1.58 13 Inter-orbital breadth Penna., modern 11.7 10.2-13.7 6 Michigan, modern 10,7 10.2-11.2 2 Moose Factory, Ont. modern 10.5 9.7-11.4 8 New Paris No. 4, late Pleistocene 10.4 1.48 9.0-11.7 1.7 1 .34 16.71 1' 3.28 13

170 THE NATIONAL SPELEOLOGICAL SOCIETY Table 34 — Measurements in millimeters, P'Mu r Ia. various species sence of both Mylobyus and Indian cul- Catalogue No, Species Sex Locality Age Length Width tural material at Hartman's Cave in eastern CM 6021 Al. rixo,u ? New Paris No. 4 Lao 3.3 1.5 Pleistocene Pennsylvania is suggestive but the associa- Nlammal No. M. RINI150 9 Pennsylvania modern 3.2 1.6 24917 tion may well have been fortuitous. It is pro- CM Mammal No .11 riv"la d Pennsylvania modern 3.3 1.6 28275 bable that Mylobyus like so many other CM Mammal No Al, erm in en 9 Pennsylvania modern 5.6 1.8 36275 Pleistocene large mammals was a victim of a long chain of ecological events initiated Order: Artiodactyla by the appearance of a new and efficient predator, the Paleo-Indian in the area. Lun- Family: Tayassuidae delius (1960, p. 34) is of the opinion that Mylobyus nasutur (Leidy) — Mylobyus is the ecological equivalent of the Long-nosed peccary Old World Sus. Its environmental niche in Material: 1 partial skeleton. CM 5860 (figs. the Appalachian area is now probably filled 25, 26). by the black bear ( Ursus americanus), anoth- Stratigraphy: Remains were scattered over a er forest omnivore. Introduced European wild vertical span of 1.5 m from the 5.2 m level boar are now feral in Tennessee and other to the 6.7 m level. wooded parts of the Appalachians but apparently cannot successfully compete with the Remarks: All other species of small verte- black bear (Stegeman, 1938). brates from New Paris No. 4 are living today. The presence of the peccary in association with the flora and fauna of boreal character presently living within the Canadian and Hudsonian life zones may come as a sur- prise. The Pennsylvania peccary has been regarded as an indicator of warm climates. In the Port Kennedy fauna Mercer (1899, p. 285) wrote: "Judged by the presence of such animals as the subtropical tapir and peccary, a winter climate milder than the present probably then prevailed." But Matthew (in Twenhofel, 1932) observed that the peccary in the Pleistocene of Pennsylvania and New Jersey may be associated with such northern Figure animals as wolverine, musk ox, and cari- 25. bou, thus signifying a cold climate. Such dis- Restored skull, juvenile Mylohyus nasutus. cord in the ecological interpretation of large CM 5860. Sinkhole No. 4, New Paris, Pa. mammals has brought their use as palaeo- Frontal bones and mandibles not recov- climatic indicators into disrepute and strength- ered. Insert upper right, adult skeleton. ened the case for a non-climatic cause for TMM 1407, Friesenhahn Cave, Texas. their extinction. Evidently Mylobus occupied The New Paris No. 4 specimen was a pig- a wide range of environments until its ex- let. All cranial sutures with the exception of tinction and disappeared along with other the lambdoidal and the sagittal were open, Pleistocene "big game" less than 11,000 years including those between the basi-, ex-, and ago. The only continent-wide environmental supra-occipital. The lambdoid suture was fused change that is known with certainty for this and its course, both externally and intern- time interval is the arrival of big game hunt- ally, could be followed only a short distance ers, the Paleo-Indians. toward lambda beyond the asterion. The sagit- This is not to imply that Myrohyus was tal suture was barely discernible. It could be necessarily a direct victim of human preda- traced from bregma back 12 mm at which tion. This has yet to be demonstrated. Pre- point it joined the barely discernible lamb- BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 171 doidal sutures. The course of the sagittal suture was obliterated internally. Neuro-central sutures and epiphyseal disks of the vertebra were unfused. The odontoid process of the axis was separate. The sacrum consisted of at least four unfused units. Ilium, ischium and pubis were separate. Epi- physes on ribs and all major limb bones were open. No trace could be seen of the proximal epiphyses of the metapodials. Fu- sion, as in other artiodactyla, took place before birth. The distal epiphyses were unfused. Metatarsals III and IV, fused in the adult into a cannon bone, were closely adpressed but not fused. The lumbar region was fortunately well preserved. There were five lumbar vertebrae as in Tuyarsu, one or two less than in Sus. (Six or seven lumbars occur with equal frequency in Sus. [Sisson and Grossman, 1938].) The mandibles were not recovered. Three lower deciduous incisors and one lower decidous canine were recovered during screening operations. They appear slightly worn. The complete deciduous upper dentition (fig. 26), 2 incisors, 1 canine and milk molars 2, 3 and 4 were recovered. dI2 is a sim- ple peg, 2.1 mm in diameter but dl' is twice as large with a spatulate-shaped crown. The lingual surface of the crown is crossed vertically by a low loph separating the tooth into two basins. They show no wear. The canines are scimitar-shaped and project 15.6 mm from their sockets. They are oval in x-section, 5.4 mm x 4.0 mm and are slightly worn. Immediately in front of each canine Figure 26. socket is a small hole 5 mm in diameter through which the tip of the developing Palate and upper deciduous dentition, My- permanent canine can be seen. An enigma- lohyus nasutus. CM 5860. Sinkhole No. 4, tic hole immediately behind each dl' pro- New Paris, Pa. In background to same scale, profile of adult skull bably represents the site of the eruption of TMM 1407, Friesen- hahn Cave, Texas (from Lundelius, 1960). a future permanent incisor. First upper permanent molars superim- The milk molars are fully erupted but show posed. no signs of wear. The first permanent molar was deep within the maxilla and still form- berland Cave, Maryland and a cast of ANSP ing. type no. 26 ( M. penns y Ivan icu s), Hartman's Two sets of My/ohyus deciduous den- Cave, Pennsylvania. Both New Paris No. 4 titions are available for direct comparison, and Cumberland dentitions show no tooth a cast of USNM 8160 ( M. exortivus), Cum- wear. Cumberland is somewhat older, M'

172 THE NATIONAL SPELEOLOGICAL SOCIETY beginning to erupt. Hartman's has M t fully dient. Huxley (1932) looks upon this as an erupted and clip' and c1134 show considerable adaptation enabling the young animal to ac- wear. dP2 is missing. company its dam as soon as possible. Proxi- mal limb elements were some 60 percent Table 35 - Measurement in millimeters, Mv/hhyns. milk dentition, adult size (taking into consideration that we Now Ports Organ - Hedrick, No. 4 Cumberland Hartman's Friosonhahn Cava, W. Va. are comparing only two animals of unknown , 41 length 10.9 10.0 age and sex from widely separated areas ), width 7.7 7.5 di.' length 13.6 12.5 13.0 12.8 12.1 while distal limb segments were some 80 per- width 10.8 9.0 10.0 9.2 10.8 cent adult size. A similar gradient was not dls° length 14.4 12.0 14.0 15.1 width 15.6 14.0 16.0 12.9 apparent in the width of limb segments. They CM 5860 USNM 8160 ANSP 26 TMM-1407 UMMP 27728 Many and Goths Lundelius Handley averaged about 80 percent of adult size 1938 1960 1956 throughout (table 37). The minor differences in size and com- Comparative measurements of the restored plexity of cusplets between these dentitions New Paris juvenile skull* and the Friesen- could be accounted for by individual varia- hahn adult skull indicated that the juvenile tion alone. Gidley and Gazin (1938, p. 86) skull was only 59 percent of adult size. But state that the amount of individual variation there were proportional differences. The post- present in any of the nominal "species" is glenoid length of the juvenile skull was 69 unknown. All were described from frag- percent adult size but the pre-glenoid length mentary material. was only 56 percent adult size. This juvenile peccary had a relatively shorter snout than the Lundelius has assigned all of the late Pleis- adult, a well nigh universal mammalian charac- tocene Mylohyus to two "species", M. nu- , teristic. There appears to be considerable dif- sutus (including M. peons) I van icus and ferential growth within the snout, however. The M. browni), a larger western and northern post-canine diastema was only 52 percent of form and M. foss//is (including M. ex- adult size while the pre-canine diaszema was ortivus) a smaller southeastern form. This is already 77 percent of adult size. Width, as admittedly a tentative classification. No ade- measured between the canines, was already quate study of differences due to age, sex, 93 percent of that of the Friesenhahn skull. or individual variation is possible until more material is available. The relatively minor dif- Table 36 - Growth Comparison, ferences that differentiate the nusutus and it yloh torn nalutn, Length, limb elements . foss//is groups may be of infra-specific nature. Adult Juvenile Percent Friesenhahn New Ports No. 4- (Juv.)/(Adulti Following Lundelius, the New Paris No. scapula length 2 i6rnm 120mm 50.8 4 specimen is referred to M. nasutus. It is pelvis lengtl, 240 145 60.4 somewhat larger than M. exortivus, USNM humerus length 219 I 37 62.5 femur length 212 150 70.7 8160 from Cumberland Cave and identical radius length 182 125 68.7 with the type of M. pennsylvanicus from ulna length 244 150 61.5 Hartman's Cave, Pennsylvania. tibia length 228 163 71.5 metacarpal 2 71.5 65 89.5 Several interesting growth gradients be- metacarpal 3 100.8 83 82.1 metacarpal 4 11)2.7 82 79.8 come apparent if corresponding elements of metatarsal 3 113 91 80.5 the juvenile CM 5860 and the adult TMM metatarsal 4 115 88 76.5 1407 from Friesenhahn Cave are compared. astragalus length 43 35 81.3 Relative lengths of limb elements were com- Data from Lundelius, 1960. pared (table 36). A proximal-distal negative *. Estimated lengths taking into comideration missing cartilage between epiphyses and di, growth gradient is apparent; distal elements physes. growing at a slower post-natal rate than proximal elements. The young of most, if not all cursorial ungulates are born with relatively * All sutures were open and the skull elements large limbs and exhibit a similar growth gra- were recovered separately.

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 173 Subjectively, the dorsal profile of the snout Wisconsin is not limited to the vertebrate fos- was more concave, the zygoma less flaring, the sil record. Frey (1953) encountered a domi- nuchal crest not as well developed, and all nance of small-sized pine pollen with a low areas of muscle attachment less defined than frequency of spruce in the Carolina coastal in the adult. plain during the Wisconsin maximum to which Whitehead (1963) has added the pollen of Table 57 —, Growth Comparison, ,111 /,,Ayte, cool temperate plants such as dwarf mistle- Width, limb elements toe ( A rceuthobium), Canadian burnet ( San- Adult Juvenile Percent guisorha canadensis), curly grass (Schi- Friesenhohn' New Paris No. 4 (Juv.}/(Adult) zaea). Lycopodium lucidulum, and other femur, prox. width 53 mm 43 mm 81 club mosses. Older Wisconsin deposits of distal width 52 42 81 tibia, prox. width 53 44 83 the Carolina Piedmont also indicate the pene- distal width 33 26 78 tration of northern species (Whitehead and am] agalus, width 22 18 81 Barghoorn, 1962). In the Pennsylvania Pied- metatarsal 3 and 4 pron. width 27 22 81 mont an alluvial pollen record dominated by distal width 32 27 84 herb and pine pollen is thought to repre- Data from Lundelius, 1960. sent Wisconsin-age tundra and taiga (Martin, 1958). Table 38 — Growth Comparison, Cranial Measurements, More difficult to establish is whether a II vlohyus rialulus greater mixing of boreal and temperate ele- Adult Juvenile Percent ments occurred in Pleistocene biotic commu- Friesenhahn New 'orb No. 4 (Juv.)/(Adult) nities than is evident today. In this and other post-canine diastema 71 mm 37 mm 52 matters concerning life during the time of pre-canine diastema 30 23 77 maximum ice advance 20,000 years ago, a width between canine alveoli 30 28 ? 93 richer fossil record is essential if we wish condyles to to reduce our present biogeographic uncer- premaxilla 357 210 59 ant, origin of tainty. zygomatic ridge to tip of premaxilla 195 110 56 During the last maximum of continental ant. origin of glaciation ice extended below 40° in latitude zygomatic ridge to condyles 152 105 69 almost to the Ohio valley. Although the exact position of the ice margin in the Appala- Data from Lundelius, 1960. chian region is uncertain, it lay close to 42° N. latitude, some 240 km north of New THE LATE-GLACIAL ENVIRONMENT OF Paris. Because of the southerly dip of the UNGLACIATED PENNSYLVANIA glacial front west of the Appalachians across Introduction: The last maximum of con- northwestern Pennsylvania and into southern tinental glaciation, presumably the extreme Ohio, the glacial border lay slightly closer to development of ice-margin climates during the New Paris on the northwest side (fig. 1). 60,000 years of the Wisconsin glaciation, Evidently at that time Sinkhole No. 4 was took place about 20,000 years ago (Flint, not open to the surface and it will not help 1962). Undated fossil remains of Canadian- to reveal the biota of full-glacial times. zone mammals such as the Arctic shrew On the basis of two radiocarbon dates we Sorex arcticus), northern bog lemming ( estimate the rate of post-glacial filling of (Synaptomys borealis), northern flying Sinkhole No. 4 at from 0.5 to 1.0 m per squirrel ( Glaucomys sabrinus), pine mar- thousand years. The latter figure is probably ten (Martes americana), and caribou too low under a late-glacial climate of more ( Rangifer) as far south as Robinson Cave intense frost action. Accepting one meter as in north-central Tennessee (lat. 36° N.) sug- a minimum estimate of the rate of infill, gest boreal conditions far south of the Wis- the lower three meters (pollen unit B) should consin terminal moraine. represent late-glacial conditions 12,000 to Evidence of boreal conditions during the more than 15,000 years ago. Unfortunately,

174 THE NATIONAL SPELEOLOGICAL SOCIETY organic material below 6 meters was scarce

and the equivocal result of a carbonate date Talpidoe from bone at 5.4 m - 6.5 m (M-1067) adds Zopodidae to our uncertainty. Nevertheless, we view the 10 l fill from 6 - 9 meters as reflecting local _0%— Sela ridae

conditions in an Appalachian valley roughly 1 0 Leporidae 5,000 years after the last glacial maximum, —0% — i 1 at a time when the continental glacial front 30 20 GrAceti nae had retreated to a point some 600 km or 1 o more to the noith of New Paris. It is surprising to find an Arctic tundra species, the io Soricidoe collared lemming, lingering at this relatively 40 late date and intriguing to find it in associa- 20 tion with boreal species plus a prairie rodent Vesperi, I ion !doe and a prairie grouse. 033_ Faanal change: Faunal changes in the 9.1 m 80 column of matrix were surprisingly well mark- 60 Micro? tnae ed despite the lack of any obvious stratifi- 40 cation and the irregular deposition of the 20 matrix. Faunal changes were not apparent until the minimum numbers of each species were plotted by .91 m (three foot) intervals. Difficulties presented themselves. Chief among them was the likelihood of contami- Figure 27. nation of older by younger matrix from above. Stratigraphic distribution of selected mam- All obvious cases were discarded but the malian families and subfamilies. Sinkhole site was excavated on the average of one week- No. 4, New Paris, Pa. end a month over a four-year period and dur- a group increased from 34 percent of the ing this time a small amount of unrecorded fauna at the 4.9 m - 5.5 m level to 80 per- slumpage may have occurred. This may ex- cent at the lowest levels. Fig. 30 presents plain such seeming anomalies as the white- the relative status of nine microtine species footed mouse ( Peromyscus leucopus) and from Sinkhole No. 4. All of the species, with the pine vole ( Pitymys pinetorum) (one the exception of the pine vole ( Pitymys mandible each) recorded from the deepest pinetorum) and the southern bog lemming levels. There was also the possibility of un- (Synaptomys cooperi), range north today a uthorized tampering with the drying racks at least to the northern edge of the boreal during periods when the site was unattended. woodland (Hudsonian life-zone). The col- Several trays of matrix, although labeled stra- lared lemming ( Dicrostonyx budsonius) tigraphically, were processed as "depth un- is now confined to the tundra areas of the known" because it was suspected that they had Arctic life-zone. Its rarity (three animals out been tampered with in just such a fashion. of a total microtine population of 1,423 The relative percentages of most mamma- individuals) indicates submarginal habitat for lian families in the Sinkhole No. 4 fauna that form. Tundra conditions may have exist- (Talpidae, Soricidae, Zapodidae, Leporidae, ed in the area (perhaps along the crest of Vespertilionidae, fig. 27) declined with depth. Chestnut Ridge, higher than the mouth of Only the Cricetinae and the Microtinae re- the sinkhole) but not around the sinkhole. mained as abundant or increased. During level 8.5 m - 9.1 m time, boreal and Microtine rodents, abundant rich in spe- grassland microtines, yellow -cheeked vole cies, and of diverse habitat requirements, fur- ( Microtus xantbognatbus), meadow vole nish the best guide to ecological fluctuation ( Microtus pennsylvanicus) an d northern during the period of infilling. Microtines as bog lemming ( Synaptomys borealis) dom- BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 175 m ated the fauna. The rock vole ( Microtus #6 chrotorrbinus), a rock-inhabiting species, Eptesicus shows a slight increase with depth. This Can- 10 • Pisnstrellus adian Hudsonian-zone form can occur in eith- 60. er timbered or grassy situations as long as it has access to rocky terrain. It is not so 40 Myotis keenii sensitive an indicator of open areas as the 20 other species. As infilling progressed the meadow and woodland forms gave way to for- —0% est species — the red-backed vole ( Clethrio- nomys gapperi) and to a limited extent the spruce vole ( Phenacomys); and southern Myotis Iscitugus forms such as the southern bog lemming (Synaptomys cooperi) and pine vole ( Pity- mys pinetorum). The woodland Cletbrio- nomys represented only four percent of the Figure microtine fauna at level 8.5 m - 9.1 m, in- 28. creased to 11 percent at level 6.7 m - 7.3 m Stratigraphic distribution of bats (Chirop- and up to 50 percent at the highest level ana- tera). Sinkhole No. 4, New Paris, Pa. lyzed, 4.9 m - 5.5 m.

This picture of gradual transition from bor- 40. eal parkland to boreal forest is also rein- 20. forced by the stratigraphic fate of other species. Bats (fig. 28) and squirrels (fig. 29) increase both in species and in individuals 30 from bottom to top. Thethirteen-lined ground 10 • 102MIEIZI1 squirrel, Cite//us tridecemlineatus, gave 0% place in the upper levels to the tree squirrels; the woodland shrew, Blarina brevicauda, a nominally southern form, does not occur in the lower levels (fig. 31), but is the commonest shrew in the higher levels , • 16'21. 22 30' 22-30' of the sinkhole. elep tn. Faunal changes were also apparent in the amphibians (fig. 33) and the reptiles (fig. Figure 29. 34). Among the snakes the Crotalidae and Stratigraphic distribution of squirrels (Sciu- the Colubrinae decreased with depth. At the ridae). Sinkhole No. 4, New Paris, Pa. deepest levels only the garter snake ( Tham- nophis) persisted. This genus ranges further north than any other North American rep- placed by B. a. americanus in the upper tile. It has been recorded from the juncture levels of Sinkhole No. 4. of the Hudsonian and Arctic life-zones of If the mean depth of each species of mam- Churchill, Manitoba (Shelford and Twomey, mal represented by more than one indivi- 1941). Both the collared lemming ( Die- dual in Sinkhole No. 4 is calculated (mini- rostonyx) and the yellow-cheeked vole ( Mi- mum number of individuals at each level t rotus xantbognaibus) have been reported multiplied by distance from surface, the en- from Churchill (Hall and Kelson, 1959). tire sum then divided by total number of The only amphibian that occurred at the individuals), and the species are then ar- bottom levels was the Hudson Bay toad ranged in order of increasing mean depth (Bufo a. copei) which apparently was re- from the surface, a pronounced trend is not-

176 THE NATIONAL SPELEOLOGICAL SOCIETY 2.■ Doom 0% 10 20 30 40 50 60 70 80 90 0 Dicrostortyx .6 30- 10 • S. cooperi 60-1r

10 S. bor. 90-

ii0- 10 • Mi rot us Ph.,. 'a 1111•sehmc Arvicola 50 10 • 170 _ Prlymys

50 • 200 40. 230 30 C leth rionOmy5 P•levIdh. 20 250

i 0 . 270- 0% —

10 • M. ehrot 0%- Figule 32. Upon. o, Stratigraphic distribution of Microtinae,Jan- kovitch Hohle, Hungary. Adapted from Jcin- ossy, 1960. 50 40 M. xanthognathus 30 Depth 20 m. ft. 10 • 0%-

I 00 E 4,

Figure 30. Stratigraphic distribution of voles and lemmings (micron-noe). Sinkhole No. 4, New Paris, Pa.

Figure 33. Stratigraphic distribution of frogs and toads (Salientia). Sinkhole No. 4, New Paris, Pa.

ed. If the maximum northern limits of individual species are then plotted (table 39) it becomes apparent that there is a correlation between increase in average depth and latitude. There appears to be a point (c. 5.9 Figure 31. m) below which maximum ranges are consis- Stratigraphic distribution of shrews (Sori- tently boreal. Above this point maximum cidae). Sinkhole No. 4, New Paris, Pa. ranges tend to be temperate, 50° N. lat. or

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 177 • ▪

less, with some conspicuous and interesting exceptions: the red squirrel Tam iasciurus budsonicus), the northern flying squir- ■00. Colubrinae rel ( Glaucomys sabrinus) the red-backed Crotalidoe mouse (Cletbrionomys gapperi), and the BO woodland jumping mouse ( Napaeozapus ).

60. These are woodland forms which become increasingly scarce as depth increases, thus 40. forcing the mean depth for these species higher in the column. Figure 34. (At left)

— Stratigraphic distribution of snakes (Ser- 4 4 . . pentes). Sinkhole No. 4, New Paris, Pa.

Table 39 — Average stratigraphic depth, various species of mammals, New Paris No. 4, Pennsylvania

Average depth reaches north latitude today in meters Species 40 45 50 55 60 65 70 4.4 Parascalops brewer/ Peromyscus leucopus 4.9 Sorex Amens 5.2 Synaptomys cooperi 5.3 Blarina brevicauda 5.4 Glaucomys comas 5.5 Glaucomys sabrinus Pipistrellus subflavus 5.6 Tam /as striatus Cletbrionomys gapperi 5.8 Tamiasciurus budsonicus 5.9 Sorex dispar Myotis keenii Pit. ymys p /net oru m Zap us budsonicus Napaeozapus ignis Eretbizon dorsatum 6.1 Myotis lucifugus Pero myscus man iculatus 6.2 Lepus americanus Condylura cristata 6.4 Microsorex boy/ Phenacomys ungava 6.7 Sorex cinereus Dicrostonyx budsonius Microtus cbrotorrbinus 6.8 Microtus pennsylvanicus Microtus xantbognatbus 7.0 .S'yortpt omys borealis 7.1 Sorex arcticus

178 THE NATIONAL SPELEOLOGICAL SOCIETY Fourteen species of mammals and one am- ( Rana sylvatica), The Hudson Bay toad phibian occurred in the bottom one meter of ( Bufo americanus copei) is the only cold- the deposit. All occur, at least in part, in the blooded terrestrial vertebrate presently rang- Canadian or Hudsonian life-zone today. ing that far north in eastern North America Table 40 — Species from the bottom one meter of New Paris No. 4, (Conant, 1958). Pennsylvania. 1. Byfo cf. americanus copei Hudson Bay toad 6-8 THE BERGMANN RESPONSE 2. Microsorex boyi pygmy shrew 1 3. Sorex cheererus masked shrew 2 Zoologists commonly find that within a 1 4. Sorex arcticus Arctic shrew given species of warm-blooded animal, body 5. Sorex palustris water shrew 1 6. CiteUses tridecemlineatus 13-lined ground squirrel 1 size will increase with latitude, an adaptation 7. Peromyscus cf. maniculatus deer mouse 16 they believe serves to decrease surface/mass 8. Synaptomys borealis northern bog lemming 12 9. Cletbrionomys gapperi red-back vole 5 ratio and thus conserve body heat in cold- 10. Pbenacomys ungana spruce vole 1 er environments. This has become known 11. Mirrotus chrotorrhinus rock vole 8 as "Bergmann's rule." The remains of many 12. MiCrOtta pennsylvanicus meadow vole 30 13. Micro/ca xanthognathus yellow-cheeked vole 69 late Pleistocene animals are larger than their 14. Napaeozapus insignis woodland jumping mouse 1 present-day counterparts (fig. 16). Since the americanus snowshoe hare 1 15. Lepus environment of periglacial North America Five of these 15 forms do not range as was profoundly modified by a decrease in far south as Pennsylvania today: nos. 1, 4, temperature it would be logical to suspect 8, 10, 13 — one (no. 6) inhabits prairie west a "Bergmann's response" from some spe- and north of the state. An additional four cies. species (nos. 2, 5, 11, 15) are either rare In some forms size is negatively corre- or occur as boreal relicts in the state. By lated with latitude (fig. 23). Lepus ameri- way of contrast 11 of the above forms oc- canus, Mustela emir/ea ( Kairten, 1960) cur or may be expected to occur at the site and many burrowing mammals (Mayr, 1963). of fig. 5 along the eastern coast of Hudson Kurten (ibid.), using mid-Pleistocene ham- Bay at Great Whale River, Ontario, Lat. 0 sters and stoats whose modern representatives 55 N. Here, the Arctic and Hudsonian life- are respectively positively and negatively cor- zones meet. The collared lemming ( Dicro- related in size relative to latitude, demon- stonyx) may be found here as well and while strated that in those deposits in which the it is a true tundra animal it may occasionally stoats were small, the hamsters were large range very slightly into the northern fringes and vice versa. He attributed this to cold/ of the Hudsonian life-zone (Harper, 1961, warm oscillations during the mid-Pleisto- p. 21). cene. Hibbard (1963) has demonstrated Of the four species not found presently at much the same situation in Blarina from Great Whale River one, Microtus xanthog- Illinoian and Sangamon deposits in the Great natbus, is found in similar habitat from the Plains. Hooijer (1947) has shown that many west shore of Hudson Bay (Churchill) to East-Asiatic mammals, such as the tiger ( Pan- Alaska. The shrews Sorex arcticus and thera tigris), underwent a diminution in Sorex palustris both characteristic Canadi- size (climatically dictated? ) from Pleistocene an life-zone species reach the James Bay re- to recent times. 0 gion (lat. 52 N.) but are not found as far A response to climatic change is also clear- north as Great Whale River. In eastern Can- ly demonstrated in the mammal fauna from ada they stop at the southern limits of the New Paris No. 4, and serves to corrobor- Hudsonian life-zone and at a lower latitude ate the conclusions of these workers. In many than in central and western Canada. Citel- species from the deposit a "Bergmann's re- /us tridecemlineatus reaches a similar lati- sponse", either positive or negative, was pre- tude in the northern Great Plains of cen- sent. tral Canada. Although a demonstrable change was pre- With the exception of the wood frog sent in the pollen frequency profile, and the

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 179 species composition of the fauna also chang- New Paris No. 4 are compared with their ed with depth, there appeared to be no mor- modern mid-Appalachian equivalents, those phological change within a given species forms with a modern positive "Bergmann's throughout the stratigraphic column. The one response" are larger while those with a mod- apparent exception, B la). in , (fig. 16) is ern negative "Bergmann's response" are believed due to modern contamination. From smaller than their representatives which live this we infer that boreal conditions prevail- in that area today. This is discussed and ed throughout the period of infilling. statistics presented for each species under individual species accounts. They are sum- When the remains of many species front marized below.

Table 41 — Mammals with a modern positive "Bergmann's response", New Paris No. 4, Pennsylvania.

Percent increase, New Paris No. 4 vs Modern central Appalachian populations Species based upon length of tooth rows Condylura cristuta star-nosed mole 1.3" Blarinu brevicauda short-tailed shrew Microsorex boyi pygmy shrew Sorex cinereur masked shrew 5.0 Tamias s ir/at/is chipmunk 7.0 G laucomys sabrinus northern flying squirrel 13.0 G I aco m ys v 0 I ans southern flying squirrel 8.0 Tamiasciurus budsonicus red squirrel 9.0 Napaeozapus ins ign is woodland jumping mouse 6.0

* Sample believed contaminated.

Table 42 — Mammals with a modern negative "Bergmann's response", New Paris No. 4, Pennsylvania.

Percent decrease, New Paris No. 4 vs Modern central Appalachian populations Species based upon length of tooth rows Synaptomys cooperi southern bog lemming 12.0 Microtus cbrotorrbinus rock vole 5.0 Microtus pennsylvanicus meadow vole 11.0 Lepus americanus snowshoe hare 10.0

Of the 13 species studied for "Bergmann's seem to imply that climatic adjustments simi- response", nine were positive, four were nega- lar to those observed today may predate the tive, exactly as they are today, and evolutionary last glaciation in both hemispheres. The New change as a function of time alone seems not Paris No. 4 record demonstrates that the late to be involved. The evidence appears over- Pleistocene boreal small mammal fauna was whelming that these species were adjusted for adapted much as is its modern sub-Arctic boreal conditions as are their modern popu- equivalent. Johnston and Selander (1964 )have lation equivalents in the present-day Hudson- demonstrated in North American populations ian Canadian life-zones of Canada. of the house sparrow ( Passer domesticus) Kurten'S and Hibbard's evidence would a positive Bergmann's response which de-

180 THE NATIONAL SPELEOLOGICAL SOCIETY veloped within less than 100 years after the fossil record are coexisting today. Recogniz- bird was introduced into the continent. ing that the late-glacial woodland environment must have had unique climatic attri- FOSSIL POLLEN, SMALL VERTEBRATES, AND butes which can no longer be matched from THE LATE-GLACIAL CLIMATE any part of North America, it appears that Below six meters the pollen record at New the present geographical center for the fauna Paris No. 4 yielded an average of 15 per- represented in New Paris No. 4 during pre- cent grass and sedge pollen, much less than Allerlid time is found in eastern Canada at that found in clays from Cranberry Glades, approximately 50° N. latitude and in cen- West Virginia, from the Marsh, Pennsylvania, tral Canada west of Hudson Bay at approxi- 0 or from the T-zones of New England, and mately 57 N. latitude. In view of the fact slightly more than would be expected with- that many of the fossil forms found in the in a closed-canopy forest. The non-arboreal deposit can be related to populations at or pollen in late-glacial deposits is generally near the northern limit of the species to- viewed as originating at least in part in up- day, our "center of abundance" maps which land areas and blowing, along with tree pol- are based on the total range of each species len, into basins or alluvial depressions which have a southerly bias. For example, in the serve as pollen traps. This assumption is sel- case of the snowshoe rabbits, flying squir- dom proved and may be challenged in the rels, and voles (to name a few), the late- case of those basins surrounded by a fringe glacial populations represented at New Par- of hygric plant communities particularly fav- is No. 4 have size attributes of populations orable to the growth of sedges and grasses. at or near the northern limits of the ranges In the absence of riparian habitats near the of these forms so that the center of maxi- sinkholes the sedge-grass pollen in New Par- mum overlap shown in map B, center leaf, is No. 4 is best attributed to an upland is prombly several degrees of latitude too far source. The abundance of microtine rodents south. A second difficulty is the fact that and other species typical of grassy meadows the maps for units above and below six within the boreal woodland (Hudsonian-zone) meters share no species in common, thus is a more substantial demonstration of the exaggerating the differences between the fau- lack of closed-canopy forest. But the rodents nas. Map A, center leaf, is more southerly do not indicate a completely treeless tundra than should be the case. or steppe-tundra. The pollen record suggests that at least a few jack pines and spruce must Above 6.0 meters a decrease in the boreal have grown in the vicinity of the sinkhole. woodland element and an increase in boreal At 8.5 m a pollen sample of 20 percent forest-temperate forest species such as Cle- NAP was taken in a bone pocket beneath thri on o mys and G laucomy r suggests a a rock which yielded the remains of 35 Mi- denser growth of trees. In matrix from this cro xantbogna thus, 10 M. p coosylva- zone there is a sharp reduction in the relative nicur, two M. cbrot orrh in us, eight Synap- abundance of grass and sedge pollen, al- tomys borealis, one Sorex palustris, and though the non-arboreal sum remains relative- six to eight Bufo americanus cf. copei. ly constant. The grass and sedges are re- The fauna from this pocket is of either grass- placed largely by composite pollen of the land or woodland species of sub-Arctic dis- subfamily Liguliflorae. At the same time fern tribution. Neither extensive closed-canopy for- spores (Polypodiaceae) increase markedly. Li- est nor a treeless tundra or prairie would guliflorae pollen is not typical of lake sedi- support such a fauna. ments, and to find more than a few percent An overlapping distribution map prepared of it strongly suggests a dense local popu- for the 12 species of mammals with their lation of one species rather than wind trans- mean abundance below six meters in the port from a distant point. The appearance of deposit is shown in map B, center leaf. Such oak and betulaceous pollen types above 6.0 a map reveals the geographical areas in which meters further suggest a forest of the type the largest number of species found in the one presently finds in southern Canada and BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 181 the northern Lake States. While the local vege- and the Carolinas. Curiously, in the late- tation, whose pollen is represented in unit glacial of Minnesota (Fries, 1962; Jelgers- B, was probably an open jackpine-spruce ma, 1962; Wright, Winter and Patten, 1963), w oodland, that of pollen unit A was likely other parts of the midwest (for example to have been a closed-canopy forest with hard- West, 1961; Kapp and Gooding, 1964; Ben- woods starting to invade the jackpine. ninghoff and Hibbard, 1961), and east to A center of abundance map plotted for the extreme western Pennsylvania (Walker and Hartman, 1960) pine pollen frequencies are vertebrates present above 6.0 meters is shown quite low until the early post-glacial "pine in map A, center leaf. In this case the center is considerably south of that shown in period". In the deeper levels of midwestern diagrams spruce rather than pine is the domi- map B, center leaf, and focuses in the north- nant tree pollen type. Wood from late-glacial ern Appalachians from West Virginia to deposits in Iowa is mainly of mesic coni- southern Vermont. The westerly component fers such as hemlock ( Tsuga), fir ( Abies), strongly marked in map B, center leaf, is tamarack [ Larix], spruce ( Picea) and yew feebly represented by a distributional high in ( Taxus ).(Ruhe, Rubin and Scholtes, 1957). Wisconsin and northern Michigan. However, Pollen of Abies, Larix and nuga domi- the center of abundance is over the north- nated below 3.6 m in Muscotah Marsh, Kan- ern hardwoods region of the eastern decidu- sas (Horr, 1955), dated at 10.8 m as 15,- ous forest at latitude 40-40° N. Again, it 1,500 B.P. (M-352). should be pointed out that while the boreal 500 woodland element has greatly diminished in The relatively poor spruce record in un- the fauna, it is still present, and the center glaciated eastern and central Pennsylvania in- of abundance map shown in map A, center vites further study. Is it possible that during leaf, does not take into consideration the the late-glacial precipitation was heavier along fact that the species ranges used to plot the the midwestern ice margin, allowing forests map include the total range of the species, of spruce, fir, yew, hemlock and tamarack to whereas the measurements of the populations extend from eastern Kansas and Iowa to west- at New Paris, Pennsylvania strongly indicate ern Pennsylvania, while east of the Alleghe- only the northerly populations of each spe- nies there was a relatively dry woodland of cies were represented in the fossil record. jackpine? Both below and above six meters the dominant pollen type, pine, is of small size. Dif- Whatever the meaning of the apparent re- ficulties in relating pine pollen size to mod- gional difference in tree distribution in the ern species have been discussed by White- late-glacial, it gives little support to the bio- head (1962) who emphasized that the pre- geographic theory of the Appalachians as a sence of small-sized pine pollen in Wiscon- Pleistocene refugium for hemlock-white pine- sin-age strata is far from proof of the pre- northern hardwoods. Not only is hemlock re- sence of jack-pine. The most that might be ported from west of the Mississippi where said is that it is ecologically reasonable to it shoudn't have been, but also pollen of visualize an association of sub- Arctic hemlock-northern hardwoods is quite rare in shrews, voles and lemmings, pines, spruces, the late-glacial deposits of unglaciated Penn- and various herbs in which the dominant sylvania where this community supposedly conifer would be jackpine. Any other pine endured during glacial times. The presence with such a fauna would be anomalous in of pine-NAP in matrix containing boreal wood- terms of modern distribution of boreal land mammals including Microtus xantbog- plants and animals. nutbas as a dominant species, Sorev arc- Pine pollen (often of small size) is the ticas, Bufo an2ericanus copei, and at least dominant tree type found outside the ice one tundra mammal — Dicrostonyx— should margin in late-glacial and full-glacial deposits help dispel doubts about major biotic changes of unglaciated Pennsylvania, West Virginia, in unglaciated Pennsylvania in the late Pleis-

182 THE NATIONAL SPELEOLOGICAL SOCIETY tocene. Only in the absence of a late-glacial from different parts of the boreal forest (Ca- fossil record was it possible to imagine that: nadian-zone) and the boreal woodland (Hud- "The Hemlock-White Pine-Northern Hard- sonian-zone), with a suggestion of tundra woods Forest of the Lake region and the and prairie. The latter element reopens an Northeast has resulted from postglacial ex- old biogeographic question — that of the pansion of a forest persisting throughout prairie peninsula. the Pleistocene in mountainous portions of the East, particularly on the Alle- THE PRAIRIE PENINSULA PROBLEM gheny Plateau and Allegheny Mountains of Pennsylvania, where this forest still Before the development of post-glacial pol- maintains itself on unglaciated land." len stratigraphy in North America Gleason (Braun, 1950: 524, italics ours). The late- (1923) invoked an eastward extension of prairie to explain the relict pattern of prairie glacial biotic community at New Paris re- plant distribution in the midwest. Gleason veals no Hemlock-White Pine-Hardwood For- envisioned tundra growing on top of the est, or indeed, any closed-canopy forest, in the ice with a narrow broken fringe of boreal Allegheny Mountains 12,000 years ago. Con- coniferous forest along the margin. After de- ditions 20,000 years ago remain to be deglaciation he postulated the advance of prair- termined, but should certainly be no more ie to the east and northeast following the favorable for hardwoods than during the late- withdrawal of coniferous forests and before glacial. The biogeographic interpretation of the invasion of deciduous forest species in temperate refugia along the ice margin during post-glacial time. Transeau ( 1935 ) mapped the glacial times can be maintained only by in- distribution of prairie relicts and postulated terpreting animal remains of rodents we be- the prairie expansion during mid-post-glacial lieve are clearly intrusive ( Pitymys, Pero- time, after, not before, the initial invasion of myscus leucopus) as actually contemporane- decidous forest. ous with sub-Arctic species. In an article describing in detail the num- Any attempt to uncover the full-glacial fos- ber of reptile-amphibian distributions that sil record from unglaciated areas east of the fit the prairie peninsula pattern, Schmidt main- Mississippi is confronted with certain diffi- tained that it was a late-glacial, not a post- culties, including: (1) a shortage of lakes glacial, event. "By analogy with the situation and bogs, the traditional sources of a rich in Europe, and indeed on theoretic grounds, sedimentary record, (2) uncertainty in how there must have been a broad belt of tundra, succeeded by steppe, parallel to the re- to, proceed when sampling and interpreting `unconventional" pollen deposits such as treating front of the great glacier in eastern flood plain alluvium or cave fill which may North America. This belt of open country, be the only local sources of late Pleistocene broadly connected with the plains region to sediment, and (3) the problem of obtain- the west, and extending eastward to Pennsyl- ing radiocarbon-dated sediments exactly con- vania and sometimes even to the Atlantic temporaneous with the time of maximum Coast, to the north of the southeastern hard- biotic displacement, to coincide with maxi- wood forest, obviously afforded a natural mum ice advance of 17,000 - 23,000 years highway for the eastward spread of a part ago. Like many others the fossil record from of the western steppe fauna." (Schmidt, 1938). New Paris No. 4 is almost surely too young With the development of American post- to represent the full-glacial climate of the glacial palynology and the initial failure to Wisconsin maximum. Nevertheless, it indi- find tundra or steppe pollen in sediments cates major climatic change. And while the from the deciduous forest region, Schmidt's New Paris No. 4 fossil record includes some idea of tundra succeeded by steppe fell into mammal species which presently do not co- eclipse. Most of the more recent discussions exist in any single area, the mixture is not of the prairie peninsula have predicated its one of deciduous forest and boreal forest being a post-glacial event (Deevey, 1949; elements. Rather it is a mixture of species Braun, 1950; Smith, 1957) with some un-

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 183 certainty about possible prairie conditions dur- Smith (1957) distinguished between "cli- ing the early post-glacial pine period (pollen matic optimum" and "xerothermic" type dis- zone B, see Deevey, 1949: 1386). tributions, the former of southern species left isolated at northerly outposts, the latter The present record of two prairie species, of prairie peninsula species isolated in east- the thirteen-lined ground squirrel (Cite/Isis ward outposts. If relict northerly distribu- tridecemlineatus) and the sharp-tailed grouse tions of southern species are to be explained ( Pedioecetes phasianellus), in the east 11,- in terms of climatic change, then the climatic 000 years ago, restores currency to Schmidt's optimum is certainly the most likely time of view. Undated records of the thirteen-lined isolation, as Smith points out. Admittedly, ground squirrel, in association with boreal climatic change may not always provide the species but presumably of late-glacial age, better explanation of the southern relicts. For also come from Bootlegger Sink in the Pied- example, rice rat ( Oryzomys palustris) re- mont of eastern Pennsylvania, Cumberland mains in archaeological sites several hundred Cave, western Maryland, Natural Chimneys kilometers north of the present limit of the in the Shenandoah Valley of Virginia, and species are not likely to be the result of warm- Robinson Cave in north-cential Tennessee. er temperatures during the climatic optimum. The western magpie ( Pica pica) and the Fossil records of rice rats are closely asso- sharp-tailed grouse ( Pediocetes pbasianel- ciated with the cultivation of prehistoric maize. lus) are known from Natural Chimneys, Vir- Furthermore, all of these records are from ginia. Cumberland Cave, Maryland also pro- Indian villages of late prehistoric age, post- duced such western forms as badger ( Taxi- dating the Climatic Optimum by several thou- dea), coyote ( Canis — subgenus Thor), sand years. and cony ( Ocbotona). Cumberland Cave fossils may be pre-Wisconsin in age. Those Thus, while some of the prairie relicts from Port Kennedy Cave in eastern Pennsyl- may trace their origin to one or several post- vania, which include badger and coyote, al- glacial warm, dry periods, and while some most certainly are pre-Wisconsin. While ear- southern species probably extended farther lier invasions of western species moving east north during the "Climatic Optimum" as may also have occurred, the evidence of a Smith concludes, at least one of the latter, late-glacial invasion leads us to re-examine the race rat ( Oryzomys), is more likely the post-glacial evidence. a cultural than a climatic extension. The only Archaeofaunal sites dating back 5000 years dramatic fossil evidence for an eastern spread to the period of the climatic optimum in- of prairie border vertebrates, Cite/Isis. clude the following: Kentucky (Webb, 1946), Pedioecetes, Pica, etc., evidently predates the Illinois (Fowler and Parmalee, 1959), West post-glacial. Virginia (Mayer-Oakes, 1955), Pennsylvania ( Guilday and Parmalee, [ 19601), and New In an unpublished account of the fossil York State (Guilday, [ 19631). These faunas pollen record and post-glacial plant geogra- are composed of birds, mammals, reptiles, phy from the midwest, Benninghoff (MS) amphibians, and molluscs found in those has reached a similar conclusion — that the areas today. No major faunal change is in- late-glacial was the last episode of impor- dicated as might be expected if there were tant prairie movement eastward. While prair- an eastward penetration of prairie into the ie plants are often difficult to distinguish deciduous forest. Minor range extensions of from tundra or woodland species in the pol- a few (not strictly prairie) species include the len record, the presence of Ambrosia-type box turtle ( Terrapene carolina) and the pollen ( "low-spine Compositae" in table 2) fox squirrel ( Sciurus niger) in central New would not be expected in a modern tundra York State at the Archaic Lamoka Site (Guil- pollen rain (Wright, Winter and Patten, 1963: day, ibid.). This represents an eastward move- 1386; Benninghoff, MS). ment of approximately 160 km from the A mixture of mid-latitude heliophytes, in- Lake Erie area during the mid-post-glacial. cluding steppe plants, with an Arctic-montane

184 THE NATIONAL SPELEOLOGICAL SOCIETY element has long been recognized in the late- Plants with Coastal Plain affinities such glacial flora of Europe (see especially, Iver- as the bushy beard grass ( Andropogon vir- sen, 1954). While some plants and animals ginicus var. abbreviatus), marsh milkwort did move north during boreal and Atlan- ( Po lygala cruciata var. aquilonia), beaked tic times of the post-glacial, such as the tur- rush (Rhynchospora globularis var. recog- tle ( Emys orbicularis, see Degerbol and nita),nut rush( Scleria triglomerata), file- Krog, 1951), the steppe invasion took place leaved aster (Aster radula) and the Alle- earlier. Analysis of stratified deposits of small gheny glade gentian ( Gentiana Saponaria mammals from cave sites spanning this peri- var. al legheniensis) are found in isolated od (see Ja'nossy, 1961, for summary) shows mountain glades of Fayette County, 150 m steppe forms immediately succeeding tundra higher and some 60 km southwest of New species such as the lemming ( Dicrostonyx) Paris. Such relicts have been looked upon and preceding woodland forms such as the as remnants of a Cretaceous flora that sur- wood mouse (Apodemus), the dormice ( Glis, vived Tertiary uplift and the rigors of the 114usc,Irdinus), and the red-backed vole ( Cle- Pleistocene in situ (see Jennings, 1953 for thrionomys). As the woodland forms in- summary). A more likely assumption is that crease in relative numbers in such deposits they are late-glacial immigrants in that area. (see records from Jankovitch-HOhle and Pet- Such mid-western prairie forms as the blaz- enyi-HOhle, Hungary, Ji.nossy, 1960), the ing star( Liatris spicata),big-bluestem grass steppe forms become correspondingly scarce ( Andropogon furcatus), gray-headed-cone- and are eventually replaced during the early flower ( Ratibidapinnata), green milkweed post-Wtirm reforestation of the area (fig. 32). (Asclepias viridflora), and false boneset ( Kubnia eupatorioides), have been regarded There is evidence for a steppe phase lead- as post-glacial "Prairie Peninsula" species in ing into the WUrm glaciation as well as one western Pennsylvania, associated with the Hyp- immediately succeeding it (Ja.nossy, 1961). sithermal (Jennings, 1953). Unfortunately, A similar sequence may be anticipated during serious botanical work in northwestern Penn- sylvania did not begin until after the country early Wisconsin time in the central Appala- chians. If steppe elements were present dur- had been cleared and planted. Colonial agri- ing both early-glacial and late-glacial time the culture created, in effect, its own "Prairie correlation of cave deposits by their faunal Peninsula," one in which wind-dispersed make-up alone will require considerable cau- seeds could easily colonize. Whether these spe- tion. cies were native to the region prior to colonial deforestation will probably never be known. Speculation as to their time of arri- As Schmidt anticipated, the post-Wisconsin val in the region can only remain that. They invasion of prairie species in eastern United may or may not be valid "Prairie Peninsula" States is the mirror image of the post-WUrm indicators in the central Appalachians. steppe invasion of Europe in which the prairie elements withdrew as forest invasion pro- ceeded under an ameliorating climate. To EARLY MAN IN THE EASTERN LATE-GLACIAL date evidence for a mid-post-glacial invasion of prairie species during "Hypsithermal" Our knowledge of early man in eastern time is more difficult to find in the fossil North America grows steadily. Characteris- record. Unless such evidence emerges it seems tic chipped stone artifacts of what are pre- futile to assign a period of isolation — either sumed to be big game hunters of the late inter-glacial, late-glacial, or post-glacial — to Pleistocene ice margin are common in the such prairie relicts as the heath hen ( Tym- East. Precise dating is not yet possible, but panuchus cup ido ), the small-headed garter in Michigan (Mason, 1958) and Ohio (Pruf- snake ( Th amn oph is brachys tom a), the mas- er and Baby, 1963) sub-types of Paleo-Indian sassauga (Sistrurus catenatus), andthechor- artifacts may be correlated with Wisconsin us frog ( Pseudacris nigrita kalmi). terminal moraines. Throughout the retreat BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 185 of the Wisconsin Ice Sheet, an evolution of mental nature of the tool industry also in- artifact styles can be associated with succes- dicate that the Shoop Site represents the oc- sively younger moraines. cupation of very early hunters of large game animals. At this time such areas of rugged "The Paleo- Indian artifacts south of the Wis- physiography as the North Mountain section consin maximum boundary theoretically could may not have been heavily wooded, and much date from a period earlier than 17,000 years of Pennsylvania may have been prairie. Such ago. The remarkably consistent radiocarbon a difference in biota would help explain the dates marking the maximum advance of the location of the Enterline industry sites on Wisconsin ice suggests that the retreat of the high ground." glacier began about 17,000 years ago. It follows that the Paleo-Indian penetration of the The location of this site is apparently in- glaciated tracts of the state cannot be earlier explicable in the light of the preferences of than that date. later cultures who utilized the area when it "A second terminus is provided by a date was heavily forested. It would be an advan- for the Wabash Moraine. The retreating ice tageous site for hunters of large grassland margin had reached that position approxi- mammals if the country had been open or mately 14,500 years ago. It follows, again, sparsely forested — a lookout from which the that the area north of the Wabash Moraine Paleo-Indian hunter could locate roving herds. line could not have been occupied by Early Sites believed to be of this time period, such Man prior to that date. as Bull Brook in Massachusetts (Byers, 1954), "Datable evidence of early human occupancy Shoop in Pennsylvania (Witthoft, ibid.), Par- south of the Wabash Moraine is lacking. Nev- rish in Kentucky (Webb, 1951), Williamson ertheless the relatively heavy concentration of in Virginia (McCary, 1951), various sites in fluted points in central and south-central Ohio, Ohio (Prufer and Baby, ibid.), Well's Creek and the known existence of Pleistocene mam- Crater Site in Tennessee (Dragoo, unpubl.); mals in the area indicate the probable pene- have been in such exposed situations. tration of Early Man at least as early as 11,- Mounting paleontological evidence, espe- 000 years ago," (Prufer and Baby, 1963, cially the integrated faunal and floral picture p. 55 ). presented by New Paris No. 4, points to an These early people are envisioned as hunt- open woodland during late-glacial times. This ers of big game that roamed the open areas agrees with the interpretations of the archeo- of the glacial fore-front. Witthoft (1952, p. 494), speaking specifically of the Paleo-In- logist. If the thirteen-lined ground squirrel dian occupation at the Shoop Site at Enter- (Cite//us tridecemlineatus) is used as a line, Pennsylvania, about 160 km northeast prairie indicator as it is today, then late Pleis- of New Paris, commented: "They were prob- tocene woodland openings extended as far ably the first thin vanguard in the settlement south as northern Tennessee (Robinson Cave), of the Northeast. Highly mobile, nomadic as far east as eastern Pennsylvania (Bootlegger hunters of large game, contemporary with Sink, in press ) and central Virginia during extinct mammals of the closing Pleistocene." Paleo-Indian times, i.e. 11,300 B.P. if the In discussing the topographical situation of New Paris No. 4 carbon date is taken at the site, he says, (p. 467): "The hill itself face value. commands a large area of Armstrong's Val- The Modoc Rockshelter in southern Illi- ley north of Dividing Ridge, but is much nois (Fowler and Parmalee, 1959) and the too elevated and in too isolated a location Sheep Rock Shelter of central Pennsylvania for one to expect an archaeological site." (unpublished faunal report by Guilday and The valley is broad and rough-floored, wall- Parmalee, 1960) demonstrate that the faunas ed by mountains on three sides and broadly of these areas have remained essentially un- open to the Susquehannah River on the west. changed during the past six to seven thous- "The weathering stage of the flints, the ab- and years. The record is especially conclu- sence of ground stone tools, and the ele- sive at Sheep Rock because the fauna is com- 186 THE NATIONAL SPELEOLOGICAL SOCIETY posed mainly of small mammals from owl If the Paleo-Indian hunters were present in pellets that accumulated on the aggrading this area during late Pleistocene times, as in- shelter floor coincident with the Indian oc- dications seem to point, they would presum- cupation. The collection, freed therefore from ably be associated with a boreal fauna such any selective bias on the part of the Indians, as that of New Paris No. 4. While direct consisted primarily of ecologically sensitive evidence of prehistoric man is lacking at rodents and insectivores of the area. At Sheep New Paris, his effectiveness as a hunter may Rock, a stratigraphic change was noted in the be reflected in the disappearance of the pec- relative numbers of cottontail rabbit (Syl- cary (Mylohyus) in Pennsylvania and else- vilagus sp.) and southern flying squirrel where in eastern North America at this time. ( Glaucomys volans) bones. According to The peculiar nature of late Pleistocene ex- Dr. Anton Kovar, Pennsylvania State Uni- tinction, which affected only large-sized ter- versity, this correlates with an increase in restrial genera, has been so often mentioned non-arboreal pollen from these levels. This that it seems unnecessary to point it out is probably due to the clearing of the area again in the case of the New Paris No. 4 by late Prehistoric peoples for the growing fossil record. It is very difficult to account for of crops during Late Prehistoric times. The the pattern of late Pleistocene extinction with- fauna, even in the oldest levels (c. 6000 years out appealing to Early Man as a major cause ago), was a Recent one. (see also Martin, 1963).

CONCL USIONS A picture of late-glacial climate and life Where critical comparisons can be made in unglaciated Pennsylvania can be gained from between fossil and modern populations with- the fossil content of nine meters of cave fill in a wide-ranging species, as in the case of in a sinkhole near New Paris, Bedford County. snowshoe hare ( Lepus americanus), north- A total of almost 3000 identified vertebrates ern flying squirrel ( Glaucomys sabrinus) associated with a pollen profile and radio- or the short-tailed shrew ( Blarina brevi- carbon date of 11,300 ± 1000 years B.P. cauda), the late-glacial population reveals size- (Y-727) indicate boreal woodland and boreal attributes of populations now living in south- forest environments. The fossil-bearing por- ern or central Canada. In those modern cases tion of the deposit can be divided into two in which Bergmann's Rule is violated, i.e. units: the lower from nine to six meters small-sized animals characterize the popula- (unit B) is dominated by woodland (Hud- tions to the north, the New Paris fossils sonian-zone) vertebrates whose modern center are also relatively small. Modern and fossil of distribution lies southeast and west of populations are indistinguishable only in the Hudson Bay in central Canada (map B, cen- case of suspected contamination ( Pitymys ter leaf). The pollen record suggests that pinetorum. Peromyscus 1eucopus), where spruce and pine (probably jackpine) were the fossil material is insufficient for critical com- only trees present with open ground separat- parisons, or in species that are now confin- ing them. The upper unit (unit A) from six ed to the Arctic or sub-Arctic ( Microtus to four meters, may correspond to the Aller- xantbognatbus, Dicrostonyx budsonius). Od of Europe and to pollen zone A-3 of New England. In it the relative proportion of Although the cave fill is dominated by forest (Canadian-zone) species of vertebrates boreal woodland or boreal forest species, increases and the center of their modern it contained the first Pleistocene record from distribution lies well to the south of the spe- North America of the Labrador collared lem- cies dominant in the lower unit (map A, ming ( Dicrostonyx budsonius), whose center leaf). In unit A the pollen of broad- modern distribution lies almost entirely with- leaved trees increases and some, such as birch in the tundra of northern Quebec. In addi- and oak, may have invaded the pine-spruce tion there were bones of two prairie spec- community. ies, the thirteen-lined ground squirrel ( Ci-

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964. 187 tellus tridecem lineatus) and the sharp-tailed dians were contemporaries of the long-nosed grouse ( Pedioectr,s phasiunellus). The col- peccary during the boreal woodland period, lared lemming may be a relict from full-gla- but no direct association was evident in the cial times which lingered on the tops of record of Sinkhole No. 4. the Allegheny Mountains during the late-gla- The New Paris fossil record is only of cial, while the ground squirrel and sharp- indirect help in establishing the climate of tailed grouse indicate a steppe invasion much the Appalachian Mountains in Pennsylvania earlier than that commonly attributed to the during the maximum advance of Wisconsin Xerothermic or Hypsithermal of the pollen ice 20,000 years ago. The involved physio- stratigrapher (pollen zone C-2). A late-glacial graphy of the region may have allowed tun- steppe invasion mirroring that known in Eur- dra and taiga to interdigitate over 200 km ope was postulated by Schmidt (1938). in a north-south direction as Martin (1958) Whether all prairie relicts in the East date concluded when he discovered a high fre- back to the late-glacial is problematic, but quency of NAP associated with small-size thus far a fossil prairie element is clearly pine pollen and spruce in a swale in the seen only in deposits which we consider Pennsylvania Piedmont. Evidently a pollen de- late-glacial or older and is not known from posit similar to that of the Piedmont occurs mid-post-glacial deposits. in basal clays at Cranberry Glades, West Vir- With the exception of the long-nosed pec- ginia. Additional evidence for major Wiscon- cary ( Mylohyus nasutus) none of the New sin-age biotic displacement in the unglaciated Paris animals identified to date are unequi- East may be inferred from the recovery of vocally extinct species. Mylobvus has been boreal plant pollen in the Carolinas (Frey, found previously in association with boreal 1953, 1955; Whitehead, 1963) boreal ver- animals in the East and its occurrence at tebrates in Tennessee (Robinson Cave), and New Paris No. 4 cannot be taken as evidence also in the record of the collared lemming for warm climates, much less for an eco- ( Dicrostonyx hudsonius) at New Paris, per- logically anomalous mixture of subtropical haps 5000 years after the glacial maximum. and boreal species. While details of the While suggestive, none of the assembled evi- sequence of events which led to the dence is conclusive proof for tundra as a extinction without replacement of large ver- distinct vegetation zone outside the ice mar- tebrates in the late Pleistocene is unknown, gin. Even the presence of lyesotundra (the Mylohyui was so wide-ranging throughout boreal woodland-tundra ecotone) cannot be eastern North America at the time (to Tex- claimed with certainty. At the same time the as and Florida) that appeal to climate as boreal aspect of the late-glacial biota from a cause of extinction seems out of the ques- New Paris No. 4 provides little support for tion. The local geography of the Shoop Site the biogeographic view that the unglaciated 160 km north of New Paris and the sus- Appalachians in this region served as an ice pected dates of man's arrival in eastern North refugium for temperate species of plant and America make it seem likely that Paleo-In- animal life.

BIBLIOGRAPHY

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188 THE NATIONAL SPELEOLOGICAL SOCIETY Bee, James W. and E. Raymond Hall Michigan. Unpubl. MS on file Dept. 1956 Mammals of northern Alaska on the Anthrop. Univ. Michigan. Arctic Sipe. Univ. Kansas Mus. Nat. Conant, Roger Hist. Misc. Publ. No. 8, p. 1 -309. 1958 A field guide to reptiles and amphi- Benninghoff, William S. bians of the U. S. and Canada east [MS.] MS. The Prairie Peninsula as a fil- of the 100th Meridian. Houghton Miff- ter barrier to post-glacial plant mi- lin Co., Boston, Mass. 366 pp. gration. Cope, Edward D. Benninghoff, William S. and C. W. Hibbard 1869 Synopsis of the extinct mammalia of 1961 Fossil pollen associated with a late- a the cave formations of the U. S., with glacial woodland musk ox in Michi- observations on some Myriapoda gan. Papers Mich. Acad. Sci. Arts found in and near the same, and on and Lett., v.46, p. 155 - 159. some extinct mammals of the caves Binford, Lewis R. of Anguilla, W. 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190 THE NATIONAL SPELEOLOGICAL SOCIETY Biol. Soc. Wash., v. 66, p. 191 — 194. Iversen, Johs 1956 Bones of mammals from West Vir- 1954 The late-glacial flora of Denmark and ginia caves. Amer. Midland Nat., v. its relation to climate and soil. Dan- 56, no. 1, p. 250— 256. marhs Geol. Undersogelse, 2nd ser- Hare, F. Kenneth ies, no. 80, p. 87— 119. 1950 Climate and zonal divisions of thebor- 1960 Problems of the early post-glacial for- eal forest formation in eastern Canada. est development in Denmark, Dan- Geogr. Rev. v. 40, no. 4, p. 615 — marhs Geol. Undersogelse, 4th ser- 635. ies, v. 4, no. 3, p. 1 — 32. Harper, Francis Jackson, Hartley H. T. 1961 Land and fresh-water mammals of the 1915 A review of the American moles. N. Ungava Peninsula. Misc. Publ. no. 27, Amer. Fauna no. 38, 100 pp. Univ. Kan. Mus. Nat. Hist. Lawrence. 1928 A taxonomic review of the Ameri- 178 pp. can long-tailed shrews. N. Amer. Fauna no. 51. 238 pp. Hay, Oliver P. - 1920 Descriptions of some Pleistocene ver- JA'NOSSY, Denes tebrates found in the United States. 1960 Nacheiszeitliche Wan dlungen der Kleiniangerfauna Ungarns. Zoologi- No. 2328. Proc. U. S. Nat. Mus. scher Anzeiger. Bd. 164, Heft 3/4, v.58, p.83 —146, pls. 3 — 11. S. 114— 121. Hibbard, Claude W. 1957 Notes on late Cenozoic shrews. Trans. 1961 Die Entwicklung der Kleinsiugerfauna Kan. Acad. Sci., v. 60, no. 4, p. 327 — Europas im Pleistoian (Insectivora, Ro- 336. dentia, Lagomorpha). Zeitschrift fiir 1963 A late Illinoian fauna from Kansas Siugertierkunde, Bd. 26, H. 1, S. and its climatic significance. Papers, 1 — 64. Mich. Acad. of Sci., Arts & Lett., JLossy, Dines and E. Schmidt v. XLVIII, p. 187 — 221. 1960 Extreme Varianten des M, der Feld- HOOIJER, D. A. maus ( Microtus arvalis Pall) in 1947 Pleistocene remains of Panthera ti- Ungarn. Vertebrata Hungaric. Musei gris (Linnaeus) subspecies from Historico-Nauralis Hungarici. Tom. Wanhsien, Szechwan, China, compar- II, Fasc. 1, p. 137 — 142. ed with fossil and Recent tigers from Jelgersma, S. other localities. Amer. Mus. Novitates, 1962 A late-glacial pollen diagram from Mad- no. 1346, pp. 1 — 17. elia, south-central Minnesota. Amer. Jour. Sci. v. 260, p. 522 — 529. Horr, W. H. 1955 A pollen profile study of the Mos- Jennings, 0. E. and A. Avinoff cotah Marsh. Univ. Kan. Sci. Bull., 1953 Wildflowers of western Pennsylvania v. 27, p. 143 — 149. and the Upper Ohio Basin. Univ. Howell, Arthur H. Pittsburgh Press. 2 vols. 574 pp. 200 1918 Revision of the American flying squir- pls. rels. N. Amer. Fauna No. 44. 64 pp. Johnston, Richard F. and Robert K. Selander 1938 Revision of the North American 1964 House sparrows: rapid evolution of ground squirrels. N. Amer. Fauna No. races in North America. Science, v. 56. 256 pp. 144, no 3618, p. 548— 550. Huxley, Julian S. Jones, J. Knox, Jr. 1932 Problems of relative growth. Methuen 1960 Absence of third upper premolar in & Co., Ltd., London. 276 pp. 105 Eutamias. Jour. Mammal., v. 41, illus. no. 2, p. 268 — 269.

BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 191 Kapp, Ronald 0. and Ansel M. Gooding 1959 How many longs make a forest? Ohio 1964 A radiocarbon-dated pollen profile Jour. Sci. 59:221 — 222. from Sunbeam Prairie Bog, Darke 1963 The last 10,000 years: a ibssil pollen County, Ohio. Amer. Jour. Sci. v. record of the American Southwest. 262,p. 259 — 266. Univ. Ariz. Press, Tucson, 87 pp. Kauffman, Nelson M. Mason, Ronald J. 1960 Climates of the states, Pennsylvania. 1958 Late Pleistocene geo-chronology and Climatography of the United States the Paleo-Indian penetration into the No. 60 — 36. U. S. Govt. Printing lower Michigan peninsula. Univ. Mich. Office, \Vashington. 20 pp. Mus. Anthrop., Anthrop. Papers no. Komarek, E. V. 11, 12 pp. 1932 Distribution of Microtus chrotorr- Mayer-Oakes, William J. hinus, with description of a new sub- 1955 Excavations at the Globe Hill Shell species. Jour. Mammal., v. 13, no. 2, Heap (46 Hk 34 — 1) Hancock Coun- p. 155 — 158. ty, West Virginia. Publ. ser. no. 3, W. Va. Arch. Soc., Inc. Moundsville. Kurten, Bjorn 34 pp. 1960 Chronology and faunal evolution of Mayr, Ernst the earlier European glaciations. So- 1963 Animal species and evolution. The cietas Scientiarum Fennica. Commen- Belknap Press of Harvard Univ. Press, tationes Biologicae XXI. 5, 1 — 62. Cambridge, Mass. 797 pp. Lehmann, U. McCary, Ben C. 1954 Die Fauna des '`Vogelherds " bei 1951 A workshop site of early man in Din- Stetten ob Lontal (Wiirttenberg). N. - 52 widdie County, Virginia. Amer. An- Jb. Geol. Paamont., Abh. 99, 33 — tiquity, XVII, p. 9— 17. 146. Stuttgart. McCrady, Allen D. and Vic Schmidt Leidy, Joseph 1963 Second Interim Report — Late Pleisto- 1889 Notice and description of fossils in cene fossils from Robinson Cave, Ten- caves and crevices of the limestone nessee. The Netherworld News, Pitts- rocks of (Monroe and Berks counties) burgh Grotto, Natl. Speleological Pennsylvania. Geol. Surv. of Pa. 2nd Soc., v. 11, no. 2, p. 19— 27. Ann. Rept., 1887, p. 1 — 20. McKeever, Sturgis [MS.] Ecology and distribution of the mam- Lensink, Calvin J. - mals of West Virginia. PhD. thesis 1954 Occurrence of Mu rot/1$ voothogno- on file, N. Car. State Coll., Raleigh, thus in Alaska. Jour. Mammal., v. 35; p. i — xv, 1 — 335. no. 2, p. 259. Mercer, Henry C. Logier, E. B. S. and G. C. Toner 1899 The bone cave at Port Kennedy, Penn- 1961 Check list of the amphibians and sylvania, and its partial excavation in reptiles of Canada and Alaska. Con- 1894, 1895, and 1896. Jour Acad. tribution No. 53, Life Sci. Div., Roy- Nat. Sci. Phila. XI (ser. 2): 269 — al Ont. Mus., Toronto. p. 1 — 91. 286. Lundelius, Ernest L., Jr. 1960 111, /ob yo■ nosoto, long-nosed pec- Merrill, Glen K. cary of the Texas Pleistocene. Bull. 1960 Additional notes on vertical shafts in Texas Mem. Mus. No. 1, 40 pp. limestone caves. Natl. Speleological Martin, Paul S. Soc. Bull., v. 22, pt. 2, July. p. 101 — 1958 Taiga-tundra and the full-glacial peri- 108. od in Chester County, Pennsylvania. Paulson, Gerald R. Amer. Jour. Sci. 256:470 — 502. 1961 The mammals of the Cudahy fauna.

192 THE NATIONAL SPELEOLOGICAL SOCIETY Papers, Mich. Acad. Sci., Arts & Lett., Ruhe, R. V., M. Rubin, and W. H. Scholtes v. 46, p. 127 — 153. 1957 Late Pleistocene radiocarbon chrono- Peterson, Olaf A. logy in Iowa. Amer. Jour. Sci., v. 1926 The fossils of the Frankstown Cave, 255, p. 671 —689. Blair County, Pennsylvania. Ann. Cam. Schaefer, H. Mus. v. 16, no. 2, p. 249 — 297. 1935 Studien an mitteleurop.iischen Kleins- Pohl, E. R. iugern, mit besonderer BerUcksichti- 1955 Vertical Shafts in Limestone Caves. gung der Rassenbildung. Archiv fiir Occ. Papers Nt'l. Speleological Soc. Naturgeschichte, N. F., Bd. 4, H. No. 2 April. p. 1 — 24. 4, p. 535 — 590. Schmidt, Karl P. Preble, Edward A. 1938 Herpetological evidence for the post- 1908 A biological investigation of the At- glacial eastward extension of the steppe habaska-Mackenzie region. N. Amer. in North America. Ecology, v. 19, Fauna no. 27. 574 pp. no. 3, p. 396 — 407. Prufer, Olaf H. and Raymond S. Baby Schrock, Alta E. 1963 Palaeo-Indians of Ohio. Ohio Hist. [MS.] A preliminary analysis of an unglaci- Soc., Columbus. 68 pp. ated bog in Pennsylvania. Upubl. PhD Rand, A. L. thesis, Univ. of Pittsburgh. 1944. 1943 Canadian forms of the meadow mouse Schwartz, Albert and E. P. Odum ( Microtus pennsylvanicus). The Ca- - 1957 The woodrats of the eastern United nadian Field-Naturalist, v. 37, no. 7, States. Jour. Mammal., v. 38, no. 2, p. 115 —123. p. 197— 206. Rhoads, Samuel N. Shelford, V. E. and A. C. Twomey 1903 The mammals of Pennsylvania and 1941 Tundra animal communities in the vi- New Jersey. Philadelphia. 266 pp. cinity of Churchill, Manitoba. Ecology, Richmond, Neil D. and Harry Rosland (sic ) v. 22, no. 1, p. 47 — 69. 1949 Mammal survey of northwestern Penn- Sisson, Septimus and J. D. Grossman sylvania. Final Report Pittman-Robert- 1938 The anatomy of the domestic animals. son Project 20-R. Pa Gam Comm., 3rd ed. W. B. Saunders Co. Phila- Harrisburg, 67 pp. delphia. 972 pp. Richmond, Neil D. and William C. Grimm Smith, Philip W. 1950 Ecology and distribution of the shrew 1957 An analysis of post-Wisconsin bio- Sorex (//spar in Pennsylvania. Eco- geography of the Prairie Peninsula re- logy, v. 31, no. 2, p. 279— 282. gion based on distributional phenomena among terrestrial vertebrate popu- Roberts, Harvey A. and Robert C. Early lations. Ecology, v. 38, no. 2, p.205 — 1952 Mammal survey of southeastern Penn- 218. sylvania. Final Report Pittman-Robert- Snyder, Dana P. son Project 43-R. Pa Game Comm., 1954 Skull variation in the meadow vole Harrisburg. 70 pp. (Microtus p pennsylvanicus) in Riirig, G. and C. 136rner Pennsylvania. Ann. Cam. Mus., v. 33, 1905 Studien Ober das Gebiss mitteleuro- art. 13, p. 201 — 234. plischer rezenter Miuse. Arb. Kais. Stegeman, LeRoy C. Biol. Anst. v. 5, p. 37 — 89. 1938 European wild boar in the Cherokee Roslund, Harry R. National Forest, Tennessee. Jour. 1951 Mammal survey of northcentral Penn- Mammal., v. 19, no. 3, p. 279 — 290. sylvania. Final Report Pittman-Robert- Stone, Ralph W. son Project 37-R. Pa. Game Comm., 1932 Pennsylvania caves. Pa. Geol. Surv., Harrisburg. 55 pp. ser. 4, Bull. G3, 2nd ed., p. 1 — 135. BULLETIN VOLUME 26, NUMBER 4, OCTOBER, 1964 193 Thenius, Erich tory in Wisconsin, particularly changes 1959 Die jungpleistaine Wirbeltierfauna associated with the Valders readvance. von Willendorf i. d. Wachau, N.O. Amer. Jour. Sci., v. 259, p. 766 — Mitt. Piahistor. Komm. Osterr. Akad. 783. Wiss. 8/9, 133 — 170, Wien. Wetmore, Alexander 1960 Die pleistainen und holozinen Wir- 1959 Notes on certain grouse of the Pleis- beltierreste der Griffener Hale, Karn- tocene. Wilson Bull., v. 71, no. 2, ten. Mitt. des Naturwissenschaftlichen p. 178 — 182. Vereines fur ICirnten, 70. bzw. 150. 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Hartman tocene deposits from western North 1960 The forest sequence of the Hartstown Carolina and South Carolina. Ecol. bog area in western Pennsylvania. Eco- Monog., v. 32, p. 347 — 369. logy, v.41, p. 461 —474. Witthoft, John 195 2 A Paleo-Indian site in eastern Penn- Webb, William S. sylvania: an ear ly hunting culture. 1946 Indian Knoll Site On 2, Ohio Coun- Proc. Amer. Phil. Soc. XCVI, no. ty, Kentucky. Univ. of Ky. Rept. An- 4, p. 464 —495. throp. and Arch., v. IV, no. 3, Pt. Wright, H. E., Jr., T. C. Winter and I., p. 115 —365. H. L. Patten 1951 The Parrish Village Site. Univ. of 1963 Two pollen diagrams from southeast- Ky. Rept. Anthrop. & Arch., v. 7, ern Minnesota: problems inthe region- no. 6, p. 407 —451. Lexington. al late-glacial and post-glacial vegeta- West, R. G. tional history. Geol. Soc. Amer. Bull., 1961 Late- and post-glacial vegetational his- v. 74, p. 1371 — 1396.

Carnegie Museum Pittsburgh, Pennsylvania

Manuscript received by the editor 4 July, 1964.

194 THE NATIONAL SPELEOLOGICAL SOCIETY