Late Cenozoic Molluscan Faunas From the High Plains
GEOLOGICAL SURVEY PROFESSIONAL PAPER 337 Late Cenozoic Molluscan Faunas From the His:h Plains
By DWIGHT W. TAYLOR
GEOLOGICAL SURVEY PROFESSIONAL PAPER 337
Description of mo Husks from nine Pliocene and Pleistocene faunas of the High Plains region and interpretation of their stratigraphic and ecologic significance
UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1960 UNITED STATES DEPARTMENT OF THE INTERIOR
FRED A. SEATON, Secretary
GEOLOGICAL SURVEY Thomas B. Nolan, Director
The U.S. Geological Survey Library has cataloged this publication as follows :
Taylor, Dwight Willard, 1932- Late Ceiiozoic molluscan faunas from the High Plains. Washington, U.S. Govt. Print. Off., 1959. iv, 94 p. illus., 4 plates, tables. 30 cm. (U.S. Geological Survey. Professional paper 337) "Description of mollusks from nine Pliocene and Pleistocene faunas of the High Plains region and interpretation of their stratigraphic and ecologic significance." Bibliography: p. 83-86. 1. Mollusks, Fossil. 2. Paleontology Cenozoic. 3. Paleontology U.S. High Plains region. I. Title. (Series)
For sale by the Superintendent of Documents, U.S. Government Printing Office Washington 25, D.C. - Price 31 (paper cover) CONTENTS
Page Page Abstract- ______1 Sanders local fauna______--_--_-_--______40 Introduction ______1 Deer Park local fauna-______41 Acknowledgments. ______2 Recent mollusks from northern Nebraska,______42 Methods. ______2 Localities- ______42 Collecting methods. ______2 Habitats_____-___------43 Environmental interpretations of sediments ______3 Species found at the northern Nebraska localities- _ _ 44 Environmental interpretations of fossil mollusks _ _ _ _ 3 Systematic descriptions.______44 Previous work______4 Class Pelecypoda.__-____--_____--_------_-----_ 45 Principles of paleoecologic interpretation-______4 Family Unionidae______45 Difficulties of paleoecologic interpretation.______5 Family Quadrulidae_---_-_------__------45 Specific identification of shells ______5 Family Sphaeriidae______45 Recognition of limiting factors______5 Class Gastropoda.______48 Inadequacy of habitat and distributional data__-__- 7 Family Valvatidae______-_---_---___ 48 Unsatisfactory taxonomy.______8 Family Viviparidae______-__--__--___ 49 Methods of faunal interpretation ______8 Family Hydrobiidae-_____-_----_------_--_- 49 Age determination ______9 Family Carychiidae.______51 Faunal dating______9 Family Lymnaeidae______52 Local faunas.______10 Family Planorbidae._____--_-___-_----___-__ 56 Comparison of faunas. ______11 Family Ancylidae______------___---___ 61 Summary of faunal sequence ______14 Family Physidae______62 Correlation of North American ages with Lyellian epochs. 18 Family Strobilopsidae-_-__------_- 65 Summary of depositional and biological sequence. __ 18 Family Pupillidae______67 Correlation ______19 Family Valloniidae______76 Description of local faunas______22 Family Cionellidae______------______77 Buis Ranch local fauna______22 Family Succineid.ae______-_-__----_--____ 77 Saw Rock Canyon local fauna______22 Family Endodontidae______-__----_-___ 79 Red Corral local fauna.______25 Family Limacidae______-_-____----_-___ 80 Rexroad local fauna..______27 Family Zonitidae______80 Bender local fauna-______31 Family Polygyridae______82 Sand Draw local fauna______32 Literature cited---______-____-_------_------_--_ 83 Dixon local fauna______36 Index.______-___--_-----_-____ 87
ILLUSTRATIONS
[Plates follow index] Page PLATES 1-4. Late Pliocene and Pleistocene mollusks from the High Plains. FIGURE 1. Summary of late Cenozoie faunal, climatic, and lithologic sequences in southwestern Kansas, and correlation of other local faunas from the High Plains-______--______--___-----_----_----_---.------15 2. Frequency distributions of Pliocene Gastrocopta cristata group______--_____-_-___------_------68
TABLES
Page TABLE 1. Abundance or occurrence of gastropods in collections of Saw Rock Canyon local fauna._-_---_---_------3 2. Stratigraphic occurrence of mollusks of some late Cenozoie faunas in the High Plains______-----_-_--- 12 3. Summary of data from table 2______---__-___----_------13 4. Faunal similarities-______--_----_--_------13 5. Distribution of thermally significant species in late Pliocene and early Pleistocene faunas of the High Plains-_ 13 6. Correlation of middle Pliocene to lower Pleistocene deposits in Italy and the High Plains. ______------21 ni IV CONTENTS Page TABLE 7. Comparison of Blancan and Recent molluscan faunas of southwestern Kansas, by habitats__ ___-_-_-_____-_-__ 24 8. Habitat indications of mollusks of Saw Rock Canyon local fauna.-_____.______-__-.--___-_--___-_ 25 9. Habitat indications of mollusks of Red Corral local fauna______26 10. Comparison of mollusks of Red Corral local fauna with Recent mollusks from Palo Duro Canyon, Tex., by habitats______-______-_-______...-_-_--_---_-_.---.-___.--- 26 11. Locality occurrence of mollusks of Rexroad local fauna______--_-----_-----______29 12. Habitat indications of mollusks of Rexroad local fauna__-______------__- 30 13. Habitat indications of Bender local fauna______-____-_-___-__--_ _ 32 14. Habitat indications of mollusks of Sand Draw local fauna______35 15. Comparison of Sand Draw local fauna and Recent mollusks from northern Nebraska.,_^______-___-______36 16. Occurrence of supposed Nebraskan index fossils.______38 17. Habitat indications of mollusks of Dixon local fauna______39 18. Comparison of mollusks of Dixon local fauna with those living in Kingman County, Kans., by habitats.______39 19. Habitat indications of mollusks of Sanders local fauna______.______------_-__---_----_ 41 LATE CENOZOIC MOLLUSCAN FAUNAS FROM THE HIGH PLAINS
By D. W. TAYLOR
ABSTRACT about in the middle of the Kansan age, and the Irvingtonian- Rancholabrean boundary about in the middle of the Illinoian Mollusks are described from nine late Cenozoic assemblages age. in the High Plains region: the late middle Pliocene Buis INTRODUCTION Ranch local fauna of northwestern Oklahoma; the late Pliocene Saw Rock Canyon, Rexroad, and Bender local faunas In the spring of 1936 M. K. Elias visited a Civilian of southwestern Kansas; the late Pliocene Red Corral local Conservation Corps quarry in Meade County, south fauna of the Panhandle of Texas; the early Pleistocene Dixon, western Kansas. Questioning the workmen about the Deer Park, and Sanders local faunas of southern Kansas; and the early Pleistocene Sand Draw local fauna of north-central possible occurrence of fossil bones;,-he was given speci Nebraska. In nearly all assemblages associated mammals mens which he brought to the University of Kansas. provide an independent age determination. The control pro Claude W. Hibbard and a University of Kansas Mu vided by fossil mammals and stratigraphie succession permits seum of Vertebrate Paleontology field party, acting evaluation of the mollusks for age determination and correla on this information, returned to the area in 1936 tion. Vertebrates and mollusks are complementary, and never antagonistic, in giving age and ecologic information. (Hibbard, 1941, p. 81). Fossils collected at this The late Pliocene and Pleistocene faunal sequence of south original locality and exploration of the Meade County western Kansas and northwestern Oklahoma is especially region in general subsequently have produced almost significant for regional and intercontinental correlation because an overabundance of paleontological riches. Intensive it is the most nearly complete sequence known from such a study by Hibbard and parties from the University small area. Collecting the great number of specimens and of Kansas (1936-46) and University of Michigan species which represent these faunas has been possible by washing large quantities of fossiliferous matrix through (1947 to date) has resulted in the discovery of the screens. Eeologic interpretation of the mollusks is combined most nearly complete sequence of late Pliocene and with other information from the vertebrates and inferences Pleistocene faunas in a small area known anywhere. from the depositional sequence to summarize the late Pliocene The washing technique developed by Hibbard (1949c) and Pleistocene climatic and faunal sequence of southwestern is largely responsible for this great success in col Kansas. Fossils from three horizons in the Rexroad formation indi lecting; the method permits rapid recovery of 90 cate a relatively long interval in which the climate was warm- percent or more of the bones and shells in a fossili temperate, equable, and relatively moist. Above these as ferous deposit. The various animal groups so ob semblages the depositional and erosional sequence, and several tained birds, mammals, salamanders, frogs, toads, faunas, indicate 4 periods of erosion and deposition of coarse lizards, snakes, turtles, fishes, and mollusks provide clastic sediments in cool climates separated by 3 periods of deposition of finer materials during warmer climates. The a basis for reconstructing in considerable detail the alternate warm and cool climates are correlated with the changes in climate and animal life during the late Mississippi Valley interglacial and glacial ages. The Ballard Cenozoic. formation, which is separated from the Rexroad formation by This paper is concerned primarily with the descrip an erosional unconformity and contains the earliest evidence tion and interpretation of the late Pliocene and early of marked climatic cooling, is correlated with the earliest Pleistocene molluscan faunas of southwestern Kansas. Pleistocene sediments of the area in central Italy selected by the 18th International Geological Congress as definitive for the Several other assemblages are included for the sake Pliocene-Pleistocene boundary. of completeness: the early Pleistocene Dixoii local The successive phases in the history of North American fauna from south-central Kansas and the Sand Draw land mammals which have been called the Blancan, Irving- local fauna from north-central Nebraska; and the tonian, and Rancholabrean ages can be correlated with the middle Pliocene Buis Ranch local fauna from north Pliocene and Pleistocene sequence of southwestern Kansas, and thus indirectly with the glacial-interglacial intervals of western Oklahoma. In ecologic and stratigraphie dis the Mississippi Valley. The Blancan is late Pliocene and cussion of the faunas both vertebrate and invertebrate early Pleistocene. The Blancan-Irvingtonian boundary is evidence is considered. Thus although the systematic LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS paleontologic description is limited to the mollusks, ing funds for sorting fossils and by contributing the faunal interpretations are synthetic. advice and criticism. Earlier work on the various faunas is summarized METHODS under the description of each. The history of this COLLECTING METHODS study may be noted here, however. As a member of a University of Michigan Museum of Paleontology The fossil mollusks reported here were collected field party under Hibbard in the summer of 1950 I almost entirely by the washing technique described had opportunity to see at firsthand the process of by Hibbard (1949c). Poorly consolidated or uncon- mass screen-washing of fossils, to learn the advan solidated sediments with surface exposures of fossils tages and disadvantages of the method, and to study were collected in as large pieces as practicable. After the local stratigraphy of southwestern Kansas and thorough drying (especially important with clay or western Oklahoma. In the summer of 1953,1 collected marl sediments) the matrix was washed through a Pleistocene and Recent mollusks and stratigraphic 16-mesh screen. It was found desirable to station data in northern Nebraska for 2 weeks and spent someone with a finer screen downstream from the another week in southwestern Kansas to see newly washers to catch small fossils which passed through discovered faunal sites. Detailed examination of the the 16-mesh screen and floated away. The small fossils fossils was made at the University of Michigan in which did not float were lost. Use of a smaller mesh 1952-53, University of California from 1953-55, and would insure recovery of all the fossils, but this slows the U.S. National Museum in 1955-56. washing and also hinders sorting by leaving more matrix in the washers. ACKNOWLEDGMENTS Adult shells of only 3 of the species considered here will pass through the 16-mesh screen: CarycJiium All landowners freely gave permission to work and exiguum, Gastrocopta Jiolzingeri, and Vertigo milium. collect on their property. For this aid thanks are due Oscar Booth, Orville Galbraith, Ray Keim, Frank In addition the shells of other species of Gfastrocopta, such as G. cristata and G. procera, may pass through Law, and Loyd Schnabel, of Ainsworth, Nebr.; H. C. Bender, Plains, Kans.; Arthur D. Sanders and the screens if the shells are slightly smaller than the mean or if there are bent wires in the screens which Jack T. Sanders, Meade, Kans.; Mrs. Gladys Dixon, create larger gaps than usual. Not all these smaller Spivey, Kans.; and Richard Dixon, Cleveland, Kans. shells will go through the screens, however, because The line drawings of plate 1 are by Miss Joan Sischo, University of California Museum of Paleon only the minor axis is less than the mesh size. Shells tology. which are kept flat on the bottom or sides of the E. S. Richardson, Jr., Chicago Natural History washers will tend to stay in them, while those which are agitated and turn perpendicular to the screen will Museum, C. W. Collinson, Illinois Geological Survey, and Henry van der Schalie, University of Michigan pass through. Museum of Zoology, loaned material for examination. In rewashing the 16-mesh concentrate in the labora H. B. Herrington, Keene, Ontario, Canada, identi tory, it was found that only CarycJiium and Gastro- fied the Sphaeriidae and gave ecologic interpretations copta Jiolzingeri passed through a 20-mesh screen. of them. Probably, therefore, the relative abundances for these The systematic treatment of several groups of ga species are low. stropods has benefited from discussions with P. F. The advantage of washing matrix is that an essen Basch and J. B. Burch, University of Michigan Mu tially complete representation of the fossil fauna is obtained. By this method Hibbard (1949c) has ob seum of Zoology, J. P. E. Morrison, U.S. National tained several faunas including amphibians, reptiles, Museum, and H. E. Walter, Liberian Institute of birds, mammals, mollusks (including snail eggs) and Tropical Medicine, Harbel, Liberia. ostracodes, as well as the zygospores of CJiara. D. E. Savage, University of California Museum In collecting it was found that the larger shells of Paleontology, contributed advice and stimulating were particularly fragile. This is partly because they discussion of the principles of correlation, strati are relatively weaker than small shells, and partly graphy, and mammalian paleontology. because they are more likely to have become cracked Most of all I am indebted to Claude W. Hibbard, or broken in the ground or in the removal of blocks University of Michigan Muecum of Paleontology. He of matrix. Accordingly, surface exposures were usu introduced me to the late Cenozoic sequence of south ally examined for the larger shells before collecting western Kansas and northwestern Oklahoma in 1950, matrix for washing. Large shells found in the wash and since that time has collaborated closely by secur ers were removed as soon as possible. METHODS
The amount of matrix needed to give a fair sample Ogallala to the Vanhem formation, each grade from of the fossils is large, probably at least 20 sacks (about gravel and coarse sand at the base to finer sediment 2,500 Ibs), as suggested by Hibbard (1949c, p. 11). at the top. The coarse basal gravels differ from one The completeness of representation may be increased unit to another, however, not only in composition but by sampling from several spots in the fossiliferous in grain size (Byrne and McLaughlin, 1948, p. 80), locality. Table 1 shows relative abundance of the although the finer sediments are similar. These differ species in University of Michigan collections from the ences suggest not only changes in stream carrying Saw Rock Canyon locality, with the forms reported power within a depositional interval, but also differ by Frye and Leonard (1952) added for comparison. ences in maximum carrying power between these in It should be noted that all collections were made in tervals. The early Pleistocene coarse sand and pebble the same way by C. W. Hibbard and that as nearly gravel units [Angell member of the Ballard forma as possible the same spot was sampled. The combined tion, and the Stump Arroyo member of the Crooked 1950-52 collections naturally have more material and Creek formation] are coarser than the sand and grit more species than those of the 1947 collection. The in the base of the Rexroad formation and include rocks chief conclusion is that no single collection contains of Rocky Mountain provenance. Such records of trans all species present, and that a large amount of matrix port of more and coarser material than is being moved should be washed. by the Cimarron River today suggest greater pre cipitation in the Rocky Mountains and probably also TABLE 1. Abundance or occurrence of gastropods in collections in the Plains. of Saw Rock Canyon local fauna The origin of caliche has been studied by Brown [A, abundant (>250); C, common (41-250); S, scarce (6-40); R, rare (1-5); X, occurrence] (1956). He concluded that in the northeastern Llano Estacado caliche recorded lime enrichment of the C- University of Reported Michigan collec by Frye horizon of pedocal soils in an eolian aggrading profile. tions and Leonard Such lime enrichment occurs in soils of both subhumid Species (1952, p. 151, and semiarid environments. Caliche is left behind 1947 1950,1952 Seward County by evaporation, therefore a warmer climate will favor locality) such precipitation but is not necessary. The more R frequent occurrence and more conspicuous develop S A X S X ment of caliche in the southern than in the northern R X S C X High Plains further strengthens such an idea. Fol C A X lowing is part of Brown's summary concerning the c A X c A X origin of caliche (Brown, 1956, p. 14) : s A X s A S X The mechanics of formation appear to be as follows: The X wind and rain bring in all the materials that form the soil R S X R S and its associated caliche and deposit these materials on the X s X soil surface. The CaCO3 and silica gel are leached downward R R A X by percokiting rainwater and deposited as a subsurface X evaporite, largely as molds around roots and organisms that R R C X separated the clastic particles. This inception of lithiflcation R C X R may be interrupted and the deposits destroyed by descending R S X water following subsequent heavy rains, but eventually they R will become permanent.
ENVIRONMENTAL INTERPRETATIONS OF ENVIRONMENTAL INTERPRETATIONS OF FOSSIL SEDIMENTS MOLLUSKS The present study is primarily paleontologic and Since almost the first days of paleontology, fossil stratigraphic, but two probable inferences from a mollusks (especially those of the Cenozoic) have been study of the sediments supplement the fossil evidence. used as a basis for inferring the conditions under These interpretations are that gravel deposits in which rocks were laid down. The early discovery southwestern Kansas imply a greater volume of stream that the distribution patterns of many living animals flow at the times when they were laid down and that and plants changed radically in the Pleistocene con caliche indicates a semiarid climate more probably tributed greatly to the theory of glaciation. The uni- mesothermal than microthermal. formitarian assumption has been made that the habitat The six Pliocene and Pleistocene formations of preferences of species have been the same throughout southwestern Kansas and western Oklahoma, from the their span of existence from then until now. Pond LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS snails of today lived in ponds of the Pleistocene; clams is a searching study of the subject which unfortunately of gravelly stream beds have always preferred these seems to have passed unnoticed by American paleon situations. This basis for interpreting the local habi tologists. The paper deals with German postglacial tat of fossil shells remains unquestioned and valuable.1 nonmarine mollusks, and attempts to answer the The story is otherwise with the inferences of past question: "Have climatic changes taken place between climates, however. Workers have disagreed as to how the Pleistocene and the present, and in what direction specifically one can interpret former climates and too have they occurred?-' (p. 73). little attention has been given to the large number Geyer's conclusions are that the climate suggested of factors which make up climate. by the fossils as a whole was moist and oceanic, for there are occurrences in central Germany of species PREVIOUS WORK now living only closer to ocean coasts. Furthermore, there are today associations of species with different Many earlier attempts at paleoecologic interpreta climatic requirements, but these associations are now tions of Pleistocene fossils and deposits have been of found in western rather than in central Europe. In dubious value, or so generalized as to be of little help general, however, a fossil assemblage can give only a to geologists and stratigraphers. This circumstance picture of local conditions of environment humidity, has been the result of uncertain stratigraphic position of fossils, inadequate data on pre-Wisconsin physical vegetation, soil, and microclimate which have noth features, and the fact that so many of the known ing to do directly with climate. Geyer's ideas are fossils represent animals of little paleoecologic value discussed in more detail later in this section. horses, bison, camels, and wide-ranging snails. Prob Baker's (1937) summary of Pleistocene nonmarine mollusks is valuable chiefly for the section on habitat ably for these reasons Flint (1947, p. 52-1) was led to the pessimistic conclusion that "* * * the ecologic interpretation. That dealing with climatic inference interpretations placed on most of the plants and ani sums up Baker's general conclusions, but lacks an ade mals in the record manifestly can never be more than quate treatment of the method involved. "Mollusca approximate at best. In consequence few major infer as indicators of time"" overrates the stratigraphic util ences as to Pleistocene climates rest on fossil evidence ity of the group. Baker recognized as species and alone. The chief value of fossil data is in strengthen subspecies shell forms which are only ecological var ing inferences drawn from evidence of quite other iants, and relied too heavily on the completeness of kinds". the geological record. Flint's conclusion is valid for most previous work Eiseley (1937), like Geyer, dealt with the applica on the Pleistocene. More detailed work on strati tion of mollusks to archeology. He discussed problems graphy and fossils, particularly in nonglaciated re of a smaller scale than those of this report, problems gions of now semiarid climates, will almost surely which arise late in the Pleistocene. yield significant information on climatic changes unob PRINCIPLES OF PALEOECOLOGIC INTERPRETATION tainable from physical features. The study and joint use of various animal groups, and collection of ade It should again be emphasized that paleontologists quate samples of material, is a prerequisite for these make an assumption that species have been influenced results, however. Reliance on only one group, such in former periods by the same limiting factors and as mollusks or mammals, cannot help but give an to the same degree as they are now. Species certainly incomplete and hence distorted picture. could have changed in this respect, and one must Only four general reviews of the ecological applica always be aware of the possibility. In the late Ceno- tions of Pleistocene nonmarine mollusks are known zoic molluscan faunas studied in this paper, evidence to me those of Shimek (1913), Geyer (1922), Baker for such changes is rare and doubtful. (1937), and Eiseley (1937). The generally accepted principles used in the inter Shimek's paper is useful for information on the pretation of late Cenozoic fossil mollusks may be depositional environment of loess, and especially for summed up as follows: Interpretations based on fossil comparison of fossil assemblages and drift shells from shells are governed by the present-day distribution modern lakes and streams. Subsequent information of the various species, and their association with each has outmoded his conclusions on stratigraphic and other and with various kinds of habitats and climates. climatic interpretations, however. Fossil shells similar to those of living forms are as Geyer's (1922) essay "Die Quartarmollusken und sumed to represent the same species. Furthermore, die Klimafrage" (Quaternary mollusks and climate) these species are believed to have been subject to the
1 For summaries see Baker (1937) and Yen (1951). same limiting: factors of distribution then as now. METHODS
DIFFICULTIES OF PALEOECOLOGIC INTERPRETATION bounds its habitat on the earth. Thus the reproduction and hence the continuity, indeed the physiognomy of whole floras Attempts to apply these principles encounter several (and faunas) depends on water. But it also decides, in an difficulties: Some groups of mollusks are not identi infinite number of cases, the significance of a species within fiable by shell alone; recognition of limiting factors an association, its dispersal ability in a region and therefore is most uncertain; habitat and distributional data are the limits of its natural distribution.' Land snails are 'moist air animals.' Their life processes go on under the direct poor for many species; taxonomy of living nonmarine influence of water. Winter temperatures are unimportant as mollusks is unsatisfactory for many groups. limiting factors; the snails hibernate in the soil like plants. During the summer warmth, however, there are interruptions SPECIFIC IDENTIFICATION OF SHELLS by drought. Whatever the sun and clouds grant to the earth does not act In most nonmarine mollusks the shell is the basis directly on the biosphere (perhaps aside from light, which for specific identification, although not for classifica affects the color of mollusks), but only through the medium tion of higher categories. In some families, such as of the soil the local habitat. It is not the amount of rain the Pupillidae, the shell is probably a much better which falls from the clouds that the organisms receive. They guide than are anatomical data. In other groups, benefit from only as much as the soil can hold and pass on. Likewise snails live not by the radiant heat of the sun, which however, such as the Zonitidae, Succineidae, and Hy- they quickly avoid in most cases, but by the warmth which drobiidae, shells are of distinctly secondary value the soil has taken up and transmits to them. Climate is the and may not be characteristic of species or even genera. source of energy for life; soil is the transmitter, storage 'bat In the late Cenozoic faunas of the High Plains the tery, and transformer. Succineidae are the only common group with this In studying the climate with the help of mollusks, then, one must turn primarily to the soil on which they lived. Soil, disadvantage. however, is just as complex a quantity as climate. Further more, these two factors, climate and soil, interact, displace RECOGNITION OF LIMITING FACTORS and supplement each other reciprocally, so that their influence on the biota is very difficult and often impossible to analyze Unfortunately there is no experimental evidence and determine separately. How much warmth and moisture determining limiting factors of distribution. Infer can tJie soil absorb for storage and transmission? That is ences as to what they may be must come from knowl the first question for all climatic research. In this way all edge of the geographic distribution of the species and other influences are considered: geographic position (latitude, from collecting experience. Isolated occurrences may elevation, and distance from the sea), formation, condition, also be especially valuable in giving clues to such sig and structure of the soil, angle of inclination and direction (exposure and insolation), irrigation, precipitation, humidity, nificant factors. wind, vegetation, cultural stage of the landscape and most re Virtually all previous workers concerned with fac cent geological history. In summary, the ecological bases of tors determining distributions of nonmarine mollusks molluscan life must be determined as far as circumstances within an accessible area have seen them in part or all permit. They may be divided into local environmental and of the interacting complex of soil, climate, and vegeta climatic factors. Only if the habitat is known can the in tion. Other factors may be significant, but there is fluence of climate be sought. no good evidence for their operation as yet. The mediating influence of the soil bars us, he said, Geyer (1922) contended that the distribution of from being able to determine much about climate. nonmarine mollusks is governed primarily by mois What we see is the resultant of many factors working ture. Occurrence and abundance of species are deter together. mined by the modifying effects 011 climate of local The local habitat demands adaptation. Stable species join environment, topography and drainage. He stated 2 it through selection, the variable ones by modification to given (P. ra-ri): circumstances. The adaptation influences the faunal com position. Thus each locality will give its assemblage a special In attempts at solving climatic problems (my own not ex- aspect according to the mixture of species and ecological cepted), the concept of climate was regarded one sidedly. forms. From the aspect of a fossil assemblage the significant It was essentially the same as temperature. Climate, how local conditions can be determined which have given this char ever, is a multi-faceted and complex quantity. Warmth is acteristic. They hare nothing to do directly ivith the climate. certainly a necessity for all life, bnt only one aspect of climate. (Geyer. 1922, p. 88.) It affects the worldwide distribution of mollusks, but in our homeland f Germany] mollusks depend more obviously on Geyer"s conclusions from the German nomiiarine water than on temperature. What Diels said about plants mollusks probably hold true for parts of North Amer may be applied to mollusks without restriction: 'Water de ica, and for many mollusks. But there are many dif termines the organism's possibility of existence. It directs its ferent patterns of distribution among the species growth and is the most important factor which influences and studied in this report, some of which seem controlled
1 Translated from the original German. by temperature extremes, others by soil and available 6 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS moisture. These patterns may be grouped most broadly Valvata tricarinata (Say) into five categories, with subdivisions for some. lewisi Currier Stagnicola palustris (Muller) The first category includes widespread species found caperuta (Say) over all or nearly all of North America; they are as Gyraulus circumstriatus (Tryon) follows: deflectus (Say) Sphaerium striatinutn (Lamarck) Armiger crista (Linnaeus) Promenetus ea-acuous (Say) lacustre (Muller) Pisidimn compression Prime unibilicatellus (Cockerell) casertanum (Poll) Physa sJcinneri Taylor nitidum Jenyns gyrina Say Apleaca hypnorum (Linnaeus) Fossaria obrussa Say dalli Baker Vallonia pulchella (Muller) Gyraulus parvus (Say) Cionella- lubrica* (Muller) Discus cronkhitei (Newcomb) Helisoma anceps (Menke) Vertigo ovata Say Nesovitrea elcctrina (Gould) Euconulus fulvtis (Muller) A modified northern distribution pattern may be Hawaiia minuscula (Binney) found in Vallonia gracilicosta of the Rocky Moun Zonitoides arboreus (Say) tains, northern Great Plains, and southwestern United The second cateory includes eastern species found States. commonly over North America east of the Rocky The fourth category is southern. The few species Mountains. Some range barely west of the Mississippi with southern distributions show considerable variation River, others are common on the Plains. in their ranges and are as follows: Six species of this second category are character Widespread throughout the southern United States: istically northeastern, found in the general area of Gastrocopta pellucida hordeacella (Pilsbry) New England and the Great Lakes. They are as Vallonia perspectiva Sterki follows: Southwestern, south-central, central, and southeast Sphaerium sulcatum (Lamarck) ern United States: Bulinmea megasoma (Say) Ferrissia rivularis (Say) Gastrocopta procera (Gould) parallela (Haldeman) Helicodiscus singleyanus (Pilsbry) Stagnicola exllis (Lea) Southwestern and south-central United States: reflexa (Say) Stagnicola bulimoides techella Haldeman The other species are more generally distributed: South-central and central United States: Sphaerium transversum (Say) partumeium (Say) Gastrocopta cristata (Pilsbry and Vanatta) securis (Prime) Erratic or poorly known. Marstonia decepta (Baker) Carychium exiguum (Say) Acroloxus coloradensis (Henderson) Ferrissia meeMana (Stimpson) Helisoma trivolvis (Say) Physa anatina Lea Planorbula armigera (Say) Pupoides inornatus Vanatta StroMlops labyrinthica (Say) Oxlyoma retusa (Lea) Gastrocopta armifera (Say) holsingeri (Sterki) The species with eastern distribution range onto tappaniana (Adams) the Plains varying distances. Some, such as Helico Pupoides alMlabris (Adams) discus parallelus, may be found in the High Plains Vertigo milium (Gould) where there are scattered patches of trees. Others, Vallonia parvula Sterki like Strobilops, are narrowly restricted to areas with Retinella wheatleyi (Bland) humus and dead wood. The distributions of these rhoadsi (Pilsbry) species are probably governed by available moisture, Stenotrema leai (Binney) soil, and frequency of drought. This is essentially The third category is made up of northern species Geyer's view. The species of northern distribution, found over northern North America. The southern however, can hardly be so governed, for otherwise they boundary of distribution is generally east-west, with would range south in the humid and geographically an extension southward in the Rocky Mountains. accessible Mississippi Valley or Eastern United States Some species of this group are Holarctic. They are generally. For this reason, as well as for others men as follows: tioned below, summer extremes of temperature prob- METHODS
ably govern the southern limits of these forms. By There is only one known joint occurrence of Cionella the same token, it would seem that species of southern lubrica and Gastrocopta pellucida hordeacella, at Cape distribution (such as Gastrocopta, pellucida hordea- May, N. J., and perhaps also in the Rexroad local ceUa) are prevented from going north by winter ex fauna (fossil) of southwestern Kansas. tremes of temperature. Another northern species with a Coastal Plain range This thesis that the species of northern and southern extension southward is Promenetus exacuous. distributions are limited by temperature extremes re Several isolated occurrences of species further sug ceives support from three other sources: ranges in gest temperature extremes as range-limiting factors. mountains, ranges in the Coastal Plain, and relict or These relict occurrences are of fresh-water species well isolated occurrences. south of their present northern distribution. The northern species (with the possible exception In Meade County, southwestern Kansas, the constant of Valvata) have southern extensions of range in the temperature and summer-cool waters of an artesian Rocky Mountains. The absence of such a pattern for spring provide a habitat suitable for Promenetus exa Valvata could be explained by rarity of the necessary cuous (Leonard, A. E., 1943, p. 235). The nearest permanent quiet water habitat. The fact that other occurrences of the species are in the Rocky Mountains aquatic species do not have such a pattern in the to the west and in northern Nebraska to the north. Appalachian Mountains can also be readily explained In Cherry County, Nebr., a similar cool spring sup by the scarcity of pond habitats. ported Valvata tricarinata and Gyraulus deflectus On the Atlantic Coastal Plain several species show (this paper). Other permanent water bodies of equal narrow range extensions. This Coastal Plain exten depth were examined without finding these species. sion of ranges is to be expected if temperature were Probably they are able to live here because the water acting as a distributional barrier. Elsewhere hot sum is warmed little if at all during summer hot spells. mers or cold winters keep these species apart, but In the High Plains these species may not live in the along the coast the ocean acts as a moderating influence United States except in unusual habitats such as this and the ranges overlap in a narrow belt. Geyer's one. Their nearest occurrences are to the northeast, interpretation of such patterns might have been that in eastern South Dakota. greater moisture was responsible, but taking the ranges For the various reasons given above, therefore, it as wholes into consideration it seems that the more seems that temperature extremes are factors limiting equable temperatures of an oceanic climate are signi the ranges of species with generally northern or south ficant rather than any possibly greater moisture. On ern distributions. Many other species, however, are the west side of a continent, as in Germany, humidity probably more influenced by a complex interaction of may change more obviously than temperature extremes. soil, vegetation, and climate. On the Atlantic Coastal Plain of the United States At least two species have distributions which do not temperature extremes are more apparent than any other seem controlled by any of these factors: Bulimnea factor in limiting ranges of these northern and south megasoma and Acroloxus coloradensis. Their ranges ern species. are not well known, but enough collecting has been Alexander (1952, p. 54) stated: done to suggest they are erratic. Why these species, A striking feature of the New Jersey land snail fauna is the formerly so widespread on the Great Plains, are now so presence of typically northern species living in association restricted is unknown. Perhaps they are largely anal with typically southern species. This anomalous combination ogous to Anisus pattersoni, which was also abundant of species can be explained by the fact that New Jersey is on the Plains during the Pleistocene but apparently located on the central part of the Atlantic coast where some northern species approach the southern limit of their distri became extinct in North America during the Wisconsin bution and some southern species reach the northern limit of their distribution. Certain species belonging to these diverse Inasmuch as these two Recent species (Bulimnea elements can be found together only in this state. megasoma and Acroloxus coloradensis) seem signifi Northern species extending southward in a narrow cantly affected by unknown influences, it is unwise to coastal strip are Pupilla muscorum and Cionella lu- use them in paleoclimatic interpretations. brica. A southern species reaching its northern limit in New Jersey is Gastrocopta pellucida hordeacella. INADEQUACY OF HABITAT AND DISTEIBUTIONAL Concerning the distribution of this last form, Pilsbry DATA (1948, p. 914) wrote: Throughout this paper only general and vague The northward extension along the Atlantic coast is very narrow, discontinuous so far as known, and perhaps mainly conclusions about local habitat and climate have been confined to the coastal islands. possible. This reflects the lack of knowledge concern- LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS ing ecology and distribution of many species. Habitat- nearby, and some Eecent species are unknown in the data are lacking or so scanty for a number of species fossil fauna. Reliability of conclusions depends upon that they have been grouped under the "uncertain" the thoroughness of collecting, and one must not read category in charts showing faunal habitat indications. too much into a poorly sampled fauna. If the Recent Further collecting, especially on the High Plains, will fauna near the fossil locality is reasonably well sam enable considerably more detailed climatic interpre pled, then the presence of fossil species living only tations than are possible now. in other areas is significant. Degree of significance Stratigraphic ranges of species also reflect the inade will vary with the accuracy to which the distributions quate amount of collection. Further work will of of the locally extinct species are known. Inasmuch course considerably advance such knowledge, particu as a fossil assemblage is usually known from one or a larly in the fluviatile groups such as Viviparidae, very few localities, the absence from it of certain Pleuroceridae, and Unionidae. At the present time in species is much less noteworthy than absence from the Great Plains, Oligocene and middle Pliocene mol- the better known Recent fauna. lusks are essentially unknown, Miocene mollusks en Comparison with a similar Recent fauna. Many tirely unknown, and lower Pliocene forms known only fossil assemblages which differ from the fauna living from the Laverne local fauna. locally are like a living assemblage elsewhere. The Extensions of the known geological range of Eecent reasonable inference can then be made that the en species have been and will be common, but it is per vironment of the fossil site was formerly like that tinent to point out also that several species described where the species are now living. This same principle as fossils have been subsequently found living. Hender- may be applied more broadly in the case of earlier sonia occulta, Discus macclintocki, and now PJiysa Tertiary or Mesozoic faunas, and comparison made skinneri were all originally thought to be extinct Pleis between genera or pairs of closely related species. This tocene species, but have later been collected alive. approach is not strictly applicable to most of the as semblages studied in this report, howrever, for there UNSATISFACTORY TAXONOMY are many fossil associations of species not living Many, perhaps most of the species dealt with in this together today. paper are well known so far as their classification and One such association, which occurs in several of the distinguishing characters are known. In many other Blancan faunas, is that of Stagnicola ca.perata and parts of North America, however, the percentage of Promenetus umbilicatellus, both northern species, with satisfactorily classified species is much less. In the the southern Stagnicola, bulimoides techella. Probably High Plains the poorly known groups are chiefly only a moderate reduction in number and duration Planorbidae, Succiiieidae, Physa,, and Fossaria* but of summer hot spells would permit the northern forms other areas can add more. Pleistocene and late Ter to move south. The association is not inexplicable, tiary paleontology is nearly always dependent upon a but does show that strict comparison with living as knowledge of Eecent species, and hence must lag semblages may be inadequate. where large parts of the living fauna are poorly An even more striking example is the association known. of Physa skinneri and Stagnicola bulimoides techella METHODS OF FATJNAL INTERPRETATION in the Sanders local fauna. The limits of distribution in the Great Plains of these two species are hundreds The preceding summaries of the principles and of miles apart. Physa skinneri is known only as far difficulties of paleoecologic interpretation show the south as southern Manitoba; Sta.gnicola bulimoides basis on which individual species can be interpreted. techella barely reaches Nebraska, where it is known Two other complementary approaches are also useful: from a single locality in the southernmost part of the comparison of the fossil assemblage with the living state. Elsewhere in North America, the two species fauna of the region around the fossil locality and comparison of the fossil assemblage with a similar have been found closer together. In Utah Physa skin living fauna. In these methods the fauna as a whole neri is known from the Hint a Mountains, in the north is considered, and the ever-present danger of over eastern part of the state; Stagnicola bulimoides te emphasizing single species can be avoided. chella is known from central Utah, but at much lower Comparison, with Recent fauna nearby. In compar elevations. Although the ranges of these species are ing the fossil and Recent assemblages from the same- closer in Utah than in the Great Plains, the contrast area, one inevitably uses negative evidence to some in habitats is still as great. The Sanders local fauna, extent. Usually some fossil species are not living containing both these now widely separated species, METHODS 9 cannot be compared closely with modern molluscan The following points summarize these differences associations. between mollusks and mammals as shown by the late AGE DETERMINATION Pliocene and early Pleistocene faunas of the High Two methods of dating faunas by fossil evidence Plains considered in this paper. can be used in the late Cenozoic. The more general Mollusks evolve more slowly than mammals. From approach is purely faunal, based on restricted or over a stratigraphic standpoint they are therefore less lapping ranges. The second method is less widely valuable, but are more useful for paleoecologic infer applicable, but of perhaps greater value for the Plei ences. In the special case of the Pleistocene epoch, stocene. By inference of former climatic conditions, when major f aunal shifts took place, some mollusks a fauna may be placed with some precision in a known may be especially valuable in refined dating within glacial-interglacial sequence. a known climatic sequence because of their sensitivity One fundamental premise should also be stated here: to climatic change. all lines of available evidence bearing on the age of a Mollusks are smaller, commoner, and occur in denser fauna must be considered. This principle would seem populations than do mammals. For this reason they too obvious to need pointing out, but nevertheless has are more abundant and more easily collected as fossils. sometimes been ignored. The shells of mollusks usually provide the chief basis for specific identifications, although not for gen FAUNAL DATING eric or familial classification. Thus the fossils in their usual state have nearly all the characters used in In using the method of restricted or overlapping determination of living forms. Mammal skeletons ranges, animals with short vertical but wide geo are readily disarticulated and broken, and many indi graphic ranges are of course most valuable. In the vidual bones may be useless for identification of present state of paleontological knowledge, this fact means that mammals are the most useful group. Mol species. Mollusks are restricted in their powers of dispersal lusks as a whole are less reliable in the faunas under and generally show endemism in major physiographic consideration, but their great abundance means they regions. The present-day f aunal regions east and west are potentially helpful. of the Rocky Mountains, for example, appear to have Associated mammals and mollusks have not yet been in existence during the Pliocene also. Correla yielded contradictory conclusions, although often one tion between these two regions is thus impracticable group gives evidence which the other lacks. An ex on the basis of specific identity of mollusks. Most ample of the complementary use of mollusks and mam mammals, more vagile and with less temporal range, mals is as follows: The Saw Rock Canyon and offer the most precise means of correlation. Rexroad local faunas can scarcely be distinguished by Dating from climatic inference. A second method mollusks, but the mammals show a considerable differ of dating Pleistocene fossil assemblages is based on ence. The mollusks indicate an ecological similarity their ecologic interpretation. This method is poten between the two faunas, while the mammals demon tially more precise than other paleontologic means, strate time difference. Southwestern Kansas prob but depends upon an established sequence into which ably had an essentially uniform climate throughout an assemblage may be fitted. This method may also the late Pliocene. The mollusks did not change visibly, be more versatile as well as more precise. For example, while the mammals did. once a sequence of climatic changes has been verified Lyell (1833, p. 47-49) was perhaps the first to rec for a given region, any group of fossils from which ognize the principal differences in stratigraphic utility of mollusks and vertebrates. He stated: adequate environmental inferences can be drawn can be placed in this sequence. The late Pleistocene and Although the bones of mammalia in the tertiary strata, and those of reptiles in the secondary, afford us instruction of the Recent climatic history of western Europe is based on most interesting kind, yet the species are too few, and con fossil pollen and inferred vegetational changes; in fined to too small a number of localities, to be of great im theory an assemblage of amphibians, snails, or beetles portance in characterizing the minor subdivisions of geolog could be dated if their ecologic implications were ical formations. * * * We shall revert in the sequel to the curious fact, that in tracing back these series of tertiary de precise enough to permit placing them in the sequence. posits, many of the existing species of testacea accompany us In the Great Plains region there is no such estab after the disappearance of all the recent mammalia, as well lished sequence of Pleistocene climatic changes. Al as the fossil remains of living species of .several other classes. though a number of climatically significant faunas We even find the skeletons of extinct quadrupeds in deposits wherein all the land and freshwater shells are of recent have been described from southwestern Kansas and secies. northwestern Oklahoma by C. W. Hibbard and co- 508945 O 60 2 10 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS workers, the record is far from complete. The relative (Kansan Cudahy fauna, Illinoian Berends and Butler ages of the described assemblages are fairly well Spring local faunas; Wisconsin Jones local fauna). known, but documentation of a sequence of climatic 2. Early interglacial age. Climate warm-temperate, changes is hardly continuous. Plotting the curve of semiarid. Summers much as today; winters much Pleistocene climate in southwestern Kansas requires milder; precipitation about the same or slightly less. more points than are now available. (Yarmouth Borchers local fauna; Sangamon Cragin Furthermore, the exact correlation of the faunas of Quarry local fauna). southwestern Kansas and northwestern Oklahoma in 3. Middle interglacial age. Climate probably warm- terms of glacial events is unknown. Whether a certain temperate, semiarid. A massive, indurated caliche fauna represents a very late glacial age, when con layer about a foot thick formed during part of each tinental glaciers were in existence but waning, for of the three interglacial ages. example, is speculative. The known stratigraphic suc 4. Late interglacial age. Summers much cooler than cession of faunas and their ecologic interpretations now; winters very much warmer; precipitation greater permits only allocation to a given glacial or inter- in amount and effectiveness (Aftonian Sanders local glacial age. fauna; Sangamon Jinglebob local fauna). Although the record is incomplete, the amount of The Sand Draw and Dixon local faunas, discussed data is sufficient to suggest climatic trends, and certain elsewhere in this paper, do not fit this pattern well. similarities between parts of the Pleistocene record Probably their geographic location some distance from are striking. A working hypothesis is warranted, the other local faunas is partly, but not entirely re as a generalization about information now available sponsible. Perhaps a smaller Rocky Mountain rain and as an aid to future work. This hypothesis is that shadow, or a different climatic regime during the Ne- the glacial-interglacial climatic changes are cyclic. braskan age, explain their peculiarities. Cyclic climatic changes. A cyclic pattern of glacial- Eelation of the present day climate to this cycle interglacial climatic changes cannot be confirmed fully is uncertain. The living molluscan fauna is poorer from the record in southwestern Kansas. The con in species than any of the known fossil assemblages, firmation is partial, however, and all of the known and suggests that the present fauna is geologically data are consistent with the hypothesis that these unusual. If some phase of the hypothetical climatic changes were cyclic from the Nebraskan to Sangamon cycle is similar to the present climate, this phase is ages. Although progressive uplift of the Eocky Moun probably somewhere between the late glacial and early tains during the Pleistocene presumably affected the interglacial stages. climate, evidence of this is not clear. LOCAL FAUNAS Consideration of the physical and climatic events The term "local fauna" is used by vertebrate paleon of the Pleistocene in southwestern Kansas and north tologists to designate an assemblage of fossils from one western Oklahoma as a cyclic pattern was prompted locality, or from several closely grouped localities by Stokes' (1955) stimulating hypothesis of oceanic which are demonstrably stratigraphically equivalent or control of glacial-interglacial alterations. Consistent nearly so. It is a group of fossils local in both time evidence from many parts of the world and of many and space. Advantages of this term are as follows: different kinds is needed to confirm Stokes' hypothesis. It provides a designation for fossils which can be The evidence from Kansas and Oklahoma is scarcely independent of formation names. A faunal name taken definitive, even though significant. Nevertheless impli from a rock unit may become either inappropriate or misleading as the fossiliferous unit is subdivided or cations of the ocean-control hypothesis fit the known restricted. Exceptions to the general policy of distinct faunal and stratigraphic succession in southwestern faunal names are those where long usage makes the Kansas better than ideas about Pleistocene climates change to a distinct name undesirable (as Barstow deduced from other theories. local fauna, Barstow formation) and those where the The following sequence of glacial-interglacial rock unit is named later than the fauna (as Rexroad changes fits the available information from south local fauna, Rexroad formation). western Kansas and northwestern Oklahoma to a Such a locally restricted name does not imply a hypothetical cycle. The cycle is intended to apply doubtful correlation. A term applied to several widely only to this local area. separated assemblages runs the grave risk of major 1. Late glacial age. Summers much cooler than now; restriction or dismemberment as more data become winters no colder, perhaps slightly warmer; precipita available. Only rarely can more widespread localities tion little if at all greater but much more effective. be shown to be contemporaneous by special methods COMPARISON OF FAUNAS 11 such as the association of the Cudahy fauna with the of some Rexroad species may be due to climatic change petrographically distinct Pearlette ash bed. also. (See table 5.) The name per se implies no special age. Leonard Last appearances of species are most apparent in (1950) called the mollusks associated with the Pearl the Bender and Sanders local Kaunas. Cooler climates ette ash bed the "Yarmouthian molluscan fauna." With associated with formation and advance of the Nebras- changed age assignment Frye and Leonard (1952) kan and Kansan continental glaciers seem reasonable termed the same assemblage the "Kansan molluscan explanations for these. fauna." A Kansan age seems probable, but neverthe Indices of faunal similarities (table 4) reveal the less the later name is still open to the objection of following significant points: The Saw Rock Canyon, special age implication. Other disadvantages are that Red Corral, and Rexroad local faunas are markedly the name cannot include other groups of animals, such similar to one another; the Sand Draw, Dixon, and as mammals, and that the name does not indicate its Sanders local faunas are markedly similar to one actual geographic restriction to the central Great another; the Bender local fa ana is most similar to Plains region. Here the noncommittal term Cudahy the Rexroad and Sanders; the Pleistocene fauna most fauna seems clearly preferable. similar to the Rexroad is the Sanders. A widespread or thick formation may have a differ Apparent ranges and occurrences of individual spe ent time range in different areas. Assemblages grouped cies may be affected by sampling error, as well as by as a fauna from separate localities in such a formation local habitat differences and by regional causes. The may be far from contemporaneous. Use of a number differences considered significant, and possible causes, of local faunas would obviate a "fauna" which is are discussed below. gradually dismembered and restricted in age as more Vertigo ovata is known from the Clarendonian La- precise information becomes available. A specific verne local fauna, Hemphillian Buis Ranch local example of a unit containing faunas of markedly dif fauna, and from numerous Pleistocene faunas. In the ferent ages is the Rexroad formation of southwestern Rexroad and Saw Rock Canyo:i local faunas, however, Kansas. Three distinct assemblages from this unit are it is replaced by V. hibbardi. This may be an example recorded in the present paper. of temporary ecological replacement, for both species Elias, too, (1942, p. 134-136) discussed this question are of similar size and both probably had similar of independent faimal and floral names, arriving at habitats. A possible explanation is that V. hibbardi opposite conclusions. The reasons set forth above arrived in the region of southern Kansas (either by seem to me to outweigh any possible advantages of migration or evolution) at the end of the Hemphillian, Elias's proposed method of a unified rock-faunal ter largely eliminated V. ovata in the area, and was in minology. Van Houten, too (1944) has discussed the turn driven out by the first glaciation. V. ovata is a question of faunal nomenclature, concluding that a eurythermal species as shown by its wide range; geographic name independent of stratigraphic termi perhaps V. hibbardi was not. nology is convenient and desirable. Gastrocopta rexroadensis and Polygyra rexroadensis COMPARISON OF FAUNAS are not known from faunas later than the Rexroad and Bender but do not necessarily indicate a late Plio Table 2 shows the occurrence and range of the spe cene age. Neither species is common enough to be cies in eight faunas. These data are summarized in table 3. Indices of taxonomic similarity are given in reliable as a stratigrahic indicator. G. rexroadensis table 4. was not found in University of Michigan collections First appearances of species show two marked dis at localities from which it is reported. continuities : Saw Rock Canyon-Rexroad and Rexroad- Twenty-nine late Pliocene species are not known Sand Draw. The first of these is considered not from earlier deposits. This ms,y be entirely the result significant because it is almost surely in large part of incomplete collecting. or entirely a result of inadequate sampling. The Rex- Gastrocopta cJiauliodonta is known only from three road local fauna is known from several localities, the faunas which are dated Nebraskan or Aftonian on Saw Rock Canyon local fauna from a single locality independent grounds. Collecting in earlier and later with little diversity of habitats. faunas seems sufficiently adequate to establish its local The Rexroad-Sand Draw discontinuity is at least restriction to the lower Pleistocene, for it is a rela partly climatic. Many species first known from the tively common species. It may be a species which Sand Draw local fauna probably represent northern lived to the north in the late Pliocene and was forced species driven south by glacial advance. Extinction southward by glaciation. 12 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
TABLE 2. Stratigraphic occurrence of mollusks of some late Ceno zoic faunas in the High Plains [Occurrences outside the faunas listed are not necessarily in the High Plains. Occurrence of specifically unidentified forms and uncertain occurrences are omitted]
Stratigraphic occurrence
Species Pre-Buis Buis Saw Red Sand Ranch Ranch Rock Corral Rexroad Bender Draw Dixon Sanders Late local local Canyon local local local local local local Pleisto Recent fauna fauna local fauna fauna fauna fauna fauna fauna cene fauna
X Promenetus kansasensis (Baker) _ X X X X X XXXXXXXXXXX X X X X Hawaiia minuscula (Binney) XXXXXXXXXXXXXXXXX X X X X X XXXXXXXXXXXXXXXXX Bulimnea megasoma (Say) XXXXXXXXXX XXXXXXXXXXXXXXX X X X X compressum Prime nitidum Jenyns. _ Promenetus umbilicatellus (Cockerell) X X X X X X X X X inornatus Vanatta _ X X Sphaerium striatinum, (Lamarck) Vallonia gracilicosta Reinhardt X Valvata tricarinata (Say) X Vertigo ovata Say _ X X xxxxxx X Zonitoides arboreus (Say) _ Strobilops sparsicostata Baker_ X X X Ferrissia rivularis (Say) X X X X X X Stagnicola caperata (Say) X X X X X X Gastrocopta rexroadensis Franzen and Leonard X xxxxxxxxxxxxxx Vertigo hibbardi Baker Gastrocopta franzenae Taylor X X Marstonia crybetes (Leonard) X Gastrocopta paracristata Franzen and Leonard xxxxxxxxxxxxxxxxx X X X Deroceras aenigma Leonard _ X X X X X Fossaria dalli (Baker) _ _ _ X X X X Gastrocopta holzingeri (Sterki) tappaniana (Adams) X X X X XXXXXXXXXXXXX xxxxxxxxxxxx Helicodiscus singleyanus (Pilsbry) X X X X Helisoma anceps (Menke) X X Marstonia decepta (Baker) Physa anatina Lea X X X X X X Stagnicola bulimoides techella (Haldeman) X X X X Strobilops labyrinthica (Say) _ _ _ Vallonia perspectiva Sterki X X X Vertigo milium (Gould) X X X X X Gastrocopta pellucida hordeacella (Pilsbry)- _ _ X xxxxxxxxxxx X Polygyra rexroadensis Taylor. X Carychium exiguum (Say) _ - X xxxx Cionella lubrica (Muller) xxxxxxx Ferrissia parallela (Haldeman) Nesovitrea electrina (Gould) X Retinella rhoadsi (Pilsbry) wheatleyi (Bland) Stagnicola exilis (Lea) X Gastrocopta scaevoscala Taylor X X cristata (Pilsbry and Vanatta) _ X X X X X X Vallonia parvula Sterki X X Gastrocopta chauliodonla Tavlor X X Acroloxus coloradensis (Henderson) X Aplexa hypnorum (Linnaeus) X Ferrissia meekiana (Stimpson) xxxxxxxxxxxxx X Fossaria obrussa (Say) Physa gyrina Say xxxx xxxxxxxxxxxxxxx xxxxxxxxxxxxxxxxxx skinneri Taylor X Planorbula armigera (Say) _ Sphaerium partumeium (Sav) sulcatum (Lamarck) transversum (Say) X Vallonia pulchella (Muller) Valvata lewisi Currier Gastrocopta procera (Gould) Helicodiscus parallelus (Say) xxxxxx Helisoma trivolvis (Say) Oxyloma retusa (Lea) Sphaerium lacustre (Muller) Stagnicola reflexa (Say) X COMPARISON OF FAUNAS 13 TABLE 3. Summary of data from table 2
Num Percent Percent Percent Num Percent Percent Percent Occurrence ber of re first last Occurrence ber of re first last species stricted appear appear species stricted appear appear ance ance ance ance
17 56 Buis Ranch local fauna.--.- ___ 5 0 60 0 Total species for all faunas _ 68 Saw Rock Canyon local fauna____ 24 0 71 0 Saw Rock Canyon, Rexroad, and Red Corral local fauna 16 0 6 0 37 5 70 35 0 23 6 Sand Draw and Dixon 52 0 37 4 17 0 18 12 54 2 35 q 38 0 34 0 Saw Rock Canyon, Red Corral, 4.c 0 13 7 40 10 73 10 20 0 0 10 Sand Draw, Dixon, Sanders, and 54 57 4 3Q 12
TABLE 4. Faunal similarities
[Calculated from data of table 2 by 100C, where C=number of species common to two faunas and Ni=number of species in smaller fauna]
Local faunas
Buis Saw Hock Red Corral Rexroad Bender Sand Draw Dixon Sanders Ranch Canyon
Buis Ranch (N = 5) _ - __ .______40 60 80 20 100 100 40 Saw Rock Canyon (N = 24)______40 81 92 71 58 67 70 Red Corral (N = 16)______.___-_.______60 81 100 50 69 69 56 Rexroad (N = 36) _ .___--____-__--______-_____ 80 92 100 83 49 60 80 Bender (N = 17)___.______. _ __.__ __ 20 71 50 83 65 71 77 Sand Draw (N = 38)-_-_-_-_-__,.______-______100 58 fiQ 49 65 82 85 Dixon (N = 45)------_-----_-__-----__-__- 100 67 69 60 71 82 85 Sanders (N = 20) ______40 70 56 80 77 85 85
TABLE 5. Distribution of thermally significant species in late Pliocene and early Pleistocene faunas of the High Plains
Local fauna Climate Species Buis Saw Red Sand Ranch Rock Corral Rexroad Bender Draw Dixon Sanders Canyon
Mesothermal: Winters relatively mild and Gastrocopta cristata (Pilsbry and X X X X short. Vanatta) . X X X X X X X X X X Stagnicola bulimoides techella (Halde- X X X X X man) . X X X X Summers mild__ X X X Vallonia pulchella (Miiller) _ X X X X Summers warm but not hot X X and dry. Promenctus umbilicatellus (Cockerell) _ X X X X X X X X X X X X X X 14 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS Nineteen species are first known in the earliest 1949) of the lower part of the Rexroad formation, Pleistocene. Perhaps all of these were living to the Seward County, Kans. Fishes, amphibians, reptiles, north during the late Pliocene. birds, mammals, and mollusks. The mammals indicate Temperature differences and similarities are shown an earliest Blancan (early late Pliocene) age. The by table 5. The Saw Eock Canyon, Eexroad, Bender, mollusks do not differentiate this fauna from the and Sanders local faunas are believed to have lived in Rexroad local fauna. Climate subhumid and meso- a climate with summers warm but not hot and dry; the thermal, with less extreme summer hot spells and Sand Draw and Dixon local faunas in one of mild slightly greater and more effective annual precipita summers. These climatic shifts probably represent con tion. Significant members: Dipoides wilsoni Hibbard, tinent-wide changes, for southwestern Kansas is well Ogmodontomys sawrockensis Hibbard, Baiomys saw- beyond the limits of glaciation. These changes are ten rockensis Hibbard, Osteoborus progressus Hibbard, tatively correlated with the glacial-interglacial se Vertigo hibbardi Baker, /Stagnicola bulimoides techella quence in the Mississippi Valley. Haldemaii, Promenetus wnbilicatellus (Cockerell). 3. Rexroad local fauna (Herrington and Taylor, SUMMARY OF FAUNAL SEQUENCE 1958; Hibbard, 1950; 1954b; 1956, p. 185; Oelrich, Figure 1 summarizes the sequence of some Pliocene 1957; Taylor, this paper; Tihen, 1955). In the upper and Pleistocene faunas in the Great Plains and their part of the Rexroad formation, below the massive stratigraphic position and age. In addition to the caliche, Meade and Seward counties, Kansas. Amphib middle Pliocene to early Pleistocene assemblages ians, reptiles, birds, mammals and mollusks. The described in detail elsewhere in this report, middle mammals indicate a considerably younger age than and late Pleistocene faunas of southwestern Kansas that of the Saw Rock Canyon local fauna, though still and northwestern Oklahoma, and other faunas men Pliocene. Climate subhumid and mesothermal, with tioned in the text are included. The following notes markedly milder winters than those today, summers present supplementary information, including refer lacking the present hot spells, and a somewhat greater ences to pertinent literature. Numbers correspond to annual rainfall. There is no evidence of a strati- those in figure 1. graphic break or of a climatic change between the The North American ages Hemphillian, Blancan, horizons of the Saw Rock Canyon and Rexroad local Irvingtonian, and Rancholabrean (Wood and others, faunas. Significant members: Dipoides rexroadensis 1941; Savage, 1951) have not been adopted for official Hibbard and Riggs, Ogmodontomys poaphagus Hib usage by the U.S. Geological Survey. bard, Bensonomys eliasi (Hibbard), Procastoroides 1. Buis Ranch local fauna (Herrington and Taylor, lanei (Hibbard), Plesippus simplicidens (Cope), Nan- 1958; Hibbard, 1954; Oelrich, 1957; Tihen, 1955). In nippus phlegon (Hay), primitive Stegomastodon, Tes- the upper part of the Ogallala formation, in Beaver tudo rexroadensis Oelrich, T. turgida Cope, Stagnicola County, Okla. Mammals, reptiles, amphibians, and bulimoides techella (Haldeman), $. exilis (Lea), Pro mollusks. Several of the mammals show close rela menetus umbilicatellus (Cockerell), Gastrocopta fran- tions with others in the Saw Rock Canyon local fauna, zenae Taylor, G. pellucida hordeacella (Pilsbry), G. in the lower part of the Rexroad formation. Occur rexroadensis Franzen and Leonard. rence in the fauna of a rhinoceros (unknown elsewhere 4. Bender local fauna (Taylor, this paper). In in North America from assemblages as young as the upper part of the Rexroad formation above the Blancan), and the stratigraphic position of the fossils massive caliche, in Meade and Seward Counties, Kans. in the Ogallala formation, are the suggestive but Mollusks only. The fauna is a little more like the somewhat arbitrary reasons for assigning this fauna Rexroad local fauna than the Sanders local fauna to the late Hemphillian rather than early Blancan. or other Pleistocene interglacial assemblages, and tends The few mollusks slightly differentiate the assemblage to favor a Pliocene age. Climate subhumid and meso from the Saw Rock Canyon and Rexroad local faunas, thermal. Significant members: Stagnicola bulimoides but how significantly is uncertain. Climate with sum techella (Haldeman), S. caperata (Say), Promenetus mers less hot and dry than those of the area today. umbilicatellus (Cockerell), Gastrocopta franzenae Significant members: Pliogeomys, Perognathus cf. P. Taylor, G. cf. G. cristata (Pilsbry and Vanatta), G. mclaugJilini Hibbard, rhinoceros, Stagnicola caperata scaevoscala Taylor. (Say), Vertigo ovata Say. 5. Unnamed assemblage (Hibbard, 1949, p. 67; 2. Saw Rock Canyon local fauna (Herrington and 1956; Savage, 1955). A few large vertebrates have Taylor, 1958; Hibbard, 1953, 1954, 1957; Taylor, this been collected from the Angell member of the Ballard paper; Tihen, 1955). In the XI member (of Hibbard, formation in Meade and Clark Counties, Kans.: Pie- PLIOCENE PLEISTOCENE SERIES
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Crooked Creek o Ballard formation TO format on Q) > 73 CO & -i O s ?; -^ O O 5T 3* 33 <£ ..-c 5' 5 * o*. Q> | CD C 3 »» > W) 'g 3 3 JTHWEST ^£ o S. o 3 > c g 5 CD D I ? S-o | ?5 * s §- a ? z H- {0 | v/W S 8- IS S- S ° 3 "*CD 1 i I?" o ! 1/ CO ° 3 x v< / m 3 CT iS o V z o'.°v';.' : ; "1 -.:; : 7 :[ r :;: ".;>: 1 lljli: Ijlj' !:: 'i'. : |''t /' ' ' !: : :i||!.: : 1 -n ;*; :jr. 7 :? V N 1 : : i i : - -'. . :{l ( 'o' '' '*' : 7 ~ :\ '" '.' < l' : r . - '. X V, ' ' ! 1 1 | ' I ! ' [ o- o/ ; | ; 1 : : > r: .'. : .' ' ' ' 1 ' 5 *1 -A -p . COQ) 1i i:. : : - : 1 ' :' ' .-'~ r'' tM 1: /) ' -T '-' ' ( ':.' o.''-'.' u : : | : J][J \ ' '." : r, j 1 : ; ; Z T ' < ' .' ' 1 '. ' ' ' ' ' - ~ 1 1 p ! ' I '.°. 01 ' '00° "^ CO ?'. c.- o F/:N ^ ° o.: :- | on : j M? ( (?.'o/°ff ;!s :c 'v' ' 33 : : ' ' ' : 33-' : " ' ' ] J ?^^-»f .". 0^7 ! M CO 3 .' : 0 C/) c p- tJ" »q <" ~\ ' O .::' ' : : : ' CD : "-'.-: f .-;.g ; ;(S : S-^ |'l- ( ('.;.°;; Z.JD ^~ : 33 T) ' 5- ; ; : ; : 5 - ? ; \ T 7 Q./ 3:--^\ ;°°' '^ j ^ CD z ra ?' = > 3" ~ 1 " : / . .°- ! ' C/5 : 3 ( ' o : ; ' ' '' ': ; ; ' S ' '' f -' : CD" / '°S ' ' : ^ v< P . i O O o M x « ~* ' - * ~ tfl II iS- 1? ( V- *^! ;I C ,: S-^r ' = : -: : j : : ! CL -_^l_: -'K ; p o- C1 '. '. . ; W ' : v °'o' ' OQ> \ ^-,'0 D* m ^ ^ ' 'T . : : [ "' '*' i ' '' ' ' i . * ^ : ' '^' "* i /'? '' : ' ; ^ ' ': o V? .' ' ' : (M : .?. i ;ij . ; i i -i^ ill \ I fK: ) V.;. ! i |l i'i ;^. °rf'. JH! ''J!! ':!! ;) ^ i ! m o co ^ ^co^ ocDCoSSjjc/j s ^ gS OKLAHOMAENT SEQUENCECLIMATIC I If II H! It (HI I! H !! it if il INFERENCE ANDBASISFOR S |.£ f.2- ? 1"^ &~ z ? ?l fl zs i'i I" ?1 ft 1 Is Sg 5 Is £5- |* o j^g i» s- o £» = ^-w g; i T T^?^ s-se? 7" g 3- T ^- &~& ~ N-^ slli' Q- 1 '
Ol vj "- . N) . M >0 IO N 10 -n i » ^ 00 m 0 f _ 3> m CD 5 DO j^ 33 CZ 33 Q. 2 O CD CO CD Q. -i Q) Q) N g r- O DO 3 Z, O CT io o ^ * D Q. & J CD o 3 | M ^ > 16 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS sip pus simpUcidens (Cope), Equus (Asinusl) sp., (1948) recognized the petrographic distinctness of Nannippus plilegon (Hay), Stegomastodon mirificus the Pearlette ash bed and listed the mollusks associated (Leidy), and camel. with it. This volcanic ash demonstrates the essential 6. Deer Park local fauna (Hibbard, 1956). In the contemporaneity of the associated fossils. The name lower part of the Missler member of the Ballard for Cudahy fauna is used in the broad sense for the entire mation, in Meade County, Kans. The mammals indi assemblage, for there seems no reason to use more cate Blaiican age, narrowed to Aftoniaii by indications than one name. Fishes, amphibians, reptiles, birds, of mild climate and by their stratigraphic position. mammals, and mollusks are known. Hibbard (1949b) From the Angell member of the Ballard formation, considered the Cudahy a glacial fauna of Kansan below the Missler member, Hibbard (1944) reported age, pointing out the boreal relations of the verte glacially striated pebbles. Significant members: Plio- brates. The snails Pupilla muscorum^ P. blandi, Ver lemmus antiquux Hibbard, Pliopotamys meadensis tigo gouldi, V. modesta, and others occur in the Great Hibbard, Procastoroides sweeti Barbour and Schultz, Plains today only far to the north. Their presence Nannippus plilegon (Hay), Plesippus simpUcidens in the Cudahy fauna as far south as Texas indicates (Cope), and large species of Testudo. radically cooler and moister summers and presumably 7. Sanders local fauna (Hibbard, 1956; Taylor, wetter, but not necessarily colder, winters. Significant this paper; Tihen, 1955). In the upper part of the members: Equus sp., Sorex, Neosorex lacustris Hib Missler member of the Ballard formation, above the bard, Microsorex pratensis Hibbard, Synaptomys massive caliche, in Meade County, Kans. Mollusks, (Mictomys] horeaUs (Kichardson), Microtus parope- amphibians, birds, and mammals. Stratigraphic posi rarius Hibbard, Pedomys llanensis (Hibbard), Pity- tion, ecologic interpretations, and faunal evidence indi mys meadensis Hibbard, Pupilla muscorum (Lin cate late Aftonian age. The fauna is distinct from naeus), Vertigo modesta (Say), Valvata tricarinata other Blaiican assemblages in its composition as well (Say). as climatic implications. Climate subhumid and meso- 10. Borchers local fauna (Hibbard, 1941b, 1949b; thermal, with more effective precipitation and sharply Tihen, 1955). In the Atwater member of the Crooked reduced summer temperatures. Significant members: Creek formation, above the Pearlette ash bed, in Meade Sigmodon, Zapus, Plesippus simpUcidens (Cope), County, Kans. Amphibians, reptiles, birds, and mam Nannippus phlegon (Hay), Gastrocopta chauliodonta mals. The small mammals indicate a climate as warm Taylor, G. paracristata Franzen and Leonard, Physa or warmer than that of present day southwestern skinneri Taylor, Stagnicola l>ulimoides techella (Hal- Kansas, and hence an interglacial age. Significant deman). members: Testudo, large species, Onychomys fossilis 8. Seger local fauna (Hibbard, 1951, 1953; Savage, Hibbard, Sigmodon hilli Hibbard, Synaptomys lan- 1955). In the Stump Arroyo member of the Crooked desi Hibbard. Creek formation, in Meade and Clark Counties, Kans. 11. Butler Spring local fauna (Herrington and Large mammals only. The few species known do not Taylor, 1958; Hibbard and Taylor, 1960; Smith, C. differentiate this fauna from earlier ones, but are dis L., 1958; Tihen, 1955). In the lower part of a section tinct from those of later assemblages. They have 110 included in the undifferentiated Sanborn group, in special climatic implications. Significant members: Meade County, Kans. Fishes, amphibians, reptiles, Plesippus. sp. cf. P. simplicidens (Cope), Nannippus mammals, and mollusks. Stratigraphic position and phlegon (Hay), Stegomastodon mirificus (Leidy). ecological interpretations indicate a late Illinoiaii age. 9. Cudahy fauna (Frye and Leonard, 1952, p. 158- Climate subhumid, differing from that of Meade 159; Herrington and Taylor, 1958; Hibbard, 1944, County today in being markedly cooler, the difference 1949b; Johnston and Savage, 1955; Leonard, 1950; Ti mostly in the summers. Significant members: Icta- hen, 1955). Includes the Tobin and Wilson Valley lurus cf. /. punctatus (Rafinesque), Perca flavescens faunules (Frye, Leonard, and Hibbard, 1943), the (Mitchill), Valvata tricarinata (Say), Probythinella "Yarmouthian molluscan fauna" (Leonard, 1950), and lacustris (Baker), Lymnaea. stagnalis jugularis Say, the "Kaiisan molluscan fauna" (Frye and Leonard, Pliysa skinneri Taylor, Pupilla muscorum (Linnaeus), 1952) in part. The Arkalon local fauna (Hibbard, Vallonia cyclophorella Sterki. 1953a) is tentatively included. Associated with the 12. Jinglebob local fauna (Herrington and Taylor, Pearlette ash bed (usually immediately below the ash) 1958; Hibbard, 1952, 1955, 1956a; Hibbard and Tay over a wide area from Iowa to Texas. In southwestern lor, 1960; Oelrich, 1953; Rinker, 1949; Taylor and Kansas it is in the At water member of the Crooked Hibbard, 1955; Tihen, 1954, 1955; van der Schalie, Creek formation. Frye, Swineford, and Leonard 1953). In the Kingsdown silt, Meade County, Kans. SUMMARY OF FAUNAL SEQUENCE 17 Fishes, amphibians, reptiles, mammals, and mollusks. those of the area today, winters not much if any cooler. Stratigraphic position and ecologic inferences suggest Significant members: Plesippus, Nannippus, Gastro- a late Sangamon age. Climate humid and mesother- copta paracristata Franzen and Leonard, G. rexroa- mal; summers were cooler and winters warmer than densis Franzen and Leonard, G. peUucida hordeacella those of the area today. Significant members: Terra- (Pilsbry), Ferrissia rivularis (Say), Promenetus um- pene llanensis Oelrich, Arctodus simus (Cope), Ory- ~bi!ica tellus ( Cockerell). zomys, Bison latifrons (Harlan), Promenetus exacu- 17. Sand Draw local fauna (Herrington and Tay ous (Say), Valvata tricarinata (Say), Laevapex kirk- lor, 1958; Hibbard, 1956, 1957; McGrew, 1944; Sav la,ndi (Walker), /Strobilops la~byrinthica (Say). age, 1955; Taylor, 1954, this paper). In an unnamed 13. Cragin Quarry local fauna (Etheridge, 1958; formation, in Brown County, Nebr. Fishes, reptiles, Hibbard, 1949b; Hibbard and Taylor, 1960). In the mammals, and mollusks. Faunal correlation and cli Kingsdown silt, Meade County, Kans. Fishes, amphib matic inferences indicate a late Nebraskan age. Climate ians, reptiles, mammals, and mollusks. Stratigraphic subhumid to humid and microthermal; summers were position and ecologic inferences suggest an early San cooler and moister than those of the area today, but gamon age. Climate semiarid and mesothermal; sum winters were no more severe. Significant members: mers were slightly less hot, winters much warmer Geomys quinni McGrew, Pliopotamys meadensis Hib than those of today. Significant members: Terrapene bard, Gastrocopta chauliodonta Taylor, G. cristata llanensis Oelrich, Testudo, medium sized, Phrynosoma (Pilsbry and Vanatta), Helicodiscus singleyanus cornutum (Harlan), P. modestum Girard, Notiosorex (Pilsbry), Physa skinneri Taylor. crawfordi (Coues), Peromyscus progressus Hibbard, 18. Dixon local fauna (Herrington and Taylor, Promenetus kansasensis (Baker). 1958; Hibbard, 1956, 1958a; Tihen, 1955). In an equiv 14. Jones Sink (lower level) (Goodrich, 1940; Her- alent of or in the Missler member of the Ballard for ringtoii and Taylor, 1958). In the Vanhem formation, mation, in Kingman County, Kans. Amphibians, birds, Meade County, Kans. Only mollusks have been re mammals, and mollusks. Faunal correlation and cli covered so far. Their Stratigraphic position close matic inferences indicate a latest Nebraskan or earliest below the Jones local fauna and climatic inferences Aftonian age. Climate mesothermal and subhumid. suggest a Wisconsin age. Climate microthermal, sub- Significant members: Sorex. PUophenacomys meaden humid. Significant members: Pisidium obtusale sis Hibbard, Pliopotamys meadensis Hibbard, Pliolem- Pfeiffer, P. lilljeborgi Clessin, P. subtruncatum Malm. mus antiquus Hibbard, Gastrocopta cliauliodonta Tay 15. Jones local fauna (Downs, 1954; Herrington lor, G. pellucida hordeacella (Pilsbry), Vallonia per- and Taylor, 1958; Hibbard, 1940, 1949; Hibbarcl and spectiva Sterki, Valvata tricarinata (Say), Physa Taylor, 1960; Taylor and Hibbard, 1955; Tihen, 1942, skinneri Taylor. 1955). In the Vanhem formation, Meade County, 19. Berends local fauna (Herrington and Taylor, Kans. Fishes, amphibians, birds, mammals, and mol 1958; Hibbard and Taylor, 1960; Mengel, 1952; Smith, lusks. Stratigraphic position and climatic inferences 1954, 1958; Starrett, 1956; Taylor and Hibbard, 1955). suggest a Wisconsin age. No assignment to a par ticular subage is yet possible. Climate subhumid, In a basin deposit, included in the undifferentiated microthermal; summer temperatures were much lower Sanborn group, in Beaver County, Okla. Faunal evi than those of southwestern Kansas today. Significant dence and climatic interpretations indicate an Illi- members: Camelops, Platygonus, Microtus pennsylva- noian age. Climate subhumid or humid, and nicus (Ord), Pisidium walkeri Sterki, P. tilljeborgi microthermal. Significant members: Castoroides, Clessin, P. ferrugineum Prime, Valvata tricarinata, Paradipoides stovalli Rinker and Hibbard, Perca fla- (Say), Lymnaea stagnalis jugularis Say, Pupil!a vescens (Mitchill), Esox masquinongy Mitchill, Physa l)landi Morse. skinneri Taylor, Valvata tricarinata (Say). 16. Red Corral local fauna (Herrington and Tay 20. Bar M local fauna (Herrington and Taylor, lor, 1958; Taylor, this paper). In an unnamed forma 1958; Hibbard and Taylor, 1960; Taylor and Hibbard, tion, in Oldham County, Tex. Mammals (principally 1955). The fauna is from two sinkhole fillings in all unpublished) and mollusks. The horse Plesippus Harper County, Okla. Mammals and mollusks. Stra indicates Blancan age, narrowed to late Pliocene by tigraphic position and climatic inferences indicate an the mollusks. The fauna is equivalent, to or slightly Illiiioiaii or Wisconsin age. Climate subhumid or older than the Rexroad, perhaps as old as the Saw humid, and microthermal. Significant members: Das- Rock Canyon local fauna. Climate subhumid and ypus belJus (Simpsoii), Microtus pennsylvanicus mesothermal. Summers were cooler and moister than (Ord), Lymnaea stagnalis jugularis Say, Physa skin- 18 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS neri Taylor, Pupilla muscorum (Linnaeus), Discus SUMMARY OF DEPOSITIONAL AND BIOLOGICAL cronkhitei ( Newcomb). SEQUENCE 21. Slaton local fauna (Herrington and Taylor, Hemphillian age. All Hemphillian fossils from the 1958; Meade, 1942, 1952). Lubbock County, Tex. southwestern Kansas area come from the Ogallala Mammals, reptiles, and mollusks. Faunal correlation formation. The grass seed Biorbia fossilia occurs in and ecologic interpretation suggest a Sangamon age. the upper part of the formation (Elias, 1942, p. 144; Climate mesothermal and subliumid or humid. Signi Hibbard, 1941b, p. 199). Three local faunas are ficant members: Neofiber leonardi Hibbard, Terrapene known: Buis Ranch, Optima, and Swayze Quarry. Uanensis Oelrich, Stagnicola caperata (Say), Prome- The Buis Ranch local fauna (late Hemphillian) is netus kansasensis (Baker), Pupoides hordaceu-s closely related to the Saw Rock Canyon local fauna (Gabb), Pupilla blandi Morse, Gastrocopta pellucida (earliest Blancan) in the basal part of the Rexroad fiordeacella (Pilsbry). formation. Deposition of the Ogallala formation was 22. Rezabek local fauna (Franzen and Leonard, apparently followed by a period of major erosion but 1947; Frye, Leonard, and Hibbard, 1943; Herrington short duration, for the Rexroad formation (Blancan) and Taylor, 1958; Hibbard, 1943). Lincoln County, is nowhere found overlying the Ogallala. Probably Kans. Fishes, amphibians, reptiles, birds, mammals, the Ogallala occurred as upland remnants when the and mollusks. Stratigraphic position and ecologic Rexroad was being laid down at lower elevations. inferences indicate Sangamon age. Abundance of Blancan age. Six superposed local faunas from broken and badly worn shells suggests some may have three formations are assigned to the Blancan. In order been reworked. Significant members: Rorex cinereus of decreasing age and successive superposition these Kerr, Microtus cf. M. pennsylvanicus (Ord), Neofiber are as follows: Saw Rock Canyon, Rexroad, Bender leonardi (Hibbard). (in the Rexroad formation), Deer Park, Sanders (Ballard formation), and Seger (Crooked Creek for CORRELATION OF NORTH AMERICAN AGES WITH LYELLIAN EPOCHS mation). The relatively fine-grained Rexroad formation is For relating the Blancan, Irvingtonian, and Ran- known to represent a large time range by the differ cholabrean ages to the Lyellian epochs Pliocene and ences between mammals of the Saw Rock Canyon and Pleistocene, southwestern Kansas and western Okla Rexroad local faunas. The Saw Rock Canyon local homa is the most significant area in North America. fauna might almost as well be termed Hemphillian as This importance is due to a combination of several Blancan; the Rexroad local fauna is more character unusual factors, principally conditions of deposition istically Blancan, containing Ple.sippus and Nannip- peculiar to this area. Oversimplifying the picture, pus. Both mammals and mollusks indicate that the one may state that from the late Pliocene through the climates in which these two faunas lived were meso present continued solution at depth of Permian salt thermal and subhumid or humid, lacking extremely and gypsum beds has caused development of many cold winters and hot summers, and with more and sinkholes and large basins. The exceptionally thick better distributed rainfall than southwest Kansas deposits "trapped" in these depressions contain the receives now. Furthermore, the deposits and faunas most complete sequence of late Pliocene and Pleisto overlying the Rexroad formation show 4 cycles of cene continental faunas yet recorded. This chapter erosion and deposition of coarse materials in a cool is intended to emphasize only the sequence of faunas climate separated by 3 intervals of mild climate with and to show their broader implications. The fore deposition of fine-grained sediments. The most rea going summary of the f aunal sequence contains a more sonable interpretation of this sequence tentatively detailed treatment, listed by individual faunas. correlates the 4 cool-climate cycles with the 4 major In the western Oklahoma and southwestern Kansas continental glaciations, and regards the Rexroad for area, the unusually detailed sequence is confined to the mation as preglacial (here, equivalent to pre-Pleisto- late Pliocene and Pleistocene. Clarendonian and Hem- cene). The upper part of the Rexroad formation, phillian faunas not of unusual significance are men above the level of the Rexroad local fauna, contains tioned below only to indicate their presence in the a massive buried caliche zone indicative of a change area. Following a summary of the depositional and to semiarid climate. Above the caliche, silt containing biological sequence is a correlation of this sequence the Bender local fauna grades into sandy silts. This with the Pliocene-Pleistocene boundary and the Blari- succession indicates a change from semiarid to moister can, Irvingtonian, and Rancholabrean ages. climate. CORRELATION OF NORTH AMERICAN AGES WITH LYELLIAN EPOCHS 19 The Ballard formation unconformably overlies the two units can be related stratigraphically to one an Rexroad formation. Its lower sand and gravel mem other with relative ease. Upper Pleistocene deposits, ber has yielded glacially striated pebbles (Hibbard, however, are of two chief types: fillings of valleys 1944), and scattered vertebrates. The higher silt mem cut into the earlier formations, and isolated sinkhole ber contains at its base the mild-climate Deer Park deposits. After deposition of the Crooked Creek local fauna, a higher presumably interglacial buried formation (that is, in post-Yarmouth time) two inter caliche zone, and the still higher Sanders local fauna vals of valley cutting and filling took place. Deposits indicative of a subhumid mesothermal climate. As of the first interval are included in the Kingsdown part of the first of four cool-climate erosional and silt, those of the second are termed the Vanhem forma depositional cycles, the lower part of the Ballard tion. The two units are correlated tentatively with the formation is considered probably Nebraskan, and the Illinoian-Sangamon and Wisconsin ages, respectively. upper part probably Aftonian. The isolated sinkhole deposits, regardless of age, are The Crooked Creek formation is a unit similar in included in the undifferentiated Sanborn group. These lithologic succession to the unconformably underlying later Pleistocene deposits usually lack the massive Ballard formation, but distinct from that formation buried caliche of earlier units, and may lack a basal in lithologic type. As part of the second of four cool- gravel. This more complicated upper Pleistocene stra climate cycles of erosion and deposition, the lower part tigraphy results in less stratigraphic control for these of the Crooked Creek formation is considered probably faunas. Warm-climate faunas are called Sangamon, and cool-climate faunas are referred to the Illinoian or Kansan. The basal gravel of this unit contains the Wisconsin, depending upon which unit they come Seger local fauna, with such characteristically Blancan from. forms as Plesippus and Nannippus. For this reason Two Illinoian faunas are from the undifferentiated the Blancan age includes the early part of the Kansan Sanborn group: the Berends and Butler Spring local age. faunas. Both Sangamon assemblages, the Cragin Irvingtonian age. Three faunas, all from the upper quarry and Jinglebob local faunas, are from the Kings- part of the Crooked Creek formation, are referred to down silt. The Jinglebob local fauna is of interest in the Irvingtonian age. These are the Cudahy fauna, containing Bison latifrons, marking the late Sangamon associated with the Pearlette ash bed; the Arkalon first appearance of Bison in the Meade County area. local fauna, equivalent to or very slightly later than The Vanhem formation has yielded the youngest the Cudahy; and the Borchers local fauna, strati- assemblage of fossils in the southwest Kansas area, the graphically the highest of these three faunas. From Jones local fauna. its position in the sequence the Crooked Creek forma CORRELATION tion is believed to be probably Kansan and Yarmouth Previous discussions of the Pliocene-Pleistocene in age. The Arkalon and Cudahy (implying micro- boundary and of the North American ages such as thermal climates) are termed Kansan; but the later Hemphillian and Blancan have been handicapped by Borchers local fauna indicates a warm, semiarid cli confusion of two essentially different categories. The mate of probable Yarmouth age. North American ages are successive broad phases in Rancholabrean age. Mammals from the younger the development of the North American mammalian deposits in the southwestern Kansas area are inade fauna. The type section of the Pliocene-Pleistocene quate to place precisely the Irvingtonian-Ranchola- boundary in central Italy is based upon changes in brean boundary in terms of glacial ages. The late marine invertebrate faunas; the Astian-Calabrian Kansan Arkalon local fauna is Irvingtonian, and the boundary is the one evidently considered definitive by the International Geological Commission. Italian Sangamon Jinglebob local fauna is Rancholabrean. mammalian assemblages no less than those from the By analogy with the Blancan-Irvingtonian boundary, United States are dated relative to the Pliocene-Plei the middle Illinoian age may include the end of the stocene boundary by indirect correlation, not defini Irvingtonian. tion. Chapters in the history of land mammals of Later Pleistocene stratigraphy in the Meade County, North America do not necessarily correspond to those Kans. area is more complex than that of the earlier of Europe, and neither of these necessarily corresponds Pleistocene, because of a change in type of deposition to chapters in the history of marine invertebrates in which occurred in about earliest Illinoian time. The the Mediterranean. lower Pleistocene Ballard and Crooked Creek forma Pliocene-Pleistocene boundary. The 18th Interna tions are widespread sheet deposits; the fossils of these tional Geological Congress recommended that the Plio- 20 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS cene-Pleistocene boundary have a type area in Italy, If one considers this critical view correct, the result is that and that the marine Calabrian stage and its conti none of the stratigraphic concepts previously applied in the nental equivalent, the Villafranchian, be considered nonmarine Quaternary (and Pliocene) of central Italy satisfy the demands which are put on stratigraphy. We may derive earliest Pleistocene (American Commission on Strati- from this the demand, that a new stratigraphy of the central graphic Nomenclature, 1949). The Pliocene-Pleisto Italian Quaternary must be provided, and this must naturally cene boundary in other parts of the world should be happen from the clarification of the local relationships them established with reference to this Italian type, but selves, lienee from some kind of local classification. application of this principle is beset with difficulties The descriptions of the stratigraphy in various local of correlation as well as ambiguities in interpretation areas of central Italy given by Liittig (1958) are of the Congress' decision. necessary for understanding the regional relationships The most recent and most careful study of the Plio of the Calabrian and Villafranchian sediments, but cene and Pleistocene stratigraphy of central Italy need not be repeated here. Liittig's discussion of the (Liittig, 1958) has shown the need for revision of pre sequence in the upper Arno Valley, however, is so vious ideas about the Villafranchian stage. The con important as a basis for correlation with the series of cept of this stage has been more broadly inclusive and faunas from the Great Plains described in this paper poorly defined than realized, and includes more than that, part of it (Liittig, 1958, p. 659-660) is quoted simply the continental equivalents of the marine Cala here (translation) : brian stage. Llittig's conclusions are so significent The Yaldarno superiore: The classic locality of Plio-Pleisto- in considering the Pliocene-Pleistocene boundary that ceue faunas and floras, which have been erroneously called the it is desirable to quote some of his statements at typical fossil associations of the Villafranchiano, includes length. After discussion of the classifications of Plio thick lacustrine clay and silt, which are truncated above by cene and Pleistocene sediments in northwestern Europe sands and fluviatile gravels. Within the lacustrine clay lies the boundary between Pliocene and Pleistocene. Consideration and in the Alps, Liittig (1958, p. 655-656) stated of the fossils shows that they can be divided into two types: (translation) : Type a: fossils of a warm to subtropical character, Type b: Both of the classifications cited above show that there is fossils of temperate climate, which therefore show that a a series of readily recognizable stratigraphic entities older cold-invasion took place during or before their dispersal. The than the Mindel glacial interval, or the part of the Quaternary first "glacial interval" of the Pleistocene corresponds to this in northwestern Europe which corresponds to it, which are in cold-invasion. The fossils of type a occur in the lower clays cluded in the "Villafraiichiano" of central Italy (under the of the Valdarno (Stellicone series); in these clays occvirs assumption that they can be differentiated also in central the greater part of the lignite, in particular the deposits of Italy). From the results of Quaternary geology in central and Santa Barbara, Gaville, Tegolaia, Monastero and San Donato northwestern Europe, the concept of the Villafraiichiano in Avane. The fossils of type b are restricted to the higher would then be put in a new light, if the observations made parts of the lake beds (Ville beds). Both sedimentary com in Italy had not already led to the demand that, the "Villa- plexes are separated by an "unconformity" according to ob franchiano" concept be applied no longer in central Italy servations by Sestini (1936). The reasoning for this uncon or at least no longer in the previous way. At the type locality, formity of Sestini, which Movius (1940) also put to use, I in the rest of northern Italy and in France, observations are cannot follow, but in effect come to the same conclusion. At accumulating which favor the view of Lona (1950), Venzo Ricasoli may be seen fluviatile gravels at an horizon which (1056), and Movius (1949) that the Villafraiichiano in the corresponds to Sestini's unconformity, and which as the strict sense is to be placed in the first warm interval of the Ricasoli beds represent the first cold time of the Pleistocene. Pleistocene, that is in the ?Tegelen interstadial or the A division of the lake beds cannot be doubted. The hypothesis Donau/Giinz interstadial. The continental Villafraiichiano of a lower warm and an upper cold "Villafranchiano" is super is in striking correlation with the marine Calabriano. The fluous ; the "warm Villafranchiano" is middle to upper Plio differentiation of a warm from a cold Villafraiichiano must be cene, the "cold Villafranchiano" corresponds to the first warm discarded, for that which has been called "warm Villafraii interval of the Pleistocene, the Ville series, and the decimation chiano" is definitely to be placed in the Pliocene, according of the fauna and flora of a warm interval which is shown in to Movius' useful summary and my own field work. The fossil-type b, occurred during deposition of the Ricasoli series. faunas and floras of the "cold Villafraiichiano" document the The overlying gravels of the so-called Sansino series are to selection of species caused by the first significant invasion of be placed in the second cold interval of the Pleistocene. cold of the Pleistocene, in which the warmth-loving "Tertiary If one wishes, the Ville series can be equated to the Villa elements" were partly decimated. The faunas and floras of fraiichiano in the strict sense. Since, however, this concept the Valdarno superiore, which have been called quite in- has been used in this area in a wider sense instead, a new admissibly typical of the "Villafranchiano," range from the usage in another sense would bring more confusion than order. (middle) Pliocene to (at least) the Tegelen interstadial. The Although some confusion in the concept of the Vil Villafraiichiano in the strict sense is only a fraction of this, lafranchian fauna may have been due to inadequate and it is not permissible to use this concept any longer for deposits which may have an age from middle Pliocene to locality information, part may also have been due to Mindel. facies changes. According to Liittig's descriptions, CORRELATION OF NORTH AMERICAN AGES WITH LYELLIAN EPOCHS 21 the Astian (late Pliocene) deposits of central Italy val, and that the faunas must be either Pliocene or are marine almost everywhere except in the upper Pleistocene. Valdarno, where all of the Piacenzian and Astian Another reason for which the Blancan age has been equivalents are continental. correlated with the early Pleistocene is the belief that Correlation between the High Plains and central the marked faunal break between the Kimball and Italy on the basis of fossil mammals has been at Broadwater formations of the Nebraska Geological tempted, but is unreliable. The number of mammals Survey does not imply a considerable time interval. known from late Pliocene deposits in the area selected (See Schultz and Stout, 1948). The Kimball has been by the International Geological Congress as the stand regarded as late Pliocene, the Broadwater as early ard of reference for the Pliocene-Pleistocene boundary Pleistocene. According to the interpretations and cor is very small indeed, and most of the mammals which relations accepted here, however, the Kimball is middle have been considered as the typical Villafranchian Pliocene, the Broadwater is Pleistocene, and the late fauna from this area are not of earliest Pleistocene Pliocene is unrepresented in the Great Plains north age (Liittig, 1958; Thenius, 1959). Furthermore, the of southwestern Kansas. latest Pliocene-earliest Pleistocene time interval in Schultz and Stout (1948, p. 556) summarized the volved is relatively short, and it seems that dispersal significant differences between the fossil mammals lag might affect mammalian correlations significantly from the Kimball and Broadwater formations. The even though it is unimportant in earlier parts of the rhinoceroses, longirostrine mastodonts, and transi Tertiary. tional PUohippus-Plesippus from the Kimball suggest The most practicable, and perhaps also the most a late Hemphillian assemblage. If the Broadwater precise correlation of the Pliocene-Pleistocene boun formation is Pleistocene, then the earlier Blancan dary between the Great Plains and the type area in interval represented by the Rexroad formation in central Italy is adoption of the earliest horizon with Kansas does not have an equivalent unit in western evidence of marked climatic cooling in temperate lati Nebraska. The correlation of the middle Pliocene to tudes as representing the earliest Pleistocene. In early Pleistocene deposits of southwestern Kansas and southwestern Kansas and northwestern Oklahoma, western Nebraska with those of central Italy (as classi where the geologic record of late Pliocene and Plei fied by Liittig, 1958) is shown in table 6. stocene times is more nearly complete than elsewhere in the Great Plains, the Pliocene-Pleistocene boundary TABLE 6. Correlation of middle Pliocene to lower Pleistocene deposits in Italy and the High Plains is on this criterion between the Rexroad and Ballard formations. These two units are separated by an North Ameri Southwestern Western Nebraska Central Italy stages erosional unconformity which represents an hiatus can ages Kansas of uncertain extent. Stump Arroyo Blancan age. The term Blancan was proposed member of Crooked Creek Broadwater ' Arno Stage (Wood and others, 1941) for a group of mammalian formation formation (Pleistocene) assemblages considered roughly contemporaneous and Ballard characterized by the association of (among others) formation Plesippus, Nannip'pus, and Borophagus. The name, Rexroad (erosion) (Pliocene) but not the entire basis for the term, was taken from formation Astian Stage the Blanco local fauna of Texas. The time charac Hem Ogallala Ogallala * Piacenzian terized by this composite faimal concept was called phillian formation group Stage the Blancan Provincial Age, and was thought to be ? late Pliocene. Since establishment of this term, evidence has ac i Terminology of Nebraska Geological Survey. cumulated that many Blancan faunas lived during the Irvingtonian and Rancholabrean ages. -Correlation early Pleistocene (Nebraskaii to early Kansaii). Other of the upper limit of the Blancan, and of the limits studies, principally by C. W. Hibbard and others in of the Irvingtonian and Rancholabrean ages with the southwestern Kansas, have shown that Blancan assem glacial and interglacial stages of the midcontinent blages lived both before and after the beginning of region is now possible only in southwestern Kansas. continental glaciation. Much of the confusion about It is hardly definite, however, for the glacial and the age of the Blancan interval comes from the tacit interglacial intervals recognized in southwestern Kan oversimplification by many paleontologists that all sas are correlated in turn with the Mississippi Valley Blancan faunas lived during a relatively short inter- area. Accepting these correlations, however, the Blan- 508945 O 60___3 22 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS can-Irvingtonian boundary falls within the Kansan Stratigraphy. Hibbard (1954) reported the faumi age, and the Irvingtonian-Rancholabrean boundary to occur in the lower part of the Rexroad formation. falls within the Illinoian age. Later fieldwork, however, has shown that it occurs in the Ogallala formation of that area (Tihen, 1955, p. DESCRIPTION OF LOCAL FAUNAS 238). Apparently both Permian rocks and Ogallala BUIS RANCH LOCAL FAUNA formation sagged, perhaps because of ground water solution of salt and gypsum at depth. The Ogallala Previous work. The Buis Eanch fossil locality was was thus kept largely below the water table, so that discovered in 1951 by a University of Michigan field weathering could not form the mortar beds so charac party under the supervision of C. W. Hibbard. Sub teristic of the formation in northwestern Oklahoma sequent collecting by Hibbard and other Michigan and southwestern Kansas. groups has yielded a small fauna, especially inter Localities. Only one locality is known, on the west esting because it shows relationships to both Hemphil- side of Buckshot Arroyo, Buis Ranch, NEV4SWV4 lian and Blancan assemblages. Previous references sec. 5, T. 5 N., R. 26 E., Beaver County, Okla. to the mollusks are by Herrington and Taylor (1958) Age. Perognathus cf. P. mclaughlini, and Plio and Hibbard (1954). geomys suggest that the Buis Ranch local fauna is Fauna. Following is a list of the Buis Ranch local of approximately the same age as the Saw Rock fauna: Canyon local fauna (earliest Blancan). The presence Class Pelecypoda of a rhinoceros tends to indicate a Hemphillian, rather Order Teleodesmacea than Blancan, age. A latest Hemphillian age thus Pisidium casertanum (Poll) seems reasonable. Class Gastropoda Order Basommatophora Mollusks are so rare that they cannot give an accu Stagnicola caperata (Say) rate picture of the fauna which lived in the area. The Ferrissia rivularis? (Say) occurrence of Vertigo ovata, unknown from the thor Order Stylommatophora oughly collected Saw Rock Canyon fauna, suggests Strobilops sparsicostata Baker a greater age for the Buis Ranch local fauna, however, Vertigo ovata Say Class Amphibia for the mutually exclusive occurrence of Vertigo Jiib- Order Caudata bardi and V. ovata suggests ecologic replacement. V. Ambystomatidae, indet. ovata is known from the Clarendonian Laverne local Order Salientia fauna and from Pleistocene faunas, but is absent from Frogs or toads, or both the Saw Rock Canyon and Rexroad local faunas. V. Class Reptilia Order Chelonia hibbardi is known only from these last two faunas. Testudo twffida Cope Paleoecology. Climate: Stagnicola caperata occurs Order Squamata in the High Plains as far South as Nebraska. It sug Snake and lizard bones gests cooler summers at the time the Buis Ranch fauna Class Mammalia Order Insectivora lived. Ferrissia requires permanent water, and implies Hesperoscalops sp. greater rainfall than northwest Oklahoma receives Order Carnivora today. Habitat: The few species and their rarity Buisnictis schoffl Hibbard suggest the possibility that the mollusks may not have Order Rodentia lived where their shells were found. Ferrissia rivul Citellus dotti Hibbard aris is a stream snail, Stagnicola caperata a temporary Perognattiiis cf. P. mclaugMini Hibbard Prodipodomys sp. pond species. Pliogeomys buisi Hibbard Cricetidae, indet. SAW BOCK CANYON LOCAL FAUNA Order Lagomorpha Previous work. The Saw Rock Canyon local fauna Leporidae, indet. Order Artiodactyla was discovered in 1943 by Thad G. McLaughlin and Giant camel Claude W. Hibbard. Subsequent fieldwork has been Order Perissodactyla by Hibbard and parties from the Universities of Kan Nannippus sp. sas (1944) and Michigan (since 1950). Hibbard Rhinoceros, indet. (1944a, b) named the fauna and described vertebrates Occurrence. The Buis Ranch local fauna is known from the deposit. Later stratigraphic studies and only from Beaver County, Okla. more vertebrate material (principal reference Hibbard, DESCRIPTION OF LOCAL FAUNAS 23 1953) have emphasized the transitional Hemphillan- Class Mammalia Blancan aspect of the fauna. Previous references to Order Insectivora Soricidae, indet. the mollusks are by Franzen and Leonard (1947), Order Rodentia Frye and Leonard (1952, p. 151, Seward County Marmota? sp. locality), Herrington and Taylor (1958), and Leonard Pliogeomys sp. (1952a). Mollusks collected by University of Michi Perognathus mclaughlini Hibbard gan parties form the chief basis for the present report. Prodipodomys sp. Dipoides ivilsoni Hibbard Fauna. Following is a list of the Saw Rock Canyon Onychomys larralteei Hibbard local fauna: Baiomys sawrockensis Hibbard Class Pelecypoda Cimarronomys stirtoni Hibbard Order Teleodesmacea Ogmodontomys sawrockensis Hibbard Sphaerium striatimtm (Lamarck) Order Carnivora sp. Osteoborus progressus Hibbard Pisidium casertanum (Poll) Buisnictis cf. B. schoffi Hibbard Class Gastropoda Order Artiodactyla Order Mesogastropoda Gigantocamelus cf. G. spatulus (Cope) Marstonia decepta (Baker) Occurrence. The Saw Rock Canyon local fauna is crybetes (Leonard) known only from Seward County, southwest Kansas. Order Basommatophora Stagnicola bulimoides techclla (Haldeman) Stratigraphy. See Hibbard, 1949a, p. 92-97. The sp. fauna occurs near the base of the Rexroad formation Fossaria dalli (Baker) in the XI member. Gyraulus parvus (Say) Localities. Only one locality is known, near the Helisoma anceps (Menke) center of the west line of sec. 36, T. 34 S., R. 31 W., Promenetus Jcansasensis (Baker) unibilicatellus (Cockerell) Seward County, Kans. Ferrissia rivularis (Say) Age. Essentially all mollusks of the Saw Rock Physa anatina Lea Canyon local fauna occur also in the Rexroad local Order Stylommatophora fauna, but considerable age difference between the two StroMlops laltyrinthica, (Say) Gastrocopta holsingeri (Sterki) is indicated by the associated mammals. Hibbard fransenae Taylor (1949a) considered the Saw Rock Canyon local fauna paracristata Franzen and Leonard as either late middle Pliocene or early late Pliocene, rexroadensis Franzen and Leonard and later (Hibbard, 1953) as early late Pliocene. The tappaniana (Adams) Rexroad local fauna (which see) is of later late Plio Pupoides alMlabris (Adams) cene age. Vertigo MJ)~ba,rdi Baker milium (Gould) Leonard (quoted by R. C. Moore, in McLaughlin, Vallonia perspectiva Sterki 1946, p. 33) stated that "fossil mollusk assemblages cf. Succinea associated with the Saw Rock Canyon vertebrates are Deroceras aenigma, Leonard clearly differentiated from those of the type Rexroad." Helicodiscus singleyanus (Pilsbry) This distinction was maintained by Leonard (1948). Nesovitreaf sp. Hawaiia minuscula (Binney) Frye and Leonard (1952) indicated that all Saw Rock Class Osteichthyes Canyon mollusks occurred in the Rexroad local fauna Small catfish also, but did not discuss the previously held difference. Class Amphibia They further believed the faunas to be of the same age, Order Caudata but did not consider the conflicting vertebrate evi Ambystomatidae, indet. Order Salientia dence. Frog and toad bones Paleoecology. Climate: Promenetus umbilicatellus Class Reptilia has a northern distribution in the High Plains, rang
Order Chelonia ing~ as far south as Nebraska. It indicates summers Turtle, indet. without the present extremes of heat. Stagnicola Order Squamata bulimoides techella is found in the High Plains only Eumeces striatiilatus Taylor Snake, indet. as far north as Kansas. It shows that winters were Class Aves very slightly, if at all, cooler than now, and may even Unstudied bird remains have been milder. 24 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Table 7 compares the Blancan and Recent molluscan TABLE 7. Comparison of Blancan and Recent molluscan faunas faunas of southwestern Kansas. There are two prob of southwestern Kansas, by habitats Continued ably significant differences between the Recent and [Recent list based on Lee (1951), Leonard (1943), Miles (1958), and personal collections in Meade County] Saw Rock Canyon faunas: The fossil fauna has four snails requiring permanent water, the Recent none. Fauna There are two fossil forms needing humus in wooded Saw- spots, but the better known Recent fauna has but one Habitat Species Rock Rex- Ben San Can road der ders Re rare species of such a habitat. In comparing the two yon local local local cent 4 local fauna fauna fauna faunas it should be noted that the Recent list is fauna slightly increased by permanent water species found X v v only in cool artesian springs, and not characteristic of Vanatta). X X the area as a whole. (Pilsbry). X Hawaiia minuscula (Bin- X X X X X TABLE 7. Comparison of Blancan and Recent molluscan faunas ney). Helicodiscus singleyanus X X X X X of southwestern Kansas, by habitats (Pilsbry). Pupoides albilabris X X X X X [Recent list based on Lee (1951), Leonard (1943), Miles (1958), and personal collections (Adams) . in Meade County] X Gould. ]Fauna Deroceras aenigma Leon X X X X ard. X v 3aw lor. Habitat Species ,ock .ex- Ben an- paracristata Franzenand X X X X Can oad der lers Re- Leonard. yon ocal ocal ocal ent X ocal auna 'auna auna Leonard. auna v X X Permanent water: v v Stream. _____ . v v Taylor. Pond or stream _ __ Anodonta grandis Say..--. X Helisoma anceps (Menke). X X v Marstonia crybetes (Leon X X atta. ard). v X decepta (Baker) __ __ X v X X Sphaerium striatinum (La X marck). Vattonia perspectiva Sterki. X X X transversum (Say). .... X X ^Ligumiasubrostrata (Say) . X Pond..-. ______v man). A former climate explaining all these data would Permanent to tempo rary: differ from that of the present by less extreme summer Permanent or sub- Helisoma trivolvis (Say) permanent. Physa anatina Lea. .. _ X X v v hot spells and rainfall not necessarily much greater but Promenetus exacuous (Say)_ v kansasensis (Baker) -..-_ X X - X more effective, probably within the 20-30 inches per Uniomerus tetralasmus- (Say). year range. The minimum changes one needs to postu Permanent or tem Gyraulus parvus (Say). _ _ X X X X porary. Stagnicola exilis (Lea) . _ . . - X late to explain the former molluscan fauna are summer sp. ______v Temporary water: rains occurring more often and more gently, rather Temporary pond or Aplexa hypnorum (Lin marginal pool. naeus) . than as occasional intense thundershowers; summer hot Phidium casertanum (Poll). X X Promenetus umbilicatellus X X X X spells less extreme; and annual precipitation perhaps (Cockerell). X v increased to 25 inches per year. Temporary pond or bulimoides techetta (Hal- X X X X X stream. deman). Habitat: Table 8 summarizes the habitat indications Ferriss) . of the mollusks of the Saw Rock Canyon local fauna. Marginal pools and Fossaria dalli (Baker)...- X X X X X wet mud. Semiaquatic, riparian v The site of deposition apparently was close to three habitat. Moister humus. _____ X sorts of aquatic habitats: a permanent, slow moving V stream, shallower water near shore, with dense vegeta Gastrocopta tappaniana X X X X X (Adams). Vertigo milmm (Gould). _. X X X X tion, and fluctuating lagoons and backwaters with X X Damp humus or grass X X shallow, muddy borders. (Sterki). v Two terrestrial habitats may be distinguished also, X bardt. moist to quite damp humus associated with dead wood V V Damp humus o X and decaying leaves, the floor of a wooded area, and a wooded area. v (Say). slightly dryer but still protected habitat, perhaps more ? X (Gould). open grassy areas. Deroceras and cf. Succinea may X have been associated in a transitional, nearly semi- v (Say). aquatic situation, but their habitat preferences are un v 7onitoides arboreus (Say). ? known for sure. DESCRIPTION OF LOCAL FAUNAS 25 TABLE 8. Habitat indications of mollusks of Saw Rock Canyon Fauna. Following is a list of the Red Corral fauna: local fauna Class Pelecypoda [A, abundant (>1,000); C, common (101-1,000); S, scarce (21-100); R, rare (1-20)] Order Teleodesmacea Habitat Species Abun Pisidium casertanum (Poli) dance Class Gastropoda Order Basommatophora Permanent water: Pond or stream. Helisoma anceps (Menke)..____-. Stagnicola, bulimoides tecJiella (Haldeman) Marstonia crybetes (Leonard)__ . Gyraulus parvus (Say) decepta (Baker)______. Sphaerium striatinum (Lamarck).... Promenetus kansasensis (Baker) Stream______Ferrissia rivularis (Say)______. Permanent to temporary water: umbilicatellus (Cockerell) Permanent or subpermanent Physa anatina Lea_____..._. Ferrissia rivularis (Say) Promenetus kansasensis (Baker)._- Permanent or temporary______Gyraulus parvus (Say)_____ - Physa anatina Lea Stagnicola sp.. ______- Temporary pond or marginal pool. Pisidium casertanum (Poli)..___- Order Stylommatophora Promenetus umHlicatellus (Cocker- ell). Strobilops sparsicostata Baker Temporary pond or stream..______Stagnicola bulimoides techella (Halde- Gastrocopta franzenae Taylor man). Marginal pools or wet mud. Fossaria dalli (Baker)...____ . pellucida hordeacella (Pilsbry) Moister humus______Vertigo mttium (Gould)._.___...__ . Damp humus or grass...___ Oastrocopta holzingeri (Sterki)..... paracristata Franzen and Leonard tappaniana (Adams)._.____. rexroadensis Franzen and Leonard Damp humus of wooded area- Nesovitreaf sp_.______. Strobilops labyrinthica (Say)....__. Pupoides inornatus Vanatta Damp to dry ground_____. Hawaiia minuscula (Binney)__ . Helicodiscvs singleyanus (Pilsbry)... Vertigo milium (Gould) Pupoides albilabris (Adams).___. Pupillidae?, indet. Uncertain- Oeroceras aenigma Leonard ____.. Oastrocopta framenae Taylor.... . cf. Succinea paracristata Franzen and Leon ard. Deroceras aenigma Leonard rexroadensis Franzen and Leon ard. Hawaiia minuscula (Binney) Sphaerium sp_..______. Class Mammalia cf. Succinea... _____.__..___---. Vallonia perspectiva Sterki_____. Order Perissodactyla Vertigo hibbardi Baker.____ . Plesippiis sp. Nannippus sp. Occurrence. The Red Corral local fauna is known Before combining these data to reconstruct the area, only from northern Oldham County, Tex. we may note that the sediments containing the fauna Stratigraphy. Essentially no data are available. are all fine grained fine sand, silt, and clay. There The fossiliferous unit in the immediate vicinity of the are no coarse elastics above the rubble incorporated in fossil locality is a west-dipping series of sand and clay. the base of the Rexroad formation. Localities. (USGS Cenozoic loc. 21040). Only one These data suggest that the Saw Rock Canyon local mollusk locality is known, included in aerial photo fauna lived in and close to a perennial, slow-moving graph DAY 10M 63. A west-dipping unit of sand stream, such as a large creek or river. There were and clay is exposed in the upper part of draws running wooded areas along the stream, and perhaps also into the west side of Rita Blanca Creek. In the basin patches of open grassland. The timbered area was of a north-northeast-trending draw about half a mile strictly riparian, not covering the uplands. south of the county line and about three-quarters of a mile north of a red corral are several Frick Laboratory BED CORRAL LOCAL FAUNA vertebrate quarries. All or most of these quarries are Previous work. In the summer of 1954 D. E. Savage about 20 feet stratigraphically below 2- to 3-foot-thick and a University of California Museum of Paleontol bed of massive caliche. Shells were collected from a ogy field party visited a group of Frick Laboratory fine to very fine compact sand immediately below a 2- vertebrate quarries southwest of Channing, Tex. In f oot bed of carbonaceous papery clay shale with abund ant crushed shells at the north end of one of the quar its brief visit the party observed mollusks and collected ries. This quarry is on the east side of the main draw, scraps of mammalian fossils. Later in the same year just south of a minor tributary gully. C. W. Hibbard went to the site to collect the mollusks. Age. Among the vertebrate material in Univer The name Red Corral local fauna is applied to the sity of California collections from a Frick Laboratory mammals and mollusks known from the quarries 10 quarry close to and at the same stratigraphic horizon miles southwest of Channing, on the west side of the as the mollusk locality are Plesippus and Nannippus, valley of Rita Blanca Creek. The only fossils pre indicative of Blancan age. viously described from the assemblage are the sphaeriid The mollusks suggest correlation of the Red Corral clams (Herrington and Taylor, 1958). and Rexroad local faunas. Virtually all species of 26 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS the Red Corral local fauna occur also in the Rexroad TABLE 10. Comparison of mollusks of Red Corral local fauna with Recent mollusks from Palo Duro Canyon, Texas, by habitats local fauna. (See tables 2 and 4.) [Recent list from Clarke (1938), modified as explained in text] Paleoecology. Climate: Two species (Gastrocopta pellucida hordeacella and Stagnicola bulimoides te- Habitat Species Red Recent Corral chella) have southern distributions in the United States. In the High Plains they range as far north as Permanent water: X Kansas. These two suggest that winters were not X X much if at all more severe than those of the Texas Pan X X handle today. Promenetus umbilicatellus, of northern Permanent to temporary water: distribution, occurs in the High Plains as far south as Permanent or subperma- X nent. X Nebraska. It suggests that the Red Corral fauna lived Promenetus kansasensis (Baker) ... X Permanent or temporary... X X in a climate with summers lacking the present extremes X Temporary water: of heat and temperature. Temporary pond or mar X ginal pool. Promenetus umbilicatellus (Cock- X Recent mollusks of the Texas Panhandle are practi erell). Temporary pond or stream. Stagnicola bulimoides techella X (Haldeman) . cally unknown. Several local lists have been published, Marginal pools and wet mud. X X but nearly all are drift collections which may be con X X taminated by Pleistocene shells. The list given by X Damp humus of wooded area. X Clarke (1938) is based partially on drift also, but at X X least there are no obvious anomalies of distribution. X X Palo Duro Creek (spelled Paladora by Clarke) has Hawaiia minuscula (Binney) ------X X X cut deeply into the eastern edge of the Llano Estacado. X Gastrocopta paracristata Franzen X Permanent water and the protected situation in a and Leonard. pellucida hordeacella (Pilsbry). _. X X deep canyon probably give an assemblage unusually rexroadensis Franzen and Leon X ard. varied for the Texas Panhandle. Clarke did not record X X the exact site of his collection, but distance and direction X X given ("* * * Paladora Creek which lies 4 miles east and 3 miles north of Canyon, Tex.") indicate Palo Duro Creek near 6-mile crossing, 10 miles upstream follows: Vallonia costata and V. perspectives, are con from Palo Duro State Park, Randall County, Tex. The sidered V. parvula following Pilsbry (1948, p. 1028). locality is about 40 miles southeast of the Red Corral Succinea avara is changed to cf. /Succinea. Other site. changes are nomenclatural. Two differences appear significant at first: The Red TABLE 9. Habitat indications of mollusks of Red Corral local fauna Corral local fauna has one permanent stream snail, the [A, abundant (>100); C, common (21-100); S, scarce (6-20); R, rare (1-5)] Recent assemblage none. The Red Corral fauna in Habitat Species Abun cludes several temporary water species, the Recent as dance semblage none. In comparing these two faunas it must Permanent stream______Ferrissia rivularis (Say). be remembered that the Recent collection is from a Permanent to temporary water: Permanent or subpermanent..- Physa anatina Lea____--______. deep canyon with a permanent stream supplied by Promenetus kansasensis (Baker),.... Permanent or temporary._... Gyraulus parvus (Say).__.____...... springs and seepage, but that the Red Corral local Temporary water: Temporary pond or marginal Pisidium casertanum (Poll)... ______pool. Promenetus umbilicateUus (Cock- fauna lived in a shallow, exposed valley. erell). Temporary pond or stream_.,. Stagnicola bulimoides techella (Halde- Habitat indications of the Red Corral local fauna man). Moister humus _.____--__-.. Vertigo milium (Gould)___._.___._.. (see table 9) suggest the presence of some woodland Damp humus of wooded area- Strobilops sparsicostata Baker..-___. Damp to dry ground...... Hawaiia minuscula (Binney). ______and moist humus, as well as seasonal ponds and a per Uncertain...... Deroceras aenigma Leonard ______Gastrocopta paracristata Pranzen and manent stream. The occurrence of /Strobilops, not now Leonard. pellucida hordeacella (Pilsbry)_._ found in the Great Plains except in woodland at the rexroadensis Pranzen and Leon ard. franzenae Taylor__--__------__ eastern edges, is especially significant. Pupoides inornatus Vanatta.-_.___. cf Succinea...... These data and inferences suggest that the Red PupUlidl. .___._____-_____-.-____-__ Corral local fauna lived in a climate with winters not much if at all more severe than those of the area today, Table 10 compares the mollusks of the Red Corra] summers lacking present extremes of heat and drought, local fauna with the Recent collection from Palo Duro and a greater rainfall of about 30 inches per year. Canyon. This list is modified from Clarke (1938) a: Habitat: Table 9 summarizes the habitat indications DESCRIPTION OF LOCAL FAUNAS 27 of the mollusks of the Ked Corral local fauna. The Class Pelecypoda four species classed as abundant suggest a shallow pond Order Prionodesmacea fLigumia subrostrata (Say) fluctuating annually, with thick, submerged vegetation. Order Teleodesmacea The other aquatic species fit such a picture, except for SpJiaerium sp. Ferrissia rivularis, which requires permanent water. Pisidium casertanum (Poll) Probably the pond was a lagoon or oxbow lake near a Class Gastropoda permanent stream. The occurrence of Strobilops in Order Mesogastropoda Marstonia crybetes (Leonard) dicates the presence of woodland and moist humus, but Order Basommatophora there are so few land snails, and most of these are so Carychtum exiguum (Say) scarce, that this habitat was probably not close to the Stagnicola bulimoides techella, (Haldeman) site of deposition. caperata (Say) exilis ( Lea) BEXROAD LOCAL FAUNA Fossaria dalli (Baker) Gyraitlus parvus (Say) Previous work. Hibbard (1938) named the Rex- Helisoma anceps (Menke) road local fauna and described vertebrates from it. Promenetus kansasensis (Baker) Subsequent field work has been by Hibbard and parties umbilicatellHS (Cockerell) Ferrissia parallela (Haldeman) from the University of Kansas (1936-46) and Univer rivularis (Say) sity of Michigan (1947 to date). The rich vertebrate Physa anatina Lea fauna has been described in numerous papers sum Order Stylommatophora marized by Hibbard (1950, 1954a). Later contribu Strobilops sparsieostata Baker tions are by Hibbard (1956, p. 185), Oelrich (1957), Gastrocopta franzenae Taylor holzingeri (Sterki) Savage (1955, p. 67-68), and Tihen (1955). The mol paracristata Franzen and Leonard lusks of the Rexroad local fauna have been studied pellucida hordeacella (Pilsbry) primarily by Baker, Franzen, and Leonard (Baker, rexroadensis Franzen and Leonard 1938; Franzen and Leonard, 1947; Frye and Hibbard, tappaniana (Adams) 1941, p. 408; Frye and Leonard, 1952, p. 151, Meade Pupoides albilabris (Adams) inornatus Vanatta County localities; Herrington and Taylor, 1958; Hib Vertigo hibbardi Baker bard, 1941, p. 95; 1941a, p. 265; Leonard, 1950, p. 38; milium (Gould) 1952a; 1957; Smith, 1940, p. 98). The most nearly Vallonia gracilicosta Reinhardt perspectiva Sterki complete lists are those of Baker (in Hibbard, 1941, Cion-ella lubrica (Muller) p. 95) and Leonard (Frye and Leonard, 1952, p. 151). cf. Succinea According to Hibbard (written communication, Helieodiscus singleyanus (Pilsbry) November 29, 1954) most of the University of Kansas Deroceras aenigma Leonard RetineHa rhoadsi (Pilsbry) material reported from the SW*4 sec. 22, T. 33 S., R. ivheatleyi (Bland) 29 W., Meade County, Kans. (collected by Hibbard Nesovitrea electrina (Gould) and Leonard), is from localities la, Ib, and Ic of the Hawaiia minuscula (Binney) Bender local fauna; some is from Rexroad locality 3. ?Zonitoides arboreus (Say) Polygyra rexroadensis Taylor Several species reported from University of Kansas Class Amphibia collections were not found by University of Michigan Order Caudata parties, and hence their stratigraphic horizon is in Ambystoma hibbardi Tihen doubt. Sphaerium sp., Marstonia crybetes (Leonard), Order Salientia and Helisoma anceps (Menke) have habitat require Scaphiopus diversus Taylor ments which suggest they came from Rexroad locality Nepscaphiopus noblei Taylor Bufo sp. 3. Gastrocopta rexroadensis Franzen and Leonard and Anchylorana moorei Taylor CioneUa lubrica, (Muller) have been found at other dubita Taylor Rexroad localities, but not in the Bender local fauna, robustocondyla Taylor and so probably came from Rexroad locality 3. Zoni- Rana fayeae Taylor toides arboreus (Say) is listed with a query under both meadensis Taylor faunas. ephippium Taylor rexroadensis Taylor Fauna. Following is a list of the Rexroad local ralida Taylor fauna: parvissima Taylor 28 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Class Reptilia Class Mammalia Continued Order Chelonia Order Rodentia Continued Testudo rexroadensis Oelrich Procastoroides lanei (Hibbard) lurgida Cope Sigmodon intermedius Hibbard Order Squamata Reithrodontomys rexroadensis Hibbard Phrynosoma cornutitm Harlan wetmorei Hibbard Sceloporus robust us Twente Peromyscus baumgartneri Hibbard Cnemidophorus bilobatus Taylor Ttansasensis Hibbard Eumeces striatulatus Taylor Parahodomys quadriplicattis Hibbard Eumecoides hibbardi Taylor Bensonomys eliasi (Hibbard) mylocoelus Taylor Baiomys Jcolbi Hibbard Heterodon plionasicus Peters rexroadi Hibbard Class Aves Onycliomys gidleyi Hibbard Order Podicipediformes Symmetrodontomys simplicidens Hibbard Podiceps sp. Ogmodontomys poaphagus Hibbard Order Ciconiiformes PliopJienacomys primaevus Hibbard Threskiornithidae, indet. Zapus rinkeri Hibbard Order Anseriforrnes Order Carnivora Anas bunkeri (Wetmore) Canis lepopliagus Johnston Bucephala albeola (Linnaeus) sp. (large) Order Falconiformes Borophagus diversidens Cope Buteo sp. Bassarisciis casei Hibbard Order Galliformes rexroadensis Hibbard Colinus hibbardi Wetmore Procyon rexroadensis Hibbard Meleagris gallopavo Linnaeus Buisnictis breviramus (Hibbard) Meleagrididae, indet. Trigondctis kansasensl
Stratigraphy. See Hibbard 1949a, 1950, and Mc- Localities Laughlin, 1946. The Rexroad local fauna occurs in Species the upper part of the Rexroad formation below the 1 2 3 4a 4b massive caliche. X 4 4 Localities. All the following localities are in Meade Pisidium casertanum (Poli) --...... X X X 4 X County, Kans.: 4 X 4 X X X Locality 1. (Big Springs Ranch; University of Kansas Meade X X 4 Fossaria dalli (Baker) ______X X X X County loc. 25). Center of south edge NW^NE 1^ sec. 18, 4 T. 32 S., R. 28 W. Exposure in the south bank of a tribu 4 4 X X tary of Spring Creek, where the Rexroad formation is un- X 3 X X X X conformably overlain by the Meade formation. Sanders 1 4 X 4 local fauna locality 1 is in this exposure also. This locality X X X was listed by Hibbard (1950, p. 174) as near the center of X X X X X X the line between sees. 17 and 18, T. 32 S., R. 28 W. paracristata Franzen and Leonard. .. X X X X X Locality 2. (Big Springs Ranch). Center of NE^NW^ sec. 2 2 2 24, T. 32 S., R. 29 W. Exposure on the south side of the X X X X X X X X main branch of Spring Creek. Nearly a mile up the Sand X X X X 4 Draw are Sanders local fauna localities 2 and 3 in the X X X overlying Meade formation. This locality is given as the X X X NW^NWi/i sec. 19, T. 32 S., R. 28 W. by Franzen and 4 4 X X X X Leonard (1947) and by Hibbard (1950, p. 174) ; and as the X X X X X X X X NW% sec. 19, T. 32 S., R. 28 W. by Frye and Leonard X 4 (1952, p. 151). 1 1 Locality 3. (Rexroad Ranch; University of Kansas Meade X X X X ?4 County loc. 3, USGS Cenozoic loc. 21171). W^SW 1^ sec. X X X 22, T. 33 S., R. 29 W. Mollusks come from a pond or bog deposit now covered by the sandy bed of a tributary of Stump Arroyo. This zone is downstream around the bend sequence in the Mississippi Valley indicates a pre- from the chief vertebrate locality, figured by Hibbard (1950, glacial (Pliocene) age for the Rexroad local fauna. pi. 3). The mollusks occur stratigraphically at or slightly above the upper limit of vertebrate occurrence at this local The Saw Rock Canyon local fauna, in the lower part ity. Mollusks were reported by Baker (1938), Hibbard of the Rexroad formation, might be assigned to the (1941, p. 95), Franzen and Leonard (1947), Leonard (1950, middle Pliocene almost as well as to the upper Pliocene. p. 38; 1952a), and Frye and Leonard (1952, p. 151). The Rexroad local fauna, in the upper part of the Locality 4. (Fox Canyon; University of Michigan loc. UM- Rexroad formation, is of late Pliocene age. A large Kl-47). Sec. 35, T. 34 S., R. 30 W. Exposures along Fox Canyon (Hibbard, 1950, pi. 5, fig. 1). Locality 4a is a time gap between these two faunas is thus indicated. pocket of fluviatile sandy silts, 17 feet below the massive Both faunas indicate a similar temperate climate, lack caliche, from which the vertebrates and a few mollusks were ing present hot, dry summers. There is no evidence of collected. Locality 4b is a lenticular bed of blue-gray clay, climatic change between these two faunas. This period 10 feet below the caliche, from which most of the mollusks were collected. Franzen and Leonard (1947), Leonard of probably uniform climate endured longer than the (1952a) and Frye and Leonard (1952, p. 151) reported glacial-interglacial ages, as indicated by mammalian mollusks from this locality. evolution. Such an interval of mild climate is more Age. Mammalian evidence for the late Pliocene age reasonably Pliocene than Pleistocene. Paleoecology. Climate: Stagnicola caperata, Pro- of the Rexroad local fauna has been summarized by menetus umbilicatellus, and Vattonia, gracilicosta occur Hibbard (1950). Additional evidence for this age is in the High Plains as far south as Nebraska. They in as follows. dicate that the Rexroad local fauna lived in a climate The Rexroad local fauna indicates a climate more lacking the hot summers of southwestern Kansas today. humid than the present and lacking such hot summers. Gastrocopta pettucida hordeacella and Stagnicola This fauna occurs below 4 formations representing 4 bulimoides techella have southern distributions in the cycles of erosion and deposition. These cycles are United States, and range north in the High Plains into represented also by four cool-climate faunas separated Kansas. They suggest the Rexroad local fauna lived in by three warmer-climate faunas. A correlation of a climate with winters slightly if at all colder than these faunas with the standard glacial-interglacial those of the region now. 30 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS Table 7 compares the Blancan and Recent molluscan TABLE 12. Habitat indications of mollusks of Rexroad local fauna faunas of southwestern Kansas. There are two prob [A, abundant (>225); C, common (41-225); S, scarce (8-40); R, rare (1-7). Based ably significant differences between the Recent and on material examined only] Rexroad local faunas: The fossil assemblage has Locality several snails requiring permanent water, the Recent Habitat Species 1 3 4a 4b none. The Recent fauna has only one rare species 2 characteristic of damp humus in wooded areas, whereas Permanent water: R there are several common species of such requirements R R R in the Rexroad local fauna. The contrast between the Permanent to temporary water: two assemblages is further shown by the much greater Permanent or subperma- S R C - A nent. number of species in the Rexroad local fauna. In Promenetus kansasensis (Bak R R er). comparing the two assemblages it should be noted that Permanent or temporary- r R A Temporary water: the Recent list is slightly increased by permanent water Temporary pond or R R marginal pool. species found only in cool artesian springs, and not Promenetus umbilicatettus A (Cockerell). characteristic of the area as a whole. Temporary pond or Stagnicola bulimoides techella A R stream. Haldeman. The association of Cionella lubrica and Gastrocopta Marginal pools and wet R R R A mud. pellucida hordeacella in the Rexroad local fauna is S C R Gastrocopta holzingeri (Sterki).. R S C especially significant, for the only known joint occur S S R Vallonia gracilicosta Rein- A rence of these two today is in southern New Jersey. hardt. Damp humus of wooded r Presumably here the ocean reduces seasonal extremes area. Strobilops sparsicostata Baker. . R "c" A R of temperature, permitting usually remote species to Hawaiia minuscula (Binney) .. S A A Helicodiscus singleyanus (Pils- c A S live in the same locality. bry). Pupoides albilabris (Adams) . . - S c C S C Hibbard (1950, p. 175-177) summarized the eco Deroceras aenigma Leonard - . s R A A R A A A logical evidence of various groups represented in the paracristata Pranzen and R S S C Leonard. Rexroad local fauna. He concluded (p. 177) : "There pellucida hordeacella (Pils- R bry). is evidence that the climate in the Upper Pliocene was s C .... R s C more equable than at present, without extremely cold C s A .... R s C winters or severely hot summers, and that there was a R S greater degree of humidity in the region than there is now.'' Locality 2 apparently had only rare surface water. Habitat interpretations (see below) of the Rexroad Scarcity of temporary-pond species shows that prob local fauna suggest the presence of timbered areas and ably habitats were unsuitable for most aquatic snails. of grassland. Such vegetation implies greater rainfall Significantly greater abundance of dry-tolerant snails than southwestern Kansas receives now. further suggests little ground moisture and perhaps no The former climate suggested by all these lines of vegetation but grass. The site of deposition may have evidence differed from that of the present in having been the bare floor of a small intermittent stream valley summers with less extreme hot spells, and a greater bordered by small shrubs and grass, but no trees. annual rainfall of about 30-35 inches. If precipitation Locality 3 is the richest and shows the greatest distribution were better, and gentle rains rather than ecologic diversity of all Rexroad localitites. Aquatic thundershowers were more frequent, one can guess that species are few in number, and only Physa anatina is perhaps rainfall was only about 30 inches per year. common. There is a rich land-snail fauna, however, Winters were markedly milder than at present. Habitat: Table 12 summarizes habitat indications of which suggests a wooded area providing damp humus the Rexroad local fauna. Only material represented and dead wood as favorable habitats for many species. in University of Michigan collections is considered. This somewhat peculiar situation a rich land as Locality 1 was apparently a temporary pond habitat. semblage but poor freshwater fauna suggests that de Rarely occurring species characteristic of permanent position may have been on land. The local area could water indicate it was perhaps a small oxbow pond well have been a well-wooded flood plain, with only close to a permanent stream. The scarcity and small minor amounts of open water. These could have been variety of land snails suggest that riparian vegetation provided by small, shaded pools, seepages, or beaver nearby was scanty. Possibly the pond was located on a canals (Hibbard, 1941, p. 97). Probably the Physa floodplain without nearby woodland, and on which lived near the area of deposition, but other freshwater only cf. Succinea thrived. shells were brought in by the stream when it over- DESCRIPTION OF LOCAL FAUNAS 31 flowed its banks. Absence of larger land snails such as Class Gastropoda Triodopsis or Mesodon probably indicates that wood Order Basommatophora land was not widespread, but restricted largely to the Stagnicola 'bulimoides techella (Haldeman) caperata (Say) stream valleys. sp. Locality 4a has yielded abundant small vertebrates Fossaria dalli (Baker) but only two species of mollusks. From a study of the Promenetus umMlicatellus (Cockerell) sediments and contained bones Hibbard (1950, p. 167- Physa anatina Lea 168) concluded: Order Stylommatophora Gastrocopta cf. G. cristata (Pilsbry and Vanatta) The deposit from which the fossils were taken in Fox franzenae Taylor Canyon was laid down by a stream that was carrying fine paracristata Franzen and Leonard sediments which consisted chiefly of silts and fine sands. The scaevoscala Taylor remains of fishes in the deposit indicate that the stream was tappaniana (Adams) probably permanent. The size of the fossils recovered shows Pupoides albilabris (Adams) the effect of stream sorting. The material does not show Vertigo milium (Gould) signs of abrasion and appears to have been transported but a Vallonia parvnla Sterki short distance, either from slope wash along the valley or cf. Succinea carried by a short tributary that headed in the upland area. Hclicodiscus singleyanus (Pilsbry) Because of the dominance of rodents, the concentration of Deroceras aenigma Leonard small vertebrates is best accounted for as an accumulation of Hawaiia minuscula (Binney) remains from the pellets or castings of owls or hawks that fZonitoides arboreus (Say) nested nearby. Polygyra rexroadensis Taylor The mollusks from locality 4a agree well with this Occurrence. The Bender local fauna is known only picture. The only two species found, Pupoides albilab- from Meade and Seward Counties, southwestern ris and Deroceras aenigma, are both of medium size Kansas. and have shells of relatively high density and strength, /Stratigraphy. This assemblage occurs in the upper compared to other mollusks in the Rexroad local fauna. most Rexroad formation above the massive caliche. Probably other shells were broken and carried away, Locality 2 is at the same stratigraphic position as the while those of only two species were concentrated at mollusks recorded by Hibbard (1949a, p. 94) in a sec the edge of the stream. tion measured close by that locality. Locality 4b represents a slough or stagnant oxbow Localities. Following are from the Bender local lake. The land snails suggest some vegetation, such fauna: as shrubs and grass, but again no extensive forest. The Locality 1. Three sites at the same stratigraphic level, in ex Physa and Stagnicola indicate a semipermanent aquat posures within about an eighth of a mile along the banks ic habitat with probably abundant vegetation. The of a tributary of Stump Arroyo, SE^SW 1^ and SW^SE 1^ rare Marstonia was perhaps washed in from a nearby sec. 22, T. 33 S., R. 29 W., Meade County, Kans. Upper stream during high water. most part of the Rexroad formation, above massive caliche bed. About 300 yards downstream from Rexroad locality 3. Locality la. NE^SE^SW 1^ sec. 22. Dark bed (soil?). BENDER LOCAL FAUNA South side of stream, upstream from locality Ib. Locality Previous work. Mollusks from above the massive Ib. NWHSW^SE^i sec. 22. East side of stream, about halfway between localities la and Ic. Just below Meade caliche in the type section of the Rexroad formation gravels. Locality Ic. SW^SW^SE 1^ sec. 22. Southwest have previously been considered part of the Rexroad side of stream, downstream from locality Ib; near section local fauna. They are segregated here as the Bender road along south side of section 22. local fauna (new name). The fauna is named after Locality 2. Seward County, Kans. Wolf Canyon, in approx Mr. H. C. Bender, Plains, Kans., who now owns the imately the center of W% sec. 12, T. 35 S., R. 31 W. Upper most part of the Rexroad formation, above massive caliche Rexroad ranch, on which the type locality is located. bed. Only mollusks are known in the fauna. Age. Since associated vertebrates are lacking, age Some species not in University of Michigan collec assignments must be based on the molluscan fauna and tions, but reported in University of Kansas material on climatic inferences from the sediments and fauna. from SW*4 sec. 22, T. 33 S., R. 29 W., Meade County, The molluscan fauna has similarities to both the may be from either Rexroad locality 3 or from Bender Rexroad and Sanders local faunas, but is slightly closer localities la, Ib, or Ic. See under Rexroad local fauna to the Rexroad. In addition, G. cf. cristata is inter for discussion. mediate between the Rexroad Gr. franzenae and the Fauna. Following is a list of the Bender local Sanders, Dixon, and Sand Draw Gr. cristata. Negative fauna: evidence, interesting but of slight or no significance, is 32 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS the absence in the Bender local fauna of Vertigo hib- quires much shelter or permanent moisture. There is bardi of the Rexroad local fauna, and the absence of no indication of vegetation heavier than grass and a Vertigo ovata, Gastrocopta chauliodonta, Physa skin- little scrub, or (in conjunction with summers slightly nerij and Aplexa hypnorum of the Sanders local fauna. cooler than present) of rainfall greater than about 20 Comparison of the Bender local fauna with early Pleis inches per year. tocene faunas besides the Sanders would be of little Habitat: Table 13 summarizes the habitat indications value. The Sanders and Bender local faunas probably of the Bender local fauna. Locality 1 probably repre lived under similar conditions, while the Sand Draw sents deposition on land, for terrestrial species are and Dixon mollusks suggest markedly different cli abundant and aquatic forms rare. The land snails are mates. all species which do not require woodland, though the Hibbard (1954a, p. 235-236) discussed the signific Polygyra may have inhabited patches of scrub. The ance of the massive caliche bed in the uppermost part few fresh water species are all able to endure drying of of the Rexroad formation. In summary, it may be their water body. The general environment suggested stated that the caliche suggests a semiarid climate, is a slow-moving temporary stream with bordering whereas the faunas above and below indicate moister climates. There is no evidence that the Bender local vegetation of grass and clumps of bushes or scattered fauna lived in a climate cooler than that of the Rex trees. road local fauna. The uppermost part of the Rexroad Locality 2 suggests similar conditions, except that formation, above the massive caliche and including conditions were more favorable for freshwater snails the Bender local fauna, might be considered Pleisto and less favorable for land snails. cene if the climatic change from semiarid to moister TABLE 13. Habitat indications of Bender local fauna were considered contemporaneous with the beginning [A, abundant (>250); C, common (41-250); S, scarce (8-40); R, rare (1-7)1 of continental glaciation, and if the Pleistocene epoch Locality were defined as beginning with the first activity of Habitat Species major continental glaciers (see Frye and Leonard, la Ib 1C 2 1953). Permanent to subper R R C manent water. The differences between the Bender and first Pleisto Permanent or temporary Stagnicola, sp...... _ . R S water. cene faunas (Sand Draw, Dixon) are much greater Temporary pond or S R C marginal pool. than those between the Rexroad and Bender, however. Promenetus umbilicatellus S S C (Cockerell). The weight of evidence favors a Pliocene age for the Temporary pond or Stagnicola bulimoides techeJla S S C stream. (Haldeman) . Bender local fauna. Marginal pools and wet S R R mud. Paleoecology. Climate: Stagnicola caperata and Vertigo milium (Gould) ... 0 R C Damp humus or grass- __ Gastrocopta tappaniana R S (Adams) . Promenetus umbilicatellus occur in the High Plains as S R Damp to dry ground, ... Gastrocopta cristata (Pilsbry A A far south as Nebraska. They indicate that the Bender and Vanatta). Hawaiia minuscula (Binney)__ A C A C local fauna lived in a climate lacking the present sum Helicodiscus singleyanus (Pils S A C R bry). mer extremes of heat. Stagnicola bulimoides techetta Pupoides albilabris ( Adams) ___ 0 C C S Deroceras aenigma Leonard ___ 0 '"s" C -.-... has a southern distribution, and ranges north in the paracristata Franzen and A A C C High Plains only as far as Kansas. Its occurrence sug Leonard. S S S gests winters only slightly if at all colder than those of S S C southwestern Kansas today. Table 7 compares the Blancan and Recent molluscan faunas of southwestern Kansas. Only 9 of the 19 spe SAND DRAW LOCAL FAUNA cies of the Bender local fauna still live in the area. Previous work. Fossils representing the Sand Draw Vertigo milium lives not too far to the east however, local fauna were first collected by M. F. Skinner for and several extinct species have ecological equivalents the Frick Laboratories in 1934. From these and later in southwestern Kansas. collections by Skinner only two species have been pub The few aquatic species suggest only subpermanent lished (Frick, 1937, p. 529; Osborn, 1936, p. 726). to temporary bodies of water. Several of the land Later collections by Bryan Patterson and J. H. Quinn snails can withstand dry upland conditions in the High of the Chicago Natural History Museum yielded many Plains today (Gastrocopta cristata, Pupoides albilab- additional vertebrates and mollusks (Baker, 1938; Mc- ris, Hawaiia minuscufa), and none of the others re Grew, 1944). In 1950 C. W. Hibbard and a party from DESCRIPTION OF LOCAL FAUNAS 33 the University of Michigan Museum of Paleontology Class Gastropoda Continued visited the area and collected at other localities not Order Stylommatophora Continued sampled by Patterson and Quinn. Additional species Helicodiscus singleyanus (Pilsbry) Deroceras aenigma Leonard found include mollusks (Taylor, 1954a) and a few Hawaiia minuscula (Binney) mammals (Hibbard, 1956, 1957). Taylor visited the Class Osteichthyes region in 1953, studied the stratigraphy of the localities let alums sp. where mollusks had been found, and collected more Class Reptilia material. This stratigraphic study and an examination Order Chelonia Pseudemys sp. of the shells collected by these persons form the basis Testudo, 2 species of the present discussion. Class Mammalia Fauna. Following is a list of the Sand Draw local Order Rodentia fauna: Geomys quinni McGrew Procastoroides siceeti Barbour and Schultz Class Pelecypoda Nebraskomys mcgrewi Hibbard Order Prionodesmacea Pliopotamys meadensis Hibbard fAnodonta grandis Say Order Carnivora Order Teleodesmacea Canis cf. C. latrans Say Sphaerittm partiimeiutn (Say) Trigonictis kansasensis Hibbard striatinum (Lamarck) Taxidea cf. T. taoeus (Schreber) sulcatum (Lamarck) Stnilodon sp. transversum (Say) Order Proboscidea Pisidium casertanum (Poll) Stegotnastodon primitivus Osborn compress utn Prime Order Artiodactyla nitidum Jenyns Platygomts sp. Class Gastropoda Gigantocamelus sp. Order Mesogastropoda Camelops sp. Valvata lew is i Currier Capromeryoe sp. tricarinata (Say) Order Perissodactyla Order Basornmatophora Plesippus simpUcidens (Cope) Carychium sp. Occurrence. All known outcrops of the beds con Stagnicola cf. 8. reflexa (Say) caperata (Say) taining the Sand Draw local fauna are in northern Fossaria dalli (Baker) Brown County, Nebr., within a northwest trending ofirussa (Say) area about 10 by 4 miles. It appears that the beds are Bitlimnea megasoma (Say) the fill in a late Pliocene valley, possibly related to an Anisus pattersoni (Baker) ancestor of the Elkhorn River. G-yraulus parvus (Say) Helisoma anceps (Menke) Stratigraphy. The deposits yielding the Sand Draw Planorbula armigera (Say) local fauna are unnamed. They rest unconformably, Promenetus kansasensis (Baker) upon the underlying Pliocene rocks, and are overlain utn'biUcatel'lus (Cockerell) unconformably by a widespread sand and gravel sheet Acrolojeus coloradensis (Henderson) which forms much of the High Plains surface in the Ferrissia meekiana (Stimpson) area. The approximate maximum thickness of the rivularis (Say) Physa anatina Lea beds is 125 feet. gyrina Say Lithology varies from fine-bedded alternating black skinner i Taylor clays and buff sands to crossbedded coarse sand and Aplexa Jtypnorum (Linnaeus) gravel. The fine-bedded sediments are extensively ex Order Stylommatophora posed along Deep Creek, and probably represent quiet- Strobilops sparsicostata Baker water deposition. The top of a thick sand and gravel Gastrocopta chauliodonta Taylor cristata (Pilsbry and Vanatta) unit is indicated at the base of the section on page 34. tappaniana (Adams) Crossbeddiiig and lenticular clay and gravel beds rep Vertigo milium (Gould) resent fluviatile deposition. ovata Say There are three principal mollusk-bearing horizons: Pupoides aJbilabris (Adams) A, 75 feet below the unconformity (loc. 1); B, 35 feet- i no mat us Vanatta Va-llonia pulchella (Mttller) 40 feet (loc. 2-3); and C, 15-22 (loc. 4-7). The cf. Succinea following section contains only the highest horizon.
508945 O 60- 34 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Section at center north side SW}i sec. 25, T. 81 N., R. %2 W., Locality 7. Near center of SW%SE% sec. 4, T. 30 N., R. 21 Brown County, Nebr. Section and photograph by McGrew (1944, W. Exposure on Bone Creek near Stegomastodon quarry. p. 84-85) are from this same exposure. Baker (1938, p. 130-131) listed mollusks from two localities, Feet one "6 miles north of Ainsworth" and the other "about 2 9. TopsoU.____---____-___.------4.0 miles west" of the former. Bryan Patterson, who collected 8. Fine sand and gravel, dark tan______1. 75 the material, in a letter (dated December 22, 1949) to C. 7. Sand, medium-grained, crossbedded, light rust-colored W. Hibbard, stated that Baker transposed the two localities. with many thin manganese-stained layers, occasional A letter from Baker to Patterson clearly shows that this fine silt lenses,______5. 25 transposition took place. McGrew (1944, p. 36) corrected 6. Coarse sand and fine gravel, dark rust color. ______2. 0 the error, but without so indicating. 5. Sand, fine white, with coarse rusty layer 1-4 inches thick at base______^______6. 25 Age. Faunal correlation and climatic inferences in Erosional unconformity dicate the Sand Draw local fauna is of Nebraskan or Formation containing Sand Draw local fauna Aftonian age. Pliopotamys meadensis is otherwise 4. Sand, fine, light tan, grading upward into silt-__-_-__ 7. 5 known only from the late Nebraskan or early Aftonian 3. Silt, tan (weathering light gray) with rodent burrows Dixon local fauna and the Aftonian Deer Park local filled by sand from unit 4; some shells (loc. 6) _____ 2. 0 fauna. This rodent is especially significant for cor 2. Sand, fine, slightly marly, gray-buff with abundant relation of the Sand Draw local fauna. The snail shells (loc. 6) ______9. 0 GastroGopta chauliodonta is known only from the Sand 1. Channel sands, fine, gray-tan, with lenses of clay and gravel ______9. 0 Draw, Dixon, and Sanders local faunas and thus tends Bed of wash (base of formation not exposed) to support the mammalian evidence. Although the mollusks of the Dixon, Sanders, and Localities. All the following localities are in Brown Sand Draw local faunas are generally similar (table County, Nebr.: 4), the Dixon and Sand Draw local faunas are related Locality 1. Center west side SE^SWVi sec. 24, T. 31 N., R. by sharing Bulimnea megasoma and Acroloxus colora- 22 W; exposure on north side of Sand Draw at mouth of densis. These two species are rare or unknown in other second wash downstream from bridge on former State High Pleistocene assemblages, and suggest a similar environ way 7; locally reddish marly spot, 75 feet below unconform ity. ment for both faunas. Locality 2. NW& sec. 14, T. 31 N., R. 23 W. (USGS Cenozoic Climatic interpretation of the fauna is somewhat loc. 19090), exposures at head of West Fork of Deep Creek; restricted because land snails are relatively underrep- sandy layer in thin-bedded sand and clay layers about 38 resented, and because the known vertebrates are mostly feet below unconformity. large mammals of little ecologic value. The large Locality 3. NE^SWii sec. 12, T. 31 N., R. 23 W; exposure on turtle Testudo indicates much warmer winters than east side of Middle Fork of Deep Creek; thin-bedded sand those of north-central Nebraska today, with minimum and dark-gray clay, 35 feet-40 feet below unconformity. temperatures rarely if ever at freezing. With the limi Locality 4. NW& sec. 33, T. 31 N.. R. 22 W; exposure on north side of Sand Draw; marly silt, 21-22 feet below unconform ted stratigraphic and paleontologic evidence available ity. no precise climatic or age interpretations are possible. Locality 5. Center of north side sec. 34 and southeast corner The mollusks alone, but especially the large Testudo, SW% sec. 27, T. 31 N., R. 22 W; exposures on small draw indicate that the assemblage did not live during a time near earth dam; marly silt, 18-20 feet below unconformity; of major glacial advance. It is not a late glacial age mollusks from this locality were listed by McGrew (1944, p. 36) as "from an exposure 2 miles west of Sand Draw." assemblage if the contrast with the late Kansan Baker (1938, p. 130) reported the snails by error as from Cudahy fauna and the late Illinoian Butler Spring and the exposure on Sand Draw. (See below.) Berends local faunas is reliable. The Sand Draw local Locality 6. North side of SW& and south side of NWVi, sec. fauna may be a transitional Nebraskan and Aftonian 25, T. 31 N., R. 22 W. (USGS Cenozoic loc. 19091). Ex assemblage, as the Dixon local fauna is thought to be, posures along first wash south of Sand Draw on former or it may date from an interstadial in the Nebraskan State Highway 7. Marly sand and tan silt (units 2 and 3 of section), 8-15 feet below unconformity. This wash is age analogous to the Donau/Giiiiz interstadial of the not the Sand Draw proper, as considered by McGrew (1944, Alps. p. 35, explanation of figure), but a small tributary to it. Paleoecology. Climate: Comparison of the mollusks The photograph and section given by McGrew (1944, p. of the Sand Draw local fauna with those living today 34-35) were taken in the center of the north side of the in north-central Nebraska (table 15) shows 2 main SW& of sec. 25. Mollusks from this locality were listed by McGrew (1944, p. 36) as from the "Sand Draw exposure," contrasts which are most simply explained in 2 dif and by Baker (1938, p. 131) by error (see below) as from ferent ways. Valvata lewisi, V. tricarinata, Physa 2 miles west of the Sand Draw exposure. skinneri, and Vattonia pulchella have a generally DESCRIPTION OF LOCAL FAUNAS 35 TABLE 14. Habitat indications of mollusks of Sand Draw that there are 13 species which live in perennial water local fauna bodies in the Sand Draw local fauna, but only 3 in the [A, abundant (>200); C, common (41-200); S, scarce (16-40); R, rare (1-15)] local living fauna. This is considered a real and Locality and abundance significant difference. The three Recent species in this habitat type are all known from a single locality where Habitat Species Level Le\el Level A B C a group of springs forms a summer-cool, permanent pond. North-central Nebraska now is probably con 1 2 3 4 5 6 7 siderably drier and warmer in the summers than for Permanent water: merly. A number of land snails, particularly those of Pond.____ _ ._ ... A croloxus coloradensis S (Henderson) . damp humus and sheltered habitats, are present in the Bulimnea megasoma S R C R R (Say). Recent fauna but not in the Sand Draw local fauna. C A Helisoma anceps C R A R (Menke). This may be explained by sampling error. Pisidium compressum A A R Prime. Only two mollusks (Gastrocopta cristata, Helicodis O R Sphaerium partumeium C cus singleyanus] in the fossil assemblage have a rela (Say). striatinum (Lamarck) . R R tively southern distribution in North America. The R S former reaches its northern known limit of range in Valvata tricarinata (Say) C A R northern Nebraska; the latter, in South Dakota. They Ferrissia rivularis (Say)._ C R A Permanent to tem suggest that winters were formerly no colder than those porary water: Permanent or sub- Planorbula armigera C R of the area today. The most significant temperature permanent. (Say). R C R S R indicator in the fauna is the large land turtle, Testudo. Promenetus kansasensis C C A C A A C (Baker). The occurrence of this species in northern Nebraska in Permanent or tem Gyraulus parvus (Say) ... A C A A A A C porary. R R dicates that winters were formerly much milder than Stagnicola cf. S. reflexa C S A C A A C (Say). now, with the lowest temperatures rarely if ever at Temporary pond or Aplexa hypnorum (Lin R R A R marginal pool. naeus) . freezing. Pisidium casertanum R S S R R (Poll). The mild winter climate inferred from the Sand Promenetus umbilicatel- R S A C A A A lus (Cockerell). Draw local fauna contrasts with the climates inferred s S R A A S Marginal pools and R R from other late glacial faunas from the High Plains, wet mud. O R such as the Cudahy fauna and the Berends, Butler Vertigo milium (GouldK R R R S S R S S C S R Spring, and Jones local faunas. The occurrence of Damp humus or grass . Gaftrocopta tappaniana R R S (Adams) . several markedly northern species suggests a climate Vallonia pulchella R R (Miiller). more likely to be glacial than interglacial, but several Gastrocopta cristata R R (Pilsbry and Van- land snails usually found in glacial faunas, such as atta). Hawaiia minuscula R R R R R Pupilla muscorum (Linnaeus), P. blandi Morse, Ver (Binney). Helicodiscus smgleyanus R tigo modesta Say, and V. gouldi (Binney), are absent. (Pilsbry). Pupoides albilabris R S Until pollen analyses or better samples of the ter (Adams) . Anisus pattersoni R C R R A C (Baker). restrial fauna are available, the former climate and Deroceras a enigma R R C R R Leonard. precise age of the Sand Draw local fauna will remain Ferrissia meekiana R R (Stimpson) . uncertain. Gastrocopta chauliodonta S R R R Taylor. Habitat: Table 14 shows the habitat indications of R C R C C S Pupoides inornatus Van- R the Sand Draw mollusks. Distribution and relative atta. Strobilops sparsicostata R R abundance of the species suggest that horizon C rep Baker. R R S R C S S resents deposition in shallower water with a smaller variety of aquatic habitats than the other levels. Such an interpretation is supported by the fine-grained non- northern distribution in North America. Physa skin- crossbedded sediments, and by the presence of rodent neri does not range south in the Great Plains as far as burrows at locality 6 (see section). Presence of Fer the United States; the others are found a little farther south, but are ordinarily found only north of the Sand rissia rivularis, Valvata tricarinata, and Helisoma an Draw fossil localities. These species suggest that the ceps at horizons A and B suggests a nearby stream. fauna lived under cooler summer conditions than those The fine-bedded, alternating black carbonaceous clays of northern Nebraska today. and light-colored sands along Deep Creek in which The other contrast between the fossil and Eecent these species occur probably indicate lacustrine deposi faunas is in the habitats represented. Table 15 showTs tion, however. 36 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
TABLE 15. Comparison of Sand Draw local fauna and Recent Horizon B is the richest of the three. The sediments mollusks from northern Nebraska and mollusks of varied habitats suggest a large, quiet lake bordered by shallow marginal pools with grass Occurrence and bushes along the shore. Absence of genera such as Habitat Species Sand Draw horizon Re Discus, Retinella. and Zonitoides may indicate there cent were no woods along the shore. Shells from the shore A B c and shallow^ water were apparently washed out to the
Permanent water: deeper areas of the lake. A connecting river probably X xxxx X X X contributed some shells, especially Ferrissia. X X X Horizon C species indicate some permanent water, X X X but apparently many shallow temporary pools were X X present. The increased percentage of temporary water X X and land species suggests a series of small pools and Pond ______Acroloxus coloradensis (Hender- son). shoreline lagoons around a basin which did not dry X X X X up, but in which water level fell annually. Grass and v Permanent to temporary a few trees and bushes lined the shore, but as in water: Permanent or subperma- horizon B. evidence of extensive woodlands is absent. nent. X xxxxixxxxx X X DIXON LOCAL FAUNA X X X Permanent or temporary- _ X X X Previous work, Frye and Leonard (1952, p. 151, X X Kingman County locality) reported mollusks from X X X what is here called Dixon locality 1. Acting on then- Temporary pond or margi X X X nal pool. X discovery, C. W. Hibbard and University of Michigan X X X Promenetus umbilicatellus (Coek- X X X X field parties have made additional large collections erell). X which yielded amphibians, birds, and mammals, be X X X Marginal pools and wet X X X sides many more mollusks (Herrington and Taylor, mud. X X Moister humus _ _ __ X 1958; Hibbard, 1956; Taylor, this paper; Tihen, 1955). X X Mollusks and a few vertebrates (Hibbard, 1958a) from X Semiaquatie, riparian habi X X a second locality in Kingman County are included in tat. X X X X the Dixon local fauna in this paper. University of Damp humus or grass. X Michigan collections are the basis of the present study. X X X X Fauna. Following is a list of the Dixon fauna: X X X Class Pelecypoda Damp humus of wooded X area. X Order Teleodesmacea X X Sphaerium laoustre (Miiller) X X partumelum (Say) X transversum (Say) X cristata (Pilsbry and Van- X X Pisidium casertanum (Poll) atta). Haicaiia minuscula (Binney) ______X X X compression Prime Helicodiscus singleyanus (Pilsbry). "x" xxxxxxxxx nitidum Jenyns X ___- Class Gastropoda X X X Order Mesogastropoda X .... Valvata tricarinata (Say) X Viviparidae, indet. X X X X X Marstonia crybetes (Leonard) Hydrobiidae, indet. Order Basommatophora Carychium exiguum (Say) Horizon A contains a relatively low proportion of Stagnicola caperata (Say) land species and a high proportion of aquatic species, reflexa (Say) especially those characteristic of permanent water. Fassaria dalli (Baker) The local habitat may have been a permanent pond Biilimnea tnegasoma (Say) Anisus pattersoni (Baker) fed by or draining into a stream. Rarity of tempor Gyro-u-lus parvus (Say) ary pond and land snails may indicate that the lo Helisoma anceps (Menke) cality is well out from the former shore. trivolvis (Say) DESCRIPTION OF LOCAL FAUNAS 37 Class Gastropoda Continued or may be part of the Missler member of the Ballard Order Basommatophora Continued formation. Planorbula armigera (Say) Promenetus kansasensis (Baker) Localities. Both in Kingman County, Kans.: nmMlicatellus (Cockerell) Locality 1: Sl/2 sec. 12, T. 29 S., K. 8 W. Most of the Acroloxus coloradensis (Henderson) material collected by Hibbard and party came from the Ferrissia meekiana (Stimpson) southeast corner of the SW^ (see Frye and Leonard, rivularis (Say) 1952, p. 62, pi. 3, fig. B). The exposure continues into Physa anatina Lea gyrina Say the southwest corner of the SEV4- Additional material skinner I Taylor was collected from the center of the SW^4, on a slope Order Stylommatophora above the northeast side of Red Creek. Locality 2: Strobilops sparsicostata Baker SEV4NE14 sec. 23, T. 30 S., R. 5 W. Gastrocopta armifera (Say) These two localities are believed to represent one chauliodonta Taylor cristata (Pilsbry and Vanatta) fauna on the basis of the mollusks alone. All other pellucida hordeacella (Pilsbry) faunas known from more than one locality which are procera (Gould) reported in this paper are defined by independent stra- scaevoscala Taylor tigraphic or mammalian evidence. tappaniana ( Adams) Table 17 shows the close similarity of the molluscan Pupoides albilabris (Adams) Vertigo milium (Gould) assemblages from localities 1 and 2. Species considered ovata Say especially significant are Acroloxus coloradensis, Bul- Vallonia perspectiva Sterki imnea megasoma, and the viviparid. In the Pleistocene Oxyloma retusa (Lea) of the Great Plains the first two are known only from cf. Sitccinea the Sand Draw and Dixon local faunas, the last from Helicodiscus parallelus (Say) singleyanus (Pilsbry) the Dixon local fauna only. It should be emphasized Deroceras aenigma Leonard that the collection from each locality gives independ Nesovitrea electrina (Gould) ently a similar and unusual ecological picture. Bar Hawalia minuscula (Binney) ring extinct species, the fossil land snail fauna is closely Zonitoides arboreus (Say) similar to that living in southern Kansas; the fossil Class Amphibia Order Caudata and Recent freshwater assemblages are markedly dis Ambystoma tigrinum (Green) tinct, however. Class Aves Mammals from locality 2 are few in number. If this Undescribed bird remains locality is correlative with locality 1, as the mollusks Class Mammalia suggest strongly, then the association of Sigmodon Order Insectivora Sorex dixonensis Hibbard with so many shrews is remarkable. Furthermore, the leahyi Hibbard occurrence of another species of shrew at locality 2, in sp. addition to the four known from locality 1, suggests Blarina sp. peculiarities of sampling error or only an approximate Cryptotis kansasensis Hibbard correlation between the two localities. Order Rodentia Citellus sp. Age. Faunal correlation and ecologic inferences in Geomys sp. dicate a latest Nebraskan or earliest Aftonian age Castoridae, indet. (Hibbard, 1956). Pliophenacomys meadensis Hibbard, Sigmodon sp. Pliopotamys meadensis Hibbard, Pliolemmus antiquus Peromyscus sp. Hibbard, and Gastrocopta chauliodonta Taylor are all Pliophenacomys meadensis Hibbard Pliopotamys meadensis Hibbard species known only from the Deer Park, Dixon, Sand Pliolemmus antiquus Hibbard Draw, and Sanders local faunas, of Nebraskan and Synaptomys r Inker i Hibbard Aftonian age. Ecologic interpretation of the mollusks Order Carnivora and mammals tends to favor a late Nebraskan age, but Mustelidae, indet. the salamanders suggest Aftonian age. The fauna is Occurrence. The Dixon local fauna is known only therefore considered latest Nebraskan or earliest from Kingman County, south-central Kansas. Aftonian. Stratigraphy. Neither the top nor the base of the Paleoecological indications (see below) are that the fine silt unit containing the Dixon local fauna is ex fauna lived in a climate with summers similar to or posed at the localities examined. On the basis of the slightly cooler than those of today, and with runoff age indicated by the fauna, the deposit is equivalent to considerably greater from winter precipitation. The 38 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS climate was not as markedly glacial as that of the water, in the Dixon local fauna. The terrestrial com Cudahy, Berends, and Bar M local faunas, nor as mild ponents of the two assemblages, however, are not as that of the Sanders or Jinglebob local faunas. Sum significantly different. In this case sampling error is mers were probably not quite so cool as those under probably not important, for the Dixon local fauna is which the Sand Draw local fauna lived, but this dif known from two fairly well sampled localities repre ference can be explained by latitude. The Dixon local senting several habitats. The absence of such char fauna seems to represent the latest Nebraskan glacial acteristically northern land snails as Pupilla, Vallonia age, when the climate had already begun to aproach gracilicosta, Discus, and some species of Vertigo is that of today. probably real. Frye and Leonard (1952, p. 148-151) distinguished The occurrence in the Dixon local fauna of some the Dixon local fauna from the Kansan Cudahy fauna. freshwater snails of generally northern distribution in They confused the late Pliocene Saw Rock Canyon and North America may be explained by their habitat in Rexroad local faunas with it, however, and gave all summer-cool, perennial water bodies. Other fresh three a Nebraskan age. This procedure was based upon water species which do not range as far west as King 12 supposed index fossils, 5 of which were found as man County also indicate a greater variety of aquatic sociated with Nebraskan till. Occurrence of these habitats in the early Pleistocene. The evidence of the species is shown in Table 16. land snails suggests that the markedly northern snails Valvata tricarinata and Physa skinneri indicate the TABLE 16. Occurrence of supposed Nebraskan index fossils persistence of cool water habitats during the summer, [After Frye and Leonard, 1952, p. 151] rather than markedly cooler summer climate. The Fauna and locality close similarity of the land snails to those living locally tends to suggest a climate not unlike that of Kingman Nebras Species Saw K ock Rexroad Dixon kan till County today. Canyon local local and local fauna fauna David The sources of water for the perennial streams which fauna City gravel contained the relatively diverse fresh-water molluscan assemblage of the Dixon local fauna are speculative. X X X X X X Presumably the mean annual precipitation did not X X X X X X X differ greatly in amount from that today, for then a X X X X X X X larger, more diverse land snail fauna would be ex X X X pected. Perhaps greater annual snowfall in the X X X X X X Rocky Mountains or the melting of glaciers were con 11. Vertigo hibbardi- ...... X X X X X X X tributing factors. The Dixon local fauna evidently represents a later stage of deglaciatioii than do the Cudahy, Berends, Numbers 1-6 are considered synonyms of living Butler Spring, or Bar M local faunas, for these as species. Number 7 is believed to be not the living Poly- semblages suggest much cooler, moister summers. It is gyra mooreana but an extinct species known only from most similar to the Sand Draw local fauna, and dif the Rexroad and Bender local faunas. Numbers 8 and ferences between the two can be explained by latitude 9 range from the late Pliocene Saw Rock Canyon local and sampling error. fauna to the Aftonian Sanders local fauna. Numbers Habitat: Table 17 summarizes the habitat indications 10 and 11 are believed restricted to the upper Pliocene; of the mollusks of the Dixon local fauna. At locality the record of 11 in the Dixon local fauna is considered 1 the area of deposition was probably close to two erroneous. Number 12 occurs in the upper Pliocene freshwater habitats: a permanent slow-moving stream and Nebraskan. The extensions in range of these or slack-water stretch, probably of at least moderate species are due to two chief reasons: difference in tax- size, and shallower, quiet water near shore, with dense onomic evaluation (whether the fossils are different vegetation. The more abundant land snails taken as a from other species or not), and use of vertebrate fossils group suggest a streamside situation of damp grass and for age determination. humus with some dead wood, but no wooded area. The Paleoecology. Climate: Mollusks of the Dixon local species characteristic of woodland are so much rarer fauna are compared with those living in Kingman that this habitat was either not important at the time, County today in table 18. The most significant and or else some distance upstream. This inference is er obvious difference is the much greater number of fresh roneous to an uncertain degree, however, because of water species, especially those requiring permanent the larger size of the species in question, whose shells DESCRIPTION OF LOCAL FAUNAS 39
TABLE 17. Habitat indications of mollusks of Dixon local fauna had Physa, Fossaria, Gyraulus, Promenetus unbilica- [A, abundant (>250); C, common (41-250); S, scarce (8-40); R, rare (1-7)] tellus, and Pisidium casertanum. On shore was some woodland, and damp grassy areas with dead wood and Abundance Habitat Species leaf mold supporting many small land snails. Loc. 1 Loc. 2 Locality 2 was generally similar to locality 1, except that the fresh water and land faunas were not quite Permanent water: Acroloxus coloradensis (Hender- R R son). so diversified. The more common fresh water snails C S S are those which live in subpermanent or temporary C R R water bodies. The land snail assemblage at locality 2 Pisidium compressum Prime.--... C R is like that of locality 1 in composition and in relative C R abundance of species, but suggests a little more grass S R R and less woodland. R C The local habitat was probably a side-channel, back Permanent to temporary water: water, or oxbow pond which dried up partly in the Permanent or subperma- R C nent. summer. The reeds and submerged vegetation sup C R C ported many Gyraulus, Helisoma, Planorbula, Pro Promenetus kansasensis (Baker) . . . A C Permanent or temporary- A A menetus, Stagnicola, Physa, and Ferrissia. In the C C Temporary pond or marginal R pool. Promenetus umbilicatellus (Cock- S R TABLE 18. Comparison of mollusks of Dixon local fauna with erell). those living in Kingman County, Kans., by habitats S C A A [Recent list based on Franzen and Leonard (1942) and personal collections] Marginal pools and wet mud. S R Semiaquatic, riparian habitat- S S R Habitat Species Dixon Recent C R C S Oastrocopta tappaniana (Adams) -. C S Permanent water: Damp humus of wooded area. S Acroloxus coloradensis (Henderson) . X R X R X R X X Damp to dry ground- ____--__ S X cristata (Pilsbry and Vanatta) . S R X R S X Hawaiia minuscula (Binney) ._. _ C c X Helicodiscus singleyanus (Pilsbry) . R A X Pupoides albilabris (Adams). _ _. S S X S X c R X Ferrissia meekiana CStimpsoii) .... R C X S Permanent to temporary pellucida hordeacella (Pilsbry) . R water: R Permanent or subper X X s S manent. X X R S X R Promenetus kansisensis (Baker) ... X Permanent or temporary. - X X X X Temporary pond or marginal X would not be transported as readily as those of smaller pool. Promenetus umbilicatellus (Cock- X erell). forms. X X The mammals from Dixon local fauna locality 1 were Temporary pond or stream. .. Stagnicola bulimoides techella X (Haldeman) studied by Hibbard (1956, p. 18). He stated "The 3 Marginal pools and wet mud. X X species of Sorex, Blarina, and the 4 species of micro- Semiaquatic, riparian hab X itat. X tines indicate a marshy habitat along the water edge, X X X X X and Blarina indicates the presence of trees and shrubs." X X X Thus the local habitat at locality 1 may have been as X Damp humus of wooded area- X X follows: A small to medium-sized river meandered X X slowly across the plain, depositing fine sand and silt. X X The quiet but more open water 2-4 feet deep had a X X X X molluscan assemblage dominated by BuJimnea mega- cristata (Pilsbry and Vanatta). X X X soma and Helisoma anceps. Driftwood, reeds, and Hawaiia minuscula (Binney) ------X X Helicodiscus singleyanus (Pilsbry) -- X X mussel shells in slow to moderate current furnished a X X X suitable habitat, for Ferrissia. The dense growth of Gastrocopta chauliodonta Taylor .... X pellucida hordeacella (Pilsbry)-. X vegetation such as Potamogeton and MyriophyUum in X X X thick patches here and there supported abundant X X Physa, Promenetus, Gyrcuulm, and Stagnicola refiexa. Ferrissia meekiana (Stimpson)...-- X X Shallow embayments and marginal pools near shore Phvsa skinneri Tavlor. . . ------40 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS deeper parts of this quiet water, and especially toward Class Mammalia Continued the nearby stream, lived Bulimnea megasoma, Helisoma Order Carnivora anceps, and Valvata. Soft mud furnished a suitable Mustela sp. Order Proboscidea habitat for the common /Sphaerium partmneium. The Mastodont, indet. vegetation on shore was grass and shrubs with some Order Artiodactyla trees, among which many small land snails lived. Camelidae, indet. Order Perissodactyla SANDERS LOCAL FAUNA Nannippus phlegon (Hay) Plesippits simplicidens (Cope) Previous work. This fauna was discovered in 1953 Occurrence. The Sanders local fauna is known only by C. W. Hibbard and a University of Michigan field from Meade County, southwest Kansas. party. Collecting in 1953 and 1954 yielded mollusks, Stratigraphy. The fauna occurs in the Missler amphibians, birds, and mammals (Hibbard, 1956; member of the Ballard formation, above the caliche. Tihen,1955). Hibbard (1956, p. 150-151; 1958, p. 56) has published Fauna. Following is a list of the Sanders local measured sections including localities of the Sanders fauna: local fauna. Class Gastropoda Localities. All the following localities are in Meade Order Basommatophora County, Kans. Stagnicola bulimoides tecliella (Haldeman) caperata (Say) Locality 1 (University of Michigan UM-K1-53) : Center of sp. south edge NW%NE& sec. 18, T. 32 S., R. 28 W. Exposure Fossaria dalli (Baker) on tributary of Spring Creek, where Ballard formation un- Gyraulus parvus (Say) conformably overlies Rexroad formation. Rexroad local Promenetus kansasensis (Baker) fauna locality 1 (Kansas University locality 25) is in this umMlicatellns (Cockerell) exposure also. PJiysa anatina Lea Localities 2 and 3 (University of Michigan UM-K2-53) : skinner i Taylor Near center of sec. 23. T. 32 S., R. 29 W. Exposures of gray Aplexa Jiypnorum (Linnaeus) silt above a tributary of Spring Creek. The two fossiliferous Order Stylommatophora localities are only a few tens of feet apart and at the same Ga,strocopta chauliodonta Taylor stratigraphic horizon. All the vertebrates have been grouped crlstata (Pilsbry and Vanatta) as coming from one locality. The more numerous shells paracristata Franzen and Leonard were kept separate to determine the extent of local environ tappaniana (Adams) mental differences evidently minor. (See table 19.) Vertigo milium (Gould) Age. The Sanders local fauna is believed to be of ovata Say Pupoides albilabris (Adams) late Aftonian age from faimal evidence, ecologic in Pupillidae, new genus and species? ferences, and stratigraphic position. The faunal evi Vallonia perspectiva Sterki dence from the restricted ranges of extinct groups cf. Succinea comes primarily from the mammals. They indicate Helicodiscus singleyanus (Pilsbry) a Nebraskan or Aftonian age. Gastrocopta chaulio- Deroceras aenigma Leonard donta* otherwise known only from the Sand Draw and Haicaiia minuscuJa (Binney) Dixon local faunas, also suggests an earliest Pleisticene Class Amphibia Order Caudata age. Ambystoma tigrinum (Green)? Ecologic inferences from the fauna suggest a meso- Class Aves thermal, subhumid climate with seasonal extremes Undescribed bird remains much less marked than those of today. The fauna as Class Mammalia a whole contrasts with the glacial assemblages which Order Insectivora Sorex sandersi Hibbard have a marked element of northern species. The in Order Rodentia ferred climate is intermediate between that of the G-eomys tobiitemiis Hibbard glacial intervals and the warm-temperate, semi arid Perognathuft cf. P. pcarlettensis Hibbard climate suggested by the caliche horizon. For these Prodipodomys sp. reasons the Sanders local fauna is called Aftonian. tiigmodon cf. #. intermedium Hibbard The stratigraphic position of the fauna in the upper Bensonomys meadentsis Hibbard Plioplienacomys meadensis Hibbard part of the Ballard formation, above a buried, massive Pliolemmus antiques Hibbard caliche layer, is interpreted to indicate an age less than Zapus sandersi Hibbard that of some part of the Aftonian. The caliche is be- DESCRIPTION OF LOCAL FAUNAS 41 lieved to be Aftonian, but what part of this interval it rence of the jumping mouse, Zapus, and the abundance represents is uncertain. of microtine rodents also indicates a moister climate Paleoecology. Climate: Both characteristically (Hibbard, 1956, p. 156). northern and characteristically southern faunal ele The former climate in which the Sanders local fauna ments occur in the Sanders local fauna. The most ex lived thus differed from the present by more effective treme contrast in distributions is between those of rainfall, and summers with much less intense hot Physa sMnneri and Stagnicola bulimoides techella. spells. Neither distribution nor amount of annual These species now live hundreds of miles apart, but rainfall was necessarily different. were associated in southwestern Kansas. The joint oc An earlier interpretation of the fauna, based on my currence of these and other species is taken to indicate misidentification of /Stagnicola bulimoides techella, a former reduction of climatic extremes. suggested that winters were considerably warmer than The southern element among the mollusks is com now (Hibbard, 1956, p. 156). The winters may indeed posed of species which in the Great Plains are re have been warmer than now, but the mollusks give no stricted to or characteristic of the southern part. These evidence for this. are Stagnicola l>ulimoides techella, (raxtrocopta cris- Habitat. Table 19 summarizes habitat indications of fata, Vallonia perspective!, and Helicodiscus single- the mollusks of the Sanders local fauna. All three lo yanus. A moderately northern element is composed of calities seem to represent the same situation: shallow Stagnicola caperata, Promenetus umbilicatellus, and sub-permanent ponds or stream backwaters with sur Aplexa, hypnorum, which range south in the Great rounding grassy areas. Discus, Euconulus, Helico Plains as far as Nebraska. The most striking element discus parallelus, Retinella, Zonitoides, and other is Physa sMnneri, which is widespread in the plains of forms usually associated with damp humus around southern Canada but unknown in the United States. dead wood are absent. This fact suggests that wood This association of species is different from that land, if present, was not all well developed, and recalls known in any other Blancan fauna. Southern and the hay meadow regions of north-central Nebraska. mid-northern species are known in the late Pliocene Here large areas of grassland have streams and ponds faunas, but with no markedly northern forms. Glacial each spring, and trees are infrequent. faunas have a much reduced southern element, or none TABLE 19. Habitat indications of mollusks of Sanders local fauna at all, and many northern species. Such a peculiar [A, abundant (>250); C, common (41-250); S, scarce (8-40); R, rare (1-7)] combination of elements is shown also by the mammals I jocalit Y of the Sanders local fauna. Hibbard (1956, p. 156) Habitat Species noted that there are some southern types of mammals: 1 2 3 the cotton rat (Sigmodon}, Bensonomys, and a kanga Permanent to temporary roo rat (Prodipodomys}. These, and the relative water: S R R scarcity of shrews, suggest a wrarm, interglacial climate, manent. Promenetus kansasensis (Baker) . "R" R R R but the cricetine rodents would then be expected to be R R R Temporary water: common. "The paucity of cricetine rodents may be due Temporary pond or mar- Aplexa hypnorum (Linnaeus) _ s c S C in part to the habitat, but it is in contrast to their Promenetus umbilicatellus (Cock- A R S erell). abundance in the warm-temperate Rexroad fauna" Temporary pond or Stagnicola bulimoides techella S R C stream. (Haldeman) . (Hibbard, 1956, p. 156). R A R A The local lenses of grayish organic muds in which R C R S the Sanders local fauna occurs suggest small, shallow A C A Vanatta) . ponds or a slow-moving, unentreiiched stream. The Hawaiia minuscula (Binney)___- A S A Helicodiscus singleyanus (Pils- S R aquatic snails and land snails of moist habitats indicate bry). Pupoides albilabris (Adams). _- -_ A s S permanent moisture and subpermaneiit standing water. C R S S Hence, the Sanders local fauna probably lived in and paracristata Franzen and S R Leonard. near shallow annual ponds or else a small, slow-moving R R R stream of local origin. The absence of genera usually C S C Vallonia perspectiva (Sterki) C R R associated with bushes or trees bordering such a habitat further suggests seasonal grassland ponds or streams. Neither of these situations necessarily implies greater DEER PARK LOCAL FAUNA rainfall than that of southwestern Kansas at the pres Previous work. Collections at the Deer Park local ent time, but only greater effectiveness of precipitation. ity have been made by C. W. Hibbard and parties Milder summers might well account for this. Occur from the University of Kansas (1936, 1944) and Uni- 42 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS versity of Michigan (1954). The mammals now in and polished bone fragments, the greater abundance cluded in the Deer Park local fauna were originally of harder enamel teeth than bones, occurrence of the referred to the Rexroad local fauna (Hibbard, 1938, relatively strong slug shells and no other mollusk re 1941a), but later recognized as coining from a higher mains, and the similarity of the deposit to the numer stratigraphic horizon (Hibbard, 1949). A number of ous artesian springs of Meade County today all indi vertebrates but only one mollusk have been discovered cate that the Deer Park quarry is in deposits of a (Hibbard, 1956). former artesian spring system. "The habitat in the Fauna. Following is a list of the Deer Park local neighborhood of these springs can be considered to fauna: have been similar to that characteristic of the same Class Gastropoda springs in Meade County today. The area of dis Order Stylommatophora charge is a death trap of boiling quick-sand, and the Deroceras aenigma Leonard entire basin is a bog, in places choked with water Class Reptilia cress and other aquatic plants that cover outlets of Order Chelonia a part of the earlier dendritic pattern. The basin Testudo, large species Order Squamata grades into a marsh that supports sedges and asso Snake, indeterminate ciated vegetation. Around the edges of the marsh Class Aves and along the banks of the outlet area are trees, shrubs, Birds, indet. and grapevines. This habitat furnishes not only a Class Mammalia home for rodents and shrews but also a watering and Order Rodentia Cynomys meadensis Hibbard feeding place for the larger grazers and browsers and Citellus sp. for some carnivores. The remains consist chiefly of Geomys quitnri McGrew those of horses, mastodons, camels, and peccaries. The Procastoroldes sweeti Barbour and Schultz large grazers and browsers had probably ventured out Ogmodontomys sp. or Mimomys (Cosomys) sp. too far from the edge of the basin while feeding" Pliopotamys meadensis Hibbard (Hibbard, 1956, p. 155-156). The slug Deroceras Pliolemmus antiquus Hibbard Order Carnivora aenigma probably lived among the semiaquatic vege Canis sp. tation around the spring, along with many other snails Mustelidae, indet. whose shells have not been preserved. Taxidea cf. T. taxus (Schreber) "The climate at the time this fauna lived is regarded Order Proboscidea as interglacial because of the presence in it of the large Stegomastodon sp. Rhynclwtherium sp. land turtle Testudo, which could have inhabited the Order Lagomorpha region only during a mild-temperate time" (Hibbard, Hypolagus sp. 1956, p. 156). Order Artiodactyla Platygonus sp. RECENT MOLLUSKS FROM NORTHERN NEBRASKA Camelidae, 2 indeterminate species Order Perissodactyla In the summer of 1953 R. D. Mitchell, A. L. Lamb, Nannippus phlegon (Hay) and D. W. Taylor collected Recent mollusks at locali Plesippus simplicidens Cope ties in north-central Nebraska while working on the Occurrence. The Deer Park local fauna is known Sand Draw local fauna. They are of interest in com only from Meade County, southwest Kansas. parison with the Sand Draw local fauna, and also Stratigraphy. The fauna occurs in the Missler because mollusks from the area have not been reported member of the Ballard formation (Hibbard, 1949, before. 1956). LOCALITIES Localities. Only one locality is known, in Meade Cherry County. Nebr.: County State Park, near the west edge SE*4 sec. 15, 1. Fort Niobrara National Widlife Refuge, 5 miles east of Valentine. Pond and seepage area at old fort T. 33 S., R. 29 W., Meade County, Kans. (University reservoir. of Kansas Meade County loc. 1). 2. North side Niobrara River flood plain % miles west Age. Ecologic interpretation and the stratigraphic of U. S. Highway 20. Nearly filled old oxbow lake position of the Deer Park local fauna indicate an with seepage and slight current. Aftonian age. 3. Sand hills pond on west side U. S. Highway 83, 7.4 miles south of junction with U. S. Highway 20. Paleoecology. The occurrence of pockets and tubes Temporary. of flour sand within the surrounding unconsolidated 4. Small creek and tributary drainage ditches 4 miles silt and silty clay of the Missler member, badly worn northeast of Simeon. Subpermanent water. RECENT MOLLUSKS FROM NORTHERN NEBRASKA 43 Cherry County, Nebr. Continued water mollusks are removed from the large streams 5. Creek at junction of dirt road and paved road to by flooding, but are abundant in small spring-fed Alkali Lake, 4.4 miles west of junction with U. S. creeks, seepages, and upland ponds and ditches with Highway 83. 6. South side Red Deer Lake. Permanent water. no flood scour. 7. Lake on west side U. S. Highway 83, 20 miles north Sand Hills. The Sand Hills region is a huge area of Thedford. Probably temporary. of western Nebraska covered by shifting sand loosely Brown County, Nebr.: held by dune grass. The hills so formed vary in shape 8. Wooded draw on south bluffs Niobrara River, on and size, but are similar in having only sparse grass State Highway 7, 0.4 mile south of bridge across river. growing on them. A large proportion of the hollows 9. Drainage ditches in northwest corner sec. 34, T. 30 in this area contain shallow ponds, many large enough N., R. 22 W. Temporary water. to be called lakes. Permanence is proportional to 10. Drainage ditch at northwest corner sec. 10, T. 29 N., size; in the worst drouth years all but the largest R. 22 W. Water probably permanent, but ditch dug lakes dry up. This large area is almost devoid of recently. 11. Seepages along stream in wooded draw 50 yards south land snails, and the freshwater fauna is drastically of Long Pine Creek at bridge 2% miles south of reduced by the temporary nature of the water bodies. Long Pine. Stagnicola caperata, S. palustris, and Physa gyrina 12. Calamus River .5 mile west of State Highway 7. occur in nearly every pond; rich localities have 7 or Seepages along south bank. 8 species. Rock County, Nebr.: Most of the Sand Hills area is undrained, but some 13. Drainage ditches along U. S. Highway 20, 2.8 miles west of Bassett. medium-sized streams flow from the area. The small 14. Pond on north side U. S. Highway 20, 2.3 miles east creeks are but little more permanent than the ponds, of Bassett. but the rivers are perennial. The streams draining Hooker County, Nebr.: this region cannot flood, for rain sinks into the sand 15. Middle Loup River 5 miles east of Mullen. as soon as it falls. There are thus no accumulations Thomas County, Nebr.: 16. Stock tank beside State Highway 2, 1 mile west of of flood debris along the banks. There is essentially Seneca. no mud or gravel anywhere, for the area is all sand. 17. Middle Loup River at south side Thedford. The only mud seen was in seepage areas along the 18. Middle Loup River, 1 mile east of Seneca. rivers, where watercress and other semiaquatic plants have produced organic debris. As in the High Plains HABITATS proper, deciduous woodland grows in the river valleys. The area studied lies on the northeast side of the Almost all of the few land snails occur in this habitat. Sand Hills region of western Nebraska, and thus The habits are tabulated below, and the mollusks includes two physiographic types, the Sand Hills and occurring in them listed in the following section. the High Plains proper. Each of these is discussed I. Aquatic habitats. below. A. Permanent water bodies. High Plains. The part of the region collected in 1. Marginal stations. At the edge of ponds, snails is typical: flat land sloping gently toward the Mis occur in a semiaquatic position, frequently as souri River. Grassland is broken rarely by an occa much out of water as in it. 2. Quiet water. sional cottonwood tree. This flat surface is largely a. Lake. Red Deer Lake (loc. 6) was the only a plain of alluviation, probably completed sometime one of this type sampled. The fauna does in the middle Pleistocene. The Mobrara River is not differentiate it from, the temporary ponds. incised well into this mantle, and its tributaries are b. Backwaters or sheltered spots along rivers. cutting rapidly headward into the poorly consolidated This habitat is peculiar to the Sand Hills for elsewhere each flood scours the banks. sediments. The widespread sands and gravels form c. Oxbow lake. The only such station visited ideal aquifers; streamside patches of springs and (loc. 2) has a permanent water supply from seepage areas are abundant. Well-developed decidu seepage through the valley fill, but is not ous woodland occurs in these valleys wherever ade cooled by it. d. Spring-fed pond. At locality 1 spring waters quate water is available; pine stands cover the drier of 15 °C. were ponded. The summer-cool slopes. Drainage and irrigation ditches on the undis- water is believed responsible for presence of sected upland dry up toward the end of summer. They some species in the area. often connect with ponds, also temporary. In this 3. Flowing water. At numerous places along streams cut into the High Plains surface small springs region land snails are largely restricted to stream val and seepages occur. The mollusks are usually leys where cover and moisture are available. Fresh among the mud and water cress. 44 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Vertigo ovata (Say), IIAl(ll) B. Subpermanent water bodies. Vallonia gracilicosta Reinhardt, IIB(1, 8) 1. Ponds. These dry up only in drier years. parvula Sterki, IIB(1, 8) 2. Sand Hills creeks. Quiet spots along such streams Cionella lubrica (Muller), IIB(8) are more nearly permanent than most Sand cf. Succinea, IIA3(7) Hills ponds, which they drain. Succinea ovalis Say, I1B(8) C. Temporary water bodies. Oxyloma retusa (Lea), IIAl(ll), IIA2(1, 2, 15, 18) 1. Ponds. This category includes the temporary Discus cronMiitei (Newcomb), IIB(2,11, 15) Sand Hills ponds as well as small ponds along Punctum minutissimwn. (Lea), IIAl(ll) ditches outside the Sand Hills. Locality 16 is Deroceras laeve (Muller), IIAl(ll), IIB(1, 2, 15) a metal stock tank. Euconulus fulvus (Muller), IIB(2, 8, 11) 2. Ditches (irrigation and drainage). These fill up Nesovitrea electrina (Gould), IIB(2,11, 15) after heavy rains but because of their low Hawaiia minuscula (Binney), IIB(1, 15) gradient flow only slowly and do not flush out Zonitoides arboreus (Say), IIB(1, 2, 8, 11, 15) the snails. Stenotrema leal (Binney), IIB(2) II. Terrestrial habitats. A. Marginal stations. SYSTEMATIC DESCRIPTIONS 1. Moist leaves around seepages. 2. Riparian station. Oxyloma was almost always Dates of genera and species given in the center within a few inches of permanent water, crawl headings have not been personally verified except in ing about on water cress or logs. the case of extinct species. Sources are Pilsbry (1940, 3. Lake shore. At locality 7 SSuccinea was found 1946, 1948) for the land snails and Baker (1911, 1928, on the wet shore several feet from the water's 1928a) for the fresh-water mollusks. edge among sparse tall grass. B. Damp ground. Among slightly damp leaves or on soil References are intended to include all records of under timber or rocks. mollusks from the faunas considered, as well as the minimum number of general references, which were SPECIES FOUND AT THE NOBTHERN NEBRASKA selected to indicate current taxonomic treatment and LOCALITIES summaries of other references and of distribution. The first symbol of letters and numbers indicates Diagnoses (and usually descriptions) are given for the habitat type, the following numbers in parentheses all the extinct species restricted to the Blancan age, the localities from pages 42-43. and for most of the other extinct species as well. Distribution is based on the most reliable source SpJiacrium lacustre (Miiller) form ryckholti (Normand) IA3 known. The planorbid distributions without source (11), IC1 (3) securis Prime, IBK14) indication are based upon my own examination of Pisidhtm casertanum (Poll), IA2c(2), IA3(11, 18) specimens, chiefly those in the University of Michigan VaJvata tricarinata (Say), IA2d(l) Museum of Zoology. Carychium exiguum (Say). IIAl(ll) Ecology is largely from published data. Personal Stagnicola palustris (Muller), IA2a(6), IB2(4), IC1(3, 7), observations made in northern Nebraska are added for IC2(13) caperata Say, IB2(4), IC1(3, 7), IC2(9, 13) comparison and supplement. Fossaria obrussa (Say), IA1U, 15, 17), IA3(11, 12) Occurrences cited are restricted to the faunas con daUi (Baker), IA2c(2), IA3U1) sidered here. Numerous late Pleistocene records, for Armiger crista (Linnaeus). IB2(5) example, are not included. Gyraulus circumstrtatus (Tryon). IA3(11, 12), IB2(4), Also under "Occurrences" are the abbreviation of parrits (Say), IA2a(6). IA2d(l), IB1(14), IB2(5), IC1(3), the name of the institution, catalog number, and num IC2(13) ber of specimens represented. This last number is not deflect us (Say), IA2d(l) the number of fragments examined, but the minimum Hclisoma anccpa (Menke), IA2d(l) number of shells required to furnish the fragments. trivolris (Say), IA2a(6), IA2d(l), IB1(14), IC2(13) Abbreviations are as follows: ANSP, Academy of Planorbula armigera (Say). IA2c(2) Natural Sciences of Philadelphia; CNHM, Chicago Promenetus exacuous (Say). IBK14), IB2(5) umbilieatPllus (Cockerell), IC1(3, 7), IC2(9) Natural History Museum; UTMNH, University of Physa anatina Lea, IA2d(l), IA3(1, 11, 12, 18) Illinois Museum of Natural History; UKMNH, Uni gyrhta Say, IA2a(6). !A2b(17), IB1(14). IB2(4, 5), IC1 (3. versity of Kansas Museum of Natural History; 7, 16), IC2(10, 13) UMMZ, University of Michigan Museum of Zoology; Aplexa Inipnornm (Linnaeus), IC2(9, 13) USGS, United States Geological Survey; USNM, Gastrocopta armifera (Say), IIB(l) crlstata (Pilsbry and Vanatta), IIB(8) United States National Museum. holsinfferi (Sterki), IIB(l) H. B. Herrington identified the Sphaeriidae and tappaniana (Adams), IIAl(ll), IIB(l) contributed notes on the specimens. The data in this SYSTEMATIC DESCRIPTIONS 45 paper are based on his notes and Herrington and donta. Thickness of shell and geographic location Taylor (1958). make A. grandis the most reasonable specific alloca Taxonomic changes: summarized below are the new tion. taxa proposed, and the disposition of species synony- Material from Sand Draw locality 2 is indeter mized or relegated to different genera. minate, and only presumptively referred to the same New: species. Phreatomenetus n. subgen.; type Promenetus umMlicatel- Occurrence and material: Sand Draw local fauna. West lus (Cockerel!) Fork of Deep Creek, UMMZ 184321 (1) ; loc. 2, UMMZ 184167 Gastrocopta (s. s.) fransenae Taylor, n. sp. (1). (s. s.) scaevoscala Taylor, n. sp. Family ftUADRULIDAE Polygyra rexroadensis Taylor, n. sp. Changed in status: Subfamily LAMPSILINAE Amnicola walkeri Pilsbry = A. lustrica Say CarycJiium perexiguum Baker = C. exiguum (Say) Genus LIGTTMIA Swainson, 1840 Lymnaea macella Leonard = Stagnicola exilis (Lea) ?Ligumia subrostrata (Say), 1831 parexilis Leonard = Stagnicola exilis (Lea) diminuta Leonard = Stagnicola bulimoides techella Lampsilis subrostrata (Say). Isely, 1925, Oklahoma Acad. (Haldeman) Sci. Proc., v. 4, p. 108. turritella Leonard = Fossaria dalli (Baker) Lampsilis subrostrata Say. Frierson, 1927, Check list of North Gyraulus enaulus Leonard = G. parvus (Say) American naiades, p. 77. Menetus kansasensis Baker transferred to Promenetus Promenetus blancoensis Leonard = P. umbilicatellus Remarks. Dr. Henry van der Schalie identified a (CockereU) fragmentary valve showing beak sculpture and worn Planorl>is circumlineatus Shuttleworth assigned to Pro pseudocardinal teeth as probably either Ligumia sub menetus rostrata (Say) or Uniomerus tetralasmus (Say), of santacrusensis (Germain) assigned to Promenetus aemginosus Morelet assigned to Promenetus which the former seems more likely. Vertigo (Angustula) Mbbardi Baker transferred to Occurrence and material: Rexroad local fauna. Rexroad Vertigo s. s. loc. 3, UMMZ 183025 (1/2). Angustula Sterki, subgenus considered of minor rank and synonymized with Vertigo s. s. Order TELEODESMACEA
Class PELECYPODA Family SPHAERIIDAE
Order PRIONODESMACEA Genus SPHAERITTM Scopoli, 1777
Family TTNIONIDAE Sphaerium partumeium (Say), 1822
Subfamily ANODONTINAE Musculium partumeium (Say). Baker, 1928, Wisconsin Geol. Nat. History Survey Bull. 70, pt. 2, p. 354, pi. 99, fig. Genus ANODONTA Lamarck, 1799 24-26. Sphaerium partumeium (Say). Herrington and Taylor, 1958, Subgenus PYGANODON" Crosse and Fischer, 1894 Michigan Univ. Mus. Zool. Occ. Papers 596, p. 7. Musculium truncatum (Linsley). Baker, 1928, Wisconsin ?Anodonta (Pyganodon) grandis Say, 1829 Geol. Nat. History Survey Bull. 70, pt. 2, p. 356, pi. 99, Anodonta grandis Say. Frierson, 1927, Check list of North fig. 21-23. American naiades, p. 14. Distribution. "Most of the United States; less com Baker, 1928, Wisconsin Geol. Nat. History Survey Bull. 70, mon in mountainous areas. Southern Canada from pt. 2. p. 152, pi. 62, fig. 5. Saskatchewan to Quebec" (Herrington and Taylor, Anodonta gigantea Lea. Baker, 1928, Wisconsin Geol. Nat. 1958, p. 8). History Survey Bull. 70, pt. 2, p. 161, pi. 62, fig. 4; pi. 65, fig. 3. Ecology. "Ponds and eddies in rivers where there is considerable vegetation and a soft bottom" (Her Remarks. Fragments referred to this species were rington and Taylor, 1958, p. 8). found as float in the West Fork of Deep Creek, sees. Remarks. None of the shells from the Sand Draw 11 and 14, T. 31 N., R. 23 W., Brown County, Nebr. local fauna is complete, but several are nearly so. They They are probably from the same stratigraphic posi compare well in every feature with a series (Herring- tion as the other material collected on Deep Creek, ton collection S-658) of S. partumeium from Baton but a more precise locality cannot be given. One frag Rouge, La., except that the Baton Rouge specimens ment shows enough of the adductor and retractor are somewhat larger but they are also larger than muscle scars to indicate it probably represents Ano- the usual northern specimens. The shells from the 508945 O 60 5 46 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Dixon local fauna are identified certainly, even though Sphaerium .striatinum (Lamarck), 1818 most are only fragments. Sphaerium striatinum (Lamarck). Baker, 1928, Wisconsin Occurrence and material: Sand Draw and Dixon local Geol. Nat. History Survey Bull. 70, pt. 2, p. 334, pi. faunas. Sand Draw loc. 1, UMMZ 186369 (77/2). Dixon loc. 98, fig. 1-5. 2, UMMZ 191501 (114/2), 191502 (17/2). Herrington and Taylor, 1958, Michigan Univ. Mus. Zool. Occ. Papers 596, p. 9. Sphaerium lacustre (Miiller), 1774 Distribution. Almost all of North America, from form ryckholti (Normand), 1844 Great Slave Lake, Northwest Territories, to Panama. Musculium ryckholti (Normand). Baker, 1928, Wisconsin Ecology. Perennial water bodies with some current Geol. Nat. History Survey Bull. 70, pt. 2, p. 359, pi. 99, action are suitable habitats. It lives in large lakes or fig. 6-9. small ones, in rivers or small streams, in a bottom of Musculium jayense (Prime). Baker, 1928, Wisconsin Geol. gravel, sand, or mud. Ponds, lagoons, and swamps are Nat. History Survey Bull. 70, pt. 2, p. 353, pi. 99, fig. 27, 28. not favorable places, probably because there is insuffi Musculium rosaceum (Prime). Baker, 1928, Wisconsin Geol. cient current to oxygenate the water well. Nat. History Survey Bull. 70, pt. 2, p. 359, pi. 99, fig. Occurence and material: Saw Rock Canyon and Sand Draw 19, 20. Sphaerium lacustre (Miiller) form ryckholti (Normand). local faunas. Saw Rock Canyon, UMMZ 186408 (1/2), an Herrington and Taylor, 1958, Michigan Univ. Mus. Zool. infant right valve. The shell is the size of an infant recently Occ. Papers 596, p. 8. expelled from the parent. Sand Draw loc. 1, UMMZ 186368 Sphaerium sp. Frye and Leonard, 1952, Kansas Geol. Survey (5/2). The only complete valve from this locality has a re Bull. 99, p. 151 (Kingman County loc.). versed hinge all the hinge characters are those of the right Distribution. Holarctic. In North America from valve, but at opposite ends. This valve perfectly matches some Recent ones of the heavy form in the Herrington collection in the Southern United States to the Northwest Terri shape of shell, striae, and shape of hinge characters, but the tories. This form of lacustre is also found in Europe cardinal is slightly more central. Sand Draw loc. 3, UMMZ and Brazil. 181125 (3/2). A fragment of the anterior end of a right Ecology. "It seems to have a preference for ponds valve has striae which seem too coarse for S. sulcatum. Frag where there is considerable vegetation and even bog ments of two more valves are referred doubtfully to S. ponds that have a bottom of muck, rotting wood, striatinum. grasses, and the like" (Herrington and Taylor, 1958, p. 9). Sphaerium sulcatum (Lamarck), 1818 Remarks. Shells from the Dixon local fauna are Sphaerium sulcatum (Lamarck). Herrington and Taylor, near the form ryckholti in shape. They are a little 1958, Michigan Univ. Mus. Zool. Occ. Papers 596, p. 9. heavier than usual but not more so than some Recent Sphaerium simile (Say). Baker, 1928, Wisconsin Geol. Nat. specimens. None are whole, but a number of frag History Survey Bull. 70, pt. 2, p. 313, pi. 96, fig. 4-7. ments are large and a few nearly whole. In beaks Sphaerium simile planatum Sterki. Baker, 1928, Wisconsin and outline these seem to be lacustre. Geol. Nat. Hist. Survey Bull. 70, pt. 2, p. 317, pi. 96, The reference by Frye and Leonard is attributed to fig. 8-10; pi. 98, fig. 29-33. this species because 8. lacustre is the commonest Sphaerium crassum Sterki. Baker, 1928, Wisconsin Geol. Nat. Hist. Survey Bull. 70, pt. 2, p. 318, pi. 96, fig. 11-13. Sphaerium in the Dixon local fauna. Sphaerium lineatum Sterki. Baker, 1928, Wisconsin Geol. Occurrence and material: Dixon local fauna; Recent in Nat. History Survey Bull. 70, pt. 2, p. 321, pi. 96, fig. northern Nebraska. Dixon loc. 1, UMMZ 186386 (31/2). 18-21. Sphaerium securis (Prime), 1851 Distribution. "Northeastern North America from Musculium securis (Prime). Baker, 1928, Wisconsin Geol. Quebec to Virginia, west to Iowa, Montana, and Al Nat. History Survey Bull. 70, pt. 2, p. 360, pi. 99, fig. berta" (Herrington and Taylor, 1958, p. 10). 10-13. Ecology. S. sulcatum has a preference for a soft Distribution. "East of the Rocky Mountains, but bottom in fairly still waters, in eddies of a creek or apparently rare in the South and wanting in the Gulf river, along shore in lakes, and even in lakes filling States; common in the Northeast; South Carolina"" up with marl. Herrington has found it associated (Baker, 1928a, p. 361). with Pisidium compressum in all these habitats. Ecology. In northern Nebraska it was found in a Occurrence and material: Sand Draw local fauna. Sand small permanent or nearly permanent pond in sandy Draw loc. 1, UMMZ 186370 (3/2). All are good-sized adults, bottom among dense vegetation in shallow water. but somewhat broken. The striae are normal. One left valve Occurrence: Recent in northern Nebraska. has both ends complete but the cardinal area is broken out. SYSTEMATIC DESCRIPTIONS 47
Sphaerium transversum (Say), 1829 Pisidium sp. Baker, 1938, Nautilus, v. 51, p. 131. Hibbard, 1941, Kansas Univ. Sci. Bull., v. 27, p. 95. Musculium transversum (Say). Baker, 1928, Wisconsin Geol. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, Nat. History Survey Bull. 70, pt. 2, p. 351, pi. 98, fig. p. 408. 22-25 [not 22-28]. McGrew, 1944, Field Mus. Nat. Hist. Geol. Ser., v. 9, p. 36. Sptiaerium transversum (Say). Herrington and Taylor, 1958, Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. Michigan Univ. Mus. Zool Occ. Papers 596, p. 8. 151 (Meade County localities). Distribution. "North America east of the Rocky Psidium sp. Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p, Mountains from Mexico to the Northwest Territories1' 265. (Herrington and Taylor, 1958, p. 8). Distribution.. "Almost cosmopolitan: Eurasia, New Ecology. "In rivers, ponds and lakes, usually in Zealand, and Australia; in the Western Hemisphere soft mud, on the surface or buried at different depths" from Patagonia to Alaska" (Herrington and Taylor, (Baker, 1928a, p. 352). 1958, p. 14). Remarks. A large specimen from the Dixon local Ecology. The wide distribution of Pisidium caser fauna seems to be transversum. A dwarf form of this tanum reflects a corresponding adaptability, greater species which occurs in the Sand Draw local fauna than any other American sphaeriid. It lives where any resembles certain living individuals that appear in other species of Pisidium can live, except for deep unfavorable habitats. water. The typical form of the species, with relatively heavy shells, lives in rivers or fairly large creeks. Occurrence and material: Sand Draw and Dixon local Thinner shells with smoother outline are found in faunas. Sand Draw loc. 3, UMMZ 186378 (43/2). Dixon loc. 1, UMMZ 186385 (10/2). ponds, swamps, lagoons, bog ponds, and similar quiet water bodies. P. casertanum is one of the few species Sphaerium species of Pisidium which can tolerate seasonal desiccation, Sphaerium sp. Frye and Leonard, 1952, Kansas Geol. Survey and sometimes it may be the only aquatic mollusk in a Bull. 99, p. 151 (Meade County localities). small, temporary stream or seepage. Sphaerium indet. Herrington and Taylor, 1958, Michigan Remarks. Previous records of Pisidium sp. from Univ. Mus. Zool. Occ. Papers 596, p. 13. the Rexroad local fauna are probably based on P. Remarks. Some specimens from several localities casertanum, for this is the only one in University of are unidentifiable specifically, because they are imma Michigan collections. The original specimens reported ture or too fragmentary. They may well represent by Baker and McGrew from the Sand Draw local the thinner-shelled species, partumeium, lacustre, and fauna have been verified as P. casertanum. transversum. Occurence and, material: Buis Ranch, Saw Rock Canyon, Red University of Michigan collections contain no Corral. Rexroad, Sand Draw, and Dixon local faunas; Re cent in northern Nebraska. Buis Ranch, UMMZ 181119 (1/2). tSphaerium from the Rexroad or Bender local faunas. Saw Rock Canyon, UMMZ 173587 (51/2), 195838 (1 + The material reported by Frye and Leonard from the 1398/2). This series shows much variation in shape, just as SWVi sec. 22, T. 33 S., R. 29 W., Meade County, Kans. one finds in Recent shells. Some of the shells are pitted, evidently by solution. Many are in fragments, but this seems probably came from Rexroad locality 3, for Bender to be the result of removal from the matrix. The specimens locality 1 has yielded few aquatic forms. show no signs of being rolled, for the laterals and cardinals are unbroken. Red Corral, UMMZ 182992 (58/2). These Occiirence and material: Saw Rock Canyon and Sand Draw shells suggest still water. They are like those found in small, local faunas; Rexroad local fauna (Frye and Leonard). Saw slow-moving creeks that go dry for part of the year, or in Rock Canyon, TTMMZ 177514 (1), 177553 (6/2). Sand Draw lakes filling up with marl. Rexroad loc. 2, UMMZ 186388 loc. 2, UMMZ 184168 (1/2); loc. 3, UMMZ 181124 (2/2), (3/2) ; loc. 3, UMMZ 186387 (11/2). Sand Draw loc. 3, 186379 (31/2) ; loc. 4, UMMZ 186373 (8/2) ; loc. 5, UMMZ UMMZ 186377 (8/2). Mostly form roperi. Loc. 4, UMMZ 186374 (18/2), 186366 (263/2); loc. 7, UMMZ 186381 (1/2), 177311 (58/2). Form roperi Sterki. Much broken. Some 177225 (5/2). have "rest periods," like capped beaks at more than one place. Loc. 5, UMMZ 186367 (32/2), 181134 (11/2). Mostly form Genus PISIDITTM Pfeiffer, 1821 roperi Sterki; the others are close. Most have strong "rest periods" or capped beaks. Loc. 6, CNHM PE3439 (2/2). Loc. Pisidium casertanum (Poll), 1791 7, TTMMZ 177226 (25/2). Mostly form roperi Sterki. Dixon Pisidium. casertanum (Poli). Woodward, 1913, Catalogue of loc. 1, UMMZ 186383 (H/2). British species of Pisidium, p. 31, pi. 1, fig. 3-6; pi. 3, Pisidium compressum Prime, 1851 fig. 3; pi. 13-18. Pisidium compressum Prime. Baker, 1928, Wisconsin Geol. Herrington, 1954, Nautilus, v. 67, p. 131. Nat. History Survey Bull. 70, pt. 2, p. 370, pi. 100, fig. 9-13. Hibbard, 1954, Michigan Acad. Sci. Papers, v. 39, p. 342. Herrington, 1954, Nautilus, v. 67, p. 135. Herrington and Taylor. 1958, Michigan Univ. Mus. Zool. Herrington and Taylor, 1958, Michigan Univ. Mus. Zool. Occ. Papers 596, p. 14. Occ. Papers 596, p. 15. 48 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Pisidium sp. Frye and Leonard, 1952, Kansas Geol. Survey Class GASTROPODA Bull. 99, p. 151 (Kingman County loc.). Subclass PROSOBRANCHIA Distribution. "North America, from Mexico to Order MESOGASTROPODA Great Slave Lake, Northwest Territories" (Herrington Family VALVATIDAE andTaylor, 1958, p. 16). Genus VALVATA Miiller, 1774 Ecology. This species inhabits only perennial water Valvata tricarinata (Say), 1817 bodies with some current action, such as lakes, rivers and creeks; it is never found in ponds, swamps, la Valvata tricarinata (Say). Baker, 1928, Wisconsin Geol Nat. History Survey Bull. 70, pt. 1, p. 11, pi. 1, fig. 1-3. goons, or bogs. Taylor, 1954, Michigan Univ. Mus. Zool. Occ. Papers 557, Remarks. The reference by Frye and Leonard is p. 12. attributed to this species because P. compression is La Rocque, 1956, Nautilus, v. 70, p. 13. Valvata tricarinata perconfusa Walker. Baker, 1928, Wiscon the most abundant Pisidium in the Dixon local fauna. sin Geol. Nat. History Survey Bull. 70, pt. 1, p. 16, pi. Occurrence and, material: Sand Draw and Dixon local 1, fig. 4. faunas. Sand Draw loc. 1, UMMZ 186371 (579/2). These Valvata tricarinata unicarinata DeKay. Baker, 1928, Wiscon clearly have the hinge charcters of compressum. The beaks sin Geol. Nat. History Survey Bull. 70, pt. 1, p. 16, pi. 1, fig. 5. are moderately narrow. A few give hints of ridges. The Valvata tricarinata mediocarinata Baker, 1928, Wisconsin striation varies considerably, but suggests compressum. The Geol. Nat. History Survey Bull. 70, pt. 1, p. 17, pi. 1, shells are rather light in heft and, on the whole, give the fig. 7. impression of rather still water, not where there is swift Valvata tricarinata basalis Vanatta. Baker, 1928, Wisconsin running water with a sandy gravel or where there is heavy Geol. Nat. History Survey Bull. 70, pt. 1, p. 17, pi. 1, wave action. Loc. 3, UMMZ 177328 (1172/2). Mostly thin, fig. 8, 9. many fractured, perhaps in recovering. All have narrow Valvata tricarinata infracarinata Vanatta. Baker, 1928, Wis beaks but only the slightest number show even a hint of a consin Geol. Nat. History Survey Bull. 70, pt. 1, p. 18. ridge. These show considerable variation in outline. Loc. 6, Valvata tricarinata simplex Gould. Baker, 1928, Wisconsin UMMZ 186372 (1/2), 177422 (3/2). Dixon loc. 1, UMMZ Geol. Nat. History Survey Bull. 70, pt. 1, p. 18, pi. 1, 186382 (240/2). Good specimens. Striae as coarse as usual; fig. 10, 14. beaks narrow, some with faint hints of ridges. A little longer Distribution. "Eastern United States west to Iowa; than the usual form found in creeks; could have come from Great Slave Lake south to Virginia and the Ohio a creek or small lake. River" (Baker, 1928, p. 14) ; an isolated spring occur rence in Cherry County, Nebr. Pisidium nitidum Jenyns, 1832 Ecology. A species found only in permanent lakes Pisidium nitidum Jenyns. Woodward, 1913, Catalogue of and rivers. In northern Nebraska this species was British species of Pisidium, p. 44, pi. 1, fig. 9; pi. 3, fig. found in a spring-fed pond of temperature 15 °C. The 6; pi. 19. snails were crawling on aquatic plants, largely Cerato- Herrington, 1954, Nautilus, v. 67, p. 132. phyllum^ Elodea, and algae, near the edge of the pond. Herrington and Taylor, 1958, Michigan Univ. Mus. Zool. Remarks. Valvata tricarinata was found consider Occ. Papers 596, p. 15. ably south of its previously known range in a spring- Distribution. Holarctic. Eurasia and North Africa; fed pond at the old fort reservoir in the Fort Niobrara in North America from Hudson Bay to Mexico, except National Wildlife Eefuge, 5 miles east of Valentine, for the southeastern United States. Cherry County, Nebr. Its existence at this locality Ecology. Found only in perennial water bodies, is considered possible because of the insulating effect such as lakes, ponds, rivers, and large creeks. of the cool spring water. This northern species is Remarks. The fossil specimens are of the form probably able to live here because the pond is warmed commonly found in lakes that are drying up or have little if at all during the summer hot spells. Other dried up. permanent ponds in Cherry County, apparently simi lar except that they lack spring sources, were examined Occurrence and material: Sand Draw and Dixon local faunas. Sand Draw loc. 3, UMMZ 186375 (347/2), 186376 without finding this species. (7/2). The hinge of these varies from moderately heavy to Occurrence and material: Sand Draw and Dixon local light. Seven valves (UMMZ 186376) are of the form paupcr- faunas; Recent in northern Nebraska. Sand Draw loc. 2, clum Sterki. One of these latter has a steep posterior slope UMMZ 184170 (65). Carinae 101, 64 specimens; carinae like a pauperculum approaching the form contortum. Loc. 001, 1 specimen. Loc. 3, UMMZ 177329 (2500). Carinae 101, 7, UMMZ 186380 (1/2). Fragmentary. Dixon loc. 1, UMMZ about 2500 specimens; carinae 100, 12 specimens; carinae 001, 186384 (8/2). 20 specimens; carinae 000, 16 specimens. Dixon loc. 2, UMMZ SYSTEMATIC DESCRIPTIONS 49 191503 (33). Carinae 111. The 17 Recent specimens are tri- Amnicolinae: verge with two functional ducts (vas carinate, 111. deferens, and accessory duct of secondary lobe) ; oper- culum corneous, paucispiral. Valvata lewisi Currier, 1868 Buliminae: verge as in Amnicolinae; operculum cal Valvata lewisi Currier. Baker, 1928, Wisconsin Geol. Nat. careous, concentric with subspiral nucleus. History Survey Bull. 70, pt. 1, p. 26, pi. 1, fig. 28-30. Valvata leivisi Jielicoidea Dall. Baker, 1938, Nautilus, v. 51, Emmericiinae: verge with three functional ducts p. 130. (vas deferens, and an accessory duct in each of two McGrew, 1944, Field Nat. Mus. Hist. Geol. Ser., v. 9, p. 36. secondary lobes). Distribution. "Northern part of the United States The second subfamily was called Bythinellinae by from the Atlantic to the Pacific oceans, northward, in Morrison (1949), but is here changed to Amnicolinae. British America, to the upper Mackenzie River. Its The term Amnicolidae is the second oldest group name southward range is not fully known" (Baker, 1928, within the Hydrobiidae, and deserves retention in some p28). form. Its application, however, involves the compli Ecology. "* * * a lake species largely" (Baker, cated question of identification of the genotype of 1928, p. 28). Amnicola (Morrison 1947, H. B. Baker 1947), Palu- Remarks. A higher spire in the former is the dina lustrica Say, 1821. principal distinction between V. lewisi and V. lewisi The original description is as follows (Say, 1821, p. helicoidea. Variation of this character in the lots 175): examined is such that separation of the two forms P. *lustrica. Shell conic, whirls slightly wrinkled, convex; does not appear practicable in this case. suture profoundly indented; aperture oval nearly orbicular; labrum with the superior edge not appressed to the preceding Occurrence and material: Sand Draw local fauna. Sand whirl, but simply touching it; umbilicus rather large, rounded. Draw loc. 5, CNHM PE3428 (125), UIMNH P6781 (26) ; loc. Length less than %o of an inch. Cabinet of the Academy. 6, UMMZ 177423 (400), 181208 (150). The smallest species I have seen. The aperture somewhat re sembles that of a Valvata, to which genus it may probably be Family VIVIPARIDAE referrible. Mr. Jessup obtained two specimens, on the shore Genus and species uncertain of Cayuga Lake. Remarks. Two small individuals represent this Distinctive features mentioned in this description family, but are inadequate for closer identification. are conical shape, very deep suture, a nearly circular aperture which simply touches the preceding whorl Occurrence and material: Dixon local fauna. Dixon loc. 1, and is not modified in outline by it, very small size. UMMZ 182193 (1) ; loc. 2, UMMZ 191504 (1). Only two species of the general region (cf. Letson Family HYDROBIIDAE 1905) approach this character combination: Amnicola loalkeri Pilsbry, 1898, and Lyogyrus granum (Say), Classification of this family is still in a primitive 1822. But Say contrasted L. granwn with Paludina, state, and nomenclature is in flux. Shells of this lustrica in his description of the former, indicating group provide less information than in perhaps any that he thought of the two as distinct. other family of freshwater snails. Snails with mark Berry (1943, p. 26) described the shell of Amnicola edly different opercula, radulae, and reproductive walkeri thus: apparatus may have similar shells; rather different About 2.5 mm high, 2.2 mm wide, thin, broadly conic in out shells may contain animals of similar anatomy. Fossil line. Four whorls increasing regularly in size. Apex obtuse, shells of the Hydrobiidae are thus often very difficult elevated, whorls shouldered, convex; sutures very deep. to classify with any conviction of certainty. Aperture almost circular (1.04 mm wide; 1.04 mm high). For many years two family names have been used: Peristome continuous, adnate to the preceding whorl for only a short distance. Umbilicus rather wide, deep. Periostracum Europeans have called the group Hydrobiidae, while light tan, lines of growth crowded but distinct, often con Americans have preferred the term Amnicolidae. "Hy- cealed with a deposit from its environment. drobiae" Troschel, 1857 (1856-1893, v. 1, p. 106), has The descriptions by Say and Berry are so strikingly priority over Amnicolidae Tryoii (1862, p. 452), and similar that it seems highly probable that they refer hence the name Hydrobiidae is preferable. to the same species. The subfamily divisions listed below are slightly Accepting Paludina lustrica Say as being the species modified from Morrison (1949) : generally known heretofore as Amnicola walkeri, we Hydrobiinae: verge with one functional duct (vas find that fortunately there are no major changes in deferens). the group concept. Berry (1943) showed the close 50 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS relationship between A. walkeri. now A. lustrica Measurements. A large series of specimens (UMMZ (Say), and A. limosa (Say), usually but erroneously 177554) from the Saw Rock Canyon site was sampled considered the type species. to study variation in several size groups selected on The classification and generic synonymy of groups number of whorls. Measurements are in millimeters. treated by Morrison (1947) implied by synonymiza- tion of Amnicola walkeri with Paludina lustrica and Number of whorls by acceptance of Morrison's (1949) outline of the 3% 4 4H 4}£ Hydrobiidae is as follows: Number of specimens., , 25 44 36 25 Family Hydrobiidae Subfamily Hydrobiinae Length: Mean.. ______2. 35 ±. 026 2. 63 ±. 033 2.95 ±.027 3. 23 ±. 028 Genus Marstonia F. C. Baker, 1926. Type (by original Standard deviation . 13 ±. 018 .22 ±.023 . 16 ±. 019 . 14 ±. 020 Width: designation) : Amnicola lustrica Pilsbry, 1890 [not Mean. ______1. 80 ±. 020 1.96 ±.027 2. 16 ±. 027 2. 30 ±. 024 of Say, 1821] = Amnicola lacustris Morrison, 1947. Standard deviation, _ _ _ . 10 ±. 014 . 18 ±. 019 . 16 ±. 019 .12 ±.017 Length aperture: Subfamily Amnicolinae 1.20 ±.018 1.33 ±.018 1. 44 ±. 017 1.51 ±.014 Standard deviation. . 09 ±. 013 . 12 ±. 013 . 10 ±. 012 . 07 ±. 010 Genus Amnicola Haldeman, 1840. Type (by original Width aperture: designation) : Paludina lustrica Say, 1821, Amni . 95 ±. 012 1. 08 ±. 017 1. 15 ±. 013 1. 24 ±. 018 Standard deviation. . 06 ±. 008 . 11 ±. 012 .08 ±.0094 .09 ±.013 cola walkeri Pilsbry, 1890. Width to length ratio: .76 ±.0058 . 74 ±. 0039 . 73 ±. 0037 .71 ±.0042 Subjective synonyms: Euamnicola Crosse and Standard deviation. . 029±. 0041 . 026±. 0028 . 022±. 0026 . 021 ±. 0030 Fischer, 1891. Length aperture to length ratio: Type: Paludina lustrica Say; the statement "(ou .52 ±.0054 .51 ±.0041 . 49 ±. 0032 .47 ±.0028 Standard deviation. . 027±. 0038 . 027±. 0029 . 019±. 0022 . 014±. 0020 Anmicola sensu stricto)" fixed the type as that of Amnicola, no matter whether or not Crosse and Fischer correctly listed the type of Amnicola. Variation. The size range of adults is uncertain Marstoniopsis Altena, 1936. Type (by original because adult shells cannot be distinguished surely designation) : Hydro'bia steinii v. Martens, 1858. from juveniles on any character other than size. Some Subfamily HYDROBIINAE variation is, however, evident from the fact that shells of 4% whorls are rare, and those of 4^2 and 41/4 Genus MARSTONIA Baker, 1926 whorls progressively more common. Variation in Marstonia crybetes (Leonard), 1952 shape, the most conspicuous variant, is shown under Plate 2, figures 1-3, 7, 8 Measurements. The last part of the body wrhorl varies in degree of appression, so that the peristome may be Amnicola crybetes Leonard, 1952. Nautilus, v. 66, p. 38, pi. 5, appressed or adnate; rarely the last part of the body fig. A. Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, whorl is disjunct. p. 151, pi. 14, fig. h. Remarks. Amnicola crybetes Leonard is referred to Types. Holotype UKMNH 3805, Saw Rock Canyon local the genus Marstonia because of its similarity to M. fauna. Paratypes UKMNH 3806. decepta (Baker), the genotype. With the latter species Diagnosis. A Marstonia shorter and wider than it shares an acute spire, undepressed nuclear whorl, other species of the genus, with ovate-pyriform aper narrow umbilicus, and ovate-pyriform aperture. Cin- ture and moderately impressed suture. cinnatia [as exemplified by C. Integra (Say) and C. Description. Shell small, ovate, conoidal, with peracuta (Pilsbry and Walker)] has a shell of similar acute spire and apex. Whorls 4 to 41/2, convex, gently shape but larger, with a larger nuclear whorl. Amni shouldered and slightly flattened peripherally, expand cola lustrica (Say), to which Leonard compared M. ing fairly rapidly; body whorl about three-fourths crybetes in describing the latter, has a slightly smaller of total shell length. Embryonic shell of about one shell with more deeply incised sutures, a nearly round whorl, not depressed to the level of the postembryonic aperture, and more conspicuous umbilicus. whorls. Suture strongly impressed, slightly below Marstonia decepta differs from M. crybetes by its periphery of whorls. Aperture simple, entire, ovate- larger size and proportionately narrower shape; other pyriform; peristome appressed to preceding whorl or wise shells of the two are similar. simply adnate, broadly rounded below, narrowly The material recorded by Frye and Leonard (1952, rounded to obtusely angular at junction of outer and p. 151) from the SWl/4 sec. 22, T. 33 S., R, 29 W., parietal lips. Umbilicus small but distinct, about one- Meade County, almost certainly came from Rexroad ninth of shell diameter. Sculpture consists only of locality 3 rather than Bender locality 1. Habitat in very fine growth lines on the postembryonic shell. ferences from Bender locality 1 species suggest only SYSTEMATIC DESCRIPTIONS 51 temporary bodies of water. Rexroad locality 3 had the Genus and species uncertain permanent water which Marstonia crybetes required. Remarks. Two fragmentary specimens represent Occurrence and material: Saw Rock Canyon and Rexroad an amnicolid with a high, narrow spire, distinct from local faunas; Dixon local fauna (Frye and Leonard). Saw Marstonia crybetes. It may be Pyrgophorus or Fonti- Rock Canyon, UMMZ 173593 (33), 177554 (500), USGS 19089-11 (100) ; Rexroad loc. 1, UMMZ 183051 (1) ; loc. 4b, gens. UMMZ 177597 (2), 181046 (2). Occurrence and material: Dixon local fauna. Dixon loc. 1, UMMZ 182194 (2). Marstonia decepta (Baker), 1928 Subclass PULMONATA Plate 2, figure 9
Amnicola lustrica decepta Baker, 1928, Wisconsin Geol. Nat. Order BASOMMATOPHORA History Survey Bull. 70, pt. 1, p. 108, text fig. 45. Amnicola lustrica Pilsbry. Baker, 1928, Wisconsin Geol. Nat. Family CARYCHIIDAE History Survey Bull. 70. pt. 1, p. 104, pi. 6, fig. 16, 17, 26, 27; text fig. 45. Genus CARYCHITTM Miiller, 1774 Amnicola (Marstonia) lustrica Pilsbry. Berry, 1943, Michigan Univ. Mus. Zool. Misc. Publ. 57, p. 29, pi. 1, fig. 4-6; pi. Carychium exiguum (Say), 1822 3, fig. 3; pi. 5, fig. 6; pi. 7, fig. 4. Amnicola lacustris Morrison, 1947, Nautilus, v. 60, p. 86. Carychium exiguum (Say). Pilsbry, 1948, Acad. Nat. Sci. Distribution. New York west to Minnesota and Philadelphia Mon. 3, v. 2, pt. 2, p. 1052, fig. 561a, b, 562. Iowa; north-south range uncertain (F. C. Baker, 1928, Harry, 1952, Nautilus, v. 66, p. 5. p. 106-107; USNM colln.). Carychium perexigutim Baker, 1938, Nautilus, v. 51, p. 128. Smith, 1940, Kansas Geol. Survey Bull. 34, p. 98. Remarks. Two specimens associated with abundant Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, M. crybetes differ from that species by their narrower 151, pi. 14, fig. j. form and smaller aperture. Their measurements in Leonard, 1957, Illinois Geol. Survey Report of Investiga mm: length 2.8, 2.7; width 1.8, 1.7; length aperture tions 201, p. 10, pi. 1, fig. 1. 1.1, 1.1; width aperture 1.0, 0.9; width/length ratio Carychium perexlguum F. C. Baker. Hibbard, 1941, Kansas 0.66, 0.64; length aperture to length ratio 0.41, 0.42; Univ. Sci. Bull., v. 27, p. 95. both have 4^ whorls. Comparison of these measure Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. ments with those given under M. crybetes shows that Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, the two narrow specimens fall more than three stand p. 408. ard deviations from the mean of the characters width, Distribution. "Newfoundland to Colorado, south to length of aperture, and the length of aperture to Mobile Bay, Alabama, and near Deming, southwestern length ratio, and close to this distance for other char New Mexico" (Pilsbry, 1948, p. 1052.) acters. It is therefore most unlikely that these two Ecology. In northern Nebraska this species was shells are only extreme variants of Marstonia crybetes. found among wet leaves in seepage areas and beside They agree well with U. S. National Museum material a spring-fed brook. of M. decepta, except for being slightly smaller, and Remarks. Examination of the type series (U1MNH are referred to that species. P6776, P6777) and of a series of topotypes (UMMZ The species nomenclature is involved. Morrison 183026) shows that C. perexiguum is a synonym of (1947) proposed Amnicola lacustris as a new name for C. exiguum. Baker thought it distinguished by "the the preoccupied A. lustrica Pilsbry, [not Say]. In shorter shell, heavier lip, and the upward bending 1928, however, F. C. Baker had added two varietal columellar lamella." The series of topotypes, as well names to the species: A. lustrica decepta and A. lus as the original measurements, shows a size range trica. perlustrica. Both of these have priority over A. within that given by Pilsbry (1948, p. 1054) for lacustris Morrison, and bar that name from use for C. exiguum. The thickening of the lip in the topo- the species as a whole (cf. Bull. Zool. Nomenclature, type series appears no different from that in C. exi v. 4, p. 240, concl. 2, 1950). The name decepta is here guum. Only two of the types were dissected by Baker, applied to the species known as A. lustrica Pilsbry; and only one of these shows the principal columellar lacustris Morrison and perlustrica Baker are available lamella adequately. The upward bend of this lamella for the ecological variants distinguished by Baker. referred to in the original description is a minor flare Occurrence and material: Saw Rock Canyon local fauna, on the ventral side of the lamella within the body UMMZ 186155 (2). whorl. This feature is certainly well within the range 52 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS of variation of C. exiguum as figured by Pilsbry Remarks. Lymnaea macella Leonard and L. par- (1948). exilis Leonard are obviously young and adult of the Occurrence and material: Rexroad local fauna; Dixon local same species. This is readily demonstrable by break fauna (Frye and Leonard) ; Recent in northern Nebraska. ing the larger L. parexilis into a smaller shell, which Rexroad loc. 8, UMMZ 183026 (63), UIMNH P6776 (4), is then L. macella. The differentiae given for L. ma P6777 (3). cella were the smaller size, heavier varix, and larger Carychium sp. nuclear whorl. All are invalid. Varix and nuclear Carychitim perexiguum F. C. Baker. Taylor, 1954, Michigan whorl are not different in the specimens examined Univ. Mus. Zool. Occ. Papers 557, p. 14 [not of Baker]. personally; if it was meant that they are relatively Remarks. The single Sand Draw specimen is surely different, the characters are a function of size. Large not C. perexiguum (=(7. exiguum), and may be new. series from the Rexroad local fauna clearly show the It is distinguished by the upper columellar fold, which intergradation of the two forms. Leonard may have is bent 90° toward the outer lip, ascends the parietal reached a similar conclusion, unless a mislabelled illus lip, and is strongly thickened. The fold thus appears tration gives an erroneous implication. Plate 6, figure as a large, transverse lamella protruding conspicuously f (Frye, Leonard, and Swineford, 1956) is captioned into the aperture. Additional material is needed to Lymnaea macella Leonard; but this photograph is determine the significance of this specimen. evidently the same illustration as those published by Leonard (1952a) and Frye and Leonard (1952) as Occurrence and material: Sand Draw local fauna. Sand Draw loc. 3, UMMZ 181126 (1). Lymnaea parexilis. The Pliocene populations from the Rexroad forma Family LYMNAEIDAE tion, distinguished by Leonard as distinct species, are referred to Stagnicola exilis on the basis of large Uni Genus STAGNICOLA Jeffreys, 1830 versity of Michigan series and of published informa Subgenus STAGNICOLA s. s. tion. The few specimens from Rexroad locality 3, Stagnicola (s. s.) exilis (Lea), 1837 the type locality of Lymnaea macella and L. parexilis, are inadequate to show the range of variation. Series Galba exilis (Lea). Baker, 1911, Chicago Acad. Sci. Spec. from Rexroad locality 4c, from which Frye and Leon Pub. 3, p. 343, pi. 35, fig. 4; pi. 36, fig. 21-22; pi. 37, fig. 1-11. ard (1952) recorded L. parexilis, agree well with Stagnicola exilis (Lea). Baker, 1928, Wisconsin Geol. Nat. Stagnicola exilis as described and figured by Baker History Survey Bull. 70, pt. 1, p. 226, pi. 14, fig. 7-11; (1911, 1928). pi. 17, fig. 16. According to Leonard (1952, p. 42) "The shell of Stagnicola reflexa (Say). Hibbard, 1941, Kansas Univ. Sci. Bull., v. 27, p. 95 [not of Say]. Lymnaea parexilis is similar in general form to that Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265 of L. exilis, but is much smaller with a relatively more [not of Say]. elongate aperture, and more intricate sculpture." The Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. only significant feature of these three is the relatively 408 [not of Say]. Lymnaea macella Leonard, 1952, Nautilus, v. 66, p. 40, pi. 5, longer aperture. Recent Stagnicola exilis, and the fig. J. University of Michigan fossil series examined, have Lymnaea macella Leonard. Frye and Leonard, 1952, Kansas a slightly smaller aperture than Lymnaea parexilis as Geol. Survey Bull. 99, p. 151 [in part: Meade County figured and described by Leonard (1952a). This char localities], pi. 14, fig. q. Lymnaea parexilis Leonard, 1952, Nautilus, v. 66, p. 41, pi. 5, acter is probably not of specific value; at any rate at fig. K. least some occurrences of L. parexilis represent Stag Lymnaea parexilis Leonard. Frye and Leonard, 1952, Kansas nicola exilis. Geol. Survey Bull. 99, p. 151 [in part: Meade County localities], pi. 14, fig. x. Occurrence and material: Rexroad local fauna. Rexroad Lymnaea palustris (Miiller). Taylor, 1954, Michigan Univ. loc. 1, UMMZ 183053 (100) ; loc. 3, UMMZ 183027 (3) ; loc. Mus. Zool. Occ. Papers 557, p. 3 [in part: Rexroad local 4b, UMMZ 177599 (150), 181048 (125). fauna. Not of Miiller]. Distribution. "Ohio to Kansas, northward to north Stagnicola (s. s.) palustris (Miiller), 1774 ern Minnesota and northern Michigan1' (Baker, 1928, Galba palustris (Miiller). Baker, 1911, Chicago Acad. Sci. p. 227). Spec. Pub. 3, p. 298, pi. 26, fig. 17-37, pi. 33, fig. 1-25; Ecology. "Exilis is an inhabitant of sloughs, ponds, pi. 34, fig. 20. and streams which dry up more or less during a por Distribution. "Circumboreal. Northern Asia and tion of the year" (Baker, 1928, p. 227). Europe. North America from the Atlantic to the SYSTEMATIC DESCRIPTIONS 53 Pacific Ocean, and from Alaska south to New Mexico" Baker (1938) identified five juvenile specimens from (Baker, 1911, p. 307). the Sand Draw local fauna as Fossaria dalli grandis. Ecology. In northern Nebraska it was found abun Four (UIMNH P6T96, CNHM PE3442) are''Stagni dant in drainage ditches and ponds, temporary and cola cf. S. reflexa, the other (CNHM PE3443) Fos perennial water bodies. saria dalli. Occurrence: Recent in northern Nebraska. Occurrence and material: Sand Draw local fauna. Sand Draw loc. 1, UMMZ 181238 (150) ; loc. 2, UMMZ 184172 (28) ; Stagnicola (s. s.) reflexa (Say), 1821 loc. 3, UMMZ 177330 (700) ; loc. 4, UMMZ 177313 (80) ; loc. 5, CNHM PE3429 (18), PE3430 (19), UIMNH P6783 (1), Galba reflexa (Say). Baker, 1911, Chicago Acad. Sci. Spec. P6784 (3), P6782 (14), UMMZ 181136 (2000), 181227 (6500); Pub. 3, p. 332, pi. 30, fig. 30, 31; pi. 35, fig. 3, 5-22, pi. loc. 6, CNHM PE3441 (6), PE3442 (1), UIMNH P6795 (4), 36, fig. 1-11; pi. 18, fig. 10. P6796 (3), UMMZ 177426 (150), 181210 (1000) ; loc. 7, UMMZ Stagnicola reflexa (Say). Baker, 1928, Wisconsin Geol. Nat. 177227 (175). Hist. Survey Bull. 70, pt. 1, p. 221, pi. 14, fig. 1-6; pi. Stagnicola (s. s.) sp. 17, fig. 15. Lymnaea macella Leonard. Frye and Leonard, 1952, Kansas Remarks. Immature and fragmentary shells from Geol. Survey Bull. 99, p. 151, in part [Kingman County several localities represent a species of the Stagnicola locality. Not of Leonard]. palustris group, such as S. exilis or S. reflexa, but are Lymnaea parexilis Leonard. Frye and Leonard, 1952, Kansas inadequate for precise identification. Geol. Survey Bull. 99, p. 151, in part [Kingman County locality. Not of Leonard]. Occurence and material: Saw Rock Canyon, Bender, and Sanders local faunas. Saw Rock Canyon, UMMZ 183655 (3) ; Distribution. "Eastern Quebec west to Nebraska, Bender loc. la, UMMZ 184119 (1) ; loc. 2, UMMZ 184149 (14) ; Manitoba south to southern Illinois and southern Kan Sanders loc. 1, UMMZ 182146 (5) ; loc. 2, UMMZ 184161 (2) ; sas" (Baker, 1928, p. 224). loc. 3, UMMZ 182176 (4). Ecology. "The typical habitat of reflexa in north ern Illinois and Wisconsin is in small pools or ponds Subgenus HINKLEYIA Baker, 1928 which may become more or less dry in summer. All Stagnicola (Hinkleyia) caperata (Say), 1829 of the Wisconsin habitats have been in swales in Stagnicola caperata (Say). Baker, 1928, Wisconsin Geol. Nat. woods or fields, none have been found in large streams History Survey Bull. 70, pt. 1, p. 260, pi. 18, fig. 43-47. or lakes" (Baker, 1928, p. 224). Baker, 1938, Nautilus, v. 51, p. 130. Remarks. Large series from the Dixon local fauna Stagnicola caperata. McGrew, 1944, Field Mus. Nat. Hist. Geol. Ser., v. 9, p. 36. show that the Stagnicola has the more convex whorls Lymnaea caperata Say. Hibbard, 1954, Michigan Acad. Sci. and impressed sutures of S. reflexa, rather than 8. Papers, v. 39, p. 342. exilis which was reported by Frye and Leonard. fLymnaea diminuta Leonard. Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. 151 [in part: Kingman Occurrence and material: Dixon local fauna. Dixon loc. 1, County locality]. UMMZ 182197 (1500) ; loc. 2, UMMZ 191505 (250). Distribution. "From Quebec and Massachusetts Stagnicola (s. s.) cf. S. reflexa (Say), 1821 west to California; Yukon Territory and James Bay south to Maryland, Indiana, Colorado and California" Stagnioola cf. reflexa (Say). Baker, 1938, Nautilus, v. 51, p. 130. (Baker, 1928, p. 263). Stagnioola, sp. Baker, 1938, Nautilus, v. 51, p. 131. Ecology. "In the Mississippi Valley this species Fossaria dalli grandis F. C. Baker. Baker, 1938, Nautilus, v. seems to almost invariably occupy intermittent streams 51, p. 131 [in part. Not of Baker]. or small pools, ponds and ditches which dry up in Stagnicola cf. reflexa. McGrew, 1944, Field Mus. Nat. Hist. the summer. In Illinois it is usually found in associa Geol. Ser. v. 9, p. 36. tion with Aplexa hypnorum and Sphaeriwn occiden- Stagnicola sp. McGrew, 1944, Field Mus. Nat. Hist. Geol. Ser., tale, either in small streams, pools or sloughs, or in v. 9, p. 36. spring pools in the woods which become completely Fossaria dalli grandis. McGrew, 1944, Field Mus. Nat. Hist. dry in late spring and summer *** In these dry ponds Geol. Ser., v. 9, p. 36 [in part]. living specimens may frequently be found by digging Remarks. Although Stagnicola is abundant in into the mud, leaves and other debris. In Wisconsin, Sand Draw local fauna collections, almost all shells caperata has been found almost invariably in small are immature or fragmentary. A few halfgrown woodland pools which become dry in summer and shells from various localities suggest S. reflexa, but fall, or in small streams which become wholly or the identification is uncertain and there may even partially dry" (Baker, 1928, p. 263). In northern be more than one species represented. Nebraska, S. caperata was found in similar, temporary 54 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS bodies of water, such as irrigation ditches, and also Wakarusa valley. It may be found in ephemeral pools in the more perennial ponds of the Sand Hills. It was where it appears in large numbers for a few weeks commonly associated with Stagnicola palustris, Pro- in the early spring. Many of the adults die after menetus umbilicatellus, and Aplexa hypnorum. the egg laying period and few adults are seen through Remarks. The habitat of this species is one ap the hot summer months; immature individuals pre parently available quite widely in the southern Plains sumably remain alive, buried in the mud" (Franzen region. It has never been recorded alive as far south and Leonard, 1943, p. 405). as Kansas, however, and its usual common occurrence Remarks. Lymnaea diminuta Leonard is a syno elsewhere and distinctive shape argue that its absence nym of Stagnicola l)ulim,oides techella. It was com is reliable. The most reasonable explanation for the pared by Leonard to F. humilis and others of the northern restriction of this species is that the hot sum genus Fossaria, but its larger nuclear whorl, simple mers to the south bar its southward dispersal. columella, and wide inner lip indicate that it belongs Frye and Leonard (1952) reported Lymnaea dimi- to the subgenus Nasonia. L. diminuta agrees with 8. nuta (=8tagnicola bulimoides techella), but not 8. bulimoides techella in having the significant characters capera.ta, from the Dixon local fauna. Because the of an acutely conic spire, obese body whorl, and two species are somewhat similar, and because Univer broadly reflected inner lip (see Baker, 1911, p. 216). sity of Michigan collections contain many 8. caperata Occurrence and material: Saw Rock Canyon, Red Corral, but no 8. bulimoides techella, their reference may well Rexroad, Bender, and Sanders local faunas. Saw Rock be based on 8. caperata. Canyon, UMMZ 183656 (14) ; Red Corral, UMMZ 182993 Hibbard's (1954) report of this species in the Rex- (175) ; Rexroad loc. 1, UMMZ 183052 (600) ; loc. 2, UMMZ road local fauna is based upon the Bender local fauna 183065 (2) ; Bender loc. la, UMMZ 184118 (32) ; loc. Ic, UMMZ 183012 (33) ; loc. 2, UMMZ 184147 (200) ; Sanders loc. specimens. None of the Rexroad local fauna localities 1, UMMZ 182145 (75) ; loc. 2, UMMZ 182164 (1), 184160 (17) ; studied carefully have yielded 8. caperata, but the loc. 3, UMMZ 182175 (50). species occurs at at least one other site represented in University of Michigan collections. Genus FOSSARIA Westerlund, 1885 Occurrence and material: Buis Ranch, Bender, Sand Draw, Fossaria obrussa (Say), 1825 Dixon, and Sanders local faunas; Recent in northern Nebraska. Buis Ranch, UMMZ 181113 (1) ; Bender loc. 1, UMMZ 177504 Galbra obrussa (Say). Baker, 1911, Chicago Acad. Sci. Spec. (4) ; loc. la, UMMZ 184146 (17) ; loc. Ic, UMMZ 183011 (1) ; Pub. 3, p. 270, pi. 26, fig. 8-13, pi. 31, fig. 20-37. loc. 2, UMMZ 184148 (100) ; Sand Draw loc. 2, UMMZ 184171 Fossaria obrussa (Say). Baker, 1928, Wisconsin Geol. Nat. (37) ; loc. 3, UMMZ 181127 (30) ; loc. 4, UMMZ 177312 (12) ; History Survey Bull. 70, pt. 1, p. 293, pi. 16, fig. 14; pi. loc. 5, CNHM PB3431 (50), UIMNH P6785 (18), UMMZ 18. fig. 14-24. 181135 (75), 181226 (150) : loc. 6, UMMZ 177424 (150), 181209 Distribution. "From the Atlantic to the Pacific (250) ; loc. 7, UMMZ 177228 (100) ; Dixon loc. 1, UMMZ 182- 195 (24) ; loc. 2, UMMZ 191506 (41) ; Sanders loc. 1, UMMZ oceans, and from Mackenzie Territory, Canada south 182226 (150) ; loc. 2, UMMZ 184159 (10) ; loc. 3, UMMZ 182174 to Arizona and northern Mexico" (Baker, 1928, p. (43). 296). Submenus NASONIA Baker, 1928 Ecology. "The normal habitat of this species is Stagnicola (Nasonia) bulimoides techella (Haldeman), 1867 in small bodies of water, as creeks, ponds, sloughs, bays, and marshy spots along river banks. It is at Galba bulimoides tecJiella (Haldeman). Baker, 1911, Chicago Acad. Sci. Spec. Pub. 3, p. 214, pi. 27, fig. 30-35; pi. 28, home on sticks, stones, and any other debris that may fig. 1-3, 8. be in the water or along its edge" (Baker, 1928, p. Lymnaea diminuta Leonard, 1952, Nautilus, v. 66, p. 39, pi. 5, 296). In northern Nebraska it was found on mud fig. B. and logs just above water level, or in the mud of Lymnaea diminuta Leonard. Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. 151 [in part: Meade County seepages by streams. localities], pi. 14, fig. s. Occurrence and material: Sand Draw local fauna; Recent in Distribution. Southwestern and south-central northern Nebraska. Sand Draw loc. 3, UMMZ 181128 (54). United States to central Mexico; from southern Cali Fossaria dalli (Baker), 1906 fornia through Utah, Colorado, southernmost Ne braska and Kansas to Missouri and Alabama; south Plate 2, figures 6, 10-27
to San Luis Potosi and Oaxaca. Galba dalli (Baker). Baker, 1911, Chicago Acad. Sci. Spec. Ecology. In northeastern Kansas "Lymnaea buli- Pub. 3. p. 251, pi. 30, fig. 13-18. moides techella is locally abundant in roadside ditches Fossaria dalli (F. C. Baker). Baker, 1928, Wisconsin Geol. * * * throughout the broad terraces of the lower Nat. Hist. Survey Bull. 70, pt. 1, p. 288, pi. 16, fig. 11. SYSTEMATIC DESCRIPTIONS 55 Fossaria dalli grandis F. C. Baker. Baker, 1938, Nautilus, Distribution. "Ohio to Northern Michigan and v. 51, p. 131 [in part]. Montana, south to Kansas and Arizona" (Baker, 1928, Fossaria dalli grandis. McGrew, 1944, Field Mus. Nat. History Geol. Ser., v. 9, p. 36. p. 288). Lymnaea turritella Leonard, 1952, Nautilus, v. 66, p. 39, pi. Ecology. "* * * inhabits wet, marshy places, gen 5, fig. 0. erally out of the water, on sticks, stones, or muddy Lymnaea turritella Leonard. Frye and Leonard, 1952, Kansas flats" (Baker, 1928, p. 289, 287). In northern Ne Geol. Survey Bull. 99, p. 151, pi. 14, fig. r. fLymnaea humilis modicella (Say). Frye and Leonard, 1952, braska it was found in seepage areas, in the water Kansas Geol. Survey Bull. 99, p. 151. or in wet leaves and debris at the water's edge.
Size-shape variation in Lymnaea turritella topotypes is as follows:
Number of whorls 3%
Number of specimens.
Length: Mean______3. 44 ± 0. 066 3. 79 ± 0. 059 4.27 ±0. 078 4. 60 ±0. 25 Standard deviation, _ _ .28 ± . 047 .27 ± . 042 .31 ± . 055 . 70 ± . 018 Width: Mean.______1. 94 ± 028 2. 09 ± . 035 2. 27 ± 040 2. 31 ± . 100 Standard deviation.- ______.12 ± 020 . 16 ± . 025 .16 ± 028 . 28 ± . 070 Length aperture: Mean.______1. 60 ± 033 1. 73 ± . 028 1. 89 ± 035 1. 92 ± . 11 Standard deviation.. _ . 14 ± 023 . 13 ± . 020 . 14 ± 025 . 30 ± . 075 Width aperture: Mean. ______.99 ± 021 1. 03 ± . 018 1. 12 ± 019 1. 14 ± . 060 Standard deviation.- ______. 087 ± 015 .081 ± . 013 . 078 ± 014 . 17 ± . 043 Width to length ratio: Mean______. 566 ± 0033 . 550 ± . 0028 . 532 ± 0023 . 504 ± . 0078 Standard deviation.______. 014 ± 0023 . 013 ± . 002 . 0093 ± 0016 . 022 ± . 0055 Length aperture to length ratio: Mean.. ______. 465 ± 0019 . 457 ± . 0016 . 443 ± 003 . 416 ± . 0053 Standard deviation,. ______. 0081 ± 0014 . 0072± . 0011 . 012 ± 0021 . 015 ± . 0038
Series from the Saw Rock Canyon, Bender, Sand Small series from the Dixon and Sanders local fau Draw, Berends, and Jinglebob local faunas are all nas, and other Rexroad local fauna localities, are closely similar in having sculpture of coarse, irregular inadequate to show similarities closer either to the growth lines and small apertures. A large series from Rexroad 4b sample or to other samples. Rexroad locality 4b differs somewhat in having a Other late Pleistocene material surely representing greater range of variation. Many shells matching what is called here Fossaria dalli, and which agrees those from the other faunas may be found, but many closely with the Saw Rock Canyon, Sand Draw, Ber others cannot be so matched. These latter specimens ends, and Jinglebob material is in the U. S. National are slightly smoother, with longer apertures; some Museum, as follows: 570570, marl, Castalia, Erie attain both greater size and a larger number of whorls County, Ohio; identified by F. C. Baker as Fossaria than occur in other samples. Such large specimens dalli grandis. 570542, loess, Fulton County, 111., iden may be referable to Fossaria parva (Lea) (Baker, tified by F. C. Baker as Fossaria parva tazewelliana. 1911, p. 243) or to F. parva sterkii (Baker) (Baker, 570573, bed of drained pond, 2 miles southeast of 1911, p. 248). The Rexroad locality 4b sample pro Northport, Wash.; identified by F. C. Baker as Fos vides no evidence that more than one population is saria perplexa. F. C. Baker (1938) identified five represented, however. This sample may be F. parva, rather than F. dalli as here considered, or F. dalli may juvenile specimens as Fossaria dalli grandis. One be a synonym of F. parva. These questions involve (CNHM PE 3443) is F. dalli, the others (CNHM PE revision of a number of Recent species, a study out 3442, UIMNH P6796) /Stagnicola palustris. side the scope of this report. The Rexroad locality Frye and Leonard (1952) reported Lymnaea hwnili* 4b material is referred to F. dalli because what is modicella from some localities represented by Univer probably a single population contains numerous indi sity of Michigan collections. The form has not been viduals morphologically indistinguishable from other recognized in this material. The reference may be samples referable to F. dalli. based upon short-spired specimens of F. dalli, however. 56 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Remarks. \Lymnaea turritella Leonard was origi (4); loc. 6, CNHM PE3440 (1), UIMNH P6794 (2), P6798 nally distinguished from F. dalli by its somewhat (fragments) ; Dixon loc. 1, UMMZ 182196 (60) ; loc. 2, UMMZ larger size and slenderer form. Comparison of a 191508 (15). Family PLANORBIDAE large series of topotoypes with other large series of fossils, and with Recent material in the U. S. National Subfamily PLANORBINAE Museum identified by F. C. Baker, shows that these Genus ANISTTS Studer, 1820 differences are not constant. The description and previous illustrations of L. turritella are based on Submenus ANISUS s. s. extremely long, narrow specimens. The accompany Anisus (s. s.) pattersoni (Baker), 1938 ing table, as well as the illustrations, shows variation in shape in the Saw Rock Canyon fauna sample. Gyraulus pattersoni Baker, 1938, Nautilus, v. 51, p. 129. Comparison of these measurements with those given Gyraulus pattersoni. McGrew, 1944, Field Mus. Nat. History by Baker reveals further that there is no significant Geol. Ser., v. 9, p. 36. Gyraulus pattersoni Baker. Leonard, 1950, Kansas Univ. difference in size. Paleont. Contr. 8, p. 20, pi. 3, fig. I. Occurrence and material: Saw Rock Canyon, Rexroad, Ben Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. der, Sand Draw, Dixon, and Sanders local faunas; Recent in 158, pi. 15, fig. g. northern Nebraska. Saw Rock Canyon, UMMZ 173595 (12), Leonard, 1957, Illinois Geol. Survey Report of Investiga 177555 (250) ; Rexroad loc. 1, UMMZ 183054 (1) ; loc. 2, tions 201, p. 12, pi. 2, fig. 4-6. UMMZ 183066 (4) ; loc. 3, UMMZ 183028 (4) ; loc. 4b, UMMZ Anisus pattersoni (Baker). Taylor, 1958, Jour. Paleont., v. 177598 (200), 181047 (125); Bender loc. Ib, UMMZ 184134 32, p. 1149. (9) ; loc. Ic, UMMZ 183013 (1) ; loc. 2, UMMZ 184150 (3) ; Types. Holotype CNHM, Sand Draw loc. 5. Para- Sand Draw loc. 2, UMMZ 184187 (6) ; loc. 6, CNHM PE3443 types CNHM (2), UIMNH P7455 (4), ANSP 169948 (1), UMMZ 177425 (7); Dixon loc. 1, UMMZ 182198 (34); (2). loc. 2, UMMZ 191507 (2) ; Sanders loc. 3, UMMZ 182177 (4). Diagnosis. An Anisus of small size (less than 5 Genus BULIMNEA Haldeman, 1841 mm diameter), with 5-6 whorls, nearly plane on both sides, subangular at junction of left side and Bulimnea megasoma (Say), 1824 periphery of whorls. Bulinmea megasoma (Say). Baker, 1911, Chicago Acad. Sci. Description. Shell small, discoidal, presumably Spec. Pub. 3, p. 184, pi. 25, fig. 1-6. ultradextral, nearly plane on both sides; whorls 5-6, Baker, 1928, Wisconsin Geol. Nat. Hist. Survey Bull. 70, flattened on the left side, subangulate at junction of pt. 1, p. 277, pi. 8, fig. 19-23; pi. 17, fig. 31, 35. left side with periphery, rounded on periphery and Lymnaea cf. stagnalis jugularis Say. Baker, 1938, Nautilus, right side, very slowly and regularly increasing; v. 51, p. 131. suture well impressed; aperture subquadrate, left side Stagnicola sp. Baker, 1988, Nautilus, v. 51, p. 131. flattened to arcuate, periphery gently rounded, right Lymnaea cf. stagnalis jugularis. McGrew, 1944, Field Mus. Nat. History Geol. Ser., v. 9, p. 36. side strongly rounded; lip sometimes thickened Stagnicola sp. McGrew, 1944, Field Mus. Nat. Hist. Geol. Ser., within; sculpture of numerous fine, irregular growth v. 9, p. 36 [in part]. lines and weak spiral striation; embryonic shell of Lymnaea megasoma Say. Taylor, 1954, Michigan Univ. Mus. about one-half whorl, spirally striate only. Zool. Occ. Papers 557, p. 14. Measurements. See table of comparative measure Distribution. "Northern New England (Vermont) ments below. west to Minnesota, Iowa, and Manitoba; northern Distribution. Pliocene in southwestern Idaho, Ohio (latitude 41°) northward to about latitude 57° northern Utah, western Wyoming; early Pleistocene in British America" (Baker, 1928, p. 282). in southwestern Idaho, Nebraska, and Kansas; middle Ecology. "Megasotna is usually an inhabitant of Pleistocene in the central Great Plains; late Pleisto small, quiet bodies of water or swamps" (Baker, 1928, cene, Ohio. p. 281). Ecology. Known only from inference, since the Remarks. The material which Baker (1938) re species is extinct. Its association with fossils which ported from the Sand Draw local fauna is fragmen represent living species suggests it lived in shallow, tary, but of the species known from this fauna can quiet waters, in the backwaters along streams or in be only B. megasoma. semipermanent ponds and sloughs. Occurrence and material: Sand Draw and Dixon local Remarks. Anisus pattersoni (F. C. Baker) seems faunas. Sand Draw loc. 1, UMMZ 181237 (21) ; loc. 2, UMMZ most nearly related to A. rotundatus (Poiret); but 184186 (1) ; loc. 3, UMMZ 177331 (100) ; loc. 4, UMMZ 177314 insufficient data for adequate comparison are available SYSTEMATIC DESCRIPTIONS 57 on A. perezi (Dupuy), to which it may be closer. A. and A. perezi, A. pattersoni possesses an internal pattersoni is smaller than A. rotundatus (see measure thickening of the lip. Formation of this callus is not ments), with fewer whorls than either A. rotundatus restricted to mature individuals; most specimens show or A. perezi. No specimens of A. pattersoni with over several such calluses, the number depending on the 51/2 whorls have been seen, whereas Germain (1931, individual's age. p. 530-531) ascribes six or seven to the European Differences in size between the two species are shown species. The left side of the whorls of A. pattersoni by the following measurements (in millimeters). Cata is more flattened than the right side, so that there is logue numbers are those of the University of Michigan an obtuse angle of the aperture above, but only a Museum of Zoology, as follows: 177231, Sand Draw curve below. In most specimens examined, the left local fauna, locality 7, Brown County, Nebr., 177290, side is broader than the right. The largest specimens, probably Cudahy local fauna, W1/^ sec. 35, T. 33 S., however, lose the subangulation on the last whorl, E. 32 W., Seward County, Kans.; 90458, Eecent, and thus approach A. rotundatus. Like A. rotundatus Menonville, France.
Differences in the two species are as follows:
Number of Diameter Height Diameter of Height of Catalog whorls aperture aperture No.
5 3. 7 0 8 0 6 0 8 177231 4!4 3. 1 7 6 7 4% 2. 9 7 5 7 5Y2 4. 6 1 2 8 9 177290 5 l/4 4. 2 8 8 8 4^ 3. 2 7 6 7 Anisus rotundatus (Poiret) 5y2 6. 3 1 3 1 0 1 3 90458 5y2 6. 3 1 3 1 2 1 3 5 4. 9 9 9 9 4K 3. 9 9 7 9 4/2 4. 2 1 1 9 1 1
Occurrence and material: Sand Draw and Dixon local Gyraulus deflectus (Say), 1824 faunas. Sand Draw loc. 2, USGS 19090-1 (6) ; loc. 3, UMMZ Gyraulus deflectus (Say). Baker, 1928, Wisconsin Geol. Nat. 177332 (150) ; loc. 4, UMMZ 177316 (6) ; loc. 5, CNHM (3), Hist. Survey Bull. 70, pt. 1, p. 370, pi. 23, fig. 15-21. UIMNH P7455 (4), ANSP 169948 (2); loc. 6, UMMZ 177427 (48), 181211 (500) ; loc. 7, UMMZ 177231 (75) ; Dixon loc. 1, Distribution. British Columbia to Nova Scotia and UMMZ 182199 (26). Mackenzie District; south in the Eocky Mountains to Montana and Idaho; eastern South Dakota and Iowa Genus GYRAULUS Charpentier, 1837 to the Atlantic Ocean; south in the Atantic Coastal Plain to the Potomac Eiver, Virginia. Gyraulus circumstriatus (Tryon), 1866 Ecology. "A species of quiet bodies of water" (F. C. Baker, 1928, p. 370). In northern Nebraska this Gyraulus circumstariatus (Tryon). Baker, 1928, Wisconsin species was found in a spring-fed pond with a tem Geol. Nat. History Survey Bull. 70, pt. 1, p. 378, text fig. 162. perature of 15 °C. The snails were crawling on aquatic plants, largely Ceratophyllum, Elodea, and algae, near Distribution. Gyraulus circumstriatus is found in the edge of the pond. a broad belt across central North America, in south Remarks. In north-central Nebraska this species is ern Canada and the northern United States between about 200 miles south of its previously recorded range. the Atlantic and Pacific oceans, southward in the Its existence in the area is considered possible because Rocky Mountains to northern Arizona. of the insulating effect of the cool spring water. Ecology. Gyraulus circumstriatus is characteristic Essentially a northern species, it is probably able to of small, seasonal water bodies, such as woods pools, live here because the pond is warmed little if at all marshes, ponds on flood plains or prairie ponds. In during the summer hot spells. Other permanent ponds, northern Nebraska it was found in seepages beside apparently similar except that they lack spring streams, and in temporary Sand Hills ponds. sources, were examined without finding this species. Occurrence. Recent in northern Nebraska. Occurrence: Recent in northern Nebraska. 508945 O 60 6 58 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Gyraulus parvus (Say), 1817 Genus ARMIGER Hartmann, 1840
Plate 4, figs. 1-13, 17, 18 Armiger crista (Linnaeus), 1758
Gyraulus parvus (Say). Baker, 1928, Wisconsin Geol. Nat. Gyraulus crista (Linn.). Baker, 1928, Wisconsin Geol. Nat. History Survey Bull. 70, pt. 1, p. 374, pi. 23, fig. 27-31, History Survey Bull. 70, pt. 1, p. 385, text fig. 164. 39, text fig. 162. Distribution. Holarctic; in North America, Gyraulus altissimus (F. C. Baker). Baker, 1938, Nautilus, v. 51, p. 130-131 [not of Baker]. north of about 41°, but south of this latitude on the Gyraulus altissimus. McGrew, 1944, Field Mus. Nat. History Pacific Coast to San Mateo County, Calif., and south Geol. Ser., v. 9, p. 36. ward in central Utah. Gyraulus cnaulus Leonard, 1952, Nautilus, v. 66, p. 43, pi. 5, Ecology. Kenk (1949, p. 53) found A. crista was fig. G. H, I. common in a temporary pond in southern Michigan. Gyraulus enaiilus Leonard. Frye and Leonard, 1952, Kansas "Armiger crista (Linnaeus) was taken in Pond 1 from Geol. Survey Bull. 99, p. 151, pi. 14, fig. u. March to June. Numerous individuals appeared also Distribution. North America, from Alaska and in the culture of a soil sample from the bottom of the northern Canada to Cuba and from the Atlantic to pond, prepared in October. No specimens were col the Pacific coast. Perhaps also in northern Eurasia. lected from December to February. The animals Ecology. "Usually in quiet bodies of water, often obviously estivated buried in the dry mud and, after of small size. * * * parvus is partial to a habitat the pond had filled, remained inactive in the soil until which has rather thick vegetation. This species is the latter part of the winter." In northern Nebraska more often found in vegetation than in any other a single specimen was found, under culvert fill stones situation" (Baker, 1928, p. 376-377). In northern at the edge of a stream. Ecologic information on this Nebraska it was found in. permanent and temporary species in America is scanty. quiet water bodies: in permanent and temporary Occurrence: Recent in northern Nebraska. ponds, in temporary water in ditches, and in quiet spots at the edge of a stream. Subfamily HELISOMATINAE Remarks. Gyraulus altissimus Baker (1919), de Genus HELISOMA Swainson, 1840 scribed from upper Pleistocene deposits in Illinois, Submenus HELISOMA s. s. was reported from the Sand Draw local fauna. The material is G. parvus rather than G. altissimus. The Helisoma (s. s.) anceps (Menke), 1830 shells from the Sand Draw local fauna are character Helisoma antrosa (Conrad). Baker, 1928, Wisconsin Geol. ized by a lateral flattening of the whorls, relatively Nat. History Survey Bull. 70, pt. 1, p. 317, pi. 19, fig. close coiling, and no deflection of the body whorl. G. 8-15. altissimus also has closely coiled whorls, but the body Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. whorl is deflected and the flattening is at an angle of 151. Helisoma anceps (Menke). Pilsbry, 1934, Acad. Nat. Sci. about 45 degrees to the axis of the coil. Philadelphia Proc., v. 86, p. 45. Gyraulus enaulus Leonard was not compared to G. Taylor, 1954, Michigan Univ. Mus. Zool. Occ. Papers 557, parvus in the original description, but is referable to p. 14. that species as represented in University of Michigan Distribution. Oregon to Maine, southward to west Museum of Zoology and U. S. National Museum col ern Mexico and Alabama. lections. Ecology. "* * * primarily a river and creek species, Occurrence and material: Saw Rock Canyon, Red Corral, not living in the large lakes" (Baker, 1928, p. 319). Sand Draw, Dixon, and Sanders local faunas; Recent in In northern Nebraska it was found in a spring-fed northern Nebraska; Rexroad local fauna (Frye and Leonard). pond with a temperature of 15 °C. Saw Rock Canyon, UMMZ 173591 (125), 177556 (1200), uncat., USGS 19089-1 (200) ; Red Corral, UMMZ 182994 (200) ; Sand Remarks. Pilsbry (1934) considered the name an Draw loc. 1, UMMZ 181239 (250) ; loc. 2, USGS 19090-2 (100) ; trosa as a junior synonym of anceps. The name anceps loc. 3, UMMZ 177333 (2250) ; loc. 4, UMMZ 177317 (300) ; loc. has been used by subsequent writers except Leonard 5, UMMZ 181228 (125), 177219 (24), CNHM PE3436 (175), (1950, p. 16, and later papers). Pilsbry (1950, p. 68) UIMNH P6790 (41) ; loc. 6, UMMZ 177428 (200), 181212 (75), discussed the reasons for acceptance of anceps. CNHM PE3444 (46), UIMNH P6797 (15) ; loc. 7, UMMZ 177- The single Recent occurrence in a summer-cool pond 232 (150) ; Dixon loc. 1, UMMZ 182200 (6500), uncat.; loc. 2, UMMZ 191509 (300) ; Sanders loc. 1, UMMZ 182147 (5) ; loc. together with Valvata and Gyraulus deflectus might 3, UMMZ 182178 (1). be interpreted to mean that its existence in the area SYSTEMATIC DESCRIPTIONS 59 is dependent upon a temperature factor. Considering chiefly of fragments and young specimens. None its wide distribution and the number of High Plains shows the apertural dentition, but all agree well in localities given by Walker (1909), I think the con size and shape with P. armigera. trolling factor is permanent water. The series from Dixon locality 2 agrees well in size, The material reported by Frye and Leonard (1952, shape, and lamellae with Recent specimens of P. armi p. 151) from the SW^4 sec. 22, T. 33 S., R. 29 W., gera. The three immature specimens from Dixon Meade County, almost certainly came from Rexroad locality 1 are referred to the same species, since they locality 3 rather than Bender locality 1. Habitat show no differences from the locality 2 series. inferences from Bender locality 1 species suggest only The single Recent Nebraska specimen differs from temporary bodies of water. Rexroad locality 3 had P. armigera as figured and described by Baker (1928) the permanent water which Helisoma anceps requires. in having a lesser axial height, five whorls, and a more Occurrence and material: Sand Draw and Dixon local open spire-pit. It differs from P. christyi (Dall) as faunas; Saw Rock Canyon and Rexroad local faunas (Frye figured by Baker (1945, pi. 119, figs. 13-15) in having and Leonard) ; Recent in northern Nebraska. Sand Draw loc. fewer whorls. The specimen probably represents nor 1, UMMZ 181240 (65) ; loc. 2, UMMZ 184173 (5) ; loc. 3, UMMZ mal variation within P. armigera. 177334 (250) ; loc. 6, UMMZ 177429 (1) ; Dixon loc. 1, UMMZ 182201 (100) ; loc. 2, UMMZ 191510 (1). Occurrence and material: Sand Draw and Dixon local faunas; Recent in northern Nebraska. Sand Draw loc. 1, Subgenus PIEROSOMA Ball, 1905 UMMZ 181241 (44) ; loc. 4, UMMZ 177318 (14) ; Dixon loc. 1, UMMZ 182203 (3) ; loc. 2, UMMZ 191512 (200). Helisoma (Pierosoma) trivolvis (Say), 1817 Genus PROMENETUS Baker, 1935 Helisoma trivolvis (Say). Baker, 1928, Wisconsin Geol. Nat. History Survey Bull. 70, pt. 1, p. 330, pi. 20, fig. 1-13, Subgenus PROMENETTIS s. s. 22, 23. Promenetus (s. s.) exacuous (Say), 1821 Distribution. "Atlantic Coast and Mississippi Menetus exacuous (Say). Baker, 1928, Wisconsin Geol. Nat. River drainages, northward to Arctic British Amer History Survey Bull. 70, pt. 1, p. 361, pi. 23, fig. 1-5. ica and Alaska and southward to Tennessee and Mis Promenetus exactions exacuous (Say). Baker, 1945, Molluscan souri. The southern distribution is not clear owing family Planorbidae, p. 182. to mixing with related species" (Baker, 1928, p. 332). Distribution. North America north of about 39°; Ecology. "Typical trivolvis is always an inhabitant south in the Colorado Plateau and Rocky Mountains of quiet, more or less stagnant water" (Baker, 1928, p. to New Mexico; south along the Atlantic Coastal Plain 332). In northern Nebraska it was found in perma to North Carolina; an isolated occurrence in cool nent ponds almost exclusively. The exception was a springs, Meade County, Kans. single specimen found in a drainage ditch where it Ecology. Promenetus exacuous is found in shallow, might have come from a permanent pond. perennial or subpernianent, quiet water bodies, such as Occurrence and material: Dixon local fauna; Recent in ponds, ox-bow lakes, marshes, and sloughs or back northern Nebraska. Dixon loc. 1, UMMZ 182202 (2, + frag waters along streams. It is usually on the submerged ments) ; loc. 2, UMMZ 191511 (72). vegetation in such habitats. The isolated occurrence in Subfamily PLANORBTILINAE Meade County, Kans., can be explained by the summer- cool habitat furnished by artesian water in which this Genus PLANORBULA Haldeman, 1842 snail lives. In northern Nebraska it was found in Planorbula armigera (Say), 1818 permanent or nearly permanent quiet water. Planorbula armigera (Say). Baker, 1928, Wisconsin Geol. Occurrence: Recent in northern Nebraska. Nat. Hist. Survey Bull. 70, pt. 1, p. 355, pi. 8, fig. 27-30. Planorbnla sp. Taylor, 1954, Michigan Univ. Mus. Zool. Occ. Promenetus (s. s.) kansasensis (Baker), 1938 Papers 557, p. 15. Plate 3, figures 7, 8, 12-30
Distribution. "New England west to Nebraska, Menetus kansasensis Baker, 1938, Nautilus, v. 51, p. 129. south to Georgia and Louisiana, north to Great Slave Menetus kansasensis F. C. Baker. Baker, 1938, Nautilus, v. Lake." (F. C. Baker, 1928, p. 359.) 51, p. 130-131. Ecology. "* * * largely a species of swales or of Hibbard, 1941, Kansas Univ. Sci. Bull., v. 27, p. 95. small and stagnant bodies of water." (F. C. Baker, Hibbard. 1941. Kansas Acad. Sci. Trans., v. 44, p. 265. 1928,. p. 358). In northern Nebraska it was found Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. 408. in a small oxbow lake in the Niobrara River valley. Baker, 1945, Molluscan family Planorbidae, p. 516, pi. 140, Remarks. Sand Draw local fauna material consists fig. 26-29. 60 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Menetus kansasensis Baker. Smith, 1940, Kansas Geol. Sur Promenetus (Phreatomenetus) umbilicatellus (Cockerell), 1887 vey Bull. 34, p. 408. Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99. p. Gyraulus umbilicatellus (Cockerell). Baker, 1928, Wisconsin 151, pi. 14, fig. t. Geol. Nat. History Surrey Bull. 70, pt. 1, p. 383, pi. 22, Leonard, 1957, Illinois Geol. Survey Report of Investiga fig. 18-21. tions 201, p. 13, pi. 2, fig. 1-3. Menetus unibilicatettus (Cockerell). Baker, 1938, Nautilus, v. Menetus kansasensis. McGrew, 1944, Field Mus. Nat. History 51, p. 130. Geol. Ser., v. 9, p. 36. Gyraulus unibllicatellus. McGrew, 1944, Field Mus. Nat. His tory Geol. Ser., v. 9, p. 36. Types. Holotype TJIMNH P6778, Kexroad loc. 3. Promenetus timUlicatellus (Cockerell). Baker, 1945, Mollus- Paratypes UIMNH P6778 (3), uncataloged (20), can family Planorbidae, p. 182, pi. 123, fig. 34-36. ANSP 169884 (not found). Promenetus Wancoensis Leonard, 1952, Nautilus, v. 66, p. 42, Diagnosis. Similar in shape and size to Promenetus pi. 5, fig. D-F. Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, exacuous, but with sculpture of numerous irregularly p. 151, pi. 14, fig. o. spaced riblets. Distribution. North America north of about lat. Distribution. Late Pliocene to late Pleistocene 41° from Nevada and Alaska east and southeast to (Sangamon) in the Great Plains. western New York; south in the Rocky Mountains to Occurrence and material: Saw Rock Canyon, Red Corral, southern Colorado; an isolated occurrence in the Ozark Rexroad, Sand Draw, Dixon, and Sanders local faunas. Saw Mountains, northeastern Oklahoma. Rock Canyon, UMMZ 173588 (75), 177557 (2500), uncat., USGS Ecology. "All specimens in Wisconsin have been 19089-2 (75) ; Red Corral, UMMZ 182995 (8) ; Rexroad loc. found in swales11 (Baker, 1928, p. 384). In northern 1, UMMZ 183055 (1) ; loc. 3, UIMNH P6778 (4), uncat. (20), Nebraska it was found common in drainage ditches UMMZ 183029 (12) ; Sand Draw loc. 1, UMMZ 181242 (100) ; loc. 2, USGS 19090-3 (48) ; loc. 3, UMMZ 177335 (1500) ; loc. and temporary ponds. 4, UMMZ 177319 (54) : loc. 5, CNHM PE 3434 (30), UIMNH Remarks. Promenetus blancoensis appears to be a P6788 (17), UMMZ 177217 (3000), 181229 (3500); loc. 6, synonym of P. umbilicatellus, easily falling within the UIMNH P6798 (6), UMMZ 177430 (100), 181213 (600); loc. range of variation of the latter. The differentiae given 7, UMMZ 177229 (125) ; Dixon loc. 1, UMMZ 182204 (1000) ; were the smaller size, lesser number of whorls (always loc. 2, UMMZ 191513 (150) ; Sanders loc. 2, UMMZ 182165 less than 4), nonstriate nucleus, absence of spiral (2) ; loc. 3, UMMZ 182179 (1). sculpture, and depressed spire. The great majority of Subgenus PHREATOMENETTTS 8 Taylor, n. subg. Recent specimens examined have less than 4 whorls and have a diameter of approximately 4 millimeters, Type. Promenetus umbilicatellus (Cockerell). thus agreeing with the measurements of P. l)lancoensis. Diagnosis. Snails fitting the definition of Promene To the "nonstriate nucleus1' not much significance tus (see Baker, 1945, p. 178-182), but distinguished should be atached. All other characters are in agree conchologically as follows: shell broadly rounded ment with those of P. umbilicatellus, and the nuclear peripherally; sides of whorls convex to slightly flat striation might easily be lost through wear. The ap tened; left side strongly convex or subangular at edge parent absence of spiral sculpture does not differentiate of spire-pit; right side sometimes strongly convex or the form from P. umbilicatellus, for this character is subangular near the suture. often very difficult to observe in unworn Recent spe Referred species. None of the species besides the cimens, and in fossils faint spiral sculpture might have type is known anatomically; others listed below are as been lost or easily overlooked. The depressed spire is signed on shell characters. also a character which may be found in Recent speci Promenetus umUlicatellus (Cockerell), 1887. Type. mens, although it is not common. Material from the Northern North America. Saw Rock Canyon local fauna, and from Rexroad local earns (Pilsbry and Ferriss), 1906. fauna localities 1 and 3 confirms the synonymization. Southwestern Texas. The specimens agree well in size and form with P. um- circumlineatus (Shuttleworth), 1854. bilicatellux, and less worn individuals show the spiral Central America to the Lesser Antilles. striation and nuclear sculpture typical of that spe (Including Planorbis santacruzensis Germain, cies. 1923; may be a synonym of Plcm,orbis aeruginosus The material reported by Frye and Leonard (1952, Morelet, 1851). p. 151) from SW4 sec, 22, T. 33 S., R. 29 W., Meade County, is probably from Bender locality 1 rather than * Gr. phrear, -atos, n., well or reservoir; in reference to the steep- sided, well-nke spire-pit of the species referred to the group. Rexroad locality 3. University of Michigan collections SYSTEMATIC DESCRIPTIONS 61 contain numerous specimens from the Bender locality, Hibbard, 1941, Kansas Univ. Sci. Bull., v. 27, p. 95. but none from the Rexroad locality. Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. Occurrence and material: Saw Rock Canyon, Red Corral, 408. Rexroad, Bender, Sand Draw, Dixon, and Sanders local Frye and Leonard, 1952, p. 151. faunas; Recent in northern Nebraska. Saw Rock Canyon, UMMZ 177515 (50), 177558 (600), USGS 19089-3 (75); Red Taylor, 1954, Michigan Univ. Mus. Zool. Occ. Papers 557, Corral, UMMZ 182996 (150) ; Rexroad loc. 1, UMMZ 183056 p. 15. (1500) ; Bender loc. 1, UMMZ 177505 (1) ; loc. la, UMMZ Ferrissia rivularis"? (Say). Hibbard, 1954, Michigan Acad. 184120 (30) ; loc. Ic, UMMZ 183014 (11) ; loc. 2, UMMZ 184151 Sci. Papers, v. 39, p. 343. (47) ; Sand Draw loc. 1, UMMZ 181243 (6) ; loc. 2, UMMZ Distribution. "Massachusetts west to Nebraska, 184174 (40) ; loc. 3, UMMZ 177336 (225) ; loc. 4, UMMZ 177320 (150); loc. 5, CNHM PE3435 (75), UIMNH P6789 (36), Ohio nortlrward to Wisconsin and Michigan" (F. C. UMMZ 177218 (125), 181230 (800) ; loc. 6 UMMZ 177431 (300), Baker, 1928, p. 399). 181214 (6000) ; loc. 7, UMMZ 177230 (250) ; Dixon loc. 1, Ecology. "Found on stones in streams" (Robertson UMMZ 182205 (9) ; loc. 2, UMMZ 191514 (6) ; Sanders loc. and Blakeslee, 1948, p. 72). 1, UMMZ 182148 (300); loc. 2, UMMZ 182166 (2), 184162 (31) ; loc. 3, UMMZ 182180 (20). Occurrence and material: Saw Rock Canyon, Rexroad, Sand Draw, and Dixon local faunas; Buis Ranch local fauna? Family AWCYLIDAE Buis Ranch, UMMZ 181114 (1, fragmentary) ; Saw Rock Subfamily ACROLOXINAE Canyon, UMMZ 173594 (24), 177559 (350), USGS 19089-9 (100) ; Red Corral, UMMZ 182997 (5) ; Rexroad loc. 3, UMMZ Genus ACROLOXTTS Beck, 1837 183031 (8) ; Sand Draw loc. 1, UMMZ 181245 (57) ; loc. 2, Acroloxus coloradensis (Henderson), 1930 UMMZ 184175 (1) ; loc. 3, UMMZ 181111 (250) ; Dixon loc. 1, UMMZ 182210 (56). Plate 1, figures 1-4
Ancyltis hendersoni Walker, 1925, Michigan Univ. Mus. Zool Ferrissia parallela (Haldeman), 1841 Occ. Papers 165, p. 1. Mozley, 1926, Nautilus, v. 40, p. 56. Ferrissia parallela (Haldeman). Baker, 1928, Wisconsin Geol. Ancylus coloradensis Henderson, 1930, Nautilus, v. 44, p. 31. Nat. History Survey Bull. 70, pt. 1, p. 395, pi. 24, fig. Taylor, 1954, Michigan Univ. Mus. Zool. Occ. Papers 557, 1-5. p. 15. Hibbard, 1941, Kansas Univ. Sci. Bull., v. 27, p. 95. Distribution. Known from only two areas in the Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. Rocky Mountains in Colorado and Alberta. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. Ecology. In high mountain lakes. "* * * only 408. clinging to stones, etc., in very tiny inlets from the Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. lake, where they were well protected from the waves, 151, pi. 14, fig. n. but in clear lake water, not in swampy ground" (Walker, 1925, p. 3). Distribution. Nova Scotia and New England west Remarks. The discovery of this rare and little- to Manitoba and northeastern Illinois (F. C. Baker, known species in the Pleistocene of the Great Plains is 1928, p. 397). remarkable. The specimens are closely similar to a Ecology. "Femssia parallela is usually found in series of paratypes, however, and also to specimens quiet water, on plants, the water in such places ranging collected in Alberta by Mozley. from .3-2 meters in depth. The animal is usually International Commission on Zoological Nomen found near the surface but may occur on the lower part clature Opinion 363 (1955) fixes the type oiAncylus as of such plants as Scirpus, near the bottom. Parallela A. fluviatilis Miiller, 1774, and of Acroloxus as Patella lacustris Linnaeus, 1758, as interpreted by Miiller, 1774. appears to be a pond or lake species, at least in Wiscon Ancylus coloradensis Henderson is accordingly to be sin" (F. C. Baker, 1928). called Acroloxus coloradensis (Henderson). Occurrence: Rexroad local fauna (Hibbard). Occurrence and material: Sand Draw and Dixon local faunas. Sand Draw loc. 3, UMMZ 181129 (28) ; Dixon loc. 1, Ferrissia meekiana (Stimpson), 1863 UMMZ 182209 (1) ; loc. 2, UMMZ 191515 (1). Gundlachia meekiana Stimpson, 1863, Boston Soc. Nat. History Subfamily FERRISSIINAE Proc., v. 9, p. 250. Genus FERRISSIA Walker, 1903 Binney, 1865, Smithsonian Misc. Coll. 143, p. 150. Pilsbry, in Heclley and Pilsbry, 1895, Nautilus, v. 9, p. 63. Ferrissia rivularis (Say), 1819 Ancylus pumilus Sterki. Walker, 1904, Nautilus, v. 18, p. 82, Ferrissia rivularis (Say). Baker, 1928, Wisconsin Geol. Nat. pi. 6. fig. 20-22. History Survey Bull. 70, pt. 1, p. 398, pi. 24, fig. 16-18. Walker, 1914, Nautilus, v. 27, p. 129. 62 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Ferrissia cf. pumila (Sterki). Taylor, 1954, Michigan Univ. ture before continuing growth. The larger is striate anteriorly. Mus. Zool. Occ. Papers 557, p. 15. (d) Four mature specimens, 3.0-3.2 mm long, nonseptate, with nearly straight sides roughly parallel or slightly diver Distribution. Probably the species ranges over most gent anteriorly. Three are anteriorly striate. of the eastern United States. It is known from scat (e) Twenty-nine specimens, immature and mature, reach tered localities in Alabama, Missouri, Virginia, the ing a size of 3.3 X 1.9 mm. The lateral outline of the shell District of Columbia, and from Iowa eastward to New varies from only slightly to moderately curved. One only is York. anteriorly striate. The apex is about one-fourth of the shell length from the posterior end and somewhat to the right of Ecology. Uncertain. It has been found in tempo the median line. The right slope is straight or gently con rary pools, ponds, and sloughs, but may occur in other cave ; the left slope straight or gently convex. The height of habitats as well. the shell varies from less to more elevated than F. pumila as Remarks. The kinds of shells called Ferrissia meek- figured by Walker (1904, pi. 6, fig. 20). Most shells are more iana in this paper have been found at several locali depressed, however, with a slightly more concave posterior slope. The anterior end of the shell is symmetrical in out ties in Pleistocene deposits of the Great Plains. Fre line, the posterior end usually so. The slight flattening of quently the very small, septate, immature shells are as- the dextroposterior margin is not as marked as in the speci associated with larger, non-septate shells similar to F. mens from Sand Draw loc. (UMMZ 181244). This group pumila. Only one sample large enough to show the agrees well with a series (UMMZ 67269) from Clear Lake, range of variation adequately has been collected; it is La Porte, La Porte County, Ind., identified as Gundlachia from Dixon locality 2 (UMMZ 191516). On the basis meekiana by Bryant Walker. of the range of variation in this fossil sample, the range Family PHYSIDAE of variation in Recent material in the University of Michigan Museum of Zoology, and the similarity of Genus FHYSA Draparnaud, 1801 the fossils to specimens identified by Walker as Grund- Fhysa anatina Lea, 1864 lachia meekiana it appears that the small, septate shells and larger nonseptate specimens represent one species, Physella anatina Lea. Baker, 1928, Wisconsin Geol. Nat. History Survey Bull. 70, pt. 1, p. 470, pi. 19, fig. 5 and that Ancylus pumilus Sterki represents the non- [not 7]. septate form of "Grundlachia" meekiana. Walker Physa species, young and fragments. Baker, 1938, Nautilus, (1914, p. 129) reached similar conclusions from study v. 61, p. 130. of Recent shells. Physa anatina Lea. Hibbard, 1941, Kansas Univ. Sci. Bull., Watson (in Connolly, 1938, p. 525-526) presented v. 27, p. 95. Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. evidence to indicate that reference of all septum-form Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, ing Ancylidae to Gundlachia is unjustified. He re p. 408. stricted Gundlachia to tropical American and West Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, Indian species with a smooth apex and an asymmetrical p. 151, pi. 14, fig. z. central tooth, and placed apically striate septum-form Physa sp. McGrew, 1944, Field Mus. Nat. Hist. Geol. Ser., v. ing species (Kincaidilla} in Ferrissia. In accordance 9, p. 36 [in part]. with this classification Gundlachia meekiana (includ Distribution. Very poorly known, on account of the ing Ancylus pumilus} is referred to Ferrissia. lack of study of the genus. Probably it will be found Occurrence and material: Sand Draw and Dixon local in most of the central and southern Great Plains. faunas. Sand Draw loc. 1. UMMZ 181244 (7), 181246 (2) ; Crandall (1901, p. 57) states, "It is plentiful around loc. 3, UMMZ 177338 (13) ; Dixon loc. 1, UMMZ 182211 (5) ; Wichita, Kansas, and has been reported from Missouri, loc. 2, UMMZ 191516 (44). Kansas, Nebraska, and Michigan. It probably inhabits Sand Draw loc. 1: Only one of the nine specimens is all the intervening territory." septate. Loc. 3: all 13 lack septa. Dixon loc. 1: Of the 5 specimens 3 are septate. Loc. 2: 7 of the 44 are septate. Ecology. This species is found in almost any sort of The series from Dixon loc. 2 is the only one large enough to perennial, shallow water body, in either flowing or give a reasonably adequate idea of variation within the standing water. In Meade County, southwestern Kan population. It may be divided into five groups, as follows: sas, it can be found in small streams, in the spring-fed (a) Five small specimens, 1.7-2.3 mm long, with the septum covering about three-quarters of the aperture. These ponds in the State Park, or in stock tanks. In northern are similar to the two small, septate specimens from Dixon Nebraska it was not found in still water, but only in loc. 1. Three are striate anteriorly. seepages or spring-fed streams. (b) Four small specimens, 2.3-2.5 mm long, nonseptate. These are similar in shape to those of group (a), and to one Occurrence and material: Saw Rock Canyon, Red Corral, specimen from Sand Draw loc. 3. All four are smooth. Rexroad, Bender, Sand Draw, Dixon, and Sanders local (c) Two immature specimens, 2.5 and 3.3 mm long. Both faunas; Recent in northern Nebraska. Saw Rock Canyon, apparently formed a septum over about one-fifth of the aper UMMZ 172364 (31), 177560 (2500), USGS 19089^ (800); SYSTEMATIC DESCRIPTIONS 63
Red Corral, UMMZ 182998 (150) ; Rexroad loc. 1, UMMZ Types. Holotype UMMZ 181292, Berends local fauna. Para- 183057 (35) ; loc. 2, UMMZ 183067 (1) ; loc. 3, UMMZ 183030 types UMMZ 177533 (31), MCZ 198177 (68), USNM 562010 (60); loc. 4b, UMMZ 177600 (3000), 181049 (1500); Bender (12). loc. la, UMMZ 184121 (3) ; loc. Ib, UMMZ 184135 (1) ; loc. Diagnosis. A species distinguished by its small size 2, UMMZ 184152 (50) ; Sand Draw loc. 2, UMMZ 184176 (1) ; loc. 3, UMMZ 177340 (55) ; loc. 4, UMMZ 177321 (8) ; loc. 5, (less than 7.5 mm), narrow ovoid form, and obtusely CNHM PB3432 (4), UIMNH P6786 (3), UMMZ 181231 (25); rounded apex. It is most similar to P. fontinalis loc. 6, UMMZ 177432 (1), 181215 (1); Dixon loc. 1, UMMZ (Linne) and P. hordacea Lea. 182206 (150) ; Sanders loc. 1, UMMZ 182149 (13) ; loc. 2, Description of types. Shell small, narrowly ovoid, UMMZ 184163 (2) ; loc. 3, UMMZ 182181 (3 + fragments). polished; apex obtusely rounded. Whorls about three, Physa gyrina Say, 1821, form hildrethiana Lea, 1841 gently and regularly convex, slowly enlarging, at first overlapping; body whorl over three-fourths of total PJiysella gyrina hildrethiana (Lea). Baker, 1928, Wisconsin shell length. Suture weakly impressed, slightly Geol. Nat. History Survey Bull. 70, pt. 1, p. 453, pi. 27, descending at first but later more strongly; above pe fig. 36; pi. 28, fig. 2-4, 7-14. Physa elliptica Lea. Taylor, 1954, Michigan Univ. Mus. Zool. riphery of first whorl, at periphery of later whorls. Occ. Papers 557, p. 15. Aperture narrowly elongate; outer lip slightly convex, Distribution. Probably found in most of North more strongly so toward base; base strongly rounded; America north of Mexico. According to Baker (1928), inner lip nearly straight or with a pronouced angle at P. gyrina is found "from the Arctic regions south to junction of columella and parietal lip; peristome usu Alabama and Texas;" the form hildrethiana, " * * * ally heavily callused in larger specimens, in which case is common east of the Mississippi River from western the suture descends slightly at the callus; parietal wall New York and Pennsylvania to Illinois and south to covered by a thin callus. Nucleus smooth; later sculp Alabama. Occurs also in Iowa and possibly in other ture of A^ery fine, indistinct growth lines occasionally states west of the Mississippi River/' cut by very faint, short segments of spiral striae be Ecology. "Hildrethiana is characteristic of swales tween which growth lines may be locally more promi nent. and summer-dry ponds, where it is deprived of moisture Measurements. The mean of 21 adult paratypes of for a large part of the year. The variety in such 3 whorls is folloAved by the range in parentheses. These localities rarely reaches maturity, the usual form being are judged to be adult because they usually have a a small, short, shell with dome-shaped spire, greatly heavy peristome callus and because specimens with resembling oleacea and elliptica. These small forms more whorls are rare. Measurements are given in mil usually show no rest marks, living only as long as the limeters. Length, 3.7 (3.0-4.3); width, 2.0 (1.7-2.5); water remains in the pool. Mature shells may often length of aperture, 2.2 (1.7-2.6) ; width of aperture, be found by digging in the mud or in cracks in the 1.1 (0.8-1.3). bottom of the pool where the mollusk has descended The measurements in millimeters of 10 Recent spe for moisture" (Baker, 1928, p. 454). In northern cimens from Utah (USNM 533484) are as follows: Nebraska this snail was common at many places, in temporary ponds and drainage ditches, permanent Length of Width of Number of Length Width aperture aperture whorls ponds and a stock tank. P. gyrina, and P. anatina were 6. 4 3. 4 4. 0 1. 8 3% never found together. P. anatina occured only in seep 6. 7 3.7 4. 4 2. 1 4 7. 3 4. 0 4. 6 2. 1 4 ages and streams. 6. 9 3.7 4. 1 2. 0 4 7. 1 3. 9 4. 3 2. 1 4 Occurrence and material: Sand Draw and Dixon local 6. 6 3. 6 3. 9 1. 9 4 faunas: Recent in northern Nebraska. Sand Draw loc. 1, 6. 7 3. 7 4. 1 2.0 4 6. 6 3. 6 3. 9 1. 9 4 UMMZ 181247 (13) ; loc. 2, UMMZ 184177 (3) ; Dixon loc. 1, 7. 1 3. 9 4. 1 1. 8 4H UMMZ 182207 (60) ; loc. 2, UMMZ 191517 (65). 6. 9 3. 7 4.0 2. 0 4H
Physa skinneri Taylor, 1954 These are the largest specimens seen, in both size and number of whorls. Plate 3, figures 5, 6, 9-11 Variation. The proportions vary slightly, as in Physa species, young of small, narrow species. Baker, 1938, dicated by the measurements, but the obtuse apex is Nautilus, v. 51, p. 130. constant. In the shorter shells the shallow suture gives Physa sp. McGrew, 1944, Field Mus. Nat. Hist. Geol. Ser., v. 9, p. 36 [in part]. an almost unbroken curve from apex to base. Some Physa skinneri Taylor, 1954, Michigan Univ. Mus. Zool. Occ. times an earlier peristomal callus is visible through Papers 557, p. 9, 15. the shell. The callus is accompanied by the usual brief 64 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS descent of the suture, which thereafter ascends and re tory (USNM 251936), and a pond at lat. 69° 40' N., sumes the former angle of descent. long., 141° W., on the coast of Yukon Territory Distribution. Alaska southward to northern Utah; (USNM 251938). eastward across southern Canada to western Ontario Physa hordacea Lea (1864, p. 116; 1866, p. 176; 1867, and northern Minnesota. p. 132; this paper, pi. 3, figs. 1-4) is similar to P. skin- Ecology. The scanty information from the Recent neri in size, sculpture and general shape, but has an occurrences suggests this species lives in shallow bodies acute apex instead of a blunt, rounded one. The only of water, either perennial or seasonal, such as tempo known material with precise locality data is the type rary ponds, sloughs, and backwaters along streams. lot from Columbia River, Vancouver Island, at Van Remarks. A number of Recent series of P. skinneri couver, Clark County, Wash. (USNM 170764, holotype are now known which demonstrate that the species is and 111 paratypes). not extinct, as was first thought, and that it has a wide The European Physa fontinalis (Linne) is perhaps distribution in North America. This living material the species most closely related to P. skinneri. It also helps to establish the range of variation in P. shares the blunt, rounded apex of the latter, but has skinneri. usually another whorl and is half again as long. It is Two of the larger series of adults from the Uinta proportionally wider, globose rather than narrowly Mountains, Wyoming and Utah, differ from the type ovoid as is P. skinneri, and thus has a wider aperture series in being slightly larger, thinner shelled and with with the inner lip markedly angular at the junction weak or absent peristomal calluses. The larger series of columellar and parietal portions. (USNM uncat., 72 specimens) is from the Heward The grouping of Physa species is very diffcult, be ranch, Uinta County, Wyo., from a slough by the Bear cause the limits of variation of the species are poorly River 1 mile above the Utah border. Numerous adult understood, and biological information is not avail shells agree well in proportions with the type series. able for more than a very few species, however they are They attain 31/2-4 whorls, and reach a length of 6 mm. delimited. The relations of P. skinneri, as inferred There are no peristomal calluses. Sculpture consists from a single shell character, may be viewed in two of weak growth lines, and many spiral series of very ways: The blunt, rounded apex of P. skinneri and P. short, axial, impressed lines. fontinalis indicates their close relation. P. hordacea is Another lot (USNM 533484, 10 specimens) is from similar only by convergence, for its acute apex indi Lost Creek, Summit County, Utah, T. 1 S., R 5 E., cates a quite different ancestry. P. hordacea and P. 6,000 feet elevation. All ten shells are adult, and skinneri are so similar that they must be closely related larger than those of the type series or those from Uinta no matter what apical differences there may be. P. County (see measurements). They are also proportion fontinalis is similar only in the not-too-significant fea ally higher spired, but not more so than the extreme ture of a rounded apex. observed in fossil material. Sculpture is as in the When anatomical data are available on the American Uinta County material. Several individuals have a species, it wrill be possible to evaluate these shell char thin callus around the peristome and another one about acters, and perhaps even to recognize subgenera of %-l whorl earlier. The suture descends slightly to Physa. the earlier callus, as it does in the type series. Types. Holotype FMMZ 181292, Berends local fauna. In the Pleistocene of the Great Plains, Physa skin- Paratypes UMMZ 177533 (31), MCZ 198177 (68), USNM neri is commonly associated with P. anatina Lea and P. 562010 (12). gyrina Say. It differs from these two species in its Occurrence: Sand Draw, Dixon, and Sanders local faunas. smaller size, narrower form, obtuse and rounded apex, Sand Draw loc. 2, USGS 19090-4 (9) ; loc. 3, UMMZ 177339 (100) ; loc. 4, UMMZ 177322 (3) ; loc. 5, CNHM PE3433 (3), regularly convex whorls, and proportionally smaller UIMNH P6787 (3), UMMZ 181137 (25), 181232 (100); loc. aperture. It apparently has no close relatives living in 6, UMMZ 177433 (4), 181216 (100); loc. 7, UMMZ 177233 the Great Plains. (20) ; Dixon loc. 1, UMMZ 182208 (36) ; loc. 2, UMMZ 191518 Physa jennessi Ball (1919, p. 20A) differs from P. (22) ; Sanders loc. 1, UMMZ 182150 (1) ; loc. 3, UMMZ 182- skinneri by being nearly twice as long, with l-l^ 182 (4). Genus APLEXA Fleming, 1820 more whorls, a deeper suture, and a proportionally longer spire and shorter aperture. The columella may Aplexa hypnorum (Linnaeus), 1758 be markedly gyrate, a feature not seen in P. skinneri; Aplexa hypnorum (Linne). Baker, 1928, Wisconsin Geol. Nat. no trace of peristomal callus is visible. The only locali History Survey Bull. 70, pt. 1, p. 473, pi. 19, fig. 1-4. ties known surely are Bernard Harbor, Northwest Ter Taylor. 1954, Michigan Univ. Mus. Zool. Occ. Papers 557, ritories (type locality), White Horse, Yukon Terri p. 15. SYSTEMATIC DESCRIPTIONS 65 Distribution. Holarctic. In North America "* * * S. labyrinthica-texasiana group; they differ from those from east of the Cascade Mountains to the Atlantic and observed in /S. sparsicostata topotypes in that the first from Alaska and Hudson Bay south to the vicinity of and second basals are not so large, the third through the Ohio River'1 (Baker, 1928, p. 474). fifth basopalatals are stronger, and the sixth basopa Ecology. ^ A pi. eve a hypnorum is a species of swales latal is sometimes absent. Six specimens examined have and intermittent streams or stagnant pools in Wiscon parietal lamellae entering half a whorl. sin, as far as present data goes. It is especially Frye and Leonard (1952) reported S. cf. labyrin abundant in woodland pools which become dry in thica in association with S. sparsicostata from the Saw summer * * *" (Baker, 1928, p. 474). In northern Rock Canyon local fauna. University of Michigan Nebraska it was locally abundant in roadside drainage material from the same locality justifies recognition ditches with temporary water. of only one species, 8. labyrinthica (Say). Occurrence and material: Sand Draw and Sanders local For discussion of the differences between #. laby faunas; Recent in northern Nebraska. Sand Draw loc. 3, rinthica and S. sparsicostata, see under the latter UMMZ 181202 (1) ; loc. -i, UMMZ 177323 (2) ; loc. 5, UMMZ species. 181138 (100). 181233 (300) ; loc. 6. UMMZ 181217 (1) ; Sanders Occurrence and material: Saw Rock Canyon local fauna. loc. 1, UMMZ 182151 (16). UMMZ 177561 (35).
Order STYLOMMATOPHORA Strobilops (s. s.) sparsicostata Baker, 1938
Family STROBI10PSIDAE Plate 1, figures 7-10 Genus STROBI10PS Pilsbry, 1893 Strobilops sparsicostata Baker, 1938, Nautilus, v. 51, p. 127. Submenus STROBILOPS s. s. Smith, 1940, Kansas Geol. Survey Bull. 34, p. 98. Leonard. 1957, Illinois Geol. Survey Report of Investiga Strobilops (s. s.) labyrinthica (Say), 1817 tions 201. p. 18, pi. 1, fig. 4-6. Strobilops sparsicostata F. C. Baker. Hibbard, 1941, Kansas Plate 1, figures 5, 6 Univ. Sci. Bull., v. 27, p. 95. Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. Strobilops labyrinthica (Say). Pilsbry, 1948, Acad. Nat. Sci. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, Philadelphia Mon. 3, v. 2, pt. 2, p. 854, fig. 463. p. 408. Strobilops cf. labyrinthica. Frye and Leonard, 1952, Kansas Strobilops sparsicosta Baker. Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. 151 [in part: Seward County Geol. Survey Bull. 99. p. 151 [in part: Meade County loc.]. loc.]. Strobilops sparsicosta Baker. Frye and Leonard, 1952, Kansas Strobilops densecostata F. C. Baker. Pilsbry, 1948, Acad. Nat. Geol. Survey Bull. 99, p. 151 [in part: Seward County Sci. Philadelphia Mon. 3, v. 2, pt. 2, p. 851 [lapsus for loc.]. sparsicostata]. Distribution. "Maine and Quebec west to Manitoba, Strobilops cf. labyrinthica. Frye and Leonard, 1952, Kansas Minnesota, Kansas and Arkansas, south to Georgia Geol. Survey Bull. 99, p. 151 [in part: Meade County loc.]. and Alabama" (Pilsbry, 1948, p. 854). Types. Holotype UIMNH P6774, Rexroad loc. 3. Paratypes Ecology. "Its usual stations are 'under loose bark UIMNH P6774 (51), P6775 (4), ANSP 169886 (not found). of logs, in half-decayed wood, among dead leaves and Diagnosis. A Strobilops (s.s.) characterized by 5- in sod at bases of trees' v (Pilsbry, 1948, p. 854). 6 strongly unequal basopalatal folds, the sixth (when Remarks. Specimens from the Saw Rock Canyon present) above the periphery, and by axial sculpture local fauna differ from S. texasiana and $. sparsico- of strong, widely spaced riblets not extending onto stata, and agree with S. labyrinthica. in that the axial the base of the shell. riblets are small, not widely spaced, and usually cover Description. Shell broadly conic, subangular at pe about half the base, and also that the parietal lamellae riphery. Whorls about 5l/2> embryonic l1/^ weakly enter only one-half whorl. They are also smaller and finely granular with faint axial striation; later (diameter 2.5 mm.) than /S. sparsicostata topotypes whorls with axial riblets, small and closely but irregu or referred specimens. These specimens are thus con larly spaced from 11^-3 whorls, becoming stronger sidered referable to S. labyrinthica. and wider spaced thereafter; last 2 whorls with strong, Sculpture on the base is variable. The riblets may irregularly and widely spaced riblets, occasionally pass over the periphery to cover the whole base, or with 1 or 2 weak riblets between. Base smooth, or with only about a quarter of a whorl behind the aperture. a few weakening riblets extending over the periphery Ten specimens were dissected for examination of the on the last quarter whorl; umbilicus contained 7 times basopalatal folds. These folds are as usual for the in shell diameter. Aperture roughly semicircular, with 66 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS a prominent, massive, parietal lamella extending for tween the first and second of the usual series, 1 between ward to fuse with a thick parietal callus; a low, incon the second and third. The most axiad supernumerary spicuous, infraparietal lamella almost reaching the fold is fused to the first basal fold, but the next two parietal callus, visible externally; lip reflected, thick are distinct. The fourth supernumerary fold is fused ened. Parietal and infraparietal lamellae entering to the second basal fold. The third basal is present three-fourths of a whorl; a weak interparietal lamella and typical, but the fourth through sixth are absent. about a quarter whorl long nearly reaching the inner (The description and the above discussion of variation ends of the other parietal lamellae; interparietal and are based on about 250 topotypes, UMMZ 183032; 15 inner halves of parietal and infraparietal lamellae specimens were dissected to observe the basopalatal with prickly nodes at intervals of about 11/2 times folds, 10 for the parietal lamellae.) their length; columellar lamella low, about half a Distribution. Middle Pliocene to lower Pleistocene whorl long, beginning at about a quarter whorl within (perhaps middle Pleistocene also) of the High Plains. aperture, increasing and decreasing in size gradually; Remarks. /Strobilops sparsicostata is closely related basopalatal folds parallel, arranged in an arc from to both 8. labyrinthica (Say) and 8. texasiana Pilsbry about two-thirds of a whorl within aperture nearest and Ferriss. It differs from 8. labyrinthica by the the columella to one-third of a whorl within at the slightly larger size, deeper penetration of the parietal periphery, 5-6 in number, the sixth above the periph lamellae, coarser and more widely spaced axial riblets, ery if present; first 2 basal folds much larger than and sometimes smoother base. It differs from 8. texas- others, the first nearly horizontal, roughly triangular iana in the slightly larger size and smooth base. or tongue shaped, the second longer, erect, lamellar; Frye and Leonard (1952) reported S. cf. labyrinthica other folds short and low, the sixth basopalatal weak in association with S. sparsicostata from Rexroad or absent. localities 2 and 3. University of Michigan material Measurements. Five randomly chosen topotypes from locality 3 justifies recognition of only 8. sparsi measure as follows, the mean being followed by the costata, and for this reason both references to 8. cf. range in parentheses. Measurements are in millime labyrinthica are assigned to 8. sparsicostata. ters. Length, 2.26 (2.1-2.5) ; width, 2.70 (2.6-2.8) ; Leonard (1950, 1952) has reported 8. sparsicostata number of whorls, 5.5 (514-5%). from middle Pleistocene deposits, but I have not Variation. The chief observed variables are axial examined material from these higher levels. ribbing and basopalatal folds. Most of the major Occurrence: Buis Ranch, Red Corral, Rexroad, Sand Draw series of riblets extend from suture to periphery, at and Dixon local faunas. least. Some are shorter, however, and reach only Discussion of material: Rexroad loc. 1, UMMZ 183058 (1). halfway to the periphery from suture, or vice versa. The single, juvenile (3% whorls) specimen shows coarse The minor series is variably developed, from none to ribbing, and may be referred to S. sparsicostata. Rexroad loc. 4b, UMMZ 181056 (1). The single specimen two small riblets being present between the larger (adult) was not dissected, but is similar to S. sparsicostata ones. The columellar lamella varies in length and topotypes in external features. size. Both first and second basal folds vary greatly Buis Ranch, UMMZ 181116 (3). The material consists of in size and shape. The first is usually subhorizontal, 3 adult body whorls only, 2 slightly broken. The axial riblets but sometimes nearly erect; almost always shorter are as strong as those in S. sparsicostata topotypes, and stronger than those of 8. labyrinthica. They are spaced more than the second basal fold, never longer; and trian widely than those of $. labyrinthica and more closely than gular to linguiform. The second basal fold varies those of ASf. sparsicostata topotypes. The one specimen on chiefly in slope of its ends; these may slope equally, which the adult diameter can be measured is 2.7 mm in di or either one may be steeper. The upper side of the ameter, larger than $. labyrinthica but within the size range fold may be parallel to the base for much of its length, of S. sparsicostata topotypes. Only one specimen retains the parietal lamella; it enters three-fourths of a whorl, as in or it may slope toward the ends from a subcentral S. sparsicostata. This material is referred to $. sparsicostata. point. The sixth basopalatal fold may be present or Red Corral, UMMZ 182999 (1). The single specimen (adult) absent. Two specimens have supernumerary baso was not dissected, but is similar to 8. sparsicostata topotypes palatal folds between the first and second (recognized in external features. Sand Draw loc. 2, UMMZ 184178 (11). A series of speci by their characteristic shape and position) of a more mens has ribs weaker than in S. sparsicostata topotypes but usual series. One specimen has a small, tubercular fold stronger than those of 8. labyrinthica. These specimens approximate to the second basal and fused to the agree with 8. sparsicostata and differ further from $. labyrin peripheral end of the first basal fold. Other basopa thica in size (diameter 2.7-2.8 mm) penetration of the parietal lamellae for a distance of three-fourths whorl, and smooth latal folds in this specimen are typical. The other ness of base. They are therefore referred to S. sparsicostata specimen has 4 supernumerary basopalatal folds, 3 be for the present. SYSTEMATIC DESCRIPTIONS 67
Sand Draw loc. 3, UMMZ 181203 (2). Two specimens (one Types. Holotype: UMMZ 183033a, Rexroad loc. 3. Hypo- adult, one subadult) agree in external features with those digm: Material from Saw Rock Canyon, Red Corral, and Rex from Sand Draw loc. 2. The inner ends of the parietal road local faunas and from Bender local fauna localities lb lamellae of the adult are broken away. and 2 in University of Michigan collections. Dixon loc. 1, UMMZ 182213 (3). The single adult specimen Diagnosis. A Gastrocopta of the G. cristata group was not dissected. It agrees with 8. sparsicostata topotypes and differs from 8. labyrinthica in the following respects: with a strong labial callus, an anguloparietal lamella large size (diameter 2.9 mm), smooth base, and widely spaced either bifid or simple; some individuals with a tubercle riblets. The strength of the riblets is, however, intermediate on the outer lip. between that of &. sparsicostata and 8. labyrinthica. Speci Distribution. Upper Pliocene of the southern High mens from the Dixon local fauna are referred to 8. sparsi Plains. costata for the present. Remarks. Late Pliocene samples of the Gastro Family PUPILLIDAE copta cristata group are unusual in that several popu lations include the diagnostic characters of distinct, Subfamily GASTROCOPTINAE Kecent species, as well as characters not found in Genus GASTROCOPTA Wollaston, 1878 these living species. For reasons set forth below, the late Pliocene populations are considered representative Subgenus GASTROCOPTA s. s. of an extinct, probably ancestral species. Gastrocopta (s. s.) procera (Gould), 1840 Gastrocopta cristata and G. procera are two wide- ranging Eastern North American land snails of simi Plate 1, figure 27 lar habitat. Each occurs separately in much of its Gastrooopta procera, (Gould). Pilsbry, 1948, Acad. Nat. Sci. range, but the 2 are also found together in a large Philadelphia Mon. 3, v. 2, pt. 2, p. 907, fig. 492:1-5. area (see under the 2 species for details of distribu Gastrocopta procera (Gould). Franzen and Leonard, 1947, tion). They are now distinct species, although closely Kans. Univ. Sci. Bull., v. 31, p. 341 [in part], pi. 18, related; and their association in the Nebraskan age fig. 3 [not fig. 6]. Dixon local fauna indicates their separate status Distribution. Maryland to South Carolina and throughout nearly the entire Pleistocene epoch. Alabama; west to South Dakota and Arizona (Pilsbry, Distinction of these two species is based chiefly on 1948, p. 907, 910). the anguloparietal lamella (straight or slightly sinuous Ecology. "This pupillid lives on timbered slopes in G. cristata, bifid in G. procera) and on the labial of streams. Its general distribution in Kansas indi callus (weak or absent in G. cristata, strong in G. cates an ability to withstand periods of summer procera). Other, less diagnostic features are the drought" (Franzen and Leonard, 1947, p. 342). stronger, more proximal crest and greater diameter Remarks. For discussion of the species and of this of G. cristata shells. Late Pliocene samples, however material see under G. fransenae. (especially those from the Kexroad local fauna), are Occurrence and material: Dixon local fauna. Dixon loc. 1, markedly more variable than Kecent series. The Plio UMMZ 182217 (1) ; loc. 2, UMMZ 191520 (11). cene samples include shells like those of G. cristata and G. procera, specimens intermediate in the degree Gastrocopta (s. s.) franzenae 4 Taylor, n. ,sp. of fusion of the anguloparietal lamella (not seen in Plate 1, figure 29 Recent or Pleistocene material), shells like G. cristata
Gastrocopta cristata Pilsbry and Vanatta. Hibbard, 1941, but with a strong labial callus as in G. procera, and Kansas Univ. Sci. Bull., v. 27, p. 95 [not of Pilsbry and shells like G. procera but with a strong tubercle on Vanatta]. the outer lip next to the sinulus. The frequency dis Hibbard, 1941, Kansas Acad. Sci. Trans. v. 44, p. 265 tributions of these character combinations suggest [not of Pilsbry and Vanatta]. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. that only one population is represented in the Kex 408 [not of Pilsbry and Vanatta]. road local fauna and at Bender local fauna localities Gastrocopta cristata (Pilsbry and Vanatta). Franzen and lb and 2. The sample from Saw Kock Canyon is inade Leonard, 1947, Kansas Univ. Sci. Bull., v. 31, p. 344 quate for analysis. [in part: Rexroad fauna material. Not of Pilsbry and Vanatta]. In figure 2, Pliocene specimens of the Gastrocopta Gastrocopta procera (Gould). Franzen and Leonard, 1947, cristata group are distributed according to arbitrarily Kansas Univ. Sci. Bull., v. 31, p. 341 [in part: Saw selected segments of the range of variation of two Rock Canyon fauna material], pi. 18, fig. 6 [not of characters: development of the labial tubercle (absent, Gould]. weak, or strong), and degree of fusion of the angulo * Named for Dr. Dorothea S. Franzen. parietal lamella (straight or slightly sinuous, strongly 68 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
= 12 N = 22 = 396 a bifid anguloparietal lamella and a strong labial 25 8 tubercle. This combination of characters is markedly (3) (1) infrequent in other samples. In this connection it should be noted that apparently all specimens from this locality available to Franzen and Leonard (1947, 67 (8) p. 341) are of this extreme type. Saw Rock Canyon The Red Corral sample is also inadequate. All N = 484 shells are similar to G. cristata, except that some have a strong labial callus. The material is believed representative of a population similar to those of the Rexroad local fauna, however; observed differences may be accounted for by geographic separation of the sample sites, possible temporal difference between the samples, small size of the Red Corral sample, and large percentage of similar shells in the Rexroad locality 4b sample. Bender local fauna samples are considered signifi cantly different among themselves. Localities la, Ib, and Ic are about one-eighth of a mile apart and at the same stratigraphic horizon. There is presumably some difference in mean age of the three samples, but it is
Bender Ib Bender 2 considered inadequate to account for the striking dif ferences observed. Material from localities la and Ic is referred to one population (G. cf. cristata}, similar Absent to G. cristata but with frequent development of a Labial strong labial callus. This strong callus occurs in a Weak smaller percentae of specimens than it does in the Tubercle Rexroad local fauna. Locality Ib material is believed Strong to represent another population, referred to G. fran Anguloparietal zenae. Bender locality 2 material is referred to G. FIGURE 2. Frequency distribution of Pliocene Gastrocopta cristata franzenae also. group. Early Pleistocene samples (Sand Draw, Dixon, San sinuous, or bifid). The groups with straight and ders local faunas) of the G. cristata group are all bifid anguloparietal lamella were established to cor typical of the Recent G. cristata and G. procera. Large respond with the range of variation in Recent G, series of G. cristata from the Sanders local fauna cristata and G. procera, respectively. have a straight or slightly sinuous anguloparietal The Rexroad local fauna has yielded ample series lamella and no strong labial callus; the same is true from three localities. Each series is considered repre of a small series from Dixon and two specimens from sentative of but one population. The slight bimodality Sand Draw. One specimen of G. procera from Dixon in the locality 4b sample is ascribed to error and (or) has a bifid anguloparietal lamella, thick labial callus, intraspecific variation. The general similarity be and no labial tubercle. tween the three populations is such that reference to Gastrocopta franzenae is a satisfactory structural one species is justified. Contemporaneity of these ancestor for both G. cristata and G. procera, for its three is established to the degree of refinement possible range of variation includes specimens similar to both by mammalian and stratigraphic correlation; but of these Recent species. Further support for a within such limits there is surely enough time that franzenae-cristata and procera lineage is the presence minor genetic changes may account for some or all at Bender localities la and Ic of a population inter of the observed differences. The holotype of G. franzenae is selected from the mediate between G. franzenae and G. cristata, but ap specimens intermediate in development of labial callus parently distinct from associated G. franzenae. Speci and in degree of fusion of the anguloparietal lamella. mens from Bender localities la, Ib, and Ic may there The Saw Rock Canyon sample, although inadequate, fore have lived shortly after the derivation of G. cri is of interest because two-thirds of the specimens have stata from G. franzenae. SYSTEMATIC DESCRIPTIONS 69 The evidence is inconclusive, but does suggest that The record by Taylor (1954a, p. 12) of this species G. franzenae gave rise to both G. procera and G. cri- in the Clarendonian Laverne local fauna is considered stata during the late Pliocene, that it became extinct erroneous. The single specimen (UMMZ 181273) shortly thereafter (perhaps with the climatic changes shows no trace of coloring on the peristome, as do all associated with the onset of continental glaciation), the other fossil Gastrocopta from the same locality. and that the two descendants have lived to the present It is almost surely a Recent specimen. The only species day. The time, place, and method of origin of the of Gastrocopta (s. s.) known in the Laverne local two younger species are uncertain. The Bender sam fauna are G. riograndensis (Pilsbry and Vanatta) ple suggests that G. franzenae had already given rise (Franzen and Leonard, 1947, p. 349, and University to a species with which it did not interbreed, and of Michigan collections) and G. lavernensis Taylor which has subsequently changed into G. cristata. G. (1954a, p. 11). procera, if already distinct, is not known from this For discussion of fossil material see under G. fran interval. zenae. One possible method of origin of the two descen Occurrence and material: Bender, Sand Draw, Sanders, and dants is by selection of their particular gene combina Dixon local faunas; Recent in northern Nebraska. Sand tions from the G. franzenae "pool". There seems no Draw loc. 5, CNHM PE3437 (1) ; loc. 6, UMMZ 177435 (1) ; need to postulate mutation, although this may have Dixon loc. 1, UMMZ 182216 (36) ; loc. 2, UMMZ 191519 (2) ; occurred. Assuming that the labial tubercle is genet Sanders loc. 1, UMMZ 182153 (400) ; loc. 2, UMMZ 182167 (125) ; loc. 3, UMMZ 182183 (600). G. cf. cristata: Bender ically produced, its genes were left behind in selecting loc. 1, UMMZ 177506 (111) ; loc. la, UMMZ 184122 (1750) ; out G. cristata and G. procera. The evolution of the loc. lc. UMMZ 183015 (2000). two species may possibly have been caused by climatic changes which "filtered" them out of G. franzenae. Gastrocopta (s. s.) paracristata Franzen and Leonard, 1947, emend. Occurrence and material: Saw Rock Canyon, Red Corral, Rexroad, and Bender local faunas. Saw Rock Canyon, UMMZ Plate 1, figures 31, 32 177516 (1), 177563 (9), 183654 (2); Red Corral, UMMZ 183- 001 (24) ; Rexroad loc. 2, UMMZ 183068 (575) ; loc. 3, UMMZ Gastrocopta paracristata Franzen and Leonard, 1947, Kansas 183033 (825); loc. 4b, UMMZ 177601 (371), 181052 (509); Univ. Sci. Bull., v. 31, p. 346, pi. 19, fig. 2, 3 [in part: Bender loc. Ib, UMMZ 184136 (31) ; loc. 2, UMMZ 184153 dextral shells only]. (175). Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. 151. Gastrocopta (s. s.) cristata (Pilsbry and Vanatta), 1900 Types. Holotype UKMNH 3929, Rexroad loc. 4b. Paratypes UKMNH. Plate 1, figures 24-26 Diagnosis. A Gastrocopta of the G. cristata group Gastrocopta cristata Pilsbry and Vanatta. Baker, 1938, with massive folds and lamellae, three equally im Nautilus v. 51, p. 130. mersed platal folds adjacent to the palatal callus, Gastrocopta, cristata. McGrew, 1944, Field Mus. Nat. Hist. and no subcolumellar lamella. Geol. Ser., v. 9, p. 36. Gastrocopta cristata (Pilsbry and Vanatta). Franzen and Description of holotype. "Shell elongate-oval, sum Leonard, 1947, Kansas Univ. Sci. Bull., v. 31, p. 344, mit obtuse; rimate, minutely perforate; suture sharply pi. 19, fig. 1; pi. 20, fig. 1. and deeply incised; whorls 5%; convex, regularly and Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, slowly increasing in size; the l1/^ nuclear whorls finely pt. 2, p. 911, fig. 493 :6, 8-12. granular; remaining whorls finely and irregularly Distribution. Northern Nebraska to southern striate; height of body whorl less than one-half of the Texas, west to Arizona. total height of the shell, constricted at the base and Ecology. "Gastrocopta cristata has, in Kansas, expanding rapidly toward the aperture; prominent, usually been found associated with G. procera (Gould) rounded crest paralleling the peristome from which on timbered slopes near streams" (Franzen and Leon it is somewhat removed; aperture rounded, slightly ard, 1947, p. 345). In northern Brown County, Nebr., oblique, expanding toward the peristome; denticles 5: it was found in a wooded ravine, among damp leaves a fused, sinuous, anguloparietal, the angular portion and dead wood. continuous with a parietal callus and outer lip; 3 Remarks. The Recent specimens from Nebraska equally entering palatal folds, connected anteriorly differ from other material examined in being paler by a low callus, a tubercular upper palatal, a higher, and in lacking a subcolimiellar tubercle. These differ elongate lower palatal, its highest point about midway ences probably represent normal variation, rather than of its length, and a low, tubercular basal fold; the specific characters. columella lamella high, elongate, entering horizontally
508945 O 6 70 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
and ascending the spire at its extreme innermost por columellar lamella, and relatively steeply inclined su tion; peristome reflected, thickened along its inner ture. border, margin sharp" (Franzeii and Leonard, 1947, Description. The shell is so similar to G. para p. 346). cristata that it can be described best by comparison. Measurements. Franzen and Leonard gave dimen It differs from G. paracristata by its sinistral coiling, sions of five specimens from the type series. The more steeply inclined suture, and gradually ascending mean of these (in millimeters) is followed by the columellar lamella. range in parentheses. Length, 3.0 (2.7-3.4) ; width, Measurements. The mean of the 11 whole speci 1.4 (1.3-1.5) ; length of aperture, 1.1 (0) ; width of mens in the type series is followed by the range in aperture, 1.0 (0.9-1.1) ; number of whorls, 5.9 (5V4~ parentheses. Measurements are in millimeters. Length, 2.82 (2.6-2.9) ; length of aperture, 0.97 (0.9- _ Variation. "The range in height of the shells of 1.0); width, 1.28 (1.2-1.3); width of aperture, 0.88 Gastrocopta paracristata is from 2.7 mm to 3.4 mm. (0.8-0.9); number of whorls, 5.7 (51/4-6). The width varies independently of the height, there Variation. G. scaevoscala seems relatively invari fore, shells vary in shape from elongate-ovate to able, but this may be an artifact of small sample size. elongate-conic. The basal fold, usually prominent, is Observed variation is like that in G. paracristata: in sometimes very low and occasionally wanting. The size (see measurements), prominence of basal fold callus of the peristome varies in prominence. A promi and crest, and in degree of fusion of the anguloparietal nent, rounded crest on the ultimate whorl, paralleling lamella. the peristome is usually present, rarely wanting, some Distribution. Latest Pliocene and early Pleistocene times low and sharp. The distance between the peri of southern Kansas. stome and the center of the crest varies from 0.5 to Remarks. The record of sinistral Gastrocopta para 0.25 mm/1 (Franzen and Leonard, 1947, p. 347). cristata in the Kexroad local fauna (Franzen and Distribution. Upper Pliocene and lower Pleistocene Leonard, 1947, p. 347) is almost certainly based on of the High Plains. G. scaevoscala from the Bender local fauna. Univer Remarks. Sinistral specimens from the SWl/4 sec. sity of Michigan collections from both assemblages 22, T. 33 S.; K. 29 W., Meade County, Kans., were contain the sinistral G. scaevoscala from the Bender included in G. paracristata by Franzen and Leonard local fauna only. University of Kansas material in their original description of the species. These reported from the SWl/4 sec. 22, T. 33 S., K. 29 W., sinistral specimens are described as G. scaevoscala Rexroad Ranch, Meade County, is largely from Taylor, n. sp., in this report. Bender localities la, Ib, and le, according to C. W. Occurrence and material: Saw Rock Canyon, Red Corral, Hibbard (written communication Nov. 29, 1954), Rexroad, Bender, and Sanders local faunas. Saw Rock who helped collect the fossils. Canyon, UMMZ 173590 (1), 177562 (35), USGS 19089-5 (1); G. scaevoscala is apparently the only sinistral species Red Corral, UMMZ 183003 (1) ; Rexroad loc. 1, UMMZ 183060 of Gastrocopta (s. s.), although the subgenera Immer- (1) ; loc. 2, UMMZ 183071 (8) ; loc. 3, UMMZ 183036 (10) ; sidens and Sinalbinula have such forms. The simi loc. 4b, UMMZ 177602 (35), 181051 (50) ; Bender loc. 1, UMMZ 177507 (11-4) loc. la, UMMZ 18-412-4 (600): loc. Ib. UMMZ larity to G. paracristata suggests it is a species derived 184138 (300) loc. le. UMMZ 18-41-45 (150) ; loc. 2, UMMZ through mutation in direction of coil, and so repro- 18-4154 (200) Sanders loc. 1, UMMZ 182154 (27) ; loc. 2, ductively isolated. The differences indicated in the UMMZ 182168 (1). description are minor, but constant, so that it is diffi cult to regard the specimens from the Bender local Gastrocopta (s. s.) scaevoscala Taylor, n. sp.B fauna simply as sinistral individuals of G. paracri Plate 1, figures 33, 34 stata. Furthermore, G. scaevoscala occurs in the Dixon local fauna unaccompanied by G. paracristata. Gastrocopta paracristata Franzen and Leonard, 1947, Kansas Univ. Sci. Bull., y. 31, p. 346-347 [in part: sinistral Occurrence and material: Bender and Dixon local faunas. specimens]. Bender loc. 1, UMMZ 184320 (1) ; loc. la, UMMZ 184125 (9, Types. Holotype UMMZ 1S4320, Bender loc. 1. Paratypes only 4 whole) ; loc. le, UMMZ 183017 (13, only 7 whole) ; UMMZ 184125 (9), 183017 (13). Dixon loc. 2, UMMZ 191522 (6). Diagnosis. A sinistral Gastrocopta of the G. Gastrocopta (s. s.) chauliodonta Taylor, 1954 cristata group with massive folds and lamellae, three Plate 1, figures 11-16 equally immersed palatal folds adjacent to the palatal (rastroeopta ehauliodonta Taylor, 1954, Michigan Univ. Mus. callus, no subcolumellar lamella, slightly ascending Zool. Occ. Papers 557, p. 12, 15. Types. Holotype LTMMZ 181120, Sand Draw loc. 6. Para- 5 L. scaei'tis, on or toward the left, scala^ flight of stairs. types UMMZ 181121 (11). SYSTEMATIC DESCRIPTIONS 71 Diagnosis. A species most closely related to the in which case the inner half turns downward abruptly, lower Pliocene Gastrocopta lavernensis Taylor and to or it may descend gradually. the Recent G. curacoana Pilsbry. It is characterized by Distribution: Lower Pleistocene of the High Plains. the massive anguloparietal lamella which almost hides Remarks. Gastrocopta chauUodonta appears most the deeply immersed palatal folds, the descending col- nearly related to G. lavernensis Taylor, 1954, from the umellar lamella, the absence of a subcolumellar la Pliocene of Oklahoma, and G. curacoana Pilsbry, 1924, mella, and the relatively broad, cylindrical form. living in the Dutch West Indies. It differs from the Description of types. Shell small, cylindric, rimate; former by its more massive lamellae and peristome apex obtuse. About 5^ whorls, convex, slowly and callus, simpler anguloparietal lamella, more deeply regularly enlarging; sutures strongly impressed; body immersed basal fold, and lamelliform upper palatal whorl less than half of total shell length, about one- fold. It differs from G. curacoana (see Pilsbry, 1922- half whorl behind aperture becoming flattened and 26, p. 202-204) by its less tapering shape and more then slightly concave in the middle of the outer conspicuous anguloparietal lamella. side, convex again at a prominent, rounded crest just The long, deeply immersed low^er palatal fold and behind the reflected peristome. Sculpture of nuclear the descending columellar lamella of Gastrocopta whorl minutely granular; postnuclear sculpture of chauUodonta, G. lavernensis, and G. curacoana might fine, irregular, oblique growth lines. Aperture be considered evidence indicating close relationship slightly oblique, rounded, with dentition of five folds with Immersidens, which sometimes has these char and lamellae, as follows: a prominent, massive, sinu acters. These species agree with Gastrocopta s. s. in ous anguloparietal lamella, gradually increasing in the differentiating characters of apertural callus and height and terminating abruptly, the angular part simpler anguloparietal lamellae, however. G. octo- continuous with the parietal lip; a deeply immersed, naria Pilsbry (1922-26, p. 204-205) is transitional small, lamelliform upper palatal fold of about equal from this species group to others of Gastrocopta s. s. length and height; a still more deeply immersed by having a subcolumellar lamella partially united (dorsal in position) long, angulate, deeply entering, with the columellar. This fact, and the greater devel lower palatal fold, of about the same height as the opment of the columellar lamella in Immersidens, sug upper palatal fold, increasing in height abruptly and gests that the descending columellar lamella of the decreasing gradually, the outer third ascending G. curacoana group may represent a fusion of colum slightly, the inner two-thirds approximately parallel ellar and subcolumellar lamellae, whereas in Immer- to the base and roof of the body whorl; a tubercular sidens the descending columellar lamella originated as basal fold immersed equally with, or slightly more a unit. than, the lower palatal fold; all palatal folds situated Occurrence and material: Sand Draw, Dixon. and Sanders well behind the palatal callus; columellar lamella local faunas. Sand Draw loc. 2, USGS 19090-5 (23) ; loc. 3, UMMZ 181204 (1): loc. 6, UMMZ 181120 (1), 181121 (11), heavy, elongate, descending the columella to its base. 181218 (1) ; loc. 7, UMMZ 177236 (2%) ; Dixon loc. 1, UMMZ Inner and outer lips heavily callused except in the 182215 (13): Sanders loc.'1, UMMZ 182152 (30). conspicuous sinulus. Measurements. The mean of the 11 whole speci Gastrocopta (s. s.) pellucida hordeacella (Pilsbry), 1890 mens in the type series is followed by the range in Plate 1, figure 28 parentheses. Measurements are given in millimeters. Height, 2.51 (2.2-2.7) ; width, 1.22 (1.1-1.3) ; height Gastrocopta pellucida liordeacella (Pilsbry)- Franzen and Leonard, 1947, Kansas Univ. Sci. Bull., v. 31, p. 349, of aperture, 0,86 (0.7-0.9) ; width of aperture, 0.87 pi. 19, fig. 8. (0.8-0.9); number of whorls, 51/2 (5-6). Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, Variation. The anguloparietal lamella varies in pt. 2, p. 913, fig. 494 :a-d, 495. shape considerably. Usually the angular part is rep Pilsbry, 1953, Acad. Nat. Sci. Philadelphia Proc., v. 105, resented by a slightly prominent ridge which gives p. 162. the lamella a sinuous appearance; sometimes this ridge Distribution. Southern and Baja California (lo is curved toward the outer lip and the lamella is cally) through southeast Colorado and Kansas to bifid, the sinulus almost completely enclosed; occa Florida; south to Sinaloa and Tainpico, Mexico; north sionally the whole lamella is nearly straight. The on the Atlantic Coastal Plain to southern New Jersey anguloparietal varies in width also, from about one- (Franzen and Leonard, 1947; Pilsbry, 1948, 1953). fourth to one-sixth of the internal width of the aper "The northward extension along the Atlantic Coast ture. The columellar lamella may enter horizontally, is very narrow, discontinuous so far as known, and 72 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS perhaps mainly confined to the coastal islands" (Pils in large numbers, it is probably overlooked because of bry, 1948, p. 914). its small size * * * In order to remain established Ecology. Uncertain. From the geographic distri in northwestern Kansas, as it is doing to some extent, bution, frequent occurrence in stream drift in the it must be tolerant of the hot, dry climatic conditions southern Great Plains, and occasional occurrence in which obtain in the summertime. G. holzingeri is samples of fossils washed from bulk collections of not one of the predominating species of the fauna of matrix it appears this species may live among the Kansas. According to its general distribution in grass roots, occasionally even on exposed slopes. So North America, it prefers decidedly cooler climates" far as known, the snail has not been collected alive in (Franzen and Leonard, 1947, p. 334-335). In northern the Plains. Nebraska it was found under logs, bark, and stones Occurrence and material: Red Corral, Rexroad, and Dixon in a moist tall-grass area around seepages. local faunas. Red Corral, UMMZ 183002 (4) ; Rexroad loc. Occurrence and material: Rexroad local fauna; Saw Rock 3, UMMZ 183037 (3) ; Dixon loc. 2, UMMZ 191521 (3). Canyon local fauna (Franzen and Leonard) ; Recent in north ern Nebraska. Rexroad loc. 1, UMMZ 183059 (1) ; loc. 2, Submenus ALBINULA Sterki, 1892 UMMZ 183070 (16); loc. 3, UMMZ 183035 (150). Gastrocopta (Albinula) armifera (Say), 1821 Subgenus VERTIGOPSIS Sterki, 1893 Gastrocopta armifera (Say). Franzen and Leonard, 1947, Kansas Univ. Sci. Bull., v. 31. p. 328, pi. 17, fig. 3-5. Gastrocopta (Vertigopsis) tappaniana (Adams), 1842 Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, Gastrocopta tappaniana (C. B. Adams). Baker, 1938, Nau pt. 2, p. 874, fig. 472 :l-4. tilus, v. 51, p. 131. Distribution. Quebec to northern Florida, west to Franzen and Leonard, 1947, Kansas Univ. Sci. Bull., v. the Eocky Mountains (Pilsbry, 1948, p. 875). 31, p. 336, pi. 18, fig. 8. Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, Ecology. "Gastrocopta armifera is a gregarious pt. 2, p. 889, fig. 477 :9. species occurring commonly on wooded slopes, near or Frye and Leonard. 1952, Kansas Geol. Survey Bull. 99, p. removed from a stream. It is to be found under dead 151, pi. 14, fig. k. wood, limestone rocks, or light cover of leaf mold or Gastrocopta tappaniana C. B. Adams. Hibbard, 1941, Kansas Univ. Sci. Bull., v. 27, p. 95. other debris. O. armifera frequently occurs under Hibbard. 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. boards or rocks in gardens" (Franzen and Leonard, Frye and Hibbard. 1941, Kansas Geol. Survey Bull. 38, p. 1947, p. 329). In northern Nebraska it was found 408. under logs, barks, and stones in moist grass around Gastrocopta tappaniana. McGrew, 1944, Field Mus. Nat. Hist. Geol. Ser., v. 9, p. 36. seepages. Distribution. "Ontario and Maine to Virginia and Occurrence and material: Dixon local fauna; Recent in northern Nebraska. Dixon loc. 1, UMMZ 182214 (10). Alabama, west to South Dakota and Kansas, south west to Arizona, but not known from the southeastern Gastrocopta (Albinula) holzingeri (Sterki), 1889 Atlantic states, Virginia to Florida" (Pilsbry, 1948,
Gastrocopta liolzingeri (Sterki). Hibbard, 1941, Kansas Univ. p. 889). Sci. Bull., v. 27, p. 95. Ecology. "Its most frequent habitat is on shaded Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. slopes near streams. However, it has been taken Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. from among the grass roots on an unshaded slope near 408. a pasture pond" (Franzen and Leonard, 1947, p. 337). Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, In northern Nebraska it was found among moist leaves pt. 2, p. 883, fig. 474:4-6, 475. Frye and Leonard, 1952. Kansas Geol. Survey Bull. 99, or under logs in moist grass near seepages. p. 151. Occurrence and material: Saw Rock Canyon, Rexroad, Ben Gastrocopta holsinoeri Sterki. Franzen and Leonard, 1947, der, Sand Draw. Dixon, and Sanders local faunas; Recent in Kansas Univ. Sci. Bull., v. 31, p. 333, pi. 18, fig. 2. northern Nebraska. Saw Rock Canyon, UMMZ 177517 (1), 177564 (23); Rexroad loc. 2, UMMZ 183072 (20); loc. 3, Distribution. "Ontario and western New York to UMMZ 183038 (27) ; loc. 4b, UMMZ 177604 (3), 181053 (1) ; Helena, Montana, south to Illinois, Kansas, and Albu Bender loc. 1, UMMZ 177508 (1) ; loc. la, UMMZ 184126 (1) ; querque and Mesilla, New Mexico" (Pilsbry, 1948, loc. Ib, UMMZ 184139 (39) ; Sand Draw loc. 3, UMMZ 181205 p. 883). (2); loc. 5, UMMZ 181251 (1); loc. 6, CNHM PE3445 (1), UMMZ 177436 (15), 181219 (9) ; Dixon loc. 1, UMMZ 182218 Ecology. ^Gastrocopta liolzingeri is found on tim (75) ; loc 2, UMMZ 191523 (32) : Sanders loc. 1, UMMZ 182155 bered slopes under leaf mold. Although it may occur (150) ; loc. 2, UMMZ 184164 (5) ; loc. 3, 182184 (17). SYSTEMATIC DESCRIPTIONS 73
Subgenus IMMERSIDENS Pilsbry and Vanatta, 1900 gobasodon (Pilsbry and Ferriss), and G. prototypus ( Pilsbry). It differs from G. cochisensis by the separa Gastrocopta (Immersidens) rexroadensis Franzen and Leonard, 1947 tion of the angular and parietal lamellae, the shorter and straight parietal lamella, absence of a basal fold, Plate 1, figure 30 and the fusion of the upper palatal fold with the pala Gastrocopta rexroadensis Franzen and Leonard, 1947, Kansas tal callus. It differs from G. oligobasodon by the Univ. Sci. Bull., v. 31, p. 338, pi. 18, fig. 4, 5. separation of the angular and parietal lamellae, obli Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. queness of the parietal lamella, larger columellar 151, pi. 14, fig. e. lamella, absence of a basal fold, and the fusion of the Types. Holotype UKMNH 3781, Rexroad loc. 3. Paratypes upper palatal fold with the palatal callus. It differs UKMNH. from G. prototypus by the separation of the angular Diagnosis. An Immersidens with a short, straight, and parietal lamellae, straight and oblique parietal oblique parietal lamella placed behind a much smaller lamella, larger and descending columellar lamella with angular lamella not fused to it, an oblique, descending no callus above, more deeply immersed, larger, and columellar lamella, no basal fold, and a deeply im oblique lower palatal fold, and larger upper palatal mersed lower palatal fold. fold fused with the palatal callus. Description of holotype. "Shell small, elongate- Franzen and Leonard (1947, p. 339) considered this conic; summit obtuse; rimate; suture sharply incised; species closely related to G. bilamellata (Sterki and 4% whorls; apical whorls strongly convex, lower Clapp). It differs from the latter in being shorter and whorls slightly convex; whorls slowly and regularly relatively broader in shell form; by the shorter and increasing in size; the l 1/^-nuclear whorls finely simpler angular and parietal lamellae, separate and granular; remaining whorls irregularly and finely straight, the parietal oblique; the absence of a striate; height of body whorl exceeding one-half of the basal fold; the oblique lower palatal; and the less total height, contracted basally and expanding rapidly deeply immersed upper palatal, fused to the palatal toward the aperture; a prominent, rounded crest callus. The differences between G. rexroadensis and G. paralleling the peristome from which it is separated by bilamellata seem greater than those between G. rex a deep, wide groove; aperture triangularly rounded, roadensis and G. cochisensis, G. oligobasodon, and G. scarcely oblique, widely expanding; denticles 5; a prototypus. prominent, oblique, arched, deeply immersed parietal Occurrence and material: Saw Rock Canyon local fauna lamella extending toward the periphery below the (Frye and Leonard) ; Red Corral local fauna; Rexroad local angular lamella; an elongate, prominent angular fauna (Franzen and Leonard). Red Corral, UMMZ 183004 lamella arising from the peristome, approximate to but (2). not fused posteriorly with the parietal lamella; the up New genus and species? per palatal fold immersed, fused with a triangular Description. Shell conical, perforate, apex obtuse. callus on the outer border of the peristome; a promi Whorls five, convex, separated by well impressed nent, shortly elongate, deeply immersed, oblique lower suture. Aperture subtrigonal, basally regularly palatal fold; a prominent columellar lamella, entering rounded, inconspicuously dentate. Folds and lamellae horizontally with the posterior half turned downward; five, weak: anguloparietal low, elongate, sinuous; margin of peristome very narrowly reflected; ends of columellar elongate-tubercular, entering horizontally; peristome approaching, connected by a thin callus basal and lower palatal both low, tubercular, and im upon the parietal wall" (Franzen and Leonard, 1947, mersed equally with the columellar; upper palatal p. 338). tubercular, slightly elongate parallel to suture, very Measurements. Franzen and Leonard gave dimen deeply immersed, dorsal in position; peristome re sions of five specimens. The mean of these (in milli flected, not thickened; a weak palatal callus meters) is followed by the range in parentheses. behind the reflection but in front of the folds. Length, 2.35 (2.25-2.4) ; width, 1.29 (1.25-1.35) ; length Umbilicus small, about one-seventh of the diameter, of aperture, 0.88 (0.8-0.9) ; width of aperture, 0.80 perforate to the apex, slightly covered by body whorl. (0.7-0.9) ; number of whorls, 5.0 (4%-5V4). First whorl minutely punctate; later whorls with many Distribution. Late Pliocene of the southern High nearly regular, low, rounded, but relatively coarse Plains. raised lines between which lie weak growth lines. Remarks. G. rexroadensis appears most closely re Length, 2.5 mm; width, 1.4; length aperture, 0.8; width lated to G-. cochisensis (Pilsbry and Ferriss), G. oli- aperture, 0.7; whorls 5. 74 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Remarks. Only one specimen of this unusual pu- Submenus ISCHNOPUPOIDES Pilsbry, 1926 pillid was found. The weak teeth and palatal callus Pupoides (Isehnopupoides) inornatus Vanatta, 1915 and unthickeiied peristome suggest that it may be not quite mature or else that it may be like Gastrocopta Plate 1, figure 35 (Privatula] or Microstele in having reduced dentition. Pupoides inornatus Vanatta. Hibbard, 1941, Kansas Univ. Sci. Of associated forms Gastroropta tappaniana is closest Bull., v. 27, p. 95. in having a conical shape, but dentition, umbilicus, and Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. sculpture are so different that it seems unlikely that, Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. the specimen is simply a monstrosity of G. tappaniana. 408. Franzen and Leonard, 1947, Kansas Univ. Sci. Bull., v. 31, Perhaps this individual represents an extinct derivative p. 374. of Gastrocopta (Vertigopsis) with an enlarged umbili Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mou. 3, v. 2, cus, paralleling Chae nay-is in that respect. More ma pt. 2, p. 926, fig. 499 :10. terial is needed for a decision. Distribution. Only two definitely Recent occur Occurrence and material: Sanders local fauna, loc. 3, UMMZ rences are known: Pikes Peak, El Paso County, Colo. 182187 (1). (Pilsbry, 1948, p. 926); Sidney, Cheyenne County, Nebr. (USNM 424028, 363054). Subfamily PUPILLINAE Ecology. Uncertain. Genus PUPOIDES Pfeiffer, 1854 Occurrence and material: Red Corral, Rexroad, and Sand Draw local faunas. Red Corral, UMMZ 183005 (1) ; Rexroad Submenus PTTPOIDES s. s. loc. 2, UMMZ 183074 (16) ; loc. 3, UMMZ 183040 (41) ; Sand Pupoides (s. s.) albilabris (Adams), 1841 Draw loc. 2, UMMZ 184180 (2).
Plate 1, figures 36. 37 Subfamily VERTIGININAE
Pupoides maryinatiis (Say). Hibbard, 1941, Kansas Univ. Genus VERTIGO Miiller, 1774 Sci. Bull., v. 27, p. 95. Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. Vertigo ovata Say, 1822 Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, Vertigo ovata Say. Baker, 1938, Nautilus, v. 51, p. 130. p. 408. Franzen and Leonard, 1947, Kansas Univ. Sci. Bull., v. Franzen and Leonard, 1947, Kansas Univ. Sci. Bull., v. 31, p. 354, pi. 19, fig. 6. 31, p. 370, pi. 21, fig. 3-4. Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, Pupoides albilabris (C. B. Adams). Pilsbry, 1948, Acad. Nat. pt. 2, p. 952, fig. 513 :l-3, 4, 7. Sci. Philadelphia Mou. 3. v. 2, pt. 2, p. 921, fig. 499:1-7. Vertiao ovata. McGrew, 1944, Field Mus. Nat. Hist. Geol. Ser., Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. 151, pi. 14, fig. b. v. 9, p. 36. Vertigo orata (Say). Hibbard, 1954, Michigan Acad. Sci. Taylor, 1954, Michigan Univ. Mus. Zool. Occ. Papers 557, Papers, v. 39, p. 343. p. 15. Vertigo hibbardi Baker. Frye and Leonard, 1952, Kansas Distribution. Maine to the West Indies, west to Geol. Survey Bull. 99, p. 151 [in part: Kingman County North Dakota, Arizona, and northern Mexico (Pilsbry, loc. Not of Baker]. 1948, p. 923). Distribution. Laborador to the West Indies, west to Ecology. "It lives under stones or at the roots of Arizona, Utah, Oregon, and Alaska (Pilsbry, 1948, p. grass, in well-drained but often sunny places; follow 953). ing rains it is sometimes found on trees a few feet Ecology. u ***on]y in moist environs afforded by from the ground" (Pilsbry, 1948). shaded slopes near streams and shores of ponds11 (Franzen and Leonard, 1947, p. 355). In northern Ne Occurrence and material: Saw Rock Canyon. Rexroad, Ben braska it was found on sticks and in wet leaves in der, Sand Draw. Dixou. and Sanders local faunas. Saw Rock seepages and immediately adjacent to a stream, asso Canyon, UMMZ 177565 (4) ; Rexroad loc. 1, UMMZ 183061 (14) ; loc. 2, UMMZ 183073 (75) ; loc. 3, UMMZ 183039 (100) ; ciated with Carychiiim, Fossa ria, and Pisidium. loc. 4a, UMMZ 177571 (22) ; loc. 4b, UMMZ 177605 (80), 181- Remarks. Frye and Leonard reported V. hibbardi, 054 (12) ; Bender loc. 1, UMMZ 177509 (23) ; loc. la, UMMZ but not V. ova fa. from the Dixon local fauna. Univer 184127 (350) : loc. Ib, UMMZ 184140 (75) ; loc. Ic, UMMZ sity of Michigan collections, however, contain about 183018 (200) ; loc. 2, UMMZ 184155 (11) : Sand Draw loc. 2, 200 specimens of V. ovata, but no V. hibbardi. Fur UMMZ 184179 (2); loc. 6, UMMZ 177437 (15), 181220 (1); Dixon loc. 1. UMMZ 182221 (21) ; loc. 2, UMMZ 191524 (16) ; thermore, V. ovata and V. hibbardi have never been Sanders loc. 1, UMMZ 182158 (500) ; loc. 2. UMMZ 182170 found together, a fact which suggests ecologic replace (53) ; loc. 3, UMMZ 182186 (150). ment since their geologic ranges overlap. In addition SYSTEMATIC DESCRIPTIONS 75 the two species are superficially similar, and careful Description. "Shell moderate in size for the genus; examination of the apertural lamellae is necessary for ovately conic; rimate; whorls typically 5, convex; their discrimination. The Dixon local fauna refer suture sharply incised; nuclear whorl finely granulose; ence to V. hibbardi, therefore, is probably based on remaining whorls glossy, and finely striate in well- V. ovata. (Compare Frye and others, 1943, p. 42, with preserved shells, regularly and rapidly increasing in Franzen and Leonard, 1947, p. 354). size; body whorl as much as three-fifths of total Occurrence and material: Buis Ranch, Sand Draw, Dixon, height; aperture biarcuate. expanding toward the peri- and Sanders local faunas; Recent in northern Nebraska. stome; peristome narrowly reflected, margins sharp, Buis Ranch, UMMZ 181115 (1); Sand Draw loc. 2, UMMZ terminations approaching and connected across pari 184181 (4) ; loc. 3, UMMZ 181131 (29) ; loc. 4, UMMZ 177326 etal wall by a thin callus; denticles 7: 2 elongate lamel (30); loc. 5, CNHM PE3438 (2), UIMNH P6791 (1), UMMZ lae on parietal wall, the parietal lamella the more 181140 (31), 181235 (100); loc. 6, UMMZ 177438 (8), 181222 (12) ; loc. 7, UMMZ 177235 (3) ; Dixon loc. 1, UMMZ 182219 deeply immersed, the innermost third deflected at a (200) ; loc. 2, UMMZ 191526 (4) ; Sanders loc. 1, UMMZ 182- right angle toward the outer lip; the angular about 157 (1). one-third as long as the parietal, less deeply immersed; suprapalatal fold, when present, tubercular; upper Vertigo hibbardi Baker, 1938 palatal fold lamelliform, high, long and sinuous; Plate 1, figures 18-22 lower palatal fold lamelliform, more deeply immersed than the upper palatal, increasing in height up to Vertigo hibbardi Baker, 1938, Nautilus, v. 51, p. 126. its midpoint and then declining, partially forked ante Smith, 1940, Kansas Geol. Survey Bull. 34, p. 98. Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, riorly ; infrapalatal low, elongate, as deeply immersed, p. 151 [in part: Meade County loc.]. and about half as long as the lower palatal; basal Leonard, 1957, Illinois Geol. Survey Report of Investiga lamelliform, elongate, increasing in height inwardly tions 201, p. 21, pi. 1, fig. 3. and terminating rather abruptly; the folds connected Vertigo hibbardi F. C. Baker. Hibbard, 1941, Kansas Univ. by a low callus; columellar lamella large, heavy, Sci. Bull., v. 27, p. 95. crescentic, ascending slightly toward the columella, Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. then turning downward and recurving slightly for 408. ward at the base; crest behind the peristome promi Franzen and Leonard, 1947, Kansas Univ. Sci. Bull., v. nent, rounded, separated from the peristome by a 31, p. 367, pi. 20, fig. 9, 10. groove; body whorl indented at level of the upper Types. Holotype UIMNH P6773, Rexroad loc. 3. Para- and lower palatals resulting in the contraction of types UIMNH P6773 (5), ANSP 169883 (not found). the body whorls at its base" (Franzen and Leonard, Diagnosis. A moderately large, ovate Vertigo with 1947, p. 367-368). seven folds and lamellae, the palatals deeply entering, Measurements. The mean of the series measured is the columellar descending and spoon shaped; the body followed by the range in parentheses. Measurements whorl flattened laterally and flaring in basal view. are in millimeters.
Number of Length Length aperture Width Width aperture whorls Rexroad loc. 3_____ 2. 00 0. 8 1. 3 0. 77 V/i (3 specimens)_ (1.9-2.0) (.8) (1.3) (. 7-. 8) (^A) Saw Rock Canyon. 2. 19 . 84 1. 41 . 84 4^ (7 specimens) _ (2.0-2.3) (. 7-. 9) (1.3-1.5) (. 8-. 9) (4^-5) Variation. The shape of the shell varies from Distribution: Late Pliocene of southwestern Kansas. ovate to conical, as shown by the measurements. The Remarks. For discussion of the reported occur folds and lamellae are relatively stable for pupillids. rence of V. hibbardi in the Dixon local fauna see The infrapalatal fold is rarely absent. The inner under V. ovata. end of the parietal lamella may be bent towards F. C. Baker and subsequent writers who have dealt either lip, or not at all. Shells from the Saw Rock with V. hibbardi have considered it as one of the Canyon local fauna are larger than topotypes (see subgeiius Angustula Sterki, 1888. The species has measurements), and their apertures are not so nearly most of the characters in the subgeneric definition closed by the dentition. Some topotypes have the given by Pilsbry (1948, p. 944), but other features base trumpetlike when viewed from below, as Baker seem to show that this resemblance is only convergent. noted, but others are less extreme. The Saw Rock Furthermore, V. milium (type of Angustula} appears Canyon specimens do not have this pronounced con to have close relationships with other species which striction in the body whorl. are not recognized by present taxonomy. 76 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS Vertigo hibbardi agrees with the diagnosis of Angu- Vertigo milium (Gould.), 1840 stula (based on apertural dentition) in all respects Vertigo milium Gould. Hibbard, 1941, Kansas Univ. Sci. Bull., save one: the inner end of the lower palatal fold does v. 27, p. 95. not curve downward. It differs radically from the Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. other species heretofore placed in Angustula by its Frye and Leonard, 1941, Kansas Geol. Survey Bull. 38, p. 408. larger size, conical or ovate-conic shape, and laterally Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. flattened body whorl. The difference in shape, not 151. necessarily significant in all gastropod genera, is Vertigo milium (Gould). Franzen and Leonard, 1947, Kansas Univ. Sci. Bull., v. 31, p. 365, pi. 20, fig. 8. important because the other species-groups in Vertigo Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, have unity in external form, the species differing pri pt. 2, p. 944, fig. 509. marily in lamellar characters. V. hibbardi is prob Taylor, 1954, Michigan Univ. Mus. Zool. Occ. Papers 557, ably most closely related to the V. ovata, group, al p. 15. though not nearly. It is similar in shape to species of Distribution. Maine to South Dakota, south to Ari this group, differing slightly by the laterally com zona, Mexico, and the West Indies (Pilsbry, 1948, p. pressed body whorl. The strongly developed folds 945). and lamellae are similar to those of V. ovata diaboli Ecology. "Moist situations afforded by timbered Pilsbry (Pilsbry, 1948, p. 953), from which it differs stream banks, or marshes. It is not found in areas of in the absence of an infraparietal lamella and in the low relative humidity or low annual precipitation and high summer tempertures * * *" (Franzen and Leon spoonshaped, descending columellar lamella. ard, 1947, p. 367). The criteria previously used for definition of Angu stula therefore seem artificial, because they associate Occurrence and material: Red Corral, Rexroad, Bender, Sand Draw, Dixon, and Sanders local faunas; Saw Rock Canyon otherwise dissimilar and apparently unrelated species. local fauna (Franzen and Leonard). Red Corral, UMMZ But it appears that these characters have also ex 183006 (27) ; Rexroad loc. 2, UMMZ 183076 (15) ; loc. 3, cluded a related species from the V. miliuin group: UMMZ 183042 (57) ; loc. 4b, UMMZ 177606 (1), 181055 (2) ; V. numellata. Pilsbry (1918-20, p. 149-150) compared Bender loc. 1, UMMZ 177510 (15) ; loc. la, UMMZ 184128 (75) ; loc. Ib, UMMZ 184141 f3) ; loc. Ic, UMMZ 183019 (75) ; V. bermudensis of the V. milium group with V. numel- Sand Draw loc. 2, UMMZ 184182 (2) ; loc. 3, UMMZ 181132 lata (then placed in its own species group) as follows: (7); loc. 4, UMMZ 177325 (2); loc. 5, UMMZ 181141 (4), "The much larger V. numellata has a similar crest, 181234 (18) ; loc. 6, UMMZ 177439 (1), 181221 (20) ; Dixon loc. 1, UMMZ 182220 (100) ; loc. 2, UMMZ 191525 (25) ; Sanders the lower palatal fold is long, also, and the angular loc. 1, UMMZ 182156 (500) ; loc. 2, UMMZ 182169 (5) ; loc. and parietal lamellae similar in position; yet it has not 3, UMMZ 182185 (500). the peculiarly shaped columellar lamella of Angu stula" By the same token as used with V. hibbardi. Family PUPILLIDAE? it seems that the strictly lamellar characters have re Remarks. A fragmentary specimen consisting of the sulted in an unnatural grouping of species. From the apical whorls only may represent a pupillid. It is similarities of both dentition and external form, the distinct from the other species listed from the local V. milium group includes V. numellata and V. bermu fauna. densis, but not V. hibbardi. It appears that a descend Occurrence: Red Corral local fauna. UMMZ 183000 (1). ing columellar lamella has developed more than once in Vertigo, Whether Angustula merits subgeneric Family VALLONIIDAE rank seems questionable, if its unique character of a Genus VALLONIA Risso, 1826 descending columellar lamella is no longer unique. It Vallonia pulchella (Miiller), 1774 seems best to synonymize it with Vertigo s. s., and to Valloma pulohella (Miiller). Pilsbry, 1948, Acad. Nat. Sci. consider V. milium and its relatives coordinate with Philadelphia Mon. 3, v. 2, pt. 2, p. 1023, fig. 545a. other species groups, such as those of V. ovata, and Taylor, 1954, Michigan Univ. Mus. Zool. Occ. Papers 557, V. modesta. p. 15.
Occurrence and material: Saw Rock Canyon and Rexroad Distribution. Holarctic. In North America east of local faunas. Saw Rock Canyon, UMMZ 173589 (10), 177566 the Rocky Mountains from Nova Scotia and Manitoba (400), USGS 39089-6 (125), Rexroad loc. 2, UMMZ 183075 south to Missouri and Kentucky (Pilsbry, 1948, p. (3) ; loc. 3, UMMZ 183041 (11), UIMNH P6773 (6). 1024). SYSTEMATIC DESCRIPTIONS 77
Ecology. "Found plentifully under wood, leaves, Family CIONELLIDAE stones, old logs, in moss and on the banks of streams" Genus CIONELLA Jeffreys, 1829 (Baker, 1902, p. 251). Cionella lubrica (Miiller), 1774 Occurrence and material: Sand Draw local fauna. Sand Draw loc. 3, UMMZ 181133 (6) ; loc. 6, UMMZ 181223 (1). Cionella lubrica (Miiller). Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, pt. 2, p. 1047. Vallonia gracilicosta Reinhardt, 1883 Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, Vallonia gracilicosta Reinhardt. Hibbard. 1941, Kansas Univ. p. 151. Sci. Bull., v. 27, p. 95. Distribution. Holarctic. "Point Barrow, Alaska, Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. and Queen Charlotte Islands to Labrador and New Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. 408. foundland, south in the East to Washington, D.C., and Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, southern Missouri; in all the Western and Mountain pt. 2, p. 1028, fig. 549. States except California; to the Mexican boundary Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. in Arizona; in the Sierra Madre of western Chihua 151, pi. 14, fig. f. hua." (Pilsbry, 1948, p. 1048; see also Pilsbry, 1953, Distribution. Minnesota to Montana, south and p. 165). southwest to Nebraska, New Mexico, Arizona, and Ecology. "(7. lubrica lives among the damp under- California (Pilsbry, 1948, p. 1029-1030). leaves in densely shaded places; under wood, such as Ecology. In northern Nebraska this species occurs under logs, bark, and stones on slightly moist leaf old board sidewalks; in chinks of stone walls and mold in wooded areas. under stones" (Pilsbry, 1948, p. 1049). In northern Nebraska it was found among damp leaves and small Occurrence and material: Rexroad local fauna; Recent in northern Nebraska. Rexroad loc. 3, UMMZ 183043 (1800). logs in a wooded ravine. Occurrence: Rexroad or Bender local fauna (Frye and Vallonia parvula Sterki, 1893 Leonard) ; Recent in northern Nebraska. Vallonia parvula Sterki. Pilsbry, 1948, Acad. Nat. Sci. Phila Family STTCCINEIDAE delphia Mon. 3, v. 2, pt. 2, p. 1027, fig. 547. Distribution. Great Plains from South Dakota to Land snails of the family Succineidae are common north-central Oklahoma; eastward to northern Mis and often conspicuous members of the Recent mol- souri, northern Ohio, southern Ontario, and western luscan fauna of the central Great Plains. Their New York. shells are likewise common as fossils in both Pliocene Ecology. In northern Nebraska V. parvula occurs and Pleistocene deposits of this region. Even the under logs, bark, and stones on slightly moist leaf best preserved fossils may be unidentifiable, however, mold in wooded areas. because the shells alone are often not specifically Occurrence and material: Bender local fauna; Recent in diagnostic. After studying the Succineidae of Kansas, northern Nebraska. Bender loc. la, UMMZ 184129 (31) ; loc. Miles (1958) found that most species can not be iden Ic, UMMZ 183020 (1). tified by shell characters, and that two of the three Vallonia perspectiva Sterki, 1893 genera are indistinguishable from their shells. Oxy Vallonia perspectiva Sterki. Pilsbry, 1948, Acad. Nat. Sci. loma, with the one species Oxyloma retusa in Kansas, Philadelphia Mon. 3, v. 2, pt. 2, p. 1033, fig. 553. may be distinguished from Succinea and Quickella; Distribution. Sparsely recorded from New Jersey Succinea ovalis and adult specimens of Succinea con- and Alabama to the Rocky Mountains. More abun cordialis can be identified by shell features; but the dant in Arizona and New Mexico; Chihuahua (Pils other species of Succinea and Quickella must be deter bry, 1948, p. 1034). mined by anatomical criteria. Ecology. Uncertain. The fossil succineid shells considered in this paper Occurrence and material: Saw Rock Canyon, Rexroad, include only one of the specifically identifiable types: Dixon, and Sanders local faunas. Saw Rock Canyon, UMMZ Oxyloma retusa. At least for the present, all others 177567 (2) ; Rexroad loc. 2, UMMZ 183077 (29) ; loc. 3, UMMZ are lumped as "cf. Succinea" Perhaps future detailed 183044 (150) ; Dixon loc. 1, UMMZ 182222 (1) ; Sanders loc. 1, UMMZ 182159 (150); loc. 2, UMMZ 184165 (1); loc. 3, study of the shells of dissected Recent snails will UMMZ 182188 (1). enable more of the fossils to be identified specifically. 78 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS Genus SUCCINEA Draparnaud, 1801 ture, but differ by their narrower spire and shallower Succinea ovalis Say, 1817 suture. They are closest to 8. luteola Gould (Pilsbry, 1948, p. 830). One other specimen from locality 4b Sttccinea ovalis Say. Pilsbry, 1948, Acad. Nat. Sci. Philadel is of similar form, but the remaining Rexroad mate phia Mon. 3, v. 2, pt. 2, p. 801, fig. 430-433. rial is indeterminate. Distribution. "Newfoundland and James Bay to Bender loc. 1, UMMZ 177511 (5) ; loc. la, UMMZ North Dakota and Nebraska, south to Alabama" 184130 (32) ; loc. Ic, UMMZ 183021 (9); loc. 2, UMMZ (Pilsbry, 1948, p. 803). 184156 (75). Seven adult specimens from locality 2 Ecology. "S. ovalis is usually found on low ground show considerable variation in proportions, but share near streams, in summer often upon the weedy herbage a high, narrow spire, moderately deep suture, and of such places, a foot or two from the ground" (Pils fairly smooth shell surface. They are higher spired bry, 1948, p. 804). In northern Nebraska it was and narrower than the Saw Rock Canyon specimens, found under logs among damp leaves, in a wooded similar to the Rexroad locality 3 material, and may ravine well above the flood-plain of the Niobrara represent Succinea luteola Gould also. Bender lo River. cality 1 material is not diagnostic. Remarks. The locality in northern Brown County, Sand Draw loc. 1, UMMZ 181248 (7) ; loc. 2, UMMZ Nebraska, recorded in this paper is farther southwest 184183 (13); loc, 3, UMMZ 181130 (34); loc. 4, than previously known occurrences. UMMZ 177324 (10); loc. 5, UMMZ 181139 (45), Occurrence: Recent in northern Nebraska. 181236 (150) ; loc. 6, UMMZ 177434 (11), 181224 (7) ; loc, 7, UMMZ 177234 (21). Almost all the material cf. Succinea is indeterminate. One specimen from locality 1 and Succinea grosvenori Lea. Hibbard, 1941, Kansas Univ. Sei. two from locality 6 (UMMZ 177434) are similar to Bull., v. 27, p. 95. the Rexroad material. Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. Dixon loc, 1, UMMZ 182223 (7); loc. 2, UMMZ Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. 408. 191527 (12). The few immature specimens collected Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. are closer to the Rexroad than to the Saw Rock 151. Canyon material. Succinea grosvenori gelidu F. C. Baker. Hibbard, 1941, Sanders loc. 1, UMMZ 182160 (500); loc. 2, UMMZ Kansas Univ. Sci. Bull., v. 27, p. 95. 184166 (10); loc, 3, UMMZ 182189 (44). About a Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. dozen adult specimens, and others nearly adult, show Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. 408. considerable variation in spire and aperture propor Succinea sp. Taylor, 1954, Michigan Univ. Mus. Zool. Occ. tions. They overlap the Rexroad locality 3 material, Papers 557, p. 15. but are mostly slightly lower spired. These also are Occurrence: Saw Rock Canyon, Red Corral, Rexroad, Ben like S. luteola Gould. Sanders locality 3 specimens der, Sand Draw, Dixon, and Sanders local faunas: Recent in are not determinable, but seem conspecific, northern Nebraska. Discussion of materials. Saw Rock Canyon, IJMMZ Genus OXYLOMA Westerlund, 1885 172363 (12), 1T7568 (200), ITSGS 19089-7 (17). Three Oxyloma retusa (Lea), 1834 fairly large specimens have short spires, elongate aper tures, and slightly impressed sutures. These shells Oxyloma retusa (Lea). Pilsbry, 1948, Acad Nat. Sci. Philadel agree well with Succinea concordialis Gould (Pilsbry, phia Mon. 3, v. 2, pt. 2, p. 785, fig. 421, 422c. Miles, 1958, Kansas Univ. Sci. Bull., v. 38, p. 1521, pi. 1, 1948, p. 833). fig. G, H: fig. 6, 7. Red Corral, UMMZ 183007 (1). The single juvenile Oxyloma haydeni (W. G. Binney). Pilsbry, 1948, Acad. Nat. specimen is indeterminate. Sci. Philadelphia Mon. 3, v. 2, pt. 2, p. 797, fig. 427e. Rexroad loc. 1, UMMZ 183062 (54) ; loc. 2, UMMZ Distribution. Ohio west to Montana, south to Kan 183078 (10) ; loc. 3, UMMZ 183045 (300) ; loc. 4b, sas. If Oxyloma siHimani is a synonym, as suggested UMMZ 177607 (6), 181057 (1). About a dozen speci by Pilsbry (1948, p. 796), the range may include mens from locality 3 may be only half grown. They parts of Nevada and Washington. differ from the Saw Rock Canyon material in having Ecology. In northern Nebraska this species was a narrow shell, relatively large pointed spire with always found within about 6 inches of permanent moderate suture, and smaller, ovate aperture. These water, crawling about on dead wood or among ripar specimens resemble S. grosvenori gelida and 8. avara ian vegetation and debris. Although not strictly in having a relatively prominent spire and small aper aquatic, its habitat overlaps that of the semiaquatic 79 Fossaria dalli. In Meade County State Park, Kansas, Ecology. "It lives on decaying wood in shady or "this snail thrives on the moist marshes and borders humid places, also on damp leaves" (Pilsbry, 1948, of the pools in the artesian basin, where it is frequently p. 627). "In eastern Kansas the species may occur found on watercress'1 (Leonard, A. E., 1943, p. 240). in grassy fields, on sparsely timbered slopes, and on Remarks. Miles (1958) has demonstrated the great rocky ledges as well as in more moist places. In the variation in shell form of Oxyloma retusa, even within more arid parts of the state, however, H. parallelus single populations. The original illustrations of Oxy- is limited to woodland cover, and usually occurs loma haydeni (Pilsbry, 1948, p. 796, fig. 427e) show around decaying timber1' (Leonard and Goble, 1952, striking resemblance to those of established 0. retusa p. 1039). (Miles, 1958, p. 1537, pi. 1, fig. H). 0. haydeni was Occurrence and material: Dixon local fauna. Dixon loc. 2, originally distinguished by shell features which are UMMZ 191529 (18). evidently invalid. Subgenus HEBETODISCUS Baker, 1929 Occurrence and material: Dixon local fauna; Recent in northern Nebraska. Dixon loc. 1, UMMZ 182212 (37) ; loc. 2, Helicodiscus (Hebetodiscus) singleyanus (Pilsbry), 1890 UMMZ 191528 (27). Helicodiscus singleyanus Pilsbry. Hibbard, 1941, Kansas Univ. Sci. Bull., v. 27, p. 95. Family ENDODONTIDAE Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. Subfamily ENDODONTINAE 408. Genus DISCUS Fitzinger, 1833 Helicodiscus sinyleyanus (Pilsbry). Pilsbry, 1948, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, pt. 2, p. 636, fig. 346. Subgenus DISCUS s. s. Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. 151. Discus (s. s.) cronkhitei (Newcomb), 1865 Distribution. NewT Jersey to Florida, west to South Discus cronkJtitei (Newcomb). Pilsbry, 1948, Acad. Nat. Sci. Dakota, Colorado, and Arizona (Pilsbry, 1948, p. Philadelphia Mon. 3, v. 2, pt. 2, p. 601, fig. 328a-d. 636). Distribution. Alaska to the mountains of southern Ecology. This species has apparently never been California; Rocky Mountains south into Arizona and collected alive in the High Plains. In southwestern New Mexico; Colorado and northern Canada east to Kansas fresh shells have been found frequently by northern Illinois, Maryland, Newfoundland, and University of Michigan field parties in the course Labrador; also Missouri (Pilsbry, 1948, p. 602-603). of collecting fossils by washing bulk matrix through Ecology. "In the east it lives in humid forest, screens. In this process any Eecent shells which may under dead wood, and among rotting leaves or grass be present are recovered along with the fossils. From in rather wet situations1' (Pilsbry, 1948, p. 604). In the localities so represented, it appears probable that northern Nebraska it occurred under sticks and logs Helicodiscus singleyanus lives among grass roots, even on moist leaf mold, always close to running water. on exposed slopes that become hot and dry during Remarks. The localities in northern Nebraska re the summer. ported in this paper are the most southern known Occurrence and material: Saw Rock Canyon, Rexroad. Ben Recent occurrences of this species in the High Plains. der, Sand Draw, Dixon, and Sanders local faunas. Saw Rock Canyon, UMMZ 181289 (2) ; Rexroad loc. 2, UMMZ 183079 Occurrence: Recent in northern Nebraska. (7) : loc. 3, UMMZ 183046 (500) ; loc. 4b, UMMZ 181287 (20) ; Bender loc. 1, UMMZ 181288 (24) ; loc. la, UMMZ 184131 Subfamily HELICODISCINAE (38) ; loc. Ib, UMMZ 184142 (300) ; loc. Ic, UMMZ 183022 Genus HELICODISCUS Morse, 1864 (100) ; loc. 2, UMMZ 184157 (5) ; Sand Draw loc. 2, UMMZ 184184 (4) ; Dixon loc. 2. UMMZ 191530 (400) ; Sanders loc. Subgenus HELICODISCUS s. s. 2. UMMZ 182171 (100) ; loc. 3, UMMZ 182190 (3).
Helicodiscus (s. s.) parallelus (Say), 1821 Subfamily PUNCTINAE Helicodiscus parallel us (Say). Pilsbry, 1948, Acad. Nat. Sci. Genus PUNCTUM Morse, 1864 Philadelphia Mon. 3. v. 2, pt. 2, p. 625, fig. 338c, 339. Leonard and Goble. 1952, Kansas Univ. Sci. Bull., v. 34, Punctum minutissimum (Lea), 1841 p. 1039. pi. 101, fig. 24, 25. Punctum minutissimum (Lea). Pilsbry, 1948, Acad. Nat. Sci. Distribution. Eastern North America, from New Philadelphia Mon. 3, v. 2, pt. 2, p. 644, fig. 350. foundland south to Georgia and Alabama; westward Distribution. Newfoundland to Florida, w^est to to South Dakota and Oklahoma (Pilsbry, 1948, p. Oregon, Colorado, New Mexico, and Puebla, Mexico 626). (Pilsbry, 1948, p. 644-645). 80 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS Ecology. "This dwarf among pygmies lives on States from Texas to North Carolina" (Pilsbry, 1946, damp leaves, around decaying logs, and is chiefly p. 236). to be obtained by sifting leaves, according to my expe Ecology. UE. fulvus lives among damp leaves in rience. Morse states that [in Maine] 'dense, hard well-shaded places, and may usually be obtained by wood growths appear to be their favorite position. leaf sifting where its presence wTould otherwise be They prefer the rotten bark of beech trees, and fre unsuspected" (Pilsbry, 1946, p. 236). In northern quently are found in the large forms of fungi, such Nebraska it was found in damp leaves or under logs as Polyporus and Boletus.'" (Pilsbry, 1948, p. 645). and bark on moist ground. In northern Nebraska it was taken among moist leaves Occurrence: Recent in northern Nebraska. beside a spring-fed brook. Occurrence: Recent in northern Nebraska. Subfamily ZONITINAE Genus NESOVITREA Cooke, 1921 Family LIMACIDAE Subgenus PERPOLITA Baker, 1928 Genus DEROCERAS Rafinesque, 1820 Nesovitrea (Perpolita) electrina (Gould), 1841 Subgenus uncertain Retinella electrina (Gould). Hibbard, 1941, Kansas Univ. Sci. Deroceras aenigma Leonard, 1950 Bull., v. 27, p. 95. Plate 4, figures 14-16, 19-26 Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. Deroceras aenigma Leonard, 1950, Kansas Univ. Paleont. 408. Contr. 8, p. 38, pi. 5, fig. E. Pilsbry, 1946, Acad. Nat. Sci. Philadelphia Mon. 3, v. 2, Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, pt. 1, p. 256, fig. 126. p. 151, pi. 14, fig. e. Leonard, 1950, Kansas Univ. Paleont. Contr. 8, p. 36, pi. Taylor, 1954, Michigan Univ. Mus. Zool. Occ. Papers 557, 5, fig. H. p. 6, 15. Hibbard, 1956, Michigan Acad. Sci. Papers, v. 41, p. 171. Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. 151, pi. 14, fig. 1. Distribution. Upper Pliocene to upper Pleistocene Nesovitrea (Perpolita) electrina (Gould). Forcart, 1957, in the High Plains (Taylor, 1954a). Archiv fur Molluskenkunde, v. 86, p. 110. Ecology. Like living species of Deroceras, D. aen Distribution. North America north of about lat igma probably favored damp riparian habitats. 38° N., southward in the Rocky Mountains to Arizona Occurrence and material: Saw Rock Canyon, Red Corral, and New Mexico. Rexroad, Bender, Sand Draw, Dixon, Deer Park, and Sanders Ecology. "Retinella electrina, is an inhabitant of local faunas. Saw Rock Canyon, UMMZ 172365 (1), 177569 (125, 2 sinistral), USGS 19089-10 (17); Red Corral, UMMZ woodlands where it lives in decaying leaves, beneath 183008 (50) : Rexroad loc. 1, UMMZ 183063 (40) ; loc. 2, loosened bark on dead trees and under sticks and fallen UMMZ 183080 (5) ; loc. 3, UMMZ 183047 (400) ; loc. 4a, logs. It is frequently associated with another wood UMMZ 177442 (500), 177572 (500); loc. 4b, UMMZ 177608 land snail (Zonitoides arboreus) of similar size and (2) ; Bender loc. 1, UMMZ 177513 (5) ; loc. la. UMMZ 184132 superficial appearance. R. electrina, is common in the (100) ; loc. Ic, UMMZ 183023 (100) ; Sand Draw loc. 1, UMMZ woodlands of eastern Kansas, where the annual rain 181249 (6) : loc. 3, UMMZ 181206 (2) ; loc. 4, UMMZ 177327 (100) ; loc. 6, UMMZ 177440 (10) ; loc. 7, UMMZ 177238 (3) ; fall is generally more than 35 inches but it declines Dixon loc. 1, UMMZ 182224 (75) ; loc. 2, UMMZ 191531 (6) ; in frequency of occurrence toward the more arid Deer Park, UMMZ 185798 (65) ; Sanders loc. 1, UMMZ 182161 Plains Border province, and is unknown in the Plains (57) ; loc. 2, UMMZ 182172 (3) ; loc. 3, UMMZ 182191 (24). province, even where timber is locally available"
Family ZONITIDAE (Leonard, 1950, p. 37). In northern Nebraska it was found only under dead wood or among leaves on damp Subfamily EUCONULINAE ground close to running w^ater. Genus EUCONULUS Reinhardt, 1883 Occurrence and material: Rexroad local fauna; Dixon local fauna (Frye and Leonard) ; Recent in northern Nebraska. Euconulus fulvus (Miiller), 1774 Rexroad loc. 3, UMMZ 183049 (75). Euconulus fuJwis (Miiller). Pilsbry, 1946, Acad. Nat. Sci. Nesovitrea? sp. Philadelphia Mon. 3, v. 2, pt. 1, p. 235, fig. 116, 117. Distribution. "Almost throughout the Holarctic Remarks. A single juvenile shell is indeterminate. realm, but wanting in the Gulf and South Atlantic It may represent one of the species of Retinella in the SYSTEMATIC DESCRIPTIONS 81
Rexroad local fauna, but perhaps more likely Nesovi- Hawaiia minuscula (Binney). Pilsbry, 1946, Acad. Nat. Sci. trea electrina. Philadelphia Mon. 3, v. 2, pt. 1, p. 420, fig. 228a, b, 229:l-3. Occurrence and material: Saw Rock Canyon local fauna, Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, UMMZ 183010 (1). p. 151, pi. 14, fig. m. Taylor, 1954, Michigan Univ. Mus. Zool. Occ. Papers 557, Genus RETINELLA Fischer, 1877 p. 15. Distribution. North America from Alaska and Subgenus GLYPHYALUS Baker, 1928 Maine to Costa Rica. "It is generally spread over Retinella (Glyphyalus) wheatleyi (Bland), 1883 every eastern and midwestern state, and in Florida as far south as Miami and Cape Sable, though not seen Retinella wheatleyi Bland. Hibbard, 1941, Kansas Univ. Sci. from the keys. It becomes rather local in the Rocky Bull., v. 27, p. 95. Mountain States, and has not been seen from Wash Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. ington, Oregon, Idaho, Nevada and Utah, and only 408. towards the south in California, where some of the Retinella wheatleyi (Bland). Pilsbry, 1946, Acad. Nat. Sci. records are probably owing to importation with Philadelphia Mon. 3, v. 2, pt. 1, p. 272, fig. 134, 141:1-3. plants" (Pilsbry, 1946, p. 421-423). Distribution. Sporadically from Rhode Island and Ecology. The wide geographic range of this com North Carolina west to Michigan, Missouri, and north mon snail is correlated with adaptability to various habitats. In northeastern Kansas, Franzen and Leon ern Alabama (Pilsbry, 1946, p. 272-273). ard (1943) found that it "* * * is a common snail in Ecology. In Knox County, Tenn., "Even in the the Wakarusa valley, where it lives in rocky ledges spring, it was rare except in a shallow valley on the or among dead leaves and under fallen logs. It west-facing (more humid) slope of the ridge, where thrives in piles of moist drift where it has been cast one or two individuals per square meter were obtained by flood waters." In Meade and Clark counties, Kan under the decaying leaves in the oak-chestnut woods" sas, "This species is widely distributed over the area, (Baker, quoted in Pilsbry, 1946, p. 273). being nowhere numerous. It withstands arid condi Occurrence: Rexroad local fauna (Hibbard). tions successfully but is most numerous in wooded places where moisture conditions are better than on Submenus GLYPHYALOPS Baker, 1928 the treeless prairies" (Leonard, A. E., 1943, p. 238). In northern Nebraska it was found under logs and Retinella (Glyphyalops) rhoadsi (Pilsbry), 1899 bark on damp ground near running water. Retinella rhoadsi Pilsbry. Hibbard, 1941, Kansas Univ. Sci. Occurrence and material: Saw Rock Canyon, Red Corral, Bull., v. 27, p. 9o. Rexroad, Bender, Sand Draw, Dixon, and Sanders local Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. faunas; Recent in northern Nebraska. Saw Rock Canyon, Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. UMMZ 173592 (1); 177570 (34); Red Corral, UMMZ 183009 408. (4) ; Rexroad loc. 1, UMMZ 183064 (11) ; loc. 2, UMMZ 183- 081 (200) ; loc. 3, UMMZ 183048 (1000) ; loc. 4b, UMMZ 177609 Retinella rhoadsi (Pilsbry). Pilsbry, 1946, Acad. Nat. Sci. (200), 181058 (125) ; Bender loc. 1, UMMZ 177512 (285) ; loc. Philadelphia Mon. 3, v. 2, pt. 1, p. 286, fig. 145. la, UMMZ 184133 (3200) ; loc. Ib, UMMZ 184143 (150) ; loc. Distribution. Maine to North Carolina, west to Ic, UMMZ 183024 (1600); loc. 2, UMMZ 184158 (60); Sand Michigan (Pilsbry, 1946, p. 286) ; Jackson County, Draw loc. 1, UMMZ 181250 (2) ; loc. 2, UMMZ 184185 (6) ; loc. 3, UMMZ 181207 (2) ; loc. 6, UMMZ 177441 (8) ; 181225 111. (Baker, 1939, p. 70). (1) ; loc. 7. UMMZ 177237 (12) ; Dixon loc. 1, UMMZ 182225 Ecology. Uncertain; presumably similar to that (49) ; loc. 2, UMMZ 191532 (150) ; Sanders loc. 1, UMMZ 182- of R. wheatleyi. 163 (600) ; loc. 2, UMMZ 182173 (26) ; loc. 3, UMMZ 182192 (300). Occurrence: Rexroad local fauna (Hibbard). Subfamily GASTRODONTINAE
Genus HAWAIIA Gude, 1911 Genus ZONITOIDES Lehmann, 1862 Zonitoides (s. s.) arboreus (Say), 1816 Hawaiia minuscula (Binney), 1840 Zonitoides arltoreiix (Say). Pilsbry, 1946, Acad. Nat. Sci. Hawaii miniscula Binney. Hibbard, 1941, Kansas Univ. Sci. Philadelphia Mon. 3, v. 2, pt. 1, p. 480, fig. 261, 262. Bull., v. 27, p. 95. Frye and Leonard, 1952, Kansas Geol. Survey Bull. 99, p. 151, pi. 14, fig. p. Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. Distribution. North America from northern Can 408. ada to Costa Rica (Pilsbry, 1946, p. 481).
50894S O 6C 82 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS Ecology. "* * * occurs in woodlands, living under whorls, slightly coarsening behind lip; umbilical side loosened bark, leaves, and stones, and other cover of whorls with numerous, raised, dotlike papillae; last affording protection from the sun, and providing at whorl with fine, close, irregular, wavy, impressed, least moderate moisture" (Leonard, 1950, p. 37). In spiral striae. Measurements in millimeters: paratype northern Nebraska it was found under logs and bark (UMMZ 183050) diameter 9.5, height 4.5, whorls 5; and among leaves on damp ground, usually in wooded holotype diameter 7.8, height 4.0, whorls 5. areas. Variation. The spire height varies slightly, but Occurrence: Rexroad or Bender and Dixon local faunas not as much as the measurements might indicate. The (Frye and Leonard) ; Recent in northern Nebraska. holotype has a relatively high body whorl and a low spire, rather than a much higher spire than the Family POLYGYBIDAE measured paratype. Umbilical width varies from a Subfamily POLYGYBINAE little more than one-third shell diameter (paratype) to one-fourth (holotype). The inner, permeable por Genus POLYGYRA Say, 1818 tion of the umbilicus varies correspondingly in width. Subgenus ERYMODON Pilsbry, 1956 Spiral striation on the slightly worn paratype is visible on only the last one-third whorl, but on the Polygyra (Erymodon) rexroadensis 6 Taylor, n. sp. holotype it is visible on all the base exposed (slightly Plate 1, figures 17, 23; Plate 2, figures 4, 5 over one whorl). On this paratype this sculpture ex tends over the whorl, but on the holotype it is re Polygyra mooreana tholus (Binney). Hibbard, 1941, Kansas stricted below the periphery. The other unworn para Univ. Sci. Bull., v. 27, p. 95 [not of Binney]. type shows the spiral sculpture but it is imperfect and Hibbard, 1941, Kansas Acad. Sci. Trans., v. 44, p. 265 [not of Binney]. the whorls cannot be counted. Frye and Hibbard, 1941, Kansas Geol. Survey Bull. 38, p. Distribution. Late Pliocene of southwestern Kan 408 [not of Binney]. sas. Polygyra mooreana (Binney). Frye and Leonard, 1952, Remarks. P. rexroadensis is most similar to P. Kansas Geol. Survey Bull. 99, p. 151, pi. 14, fig. v [not texasiana texasensis and P. chisosensis. It differs from of Binney]. the former by the weaker axial sculpture, the spiral Types. Holotype UMMZ lTT610a, Kexroad loc. 4b. striae on the last whorl, the umbilical papillae, and Paratypes: UMMZ 177610 (1) ; 183050 (4), loc. 3; the shape of the parietal tooth. In P. t. texasensis 183082 (2), loc. 2. this tooth is roughly v-shaped, and both branches Diagnosis. A Polygyra of the P. texasiana group descend gradually toward their distal ends. In P. characterized by its weak axial sculpture, spiral striae, rexroadensis this tooth is asymmetrical, wTith a long, high and asymmetrical parietal tooth, and absence of high, columellar branch which descends sharply at columellar tubercle. its distal end. P. chisosensis has minutely granular Description of types. Shell depressed, plane or with sculpture and a columellar tubercle; both of these low spire above; periphery rounded; whorls 5, characters are absent in P. rexroadensis. P. chiso strongly descending to aperture; umbilicus at first sensis also lacks the umbilical papillation and spiral steep sided, perforate to apex, expanding in last whorl striation of P. rexroadensis. P. mooreana and P. tho to about one-third of shell diameter, with a shallow lus have stronger sculpture, a columellar tubercle, a groove parallel to suture for last half whorl; aperture V-shaped parietal tooth, and lack the spiral striae and semicircular except for dental barrier; outer lip thick umbilical papillae. ened, reflected following marked constriction of body Material from the Bender local fauna consists only whorl, bearing 2 laterally compressed teeth of equal of small fragments. These show the spiral striae and size separated by deep sinus; from each tooth a broad, umbilical papillae of P. rexroadensis^ but suggest pal thick, ridge extends toward lip insertions; parietal atal teeth of subequal size. This difference may be callus bearing a large, asymmetrical biramose tooth a character of immaturity, however, and the specimens opposite palatal teeth; branch toward columella about are considered referable to the Rexroad species. 4 times as long as that toward the upper lip insertion, Occurrence and material: Rexroad and Bender local faunas. straight, high and even for nearly two-thirds its Rexroad loc. 2, UMMZ 1S3082 (8. including 2 fragmentary sub- length, descending sharply toward columella; no col adult paratypes) ; loc. 3, LTMMZ 183050 (67, including 1 adult umellar tubercle; first (embryonic) whorl with fine paratype (measured) and 3 subadult paratypes) ; loc. 4b, growth lines, becoming coarse and irregular on later UMMZ 177610 (6, including holotype and 1 fragmentary sub- adult paratype), 181059 (3); Bender loc. Ib, UMMZ 184144 6 Named for its occurrence in the Rexroad local fauna. (10). LITERATURE CITED 83
Genus STENOTREMA Rafinesque, 1819 1945, The molluscan family Planorbidae; collation, re vision and additions by H. J. van Cleave: Univ. Illinois Stenotrema leal leai (Binney), 1840 Press, 530 p. Baker, H. B., 1947, Amnicola and Eu-amnicola: Nautilus, v. Stenotrema monodon (Rackett). Pilsbry, 1940, Acad. Nat. Sci. 60, p. 105-106. Philadelphia Mon. 3, v. 1, pt. 2, p. 676, fig. 421a, b. Berry, E. G., 1943, The Amnicolidae of Michigan: distribution, Leonard, 1950, Kansas Univ. Paleont. Contr. 8, p. 36. ecology, and taxonomy: Michigan Univ. Mus. Zool. Misc. Stenotrema leai (Binney). Pilsbry, 1948, Acad. Nat. Sci. Publ. 57. 68 p., 9 pi. Philadelphia Mon. 3, v. 2, pt. 2, p. 1099. Brown, Charles N., 1956, The origin of caliche on the north Distribution. New York to Maryland, west to eastern Llano Estacado, Texas: Jour. Geology, v. 64, p. South Dakota and Nebraska. 1-15. Ecology. "$. monodon is a snail of damp places Byrne, F. E., and T. G. McLaughlin, 1948, Geology and ground- water resources of Seward County, Kansas: Kansas Geol. near the water" (Pilsbry, 1940, p. 678). "Stenotrema Survey Bull. 69, 140 p., 4 fold. pi. monodon thrives in rather humid forests, and may be Clarke, W. T., Jr., 1938, List of molluscs from drift debris of found on low terraces and flood plains of streams, Paladora Creek, Texas: Nautilus, v. 52, p. 14-15. under leaves, logs, and stones" (Leonard, 1950, p. 36). Connolly, M., 1939, A monographic survey of South African In northern Nebraska it was found under boards on nonmarine Mollusca: South African Mus. Annals, v. 33, p. 1-660, pi. 1-19. damp ground, on the floodplain of the Niobrara River. Crandall, A. O., 1901, The American Physae: Nautilus, v. 15, Remarks. The Nebraska specimens have the open p. 25-30, 42-45, 54-58, 69-71, pi. 2, fig. 5-7. umbilicus and broader aperture of Stenotrema, leai Dall, W. H., 1905, Land and fresh water mollusks: Harriman leai, rather than S. leai aliciae. The occurrence in Alaska Exped., v. 13, p. 1-171, pi. 1-2. northern Cherry County, Nebr., reported in this paper 1919, Mollusks, recent and Pleistocene: Canadian Arctic is farther west than previously known localities. Exped. 1913-18 Report, v. 8, part A, 29 p., 3 pi. Downs, T., 1954, Pleistocene birds from the Jones fauna of Pilsbry (1940, p. 678) listed two Kansas records Kansas: Condor, v. 56, p. 207-221. for this form, but Leonard (1950, p. 36) referred all Eiseley, L. C., 1937, Index Mollusca and their bearing on cer Kansas material to 8. leai aliciae. tain problems of prehistory; a critique: Philadelphia Anthropol. Soc. Pub., v. 1, p. 77-93. LITERATURE CITED Elias, M. K., 1942, Tertiary prairie grasses and other herbs from the High Plains: Geol. Soc. America Spec. Paper Alexander, R. C., 1952, Checklist of New Jersey land snails: 41, 176 p., 17 pi. Nautilus, v. 66, p. 54-59. Flint, R. F., 1947, Glacial geology and the Pleistocene epoch: American Commission on Stratigraphic Nomenclature, 1949, New York, John Wiley and Sons, 589 p., 6 pi. Note 9 The Pliocene-Pleistocene boundary: Am. Assoc. Forcart, Lothar, 1957, Taxionomische Revision palaarktischer Petroleum Geologists Bull., v. 33, p. 1276-1280. Zonitinae, I: Archiv fur Molluskenkunde, v. 86, p. 101-136. Antevs, Ernst, 1928, The last glaciation, with special reference Franzen, D. S., and A. B. Leonard, 1942, A preliminary sur to the ice retreat in northeastern North America: Am. vey of the Mollusca of Kingman County, Kansas: Kansas Geog. Soc. Research Ser. 17, 292 p., 9 pi. Acad. Sci. Trans., v. 45, p. 334-343. Baker, F. C., 1902, The Mollusca of the Chicago area, the 1943, The Mollusca of the Wakarusa River valley: Gastropoda: Chicago Acad. Sci. Nat. Hist. Survey Bull. Kansas Univ. Sci. Bull., v. 29, p. 363-439. 3, part 2, p. 133-410, pi. 28-36. 1947, Fossil and living Pupillidae (Gastropoda = 1911, The Lymnaeidae of North and Middle America, Pulmonata) in Kansas: Kansas Univ. Sci. Bull., v. 31, p. recent and fossil: Chicago Acad. Sci. Spec. Pub. 3, p. 311-411. i-xvi, 1-539, pi. 1-58. Frick, Childs, 1937, Horned ruminants of North America: 1919, Description of a new species and variety of Am. Mus. Nat. Hist. Bull., v. 69, p. i-xxvii, 1-669. PlanorUs from postglacial deposits: Nautilus, v. 32, p. Frierson, L. S., 1927, A classified and annotated check list of the North American naiades: Waco, Texas, Baylor Univ., 94-97, pi. 7, figs. 1-10. Ill p. 1928, The fresh water Mollusca of Wisconsin, part I, Frye, J. C., 1942, Geology and ground-water resources of Meade Gastropoda: Wisconsin Geol. Nat. Hist. Survey Bull. 70, County, Kansas: Kansas Geol. Survey Bull. 45, 152 p., 3 part 1, p. 1-507, pi. 1-28. maps. 1928a, The fresh water Mollusca of Wisconsin, part II, Frye, J. C., and C. W. Hibbard, 1941, Pliocene and Pleistocene Pelecypoda: Wisconsin Geol. Nat. Hist. Survey Bull. 70, stratigraphy and paleontology of the Meade basin, south part 2, p. 1-495, pi. 29-105. western Kansas: Kansas Geol. Survey Bull. 38, p. 389- 1937, Pleistocene land and fresh-water Mollusca as in 424. dicators of time and ecological conditions, in G. G. Mac- Frye, J. C., A. B. Leonard, and C. W. Hibbard, 1943, West Curdy fed.]. Early Man: Philadelphia, J. B. Lippincott, ward extension of the Kansas "Eqiius beds": Jour. Geol p. 67-74. ogy, v. 51, p. 33-47. Frye, J. C., A. Swineford, and A. B. Leonard, 1948, Correla 1938, New land and freshwater Mollusca from the upper tion of Pleistocene deposits of the central Great Plains Pliocene of Kansas and a new species of Gyraulus from with the glacial section: Jour. Geology, v. 56, p. 501-525, early Pleistocene strata: Nautilus, v. 51, p. 126-131. pi. 1-2. 84 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Frye, J. C., and A. B. Leonard, 1952, Pleistocene geology of 1949a, Pliocene Saw Rock Canyon fauna in Kansas: Kansas: Kansas Geol. Survey Bull. 99, p. 1-230, pi. 1-19. Michigan Univ. Mus. Paleont. Contr., v. 7, p. 91-105. 1953, Definition of time line separating a glacial and in- 1949b, Pleistocene vertebrate paleontology in North terglacial age in the Pleistocene: Am. Assoc. Petroleum America, hi Pleistocene research; a review by the mem Geologists Bull., v. 37, p. 2581-2586. bers of the committee on interrelations of Pleistocene Frye, J. C,, A. B. Leonard and Ada Swineford, 1956, Stratig research, National Research Council: Geol. Soc. America raphy of the Ogallala formation (Neogene) of northern Bull., v. 60, p. 1417-1428. Kansas: Kansas Geol. Survey Bull. 118, 92 p., 9 pi. 1949c, Techniques of collecting microvertebrate fossils: Germain, L., 1931, Mollusques terrestres et fluviatiles Michigan Univ. Mus. Paleont. Contr., v. 8, p. 7-19, pi. 1-4. (deuxieme partie) : Faune de France, v. 22, p. 479-897, 1950, Mammals of the Rexroad formation from Fox pi. 14-26. Canyon, Meade County, Kansas: Michigan Univ. Mus. Geyer, David, 1922, Die Quartarmollusken und die Klimafrage: Paleont. Contr., v. 8, p. 113-192, pi. 1-5. Paleont. Zeitschr., v. 5, p. 72-94. 1951, Vertebrate fossils from the Pleistocene Stump Goodrich, C., 1940, Mollusks of a Kansas Pleistocene deposit: Arroyo member, Meade County, Kansas: Michigan Univ. Nautilus, v. 53, p. 77-79. Mus. Paleont. Contr., v. 9, p. 227-245, pi. 1-6. Harry, H. W., 1952, Carycliium exiguum (Say) of lower 1952, New Pleistocene interglacial (Sangamon?) local Michigan: morphology, ecology, variation and life history fauna from Kansas and its climatic significance (ab (Gastropoda, Pulmonata) : Nautilus, v. 66, p. 5-7. stract) : Geol. Soc. America Bull., v. 63, p. 1262. Hedley, Charles, and H. A. Pilsbry, 1895, On the Australasian 1953, The Saw Rock Canyon fauna and its stratigraphic G-undlachia, with note on American forms: Nautilus, v. 9, significance: Michigan Acad. Sci. Papers, v. 38, p. 387-411. p. 61-68. 1953a, Eqmis (Asinus) calobatus Troxell and associated Henderson, J., 1930, Ancylvs coloradensis, new name for A. vertebrates from the Pleistocene of Kansas: Kansas Acad. Jienflersoni Walker, 1925, not 1908: Nautilus, v. 44, p. Sci. Trans., v. 56, p. 111-126. 31. 1954, A new Pliocene vertebrate fauna from Oklahoma: Herrington, H. B., 1954, Pisidium species and synonyms, North Michigan Acad. Sci. Papers, v. 39. p. 339-359. America, north of Mexico: Nautilus, v. 67, p. 97-104, 131- 1954a, Second contribution to the Rexroad fauna: 138. Kansas Acad. Sci. Trans., v. 57, p. 221-237. Herrington. H. B., and Taylor, D. W., 1958, Pliocene and 1954b, A new Synaptomys, an addition to the Borchers Pleistocene Sphaeriidae (Pelecypoda) from the central interglacial (Yarmouth?) fauna: Jour. Mammalogy, v. 35, United States: Michigan Univ. Mus. Zool. Occ. Papers 596, p. 249-252. p. 1-28, pi. 1. 1955, The Jinglebob interglacial (Sangamon?) fauna Hibbard, C. W., 1938, An upper Pliocene fauna from Meade from Kansas and its climatic significance: Michigan Univ. County, Kansas: Kansas Acad. Sci. Trans., v. 40. p. 239- Mus. Paleont. Contr., v. 12, p. 179-228, 2 pi., 1 chart. 265. 1956, Vertebrate fossils from the Meade formation of 1939, Notes on some mammals from the Pleistocene of southwestern Kansas: Michigan Acad. Sci. Papers, v. 41, Kansas: Kansas Acad. Sci. Trans., v. 42, p. 463-479, pi. p. 145-200, 2 pi. 1-5. 1956a. Microtus pennsylvanicus (Ord) from the Hay 1940, A new Pleistocene fauna from Meade County, Springs local fauna of Nebraska: Jour. Paleont., v. 30, Kansas: Kansas Acad. Sci. Trans., v. 43, p. 417-425. p. 1263-1266. 1941, Paleoecology and correlation of the Rexroad fauna 1957, Two new Cenozoic microtine rodents: Jour. from the upper Pliocene of southwestern Kansas, as in Mammology, v. 38, p. 39-44. dicated by the mammals: Kansas Univ. Sci. Bull., v. 27, 1958, New stratigraphic names for early Pleistocene p. 79-104. deposits in southwestern Kansas: Am. Jour. Sci., v. 256, 1941a, Mammals of the Rexroad fauna from the upper p. 54-59. Pliocene of southwestern Kansas: Kansas Acad. Sci. 1958a, Notes on late Cenozoic shrews: Kansas Acad. Sci. Trans., v. 44, p. 265-313. Trans , v. 60, p. 327-336. 19411), The Borchers fauna, a new Pleistocene inter- Hibbard, C. W., and Taylor, D. W., 1960, Two late Pleistocene faunas from southwestern Kansas: Michigan Univ. Mus. glacial fauna from Meade County, Kansas: Kansas Geol. Paleont. Contr., (in press). Survey Bull. 38, p. 197-220. Hubendick, B., 1951, Recent LymnaeMae; their variation, 1943, The Rezabek fauna, a new Pleistocene fauna from morphology, taxonomy, nomenclature, and distribution: Lincoln County, Kansas: Kansas Univ. Sci. Bull., v. 29, Kungl. Svensk. Vetenskapsakad. Handl., ser. 4. v. 3, p. p. 235-247. 1-223, pi. 1-5. 1944, Stratigraphy and vertebrate paleontology of Isely, F. B., 1925, The fresh-water mussel fauna of eastern Pleistocene deposits of southwestern Kansas: Geol. Soc. Oklahoma : Oklahoma Acad. Sci. Proc., v. 4, p. 43-118. America Bull, v. 55, p. 707-754, pi. 1-3. Johnston, C. S., and D. E. Savage, 1955, A survey of various 1948, Late Cenozoic climatic conditions in the High late Cenozoic vertebrate faunas of the Panhandle of Texas, Plains of western Kansas, in Colbert, E. H. [ed.], Pleisto part I: Introduction, description of localities, preliminary cene of the Great Plains: Geol. Soc. America Bull., v. 59, faunal lists: California Univ. Pubs. Geol. Sci., v. 31, p. p. 592-597. 27-50, frontis, 1 map. 1949, Pleistocene stratigraphy and paleontology of Kenk, R., 1949, The animal life of temporary and permanent Meade County, Kansas: Michigan Univ. Mus. Paleont. ponds in southern Michigan: Michigan Univ. Mus. Zool. Contr., v. 7, p. 63-90, pi. 1, map. Misc. Pub. 71, 66 p., 3 pi. LITERATURE CITED 85
La Rocque, Aurele, 1956, Variation of carinae in Valvata 1952, The water rat in the Pleistocene of Texas: Jour. tricar India: Nautilus, v. 70, p. 13-14. Mammalogy, v. 33, p. 87-89. Lea, Isaac, 1864, Descriptions of twenty-four new species of Mengel, R. M., 1952, White Pelican from the Pleistocene of Pliysa of the United States and Canada: Acad. Nat. Sci. Oklahoma: Auk, v. 69, p. 81-82. Philadelphia Proc., 1864, p. 114-116. Miles, C. D., 1958, The family Succineidae (Gastropoda: 1866, New Unionidae, Melanidae, etc., chiefly of the Pulmonata) in Kansas: Kansas Univ. Sci. Bull., v. 38, p. United States: Acad. Nat. Sci. Philadelphia Jour., v. 6, p. 1499-1543. 113-187, pi. 22-24. Morrison, J. P. E., 1947, One hundred six years of Amnicola: 1867, Observations on the genus Un-io * * *, v. 11, 146 Nautilus, v. 60, p. 84-87. p., 24 pi. 1949. The cave snails of eastern North America: Amer. Lee, C. B., 1951, Succinea vaffinacontorta (section Calcisuc- Malacol. Union Ann. Rep., 1948, p. 13-15. cinea), a new amber snail from Kansas: Michigan Univ. Mozley, A., 1926, Preliminary list of the Mollusca of Jasper Mus. Zool. Occ. Pap. 533. 7 p., 2 pi. Park, Alberta: Nautilus, v. 40, p. 53-56. Leonard, A. B., 1948, Invertebrates of the Blancan, in Colbert, Oelrich, T. M., 1953, A new Boxturtle from the Pleistocene of B. H. [ed.], Pleistocene of the Great Plains: Geol. Soc. southwestern Kansas: Copeia, 1953, p. 33-38, pi. 1-2. America Bull., v. 59, p. 589-591. 1957, The status of the upper Pliocene turtle, Testudo 1948a, Five new Yarmouthian planorbid snails: Nauti titrrjida Cope: Jour. Paleont., v. 31, p. 228-241. lus, v. 62, p. 41-47, pi. 2. Osborn, H. F., 1936, Proboscidea, v. 1: New York, Am. Mus. 1950, A Yarmouthian molluscan fauna in the midcon- Nat. Hist., 802 p., frontis., 12 pi. tinent region of the United States: Kansas Univ. Paleont. Pilsbry, H. A., 1918-20, Pupillidae (Gastrocoptinae, Vertigini- Contr. 8, 48 p., 6 pi. nae) : Manual of Conchol., ser. 2, v. 25, p. 1-401, pi. 1-34. 1952, Illinoian and Wisconsinan molluscan faunas in 1922-26, Pupillidae (Orculinae, Pagodulinae, Acanthin- Kansas: Kansas Univ. Paleont. Contr. 9, 38 p., 5 pi. ulinae) : Manual of Conchol., ser. 2, v. 27, p. 1-369, pi. 1952a, New gastropods from the Blanco formation 1-32. (Nebraskan age, Pleistocene) in Kansas: Nautilus, v. 1934, Review of the Planorbidae of Florida, with notes 66, p. 37-45, pi. 5. on other members of the family: Acad. Nat. Sci. Philadel 1953, Molluscan faunules in Wisconsinan loess at phia Proc., v. 86, p. 29-66, pi. 7-11. Cleveland, Ohio: Am. Jour. Sci., v. 251, p. 369-376, pi. 1. 1940, Land Mollusca of North America (north of 1957, Types of late Cenozoic gastropods in the Frank Mexico) : Acad. Nat. Sci. Philadlephia Mon. 3, v. 1, pt. 2, Collins Baker collection, Illinois State Geological Survey: p. 575-994, i-ix. Illinois Geol. 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O., 1944, An early Pleistocene (Blancan) fauna of the Niagara frontier region and adjacent territory: from Nebraska: Field Mus. Nat. Hist. Geol. Ser., v. 9, Buffalo Soc. Nat. Sci. Bull., v. 19, no. 3. 191 p., frontis., p. 33-66. map. 1948, [Discussion of Sand Draw fauna], In Colbert, E. Savage, D. E., 1951, Late Cenozoic vertebrates of the San H. [ed.], Pleistocene of the Great Plains: Geol. Soc. Amer Francisco Bay region: California Univ. Pub. Bull. Dept. ica Bull., v. 59, p. 622. Geol. Sci., v. 28, p. 215-314. McLaughlin, T. G., 1946, Geology and ground-water resources 1955, A survey of various late Cenozoic vertebrate of Grant, Haskell, and Stevens Counties, Kansas: Kansas faunas of the Panhandle of Texas, part II: Proboscidea: Geol. Survey Bull. 61, 221 p. California Univ. Pub. Geol. Sci., v. 31, p. 51-72, 2 pi. Meade, G. E., 1942, A new species of Capromeryx from the Say. Thomas. 1821, Descriptions of univalve shells of the Pleistocene of West Texas: Texas Archeol. Paleont. Soc. United States: Acad. Nat. Sci. Philadelphia Jour., v. 2, Bull., v. 14, p. 88-96. p. 149-179. 86 LATE CENOZOIC MOLLUSCAN FAUNAS FROM HIGH PLAINS
Schultz, C. B., and T. M. Stout, 1948, Pleistocene mammals Tihen, J. A., 1942, A colony of fossil neotenic Ambystoma and terraces in the Great Plains: Geol. Soc. America tigrlnum: Kansas Univ. Sci. Bull., v. 28, p. 189-198. Bull., v. 59, p. 553-587. 1954, A Kansas Pleistocene herpetofauna: Copeia, 1954, Shimek, Bohumil, 1913, The significance of Pleistocene mol- p. 217-221. lusks: Science, new ser., v. 37, p. 501-509. 1955, A new Pliocene species of Ambystoma,, with re Smith, C. L., 1954, Pleistocene fishes of the Berends fauna of marks on other fossil ambystomids: Michigan Univ. Mus. Beaver County, Oklahoma: Copeia, 1954, p. 282-289. Paleout. Contr., v. 12, p. 229-244, 1 pi. Troschel, F. H., 1856-1893, Das Gebiss der Schnecken: Berlin, 1958, Additional Pleistocene fishes from Kansas and 2 v. Oklahoma: Copeia, 1958, p. 176-180. Tryon, G. W., 1862, Notes on American fresh water shells, Smith, H. T. U., 1940, Geological studies in southwestern with descriptions of two new species: Acad. Nat. Sci. Kansas: Kansas Geol. Survey Bull. 34, 212 p., 34 pi. Philadelphia Proc., 1862, p. 451-452. Starrett, Andrew, 1956, Pleistocene mammals of the Berends Van der Schalie, H., 1953, Mollusks from an interglacial de fauna of Oklahoma: Jour. Paleont., v. 30, p. 1187-1192. posit (Sangamon? age) in Meade County, Kansas: Nauti Stimpson, William, 1863, Malacozoological notices, no. 1: Bos lus, v. 66, p. 80-90. ton Soc. Nat. Hist. Proc., v. 9, p. 249-252. Van Houten, F. B., 1944, Stratigraphy of the Willwood and Stokes, W. L., 1955, Another look at the Ice Age: Science, v. Tatman formations in northwestern Wyoming: Geol. Soc. 122, p. 815-821. America Bull., v. 55, p. 165-210, 7 pi. Taylor, D. W., 1954, Nonmarine mollusks from the upper Walker, Bryant, 1904, Notes on Eastern American Ancyli. Miocene Barstow formation, California: U. S. Geol. Sur Ill: Nautilus, v. 18, p. 75-83, pi. 5-6. vey Prof. Paper 254-C, p. 67-80, pi. 20. 1909, Notes on Planorbis, II; P. Mcarinatus: Nautilus, 1954a, A new Pleistocene fauna and new species of fos v. 23, p. 1-10, 21-32, pi. 1-3. sil snails from the High Plains: Michigan Univ. Mus. 1914, Notes on the Ancylidae of North Africa: Nautilus, Zool. Occ. Paper 557, 16 p. v. 27, p. 113-117, 124-131. 1925, New species of North American Ancylidae and Taylor, D. W., and C. W. Hibbard, 1955, A new Pleistocene Lancidae: Michigan Univ. Mus. Zool. Occ. Paper 165. 13 p. fauna from Harper County, Oklahoma: Oklahoma Geol. Wood, H. E., 2d, and others, 1941, Nomenclature and correla Survey Circular 37. 23 p. tion of the North American continental Tertiary: Geol. 1956, Pliocene mollusks from Jackson Hole, Grand Val Soc. America Bull., v. 52, p. 1-48, 1 pi. ley, and Star Valley, Wyoming and Idaho: Wyoming Yen, T. C., 1946, Eocene nonmarine gastropods from Hot Geol. Assoc. Guidebook 11, p. 123-125. Spring County, Wyoming: Jour. Paleont., v. 20, p. 495-500. Thenius, Erich, 1959, Wirbeltierfaunen: Handbuch der strati- 1951, Fossil fresh-water mollusks and ecological inter graphischen Geologie, v. 3, pt. 2. 328 p. pretations: Geol. Soc. America Bull., v. 62, p. 1375-1380. INDEX
[Italic numbers Indicate descriptions]
Page Acknowledgments. Armiger.--.-... _-.._.-._------_------..._...... ____ 58 Acroloxinae...... _. -_ - ...... 61 crista.------6,36,44,58 Acroloxus...... 61 armigera, Planorbula ------6,12,33,35,36,37,39,44,59 coloradensis...... ----- 6, 7,12,33,34,35,36,37,39, 61, pi. 1 Artiodactyla...... 22,23,28,33,40,42 adamsi, Blarina. -- 28 (Asinus) sp., Equus-.-.------_------....__ 16 Mammut...... ------. 28 Astian (late Pliocene) deposits.-.. .__ ....__ .....__.....___. 21 aenigma, Deroceras 3, 12,23,24,25,26, 27,29,30,31,32,33, 35, 36,37,39, 40,41,42, 80, pl.4 Atlantic Coastal Plain_------_- 7 aeruginosus, Planorbis...... 60 Atwater member. -_ -- -- _- - - - - 15 Promenetus-...... 45 avara, Sucdnea.------... .. ------_._ .._- 26,78 Age determination 9 Aves. .__ - - - - 23,28,37,40,42 albeola, Bucephala...... 28 albilabris, Pupoides. ... 3,6,12,23,24, 26, 27, 29,30,31,32,33,35,36,37,39,40,41, 74, pi. 1 B Albinula...... 72 Baiomys kolbi.------...... _...... ___. 28 (Albinula) armifera, Gastrocopta...... _ .. __ 7# rexroadi ------...... __..___------___..... 28 holzingeri, Gastrocopta...... ___ 72 sawrockensis.----... _._--._...... _..__...._...._...... 23 altissimus, Gyraulus...... ___ 58 Ballard formation.------__.....______...... 15 Ambystoma hibbardi,~-...... ___ 27 barbouri, Paenemarmota..------... - . ------____--._...__...... 28 tigrinum...... 37,40 Bar M local fauna._ ...... __ _ . .._.__...... 17 Ambystomatidae _. 22,23 basalts, Valvata tricarinata------.....--.--.-...... -..-,-...... -...... 48 americana, Fulica...... __ 28 Basommatophora.... - .. - ... 22,23,25,27,31,33,36, 40,51 Amnicola...... 50 Bassariscus casei... - _- _ - _- _-_- - __ ...... 28 crybetes...... __.. 38,50 rexroadensis .------_. 28 lacustris...... __ 50,51 baumgartneri, Peromyscus--. ------___...... 28 limosa...... ___ 50 bettus, Dasypus------.._...... 17 lustrica...... 45,50,51 Bender local fauna____ .___ __.___. __.______11, 14,31 decepta...... 51 Bensonomys eliasi ...... _.__....._------___...... 28 perlustrica. __ 51 meadensis-.--.----...... ------...... _..- 40 walkeri...... -. ... ___-______._ 45,49.50 Berends local fauna._. . _--,---._------____--._____... 17 (Marstonia) lustrica ...... 51 bermudensis, Vertigo-...... ------... 76 Amnicolinae____ - ______49,50 bicalcaratus, Playgonus. ------... ..-_-_...__------_.. 28 Amphibia...____.__...... __...._.__.__ 22,23,27,37,40 bilamellata, Gastrocopta-.--.---.... _------_------___...... 73 Anasbunkeri...... _____ 28 bilobatus, Cnemidophorus ------.... -_...... ------.._. 28 anatina, Physa-.... 3,6,12,23,24,25, 26,27,29,30,31,32,33,35,36,37, 39,40, 41,44, 68, 64 Biorbia fossilia...... -._------_------_____ ------18 Physella...... -...... _.. 62 Bir ds.------.-. .-.-.-.-...... - . -.-..-...- .-.-- . _ 42 anceps, Helisoma ...... 3,6,12,23,24,27,29,33,35,36,39, 44,58, 59 Bison latifrons------... ------...... 17,19 Anchylorana dubita. _____ 27 Blancan age- _------__-...-.-.------... 18,21 moorei .. 27 blancoensis, Promenetus ------... -...--.-.-.-...__-___....___ 38, 45,60 robustocondyla..-. ------...___ 27 Tanupolama------...... ^...... ___---.-.-.---.-..._-..... 28 Ancylidae_.... . __ 61 blandi, Pupilla ...... -______-_____...__.__ .... 16,17, 18,35 Ancylus fluviatilis...... ___ 61 Blarina... ------_..---_.----.---_-.-...... 39 hendersoni...... _____ 61 adamsi ----- ...... ------...... 28 pumilus-....-...... _____ 61,62 sj?...... 37 Angell member of the Ballard formation.._.______3,15 Borchers local fauna.. ..-.-.-...-.-.-...... ___...... _. 16 Angustula...... 45,76 borealis, Synaptomys (Mictomys)..... ___------_-_------_...... _. 16 (Angustula) hibbardi, Vertigo...... __ ....______45 Borophagus.------_------_- 21 Anisus...__ ...... ___ 56 diversidens------.... . __.__._-___-_-_ ...... 28 pattersoni...... 7,12,33,35, 36,39, 58, 57 brachyodontus, Odocoileus.------... ------_ __...... _... 28 perezi______-.-.__.______57 Brachyopsigale dubius.------... -.-.-.-...-.-.-...-.-.---.-.....-.... 28 rotundatus- _--- _.. -.--.-.--.-.-..-..-..--.-...-...... -..... 56,57 breviramus, Buisnictis.... ------_- .__-.-.-.-..__.-.._------... 28 Anodonta.----- _ __ ...... _... 45 Bucephala albeola------...... 28 gigantea_------._-.__...... 45 Buckshot Arroyo..------_-.-..-.---.------_--_------_----- 22 ffraniiw------24,33,35,36,45 Bufosp ,------...... ---...... 27 (Pyganodon) grandis...-- ...... ______46 Buis Ranch local fauna - , 1, 14,22 Anodontinae. ___.------_ 45 buisi, Pliogeomys_.---.-.-.-.-...-.-.-.-.-.-.-.-.-.-.....-.---.-.-...-.-.-.. 22 Anseriformes ------___.__-.___-,---_-.. 28 Buisnictis breviramus------.... ------.--.-.-.-.----.------28 Antilocapridae__- __ __ --_ _ ___- -_ __...... 28 22,23 antiquus, Pliolemmus.-...... -...... -...-.... __...______17,37,40,42 Buliminae------_.._------_------__...._.___.__...... 49 antrosa, Helisoma..------... ------_--.....-.... 58 Bulimnea... ------...... -..._.._ 56 40,56 Aplexa------__.----_--__--_-_----_--.-._--.___.. 64 megasoma...... -____....__------_----- 6,7,12,33,34,35,36,39, bulimoides techella, Galba.------... -----.----.-----.--. 54 hypnorum- ...... 6,12,13,24,32,33,35,36,40,41,44,53,54,64 techella, Stagnicola-...... 3,6,12,13,16,23,24,24,25,26,27,29,30,31,32,40,41. 45,54 arboreus, Zonitoides ------6,12,24,27,29,31,36,37,39, 44, 80, 81 Stagnicola (Nasonia)------... ------54 Arctodus simus-...... _...... 17 bunkeri, Anas.... __ - _- . - _..-.-.._------28 armifera, Gastrocopta...... _...._____- 6,12,24,26,36,37,39,44,72 Buteo sp- _-.. _ __._ -_ ._ _- 28 Gastrocopta (Albinula).... ----_------._---_.___.-__------_ 72 Butler Spring local fauna.__ . - .-_-_ _.___...... __-...... 16 87 88 INDEX
Page Page cyclophorella, Vallonia...... 16 Caliche, origin of-.. ------_- 3 Cynomys meadensis. ... 42 Camel, giant------. _ __ 22 Camelidae______.---_- .-._____ 40,42 D Cametops------17,33 sp. 33 dalli, Fossaria 3, 6,12,23,24,27,29,30,31,32,33,35,36,39,40,41,44,45,54,56,79, pi. 2 Canis latrans-.--...... _-_ ------33 Galba ...... 54 lepophagus...... - - 28 », Fossaria...... 53,55 sp_ 28,42 _...__-__ _ 17 caperata, Stagnicola-..... 6,12,13,18,22,24,27,29,31,32,33,35,36,39,40,41,43,44, 53, 54 decepta, Amnicola lustrka 51 Stagnicola (Hinkleyia)-...... - . _ ....__ 53 Matt-onto-.. ------3,6,12,23,24,50, 61, pi.2 Capromeryx sp__-___-_------_--- - . .... 33 Deer Park local fauna... 16,4/ Carnivore.. - - 22,23,28,33,37,40,42 deflectus, Gyraulus...... 6,7,36,44,57 carus, Promenetus------_ - __ _._. 60 densecostata, Strobilops...... 65 Carychiidae-.--__ ...... _ 51 Depositional and biological sequence, summary of. 18 Carychium... __-__-__-_-_------...... 51,74 Deroceras...... 25,80 exiguum-...... 2,6,12,24,26,27,29,30,36,39,44,45,51,52 aenigma. .. 3,12,23,24,25,26,27,29,30,31,32,33,35,36,37,39,40,41,42,80, pi. 4 perexiguum------... ------...... 45,51,52 laeve ... 24,36,44 sp- .. . 33,35,36,52 diaboli, Vertigo ovata...... 76 casei, Bassariscus.--.------.------.------.---.--.-.------...... ---.- 28 diminuta, Lymnaea...... 38,45,53,54 casertanum, Pisidium... 6,12,22,23,24,25,26,27,29,30,33,35,36,39,44,47 Dipoides wilsoni...... 23 Castoridae______------_ ...--..-.-.-_____...... ____ 37 Discus-. . - 79 Castoroides -. - .. .. __.______17 cronkhitei.... . 6,18,36,44,79 Caudata...- ...... 22,23,27,37,40 macclintocki...... -. 8 centralis, Liomys...... _.__-....______... 28 diversidens, Borophagus.. 28 Ceratophyttum...... 48,57 diversus, Scaphiopus...... 27 Chaenaxis...... ------...... ______.. 74 Dixon local fauna...... 1,11,17, S6 Charadriiformes ______.______28 dixonensis, Sorex...... 37 chauliodonta, Gastrocopta-...... 11,12,16,17,24,32,33,34,35,36,37,39,40,41, 70, pi. 1 dotti, Ctiettus ...... 22 Chelonla...... 22,23,28,33,42 dubita, Anchylorana...... 27 Chiroptera..------...... -.... ------______-___...__. 28 dubius, Brachyopsigale...... 28 chisosensis, Polygyra...... -____..______82 E Ciconiiformes______.-.______-______.__ 28 Edentata ..._____ 28 Cimarron River.--.______.___ 3 elecfrina, Nesovttrea ...... 6,12,24,27,29,30,36,37,39,80 Cimarronomys stirtoni ...... ______.._._... 23 Ntsovitrea (Perpolita)...... SO Cincinnatia integra...... -...-..__.______.__...... _ 50 eliasi, Bensonomys.... 28 peracuta...... ______...... 50 elliptica, Physa...... 63 cinereus, Sorex...... _____.______,_ 18 Elodea ______~_____ 48,57 Cionella...... ______77 Emmericiinae ___.. 49 lubrica...... 6,7,12,24,27,36,44,77 enaulus, Gyraulus ...... -...... 38,45,58 Cionellidae.______.______77 Endodontinae.. 79 circumlineatus, Planorbis ...... ______..______45 ephippium, Rana...... 27 Promenetus...... --_.______...______60 Equus (Asinus) sp 19 circumsfriatus, Gyraulus...... __._____.______6,36,44,57 so.. ______-__--____---______--____-- 16 Citellus dotti...... 22 Erymodon.------...... -. - 82 howelli. ------._-- . 28 Esox masguinongy...... -...... -- - 17 reiroadensis...... _____...______* Euamnicola...... 50 sp...... 37,42 Euconulinae_ ... .. 80 Cnemidophorus bilobatm...... ______28 Euconulus -. --- . ______. ___ 80 cochisensis, Gastrocopta_....____..______73 fultms...... 6,36,44,80 cochrani, Pliauchenia...... ______....____.______28 Eumeces striatulafus---- _ _ . ------__ 23,28 cockerelli, Stagnicola...... _...._.____....__.____...... 24 Eumecoides hibbardi. ------_____-__-----__------_._------_. 28 Colinus hibbardi______. ______28 mylocoelus------... -----. ------28 Collecting methods_____.______2 exacuous, Menetus------...... _------_------_------59 coloradensis, Acroloxus... ______....______6, 7,12,33,34,35,36,37,39, 61, pi. 1 exccuous, Promenetus--- .... - -. -----_-- -.- .------59 Cclimbiformes------_---..-...._-______...... 28 Promenetut ------6,7,17,24,36,44,59 Comparison of faunas__...__.-._____... _ ___....._...... 11 exiguum, Carychium...... 2, 6, 12, 24, 26, 27, 29, 30, 36, 39, 44, 45, 61, 52 compressum, Pisidium.-...... ______.__ 6,12,26,33,35,36,39,46,47 exilis, Galba.. -.. .-...... ^ . 52 concordialis, Succinea...... ______...... ____..._...... ___...... 24,39,77 Stagnicola...... 6,12,24,27,29,30,45.5^ contracta, Gastrocopta-...... ____....______39 cornutum, Phrynosoma...... ______._..._____. ______17,28 Correlation of North American ages with Lyellian epochs______18 Falconiformes------__ _ -... _ _ - --- 28 costata, VaZ.oiM'a.______-___-_-___.___------_-_-_--_-_----.-______26 29 Faunal dating...... _ _ ...... ------9 Cragin Quarry local fauna______17 Faunal interpretation, methods of. ____---__._----_ ------8 crassum, Sphaerium.-...... _____.._-______46 Faunal sequence, summary of.-..------14 crawfordi, Notiosorex...... ______.______17 Faunas, comparison of. ______.______.__._.-___ 11 Cricetldaeindet.______22 fayeae, Rana------... ------.....~. .------27 crista, Armiger...... -.. -..---______6,36,44,58 ------28 Gyraulus...... 58 sp.------28 cristate, Gastrocopta----. 2, 6,12,13,17,24,26,31,32,33,35,36,37,39, 40,41,44,67, 69, pi. 1 Ferrissia.------... ------__.. . ------61 cronkhitei, Discus-.-...... 6,18,36, 44, 79 meekiana. -...-...... --...... -----...-. 6,12,33,35,36,37,39, 61, 62 Crooked Creek formation.....______15; 19 parallela.. 6,12, 24,27,29, 61 crybetes, Amnicola...... ______38,50 pvmila...-.------.------.- .----.------62 Marstonia...... 3,12,23, 24, 27, 29,30,36,39, 50, 51, pi. 2 rivularis------3,6,12,17, 22, 23, 24, 25,26, 27, 29,30,33,35, 36,37,39, 61 Ferrissiinae-.------61 Cryptotis kansasensis...... _...____.__ 37 ferrugineum, Pisidium---.------... ------17 meadensis...... _.__.__... 28 flavescens, Perca.------... - - - - - 16,17 Cudahy fauna______llt 15 fluviatilts, Ancylus.------... - __ ------61 curacoana, Gastrocopta...... ______...... 71 Fontigens...-...... --.-...-...... --..-...... ------51 Cyclic climatic changes______.______.__ 10 fontinalis, Physa.------. . ------_.. . ..-- 64 INDEX 89
Page Page Fossaria.... ----.. ..------.-..-. 8,39,54,74 Qyraulus C ontinued dalli-...... 3,6,12, 23, 24, 27,29,30,31,32,33,35,36,39, 40, 41, 44, 45, 64, 56,79, pi. 2 pattersoni... ______...______56 grandis------,------.-.------. ------53,55 vmMlicatellus.... __..__-______._-_-__.___.-.______60 obnssa.---.-... ---.------.------.---- 6,12,24,26,33,35,36,44,54 gyrina, Physa. ______6,12,26,33,35,36,37,39,43,44,63,64 parva. ______55 hildrethiana, Physella.------______-.-______63 sterkii ...... -. ..-...... -...... 55 tazeweltiana ...... _____ ... ___ ... _ ., ______55 H perplexa..... ------...... ______55 Fossil mollusks, environmental interpretations. . ___ . __ _.-___.___-_ 3 Habitat and distributional data, inadequacy of..-----_-______7 fossilia, Biorbia...... 18 Hawaiia -____._.______.______81 fossilis, Lasmrus ...... 28 minuscula- ______3, 6,12, 23, 24, 25, 26,27, 29,30,31,32,33,35,36,37,39,40,41,44, 81 Onychomys. ------... -___ _ ...- _ -- _-_-____--_-_---_-_--.__. 16 haydeni, Oxyloma... ______78,79 foxi, Martes. -----.------.------.-..-...- ...... _-.--_--_--_-_ _ -- 28 Hebetodiscus...... ______79 franzenae, Gastrocopta...... 3, 12, 23, 24, 25, 26, 27, 29, 30, 31, 32, 45, 67, pi. 1 Helicodiscinae______-____ _-__-_-______79 fraternum, Stenotrema ...... ______...... _.._. _ .. 39 Hehcodiscus------______79 Fulica amencana ______.._.._.__ _ _.__._._..______... ______. 28 (Hebetodiscus) singleyanvs...... ______79 fulvus, Eueonulus ____ .______. ____ ._ 6, 36, 44. SO parallelus------..--. ...------6,12,24,26,37,39,41,79 singleyanus. ______3, 6,12,13,17, 23,24, 27, 29,30,31,32,33,35,36,37,39,40,41, 79 helicoidea, Valvata lewisi--.... ______49 Galba bulimoides techella. ------.------.--- 54 Helisoma.------______-.______68 dalli-. .-. -______-_-__-______-____-____-__-_.___-______54 o.wx'pn ------3,6,12,23,24,27,29,33,35,36,39,44,55,59 exilis...... _-____- ___.______-____-______-_____-___-___ 52 antrosa______58 obrussa.- ______5. (Pierosoma) Mvolvis.... ______59 palustris- -____-_____-_-______-__-__-___-___-_..-__ 52 trtoolois.-- - - .. ..- . 6,12,24,26,36,39,44,59 reflexa. ______53 Helisomatinae______---.____ 58 Galliformes______.______. 28 Hemp hillian age.______18 gallopavo, Meleagris... ______..____ 28 hendersoni, Ancylus,... ______-_--__-__-______---____.-_____--______61 Gastrocopta.------_-____-.______.______67 Hendersonia occulta..... ______-_--__-_____--_-_--_--______-____-____ 8 armijera.------6,12,24,26,36,37,39,44,7.? Hesperoscalops rexroadi..------_-_--______-__---..______-_--__-___ 28 bilamellata ______..______. 73 sp__------______--______-_-______-______--_____-.______-______22 chauliodonta. ______11,12,16,17,24,32, 33,34,35,36,37,39,40,41, 70, pi. 1 Heterodon plionasicus..... --_-__-_____-______-_____-____-__--____-______28 cochisensis-... ______.__-______--_-____-----_-----_____ 73 hibbardi, Ambystoma. ______27 (Albinula) armifera. ______7% Colinus---..--...... ------.-. ______28 holzingeri----.----. --- _---_-__-___--___--_-__-_.___.__----__---_ 73 Eumecoides______-----_____-__-___---__,-______--______. 28 contracta...... _.______39 Vertigo...... 3,11,12,22,23,24, 27,29,30,32,38,45, 74,75, 76, pi. 1 cristata.... -- 2, 6,12,13,17, 24, 26,31,32,33,35,36,37,39,40, 41,44, 67, 69, pi. 1 hildrethiana, Physella gyrina... ______...______._ 63 curacoana... ______71 hilli, Sigmodon ______16 Jranzenae..- --- ______3,12, 23, 24, 25, 26, 27,29,30,31,32,45, 67, pi. ] Hinkleyia... ______53 holzingeri------2,3,6,12,23,24,27,29,30,36,44,73 (Hinkleyia) caperata, Stagnicola___.--_.__._-_-_____.--__-____----___ 53 (Immersidens) rexroadensis-.-- ______73, pi. 1 holzingeri, Gastrocopta ______2,3,6,12,23,24,27,29,30,36,44,7.? lavernensis ------___._-___--__-_-______71 Gastrocopta (Albinula)-- ______.-.- 72 oligobasodon..------_._--_-______-.__.-__--.___._____ 73 hordacea, Physa...... ______--.- 64,pi.3 paracristata- ______3,12,16,17, 23, 24, 25, 26,27, 29,30,31,32,38, 40, 41, 9, pi. 1 hordaceus, Pupoides ______-_____-_-______._-_-_.______--._ 18 pellucida hordeacella... .. 6, 7,12,13,17,18, 24, 25, 26, 27,29,30,37.39, 71, pi. 1 hordeacella, Gastrocopta pellucida.... 6,7,12,13,17,18,24,25,26,27,29,30,37,39, 71, pi. 1 procera. ______2, 6,13, 24, 26,37,39, 67, pi. 1 howelli, Citellus------.------.------28 prototypus-.-- ______.______73 hvmilis modicella, Lymnaea....-- _---_-_--_____-_-_-_-__----.____----_---- 55 rexroadensis------. 3,11,12,17, 23, 24, 25, 26, 27, 29, 38,73,pi. 1 Hydrobia steinii... ______._-_-_.______-_--.--- 50 scaevoscala..------.-. ______12, 24,31,32,37,39,45, 70, pi. 1 Hydrobiidae..--.______5,36,39,49,50 tappaniana. ______3,6,12, 23, 24, 27, 29,30,31,32,33,35, 36,37,39,40, 41,44, 7S, 74 Hydrobiinae....______49,50 Gastrocoptinae.---._-_--_-____-_____.-_-____.___--__.______67 hypnorum, Aplexa. ______6,12,13,24,32,33,35,36,40,41,44,53,54,64 Gastrodontinae______._-__.______81 Hypolagus regalis------______.___._--- 28 Gastropoda...... ______22,23,25,27,31,33,36,40,42,48 Hypolagus sp____.___---_____-----_---_-_.___-_-______42 gelida, Succinea grosvenori------__---______---__.--_____--______-__ 78 Geomysguinni______17,28,33,42 tobinensis. ______.----______40 Ictalurus punctatus. ._ .- ______- __.____._-______. ______16 sp. ______37 sp. .______- _-_-______,_ _-__-_ -_._ 33 Geyer, quoted_____-____--__.______..______5 Immersidens...... -______--.-.______._ 70,73 gidleyi, Onychomys.--- ____.______...______28 (Immersidens) rexroadensis, Gastrocopta...... _._-_____.__ 73,pi. 1 gigantea, Anodonta.. ______45 indentata, Petinetta... ______.--.______._-_____.-._.. 39 Gigantocamelusspatulus--.- ______.______23.28 infracarinata, Valvata _ncara?a.a______..__..___ 48 sp______33 inornatus, Pupoides... ______6,12, 24,25,26,27,29,30,33,35,36,74, pi. 1 Glyphyalops------.... ______81 Pupoides (Ischnopupoides).. __._.______74,pi. 1 OJyphyalus.-.-.--.....--... .-...-- .--.------..-.-..... ______81 Inseetivora.______22,23,28,37,40 (Glyphyalvs) wheatleyi, Betjnella....-...... 81 Integra, Cindnnatia.. ______50 gouldi, Vertigo...... ______.______16.35 intermedius, Sigmodon------______...______28, 40 gracilicosta, Vallonia. ______.__6,12,13, 24,27, 29, 30,36,44, 77 International Geological Congress__-_--_-_-_-.-______.__ 19 grandis, Anodonta... ______24.33,35,36,45 Irvingtonian age______.____.______..__._. 19,21 Anodonta (Pyganodon)-... -__-_____-______----___.--__--______-_____-_____ 45 Fossaria dalli----.--.--- _. ______53,55 Ischnopupoides____-----______------__.__ 74 granum, Lyogyrus.--- .____-____---____-_--____-__.______49 (Ischnopupoides) inornatus, Pupoides______74, pi. 1 grosvenori, Succinea...... 78 gelida, Succinea. ______78 J Gruiformes_-_-______28 jacksoni, Notiosorex. --__-_-_----_.______-_-----_-._-__ 28 Gundlachia meekiana. ______61, 62 jayense, Musculium______.______46 Gyraulus...... ______57 jennessi, Physa...... _.______._._-_.______64 altissimus... ______58 circumstriatus-- ______6,36,44,57 Jinglebob local fauna.______16 crista- ______58 Jones local fauna.--___-______---. 17 deflectus. ______6,7,36,44,57 Jones Sink______.-._____._. 17 parvus.... ______3, 6,12, 23, 24, 25, 26, 27, 29,33,35,36,39,40,41,44,45, 68, pi. 4 jugularis, Lymnaea stagnalis ______16,17, 56 90 INDEX
Page Page K mcgrewi, Nebraskomys...... 33 kansasensis, Cryptotis- . _---....___------______37 Machairodus sp 28 Menetus...... ____.---.______38,45,59 mclaughlini, Perognathus...... 22,23 Peromyscus... ___.--.___--__-_---_--.__...... _.__._.__...... 28 Mammalia... . 22,23,25,28,33,37,40,42 Pratilepus...... _____.______._---_.___.___...... _.... 28 Mammut adamsi...... 28 Promenetus . 3,12,17,18, 23, 24,25,26,27,29,30,33,35,36,37,39,40,41, 59, pi. 3 marginatus, Pupoides...... 74 Trigonictis______...__...___.______.______28,33 Marmota sp.... 23 Kincaidilla.-.... - -. ------....-...... --.-.--.-----_..-.. 62 Marstonia...... ------50 Kinfsdown silt.-_.--.-._.______..__--..._._-.__.__...___.... 15 crybetes...... - 3,12,23,24,27,29,30,36,39,50,51, pi. 2 kirldandi, Laevapex.------... ___._._....._...... 17 decepla...... 3, 6,12,23, 24, 50, 51, pi. 2 kolbi, Baiomys.------_------_-___---______...... ___.... 28 Marstoniopsis...... 50 Martesfoxi .. -...... -...... 28 masquinongy, Esox...... 17 labyrinthica, Strobilops- . 3, 6, 12, 17, 23, 24, 65, pi. 1 Mastodont.-... 40 lacustre, Sphaerium ______.... ______... __ 6,12,36,39,44,46 meadensis, Bensonomys...... 40 lacustris, Amnicola ...... _ ._. _ __ - ____ . __ ... __ .. _ . _ 50, 51 Cryptotis...... 28 Felis ...... --- . ______.. 28 Cynomys...... -- 42 Neosorex ...... __ .__-_. ___ - ______. ______. ______16 Pitymys...... 16 Patella . ____ . ______. ______61 Pliophenacomys-...... 17.37,40 Probythinella ...... __ ... _ ...... ___ ..... ______. _ _ ._. 16 Pliopotamys...... -...... 16,17,33,34,37,42 Laevapex kirklandi...... ------...... __ __ . ___ .. _ ..... 17 Rana...... -...... 27 laeve, Deroceras ...... - _ ... _ .... _ ...... _ ...... _ ... __ .___.. 24, 36,44 mediocarinata, Valvata tricarinata...... 48 Lagomorpha ___ - ______- ______. __ ... ______22, 28,42 meekiana, Ferrissia...... 6,12.33,35,36,37, 39, 61, 62 Lampsilinae _ ... _ _ ------_ ...... __ --.--...- ___ .. _ _ ..... 45 Oundlachia...... 61,62 Lampsilis subrostrata ...... -----.-.-... _ _ ...... ______-.---.... 45 Megalonyx, leptostomus...... 28 landesi, Synaptomys __ ... _ ...... _ __ ... _ ...... _ ... _ _ ------16 megasoma, Bulimnea.---...... 6, 7,12,33,34,35,36, 39, 56 lanei, Procastoroides ... ------__ ...... __ ..... __ _. __ .._____ 28 Meleagrididae indet-- _. 28 larrabeei, Onychomys ... ------___ _ _ ... __ ._._. __ _ . _ .._____ 23 Meleagris gallo,pavo...... -- ---. ---.-...... ---...... 28 Lasiurus fossilis .... ._.. __ . _ _. __ ...... _ ..... _ . ______..... 28 Menetus exacuous...... 59 latifrons, Pison ...-. . ..--.------...... _ _ ...... _ ... _ _ _ 17, 19 kansasensis...- ...... 38,45, 59 latrans, Canis ... . __ ..----.--.-.-...... _ .... _ .... _ _ .---..-.-...._. 33 umbilicatellus---- 60 lavernensis, Oastiocopta ...... __ _ ...... _ ... _ ...... _ .. ______71 Mephitis rexroadensis-...... 28 leahyi, Sorex ------__------.---.____._ _ .--.---.-...... ______37 Mesodon...... ------...... 31 leal, Stenotrema ..... -. _ __.--___ _ _ .______...___. __ 6,36, 44,83 Mesogastropoda.. -. 23,27,33, 36,48 leai, Stenotrema.-...... _ __ _ ...... __ . ______...... _ __ ..... 83 Methods.--._ _ - __--__ . ______2 leonardi, Neofiber...... --.-...... _ --.-...... _ ._ _ ...... 18 Microsorex pratensis....------... ..______---_-- 16 lepophagus, Canis ...... -_-.. _ ... _ ._.. ______..... ___ ... _ 28 Microstele...... - -- . ------74 Leporldae indet...... ______..... _ ... _ ... _____ .______.... 22 Microtines_____----_-_-___-_- __.______-__--______-_-_--- -_._. 39 leptostomus, Megalonyx...... ___ .... _ __ _ . ______28 Microt us paroperarius...------16 lepmculus, Notolagus...... ______. ______28 pennsylvanicus.... ______.-_-_---_------_ ------_ -- 17,18 lewisi, ralvata...... 6, 12, 13, 33, 34, 35, 36, 48 (Mictomys) borealis, Synaptomys..---- ... _.__--______.. 16 helicoidea, Valvata _____ ... _ ...._._ _ _ ..._._ __ .. ______...... 49 milium, Vertigo. ______,. 2,3,6,12,23, 24, 25, 26, 27, 29, 30,31,32,33,35,36,37,39,40, 41,76 Ligumia....-- ...... _ _ _ _ .. __ . _ ... 45 minor, Nerterogeomys.------28 subrostrata- ...... __ . ______.. __ 24,27,29, 30,45 minuscula, Hawaiia.... 3, 6,12, 23, 24, 25, 26, 27,29,30, 31,32,33, 35,36,37,39, 40,41, 44,81 lilljeborgi, Pisidium...... __ ...... _. _ .... ___ _ _ ...... 17 minutissimum, Punrtum... ______--_------______--_----_ _.-___.-__- 36, 44, 79 Limacidae ______..... _ __ ._._ _ . ______...... 80 mirificus, Stegomastodon...... ------16 Limiting factors, recognition of...._ __ _ _.._._... __ .. _ . _ ...... 5 Missler member.____--_-______-__-__-_____---_------_ 15 limosa, Amnicola----.-.. ______.___-_ _ . _ ...... ___ .__ ___ ... __ 50 modesta, Vertigo....------16,35 lineatum, Sphaerium ...... ______... _ _ _ ... _ _. _ _ 45 modestum, Phrynosoma. ______....------17 Liomys centralis ...... _. ....______.... _ . ______...... 28 modicella, Lymnaea humilis------__-____------_---- 55 Literature cited __ ._--_-___-___-______. __ ._. _ _ _ _ ... ______83 monodon, Stenotrema...------...... -- 83 llanensis, Pedomys.---.--.- ... _ ------_ _ _ _. __ __ _ ...... __ _..______16 mooreana, Polygyra.. ______...... 38,82 17,18 tholus, Polygyra, ____ ------____._-____-_---- 82 Llano Estacado.. ______.. ______26 moorei, Anchylorana,------27 caliche _ -.__--_-_-_-_-______._-__-_-______. __ ...... 3 muscorum, PupUla... _._ ------... --______..... 7,16, 18,35 Local faunas ______...... ___ .. _ ... 10 Musculiumjayense...... ______.... --. .. -______---_ ___ 46 description of...... __ _--______----______..- ______22 partumeium------...._-_.------45 lubrica, Cionetta...... ______6,7, 12, 24,27,36,,44,77 rosaceum..... ______-_------_------_-___--- -_-- 46 lustrica, Amnicola ______.._...______45,,50, 51 ryckholti.------46 decepta, Amnicola .-. __ __._____. _ ___-__--_.____._. _ _ ... ______51 securis------46 Paludina...... _ --._.____-___ _ .... ______. ___ ._.__-___ 49, 50 transversum....------47 perlustrica, Amnicola.. .__...._.__.-_.______.__..-.__--____ 51 truncatum.------45 luteola, Siiccinea. ------_-___-____.______.____.______73 Mustela rexroadensis------______-_------_ ------28 Lutra piscinaria ______28 sp. 28,40 Liittig quoted..... ______. _ _ _ 20 Mustelidae _--___ ------37,42 Lyell, quoted.. ______9 myJocoelus, Eumecoides.-.... 28 Lyellian epochs, correlation of North American ages with ______. 18 Myriophyllum...... 39 Lymnaea diminuta ______. ______38, 45,, 53, 54 humilis modicella .... ______. 55 macella...... 38,45,,52,53 Nannippus. ------17, 18, 21 palustris ------...... _ _. 52 phlfgon -. 16,28, 40,42 parezilis...... 38,45, 52,53 SD *** *" stagnalis jugularis ...... ______._ __ . 16, 17, 56 Nasonia. -.- .... --_____-___.__- _ 54 turritella...... 38, 45, 55, 56 (Nasonia) bulimoides techella, Stagnicola. 54 Lymnaeidae ______52 Nebraskomys mcgrewi------.. 33 Lyogyrus granum...... ___._ 49 Nekrolagusprogressus.------2S M Neofiber leonardi------18 macclintocki, Discus ..... ______.__.._. 8 Neoscaphiopus noblei ... _._------____ ._ 27 macella, Lymnaea ...... ______. 38,45,52, 53 Neosorex lacustris------INDEX 91
Page Page Nerterogeomys minor.---...--,...-...... 28 Peromyscus baumgartneri. 28 Nesotiitrea_____...____._.___...____...______80 kansasensis...... 28 electrina...... 6,12,24,27,29,30,36,37,39,, 44, 80 progressus-.-.-...... 17 sp. 23,80 SP---.- .... - - . - -- . - ...... 37 nitidum, Pisidium._...... 6,12,33,35,36, ,^5 perplexa, Fossaria__...... -...... 55 noblei, Neoscaphiopus,__.. ,______27 Perpolita...... _____...... Notiosorex crawfordi...... ______17 (Perpolita) electrina, Nesovitrea...... jacksoni__...._...___...... 28 perspectiva, Vallonia...... 3,12,13,17,23, 24,26,27,29,30,37,39,40, 41,77 Notolagus lepusculus...... _____ 28 phlegon, Nanmppus...... 16,28: 40,42 numellata, Vertigo...... 76 Phreatomenetus...... ------.... 45,60 (Phreatomenetus) umbilicatellus, Promenetus..--...._...... 60 O Phrynosoma cornutum...... 17,28 obrussa, Fossaria-...... 6,12,24,26,33,35,36,44,54 modestum ....--..-..-----..-----..----.--..----..--.-----.---.-..-..... 17 obtusale, Pisidium...... 17 occidentale, Sphaerium...... ___...__ 53 anatina. ____..._ 3,6,12,23,24,25, 26, 27,29,30,31,32,33,35,36,37,39,40,41,44, 62, 64 occulta, Hendersonia.___...... ______8 ettiptica...... 63 Odocoileus brachyodontus...-...... _. 28 fontinalis...... 64 Ogallala formation_____...... 3,15,22 gyrina...... -... - 6,12,26,33,35,36,37,39,43,44,63,64 Ogmodontomys poaphagus...... _. 28 hordacea...... 64,pl.3 sawrockensis...... _____...______23 _ __ - - - 64 sp...... ___- ______42 skinneri...... 6,8,12,13.16,17,24,32,33,34,35,36,37,39, 40,41, 63, pi. 3 oligobasodon, Qastrocopta...... 73 SP__ ------._-- - -__-_ -_._.__- .--.. 62 Onychomys fossilis...... - .--______.______16 Physella anatina..__.._____.__...... 62 ..______,______28 gyrina hildrethiana...... -_ . - . 63 larrabeei _.._._...... __...... ______.. 23 Physidae.____...._...... 62 Oryzomys___..__...... 17 Piaeenzian deposits____ . .-_ . _ . 21 Osteiehthyes.. ______-_____. ______23,33 Pierosoma...... _ _ .. .. 59 Osteoborus progressus.______...... 23 (Pierosoma) trivolvis, Helisoma. - ...... 59 ovalis, Succinea_...... __.__.____._____ 36,44,77,75 piscinaria, Lutra.______...... 28 ovata diaboli, Vertigo...... 76 Pisidium...... ______. 47 Vertigo...... 6,11,12,22.24,26,32,33,35,36,37,40,41,44,74,76 casertanum...... 6, 12, 22, 23, 24, 25, 26, 27, 29,30,33,35, 36, 39, 44, 47 Oiyloma._..__...... 78 compressum...... 6,12,26,33,35,36,39,46.4? haydeni...... 78,79 ferrugineum ...... -...... ------...... 17 retusa...... 6,12,24,36,37,39,44,77, 78, 79 Wljeborgi-... . - - --- .-- -. 17 siUimani...... 78 nttidum....------.--...... 6,12,33,35,36,39,48 ------17 Paenemarmota barbouri...... ___ __.___. 28 svbtruncatum. ------17 Paladora Creek______....______.______26 walkeri...... --...... 17 Paleoeeologie interpretation, principles of.___...__.______, 4 sp__ ...... 47.48 Palo Duro Creek. ______.______26 Pity mys meadensis-.-----.- .... __-_ ------_ -___- _--- . 16 Paludina lustrica...... ______. ______. 49, 50 planatum, Sphaerium simile_.. ----- 46 palustris, Oalba...... _. 52 Planorbidae------_ ___ _.------_ _------8,56 Stagnicola.___...... 6,36,39,43,44,52,54 Planorbinae.--___ ._ - - 56 paracristata, Oastrocopta___... 3,12,16,17,23,24,25,26,27,29,30,31,32,38,40,41, 69, pi. 1 Planorbis aeruginosus...... ------..-. ------60 Paracryptotis rex._...... _.______28 circumlineatvs.... ------. .. 45 Paradipoides stovatti...... _...... _. 17 santacruzensis...... ___.__ _------_------_ _-- 45,60 Parahodomys guadriplicatus...... _...... _. 28 Planorbula...... - 59 parallela, Ferrissia...... ______6,12,24,27,29,61 ..____ .__ __ - .- - 6,12,33,35,36,37,39,44,59 parallelus, Helicodiscus...... 6,12,24,26,37,39,41,79 sp- ____ _ - - 59 parexilis, Lymnaea______38,45,52,53 Planorbulinae.. - - paroperarius, Microtus---...... ______16 tygonus...... partumeium, Musculium____...... 45 bicalcaratus.... Sphaerium. .-. -- .- - . __.-____._____ 6,12,33,35,36,39,45 sp.-- - 33,42 parva, Fossaria...... ____.__._____. 55 Plesippus...... -. - . 17,18,21 sterkii, Fossaria...... _. 55 simplicidens .------16,28,33,40,42 tazewettiana, Fossaria...... _ 55 sp. ------______-_- .-____-_ _-_____---____- 25 parvissima, Rana...... _.__ 27 Pleuroeeridae_____.-- ...... ------.... ----_-..... 8 parvula, Vattonia...... _. 6,12,24,26,31,32,36,44,77 Pliaucheniacochrani.------...------28 parvus, Gyraulus...... 3,6,12,23,24,25, 26, 27, 29,33,35,36,39,40,41,44, 45, 68, pi. 4 Pliocene-Pleistocene boundary.... _--...... - 19 Passerifonres_--_------____....____...____.-.._____ 28 Pliogeomys buisi...... ------... ------22 Patella lacustris.______...... __.__.___....___ 61 sp...... -...... 23 pattersoni,Anisus _...... 7,12,33,35,36,39,56,57 Pliohippvs-Plesippus...... ---...... ------. _------21 Oyraulus...... ______..______56 Pliolemmus antiquus.-...... - ...- ...... 17,37,40,42 Pearlette ash bed_____....______.______11,15 plionasicus, Heterodon...... ----....-----.....-.-.-...... 28 pearlettensis, Perognathus...... 40 Pliophenacomys meadensis. ------.------17,37,40 Pedomysttanensis ...... 16 primaecus.. 28 Pelecypoda.... - . - - _ _.______22,23,25,27,33,36,45 Pliopotamys meadensis...... ------16,17,33,34,37,42 pellucida hordeacetta, Gastrocopta.,.. 6,7,12,13,17,18,24,25,26,27,29, 30,37,39, 71. pi. 1 poaphagus, Ogmodontomys..------..---.------28 pennsylvanicus, Microtus...... __.______.__...____ 17,18 Podicepssp...... ------28 peracuta, Cincinnatia...... ______...__.______50 Podieipediformes------28 Perca flavescens...... __..______._...... ___ 16,17 Polygyra...------82 perconfusa, ValvataMcarinata____...... _.....______..______.______... 48 82 pereiiguum, Carychium...... _...... 45,51,52 38,82 perezi, Anisus...... _.______..... ______57 tholus...------.----_-_____-_------82 Perissodaetyla.______.______,______22,25,28,33,40.42 rexroadensis-...... - 11,12, 24, 27, 29,30,31,32,45, 82, pis. 1, 2 perlustrica, Amicola lustrica...... 51 texasiana...... -...... ----...-.^.--.------.--.------82 Perognathus gidleyi...... _..... 28 terasmsz'S------82 mclaughlini___...... _.__...______.... _ 22,23 Polygyridae.------82 pearlettensis-...... ______.______.____...... 28,40 Polygyrinae.------. _--______-_ . - 82 rexroadensis__...... ______.______...__..._ 28 39 92 INDEX
Page Page pratensis, Microsorex ... ___-_ ------__--. --.______16 Reptilla...... 22,23,28,33,42 Pratilepus kansasensis. ------__-_-----_----__--.-_. __ 28 Petinella...... 81 prenticei, Rallus. .... ------____- - _ __- ___ __-_---_-_--__-__.____ 28 indentata ...... 39 Previous work _._ __ --___ -_-_-. _ _-. ----.__ __ _ --_-__-___-___ 4 rhoadsi...... 6,12, 24,27,29, & primaevus, Pliophenacomys. __-_---__-____-_---- -______28 vheatleyL. --______6,12,24,27,29, & primitwus, Stegomastodon ... --...- _ _-._ __ ----..______33 (Glyphyalops) rhoadsi.-- 81 Prionodesmacea__-______... ______.. 27,33,45 (Glyphyalus) wheatleyi...... 8/ Privatula------... __-___-__._-----_-__---_-_-____-_----. _ ...... 74 retusa, Oxyloma...... _ 6,12,24,36,37,39,44,77,78,79 Proboscidea. ------.------_---_---_---_------__ 28,33,40,42 rex, Paracryptotis...... 28 Probythinella lacustris ... _____--_---______-___----.______-..___ 16 Rexroad formation 3,9,15 Procastoroideslanei.. .-- . ------...... _ _ _ ...... 28 Rexroad local fauna..__ 7,11,14,07 sweeti...... -.----.-. . __ ...... __ _._._.______. _ ____ . 16,33,42 rexroadensis, Bassariscus...... 28 procera, Gastrocopta- ______2, 6, 13, 24, 26, 37, 39, 67, pi. 1 Gastrocopta...... 3,11,12,17,23,24,25,26,27,29,38,73,pi. 1 Procyon rexroadensis ..______------______28 (Immersidens) -...... -..- ---...... 73, pi. 1 Prodipodomys rexroadensis .... ______28 Mephitis...... 28 sp... . 22,23,40 Mu^ela...... 28 progressus, Nekrolagus--..-.------. .------. 28 Perognathus...... -...... 28 Osteoborvs.- .. __ -.--._-. ______...... ______..__ 23 Polygyra...... 11,12,24,27,29,30,31,32,45,80, pi. 1, pi. 2 Peromyscus- ..______---.--.___.______-_--__... ______. _ ..__ 17 (Erymodon)- .....- . - - 82, pi. 1 Promenetus--- ______....______. __ . ______45,59,60 Procyon.-...... -. .. 28 aeruginosus------.__---.____--__ _ __-_-----__--______-___ _ ... 45 Prodipodomys...... 28 blancoensis ------__ . _ . _ .-______._.. __ ...______. _ 38,45,60 Pana...... -...... 27 carus- ..------.-.. _ ___-_-_--_-_--._-_-_-____-_-______---.-.-____-_..____ 60 Peithrodontomys.---...... 28 circumlineatus. ------___ __._... _ _ _._.-__ 60 Sorex...... 28 exacuous- .... - - _.. _ __ _.. _ . 6,7,17,24,36,44,59 Testudo.. 28 exacuot/s------.--.-,------.- ... _----__------_- _ ------59 rexroadi, Baiomys...... 28 kansasensis- ...... 3, 12, 17, 18, 23, 24, 25, 26, 27, 29, 30, 33, 35, 36, 37, 39, 40, 41, 59, pi. 3 Hesperoscalops..... 28 umbilicatellus-- ______. ____ ._. ______-... ____ .... 3,6, Spilogale...... 28 12, 13, 17, 23, 24, 25, 26, 27, 29, 30, 31, 32, 33, 35, 36, 37, 39, 40, 41 , 44, 45, 54, 60 Rhinoceros indet_ 22 (Phreatomenetus) umbilicatellus- .---.-.--.--.--... _ .. _ .___._._____ 60 rhoadsi, Petimlla...... - 6,12,24,27,29, 81 Prosobranchia_---__ _. ______.______. ______48 Phynchotherium sp_. 42 prototypus, Gastrocopta. _ --- -.. ___ _ ..______...___ 73 rinkeri, Synaptomys 37 Pseudemys sp_ ..-_-__.__----__-.-...______..._____.. 33 Zapus...... 28 Psittacidae, indet __ -----_-__--_--____-__._-___-______-___...__.._ 28 Rita Blanca Creek.. 25 Psittaciformes. _------__- ___--__-______.______..._____.. 28 rivulans, Ferrissia-._____._ 3,6,12,17,22, 23,24,25, 26,27,29,30,33,35,36,37,39, 61 pulchella, Vallonia.. _.._____._..______. ______._ 6,12,13,33,34,35,36,76 robustocondyla, Anchylorana 27 Pulmonata.._ _--_-___-___-______-.______.___.____.______...__.______51 robustus, Sceloporus- 28 pumila, Ferrissia ..... ___.--______--___.______-_-__.._____...... ______. 62 Rodentia...... 22,23,28,33,37,40,42 pumilus, Ancylus . ... ------.-.--...... ______61,62 rosaceum, Musculium 46 punctatus, Ictalurus _ . _- --.._ ._._._ --.____. ______16 rotundatm, Anisus...... - - - - 56,57 Punctinae._ __ ... __ ...... __ . ______.______.. ______79 ryckholti, Musculium 46 Punctum.- _--_-_-__-__.__-__.----______79 simum, ______36,44, 79 Pupilla blandi...... ____.______.__...... __._.______16,17,18,35 Salientia 22,23,27 muscorum.-- ______._..______7,16,18,35 Sand Draw local fauna....--. . 1,11,17,30 Pupillid.._._-_ . - - _ 26,41 Sanders local fauna..__.. H, 16, Ifi PupiUidae..------.-.------_---.-.-----.--.--...... _... 5,24,25,40,67,76 sandersi, Sorex_____ . 40 Pupillinae__---- ______74 santacruzensis, Planorbis 45,60 Pupoides. _.__.-----_-_-_.______._ 74 Saw Rock Canyon local fauna...... 3,9,11,14,00,25 albilabris ...... 3, 6,12, 23, 24, 26,27,29, 30, 31,32, 33,35, 36, 37, 39, 40,41, 74, pi. 1 sawrockensis, Baiomys 23 hordaceus-...... ______..______.______18 Ogmodontomys... __.- 23 inornatus-- ...... 6,12, 24, 25, 26,27, 29,30,33,35,36, 74, pi. 1 scaevoscala, Gastrocopta...-...... - 12,24,31,32,37,39,45, 70 marginatus-.--...... ______74 Scaphiopus diversus. __... .. - 27 (Ischnopupoides) inornatus______74, pi. 1 Sceloporus robustus - . 28 Pyganodon...... 45 ., Buisnictis...... 22,23 (Pyganodon) grandis, Anodonta....-...... ______45 Scirpus...... 61 Pyrgophorus-- ______51 Scolopacidae indet__ . 28 securis, Musculium...... 46 Q Sphaerium ...... 6,36,44, 46 guadriplicatus, Parahodomys. .______28 Sediments, environmental interpretations.... 3 Quadrulidae..______...... 45 Seger local fauna____. .. 16 Quicketta----...... -...... 77 Shells, specific identification of..____ . 5 vagans--. ....______..______24 Sigmodon__.._ 16 guinni, Oeomys...... ______17,28,33,42 hilli...... - .- . . 16 Mermedius. ______28,40 Rallus prenticei.. sp_ 37 Rana ephippium- ______.______27 sillimani, Oxyloma.-.. .. . 78 fayeae ..... ______.______27 simile planatum, Sphaerium .. ... 46 meadensis...______27 simplex, Valvata tricarinata______48 parvissima.-- ______27 simplicidens,PlesippuS--.--. .- ...... - . 16,28,33,40,42 rexroadensis ______27 Symmetrodontomys______.. 28 valida.... __ ....______27 simus, Arctodus---. .. 17 Rancholabrean age...______...___.___...____. 19,21 Sinalbinula. ______._. _ - 70 Recent mollusks from northern Nebraska______....._____ 42 Red Corral local fauna______._____.______11,17,05 singleyanus, Helicodiscus.. 3, 6,12,13,17, 23,24,27, 29,30, 31, 32,33, 35, 36,37, 39, 40, 41, 79 Helicodiscus (Hebetodiscus) __.______. 79 reflexa, Galba-..-. ______....___..______53 Stagnicola.. _.._...._..______... 6,12, 33, 35, 36, 39, 52, 53 skinneri, Physa...... 6, 8,12,13,16,17,24, 32, 33,34,35,36, 37, 39, 40,41,63, 64, pi. 3 regalis, Hypolagus______..______.______28 Slaton local fauna______... 1 Reithrodontomys rexroadensis-- ...______...______-__ 28 Smilodon sp...___._.. .. 33 wetmorei______28 Snake..- . 23,42 INDEX 93
Page Page Sorex...... 16,17,39 Synaptomys landesi. 16 cinereus ...... _. .. ___ _ . ______. 18 rinkeri.-----.... 37 dixonensis.----...... _____ . __ . 37 (Mictornys) borealis.. 16 leahyi ...... ______37 Systematic descriptions. 44 rexroadensis . ._ ...... -...... 28 sandersi ...... - .. . _.__ 40 T taylori...... _ __ 28 Tanupolama blancoensis...------28 sp...... -_.. . . 37 tappaniana, Gastrocopta.. 3,6,12, 23, 24, 27, 29,30, 31,32,33,35,36,37,39,40,41,44,72, 74 Soricidae ______. ______23 Gastrocopta (Vertigopsis)...... -...... ___.. 72 sparsicostata,Strobilops.- -...... 12, 22, 24, 25, 26, 27,29,30,33,35,36,37, 39, 65, pi. 1 Taxideataxus.....-.... ._.--.--..---.------.------.. - 28,33,42 spatulus, Gigantocamelus...... 23,28 Taxonomy, unsatisfactory...... -..._..-._...... -.--..-----...... 8 Sphaeriidae...... ______45 taxus, randea------...... ------. .. 28,33,42 Sphaerium...... 45 taylori, Sorex... 28 crassum...... 46 tazewelliana, Fossaria parva..____.______. _-. ._. . .__ 55 lacustre...... 6,12, 36, 39, 44, 46 techella, Oalba bulimoides ______...... 54 lineatum...... ______.______._ 46 Stagnicola bulimoides..____._ 3, 6,12,13,16, 23, 24, 25, 26,27,29,30,31,32, 40, 41, 45, 64 occidental..-...... ______....______... 53 (Nasonia) bulimoides....___-.------_------__ __-_ 54 partumeium...... 6,12,33,35,36,39,45 Teleodesmacea..--...... -...... 22,23,25,27,33,36,45 securis ------...... 6,36,44,46 Terrapene llanensis.------...... 17,18 simile...... _____-______46 Testudo ------16,17,33,34,42 planatum _----.-----...... __...... _.. 46 rexroadensis---...... _...... -...._.------..... 28 striatinum...... 6,12,23,24,33,35,36,46 turgida..-...... 22,28 sulcatum...... 6,12,26, 33, 35,36, 46 tetralasmus, Uniomerus---.---..-.------.------.------24, 26, 45 transversum.-...... ______6,12,24,26,33,35,36,39,47 texasensis, Polygyra texasiana----...... 82 sp...... - .. . .. 23,24,27,29,46,47 texasiana, Polygyra--..____...... ------. ----- .. ... 82 Strobilops-... ..__.__.__-_...... _-__...... ____------65 Spilogale rexroadi...... 28 texasensis, Polygyra...___-._... ------82 Squamata______.______22,23,28,42 tholus, Polygyra mooreana..------___ _------82 stagnalis jugularis, Lymnaea...... _. 16,17,56 Threskiornithidae, indet -_... .------. 28 Stagnicola .______39,52 tigrinum, Ambyttoma...... -..----. . 37, 40 bulimoides techella...... 3,6,12,13,16,23,24,25, 26, 27, 29, 30, 31, 32, 40,41,45, 54 tobinensis, Oeomys...... -_ _-_ _ 40 caperata...... 6,12,13,18, 22, 24, 27, 29, 31, 32,33,35, 36,39, 40, 41,43, 44, 53, 54 transversum, Musculium...... 47 cockerelli...... __..____.__ 24 Sphaerium ______...... 6,12,24,26,33,35,36,39,47 exilis...... 6,12, 24,27, 29, 30,45, 51. tricarinata, Valvata...... 6,7,12,13,16,17,33,34,35,36,39,44,48 palustris ...... -- ...... 6,36,39,43,44,52,54 basalis, VoJvata.-...... --....------..--...... 48 reftexa...... 6,12,33.35,36,39,52,53 infracarinata, Valvata.------_-______--_--__--__--_------48 sp...... 3,23,24,31,32,40,41,53,56 mediocarinata, Valvata-...... ______..___..... 48 (Hinkleyia) caperata... ______...______...__.__.... 53 perconfusa, Vateata...... 48 (Nasonia) bulimoides techella...... __...__. 54 simplex, Valvata...------______-_._..--_------_- 48 Stegomastodon mirificus...... 16 unicarinata, Valvata- -__ -_ . _ ____-_-_- -__ 48 primitivus_____.______...______.____ 33 Trigonictis kansasensis..----.------___ ------__ 28,33 sp...... 28,42 Triodopsis------__------_---._-_-_-______...... 31 steinii, Hydrobia... ______50 truncatum, Musculium...--___...... _ . ------.---_----- ..__ .. 45 Stenotrema.... ______r ______83 turgida, Testudo ...... 22,28 fraternum.._------.----- .------...... __.__. 39 turritella, Lymnaea...... ------...... _ .__ . 38,45, 55,56 feat.. - - - _ _. _ __._ __ _ 6,36,44,83 Turtle .- ... - - . -- - -- 23 leai______...... - 83 U monodon...---_-__-_-__--___-_---_-______.._.______83 umbilicatellus, Gyraulus---.------.------60 sterkii, Fossaria parva_____.______...... _._...._ 55 Menetus_-.._ __--.------..-----.--..-...------.-.-.--.------.---- 60 Sterna'. sp_ 28 Promentus- __....______...... _ ...... 3, stirtoni, Cimarronomys------_--_--___-__-______._.__-___ 23 6,12,13,17, 23, 24, 25, 26,-27, 29, 30, 31,32,33, 35,36,37,39,40, 41,44,45, 54, 60 stovalli, Paradipoides------...--.--...... _...... _...... _...... 17 (Phreatomenetus)...... 60 striatinum, Sphaerium--- __. .. -_ 6,12,23,24,33,35,36, 46 unicarinata, Valvata tricarinata----. ._____- ______-__ _-- __-_ - .... 48 striatulatus, Eumeces..._ - .-- ______. ____ _ 23, 28 Uniomerus tetralasmus.- __------_- --___-_- ---_- _-__ -_ 24, 26, 45 Unionidae______--____-__-__---_---_.__--..-__----_--_._--_--- 8,45 Strobilops.- , -- . ______65 trivolvis, Helisoma . -- - ...... 6,12,24,26,36,39,44,59 densecostata_._.______eg Helisoma (Pierosoma)___ _._..------_--__----._____------_- _--_-__--__ 59 Zafcyrmttjco...... 3,6,12,17,23,24,65, pi. 1 sparsif-ostata ...... 12, 22, 24, 25, 26,27, 29,30,33,35,36,37,39, 65, pi. 1 V texasiana. __------.______._ 65 vagans, Ouickella.-...... 24 Strobilopsidae_--_-______....----...... __._..._. 65 vaginacontorta, Succinea....___... -______-_ __ ...... 24 Stylommatophora. ______22,23,25,27,31,33,37,40,42,65 valida, Rana.-...... ------...-.-...... --..--.-.-.-.-.-... .----.------27 Stump Arroyo member of Crooked Creek formation... ___ 3,15 Vallonia ------76 subrostrata, Lampsilis... ______.______.-_...... _.. 45 costata...... - -.--. -.--- --.-- -.--- 26,29 Ligumia. ______24,27,29,30,45 cyclophorella------16 subtruncatum, Pisidium______17 gracilicosta...... 6,12,13,24,27,29,30,36,44,77 Succinea...... 3, 23, 24, 25, 26, 27, 29,30,31", 32,33, 35, 36,37,39,40, 41,44, 77, 78 parvula------6,12,24,26,31,32,36,44,77 avara...... ------26,78 perspective -- - .- 3,12,13,17,23,24,26,27,29,30,37,39,40,41,77 pidcheUa . 6,12,13,33, 34, 35, 36, 76 concordialis _ -.. 24,39,77 Valloniidae 76 grosvenori. _. ------78 Valvata. .- - -.-. -- . .. 48 gelida_ _ _ ------78 lewiii . 6,12,13,33,34,35,36,45 luteola.-..__ ------78 ovalis- -.---...--....-..---..-_...... -.....-...... _..... g6, 44, 77, 78 helicoidea...... -.-_----..------.---.-.--.--.. 49 vagtnacontorta_ ------_---. - ...... -...... ______._ 24 tricarinata-.. .. 6,7,12,13,16,17,33,34,35,36,39,44,48 sp-.._- .-- ..-...... _..._ ...... -.-...... _...... _...._... 78 basalis..------48 infracarinata.--.-...... ---.-----.------.------.------....--.- 48 Succineidae_ ---..--... - ._....._-... ______5,8,77 mediocarinata.------48 sulcatum, SpAaeriMm...______6,12,26,33,35,36,46 perconfusa.. ____... ___ ....__...-....- 48 sweeti, Procastoroides. ..._.------.---- -__ --_...... __...... 16,33,42 simplex.--.- _____...._-...__------_--.--- . 48 Symmetrodontomyssimplicidens. ______28 unicarinata.-- __.____..------.--.-_--.__- . . 48
508945 O 60- 94 INDEX
Page W Valvatidae.------_------.----....._...._..._...... 48 walkeri, Amnicola--.------.------.------.------45, 49, 50 Vanhem formation-..------.-.---__-.-...... __._____...... 3,15 Pisidium--.---...... -__--. ___-_-. ---..-_ ------17 Vertigininae_.___------__--.------._---.------....___. 74 Washing technique...---_--_._-. .____. ------_.-.--._-.----- 1,2 Vertigo.-- _...._..._..._-...__.--_-----.-__--.__...... _..__. 74 wetmorei, Ffeithrodontomys..... ___-____.-__.-.-.-.--__...... 28 bermudensis______------_...-_------____--_---....__.. 76 wheatleyi, ffettraeHa. . 6,12,24,27,29,81 gouldi ------. - ...... 16,35 Petinella (Glyphyalus)---...... -__- - 81 hibbardi- ...... 3,11,12, 22,23,24,27, 29, 30, 32, 38, 45, 74, 76, 76, pi. 1 wilsoni, Dipoides... _.-.-_----_.-_-_ __._-. ------23 milium. ..--..-....- 2,3,6,12, 23,24,25, 26,27, 29, 30, 31, 32,33, 35,36, 37, 39, 40,41, 76 modesta...... ____..____.______.______16,35 Zapus- .-...... 16 numellata... .._._..._..._._--.-._..__.....____..___. 76 rinkeri- - - - - 28 ovata ...... 6,11,12, 22, 24, 26, 32, 33, 35, 36, 37, 40, 41, 44, 74, 76 Zenaidura macroura__------.------_--_------_------28 diaboll...... 76 Zomtidae------5,80 (Angustula) hibbardi...... _.------...... 45 Zonitinae..__-__---..-____._.. .___---.._ . _.-.-- 80 Vertigopsis_..._._---._...... _..__------_-__-._..__-..... 72 Zonitoides-...... 81 Villafranchian fauna._..__-.------.----_....------.-.----...... 21 arboreus.-- .....-- 6,12, 24,27, 29, 31,36, 37, 39,44,80,8/ Viviparidae ...... 8,36,39,49 Zygospores of Chara...---- _- _--.-__..._------.. 2 o PLATES 1-4 PLATE 1 FIGFRES 1-4. Acroloxus color ad ensis Henderson, X 1% (p. 61). Sand Draw local fauna. 1, 2. UMMZ 181129a. 3, 4. UMMZ 181129b. 5, 6. Strobilops labyrinthica (Say), X 7^ (p. 65). Saw Rock Canyon local fauna. 5. UMMZ 177561b. 6. UMMZ 177561a. 7-10. Strobilops sparsicostata Baker, X 1% (p. 65). 7. Sand Draw local fauna. UMMZ 184178a. 8. Topotype. Rexroad local fauna. UMMZ 183032a. 9. Topotype. Rexroad local fauna. UMMZ 183032c. 10. Topotype. Rexroad local fauna. UMMZ 183032b. 11-16. Gastrocopta chauliodonta Taylor (p. 70). 11. Sanders local fauna. Basal view, X 22^, of dissected specimen showing columellar and anguloparietal lamellae. UMMZ 182152e. 12. Paratype. Sand Draw local fauna. X 15. UMMZ 181121a. 13. Sanders local fauna. Basal view, X 22^, of dissected specimen showing columellar and anguloparietal lamellae. UMMZ 182152e. 14. Sanders local fauna. Spiral view, X 15, of body whorl of dissected specimen showing the very deeply im mersed lower palatal fold. UMMZ 182152a. 15. Sanders local fauna. Lateral view, X 22^, of dissected specimen showing columellar lamella. The basal part is incompletely joined to the rest of the lamella, and gives the appearance of a subcolumellar lamella. UMMZ 182152c. 16. Sanders local fauna. Basolateral view, X 22^, of dissected specimen showing anguloparietal and colu mellar lamellae. UMMZ 182152b. 17, 23. Polygyra rexroadensis Taylor, n. sp. (p. 82). Holotype, X 1%. Two views of aperture. UMMZ 177610a. 18-22. Vertigo hibbardi Baker, X 15 (p. 75). 18. Topotype. Rexroad local fauna. UMMZ 183041a. 19. Saw Rock Canyon local fauna. UMMZ 177566b. 20. Saw Rock Canyon local fauna. Basolateral view of dissected specimen, showing angular, parietal, and columellar lamellae. UMMZ 177566c. 21. Saw Rock Canyon local fauna. UMMZ 177566a. 22. Topotype. Rexroad local fauna. UMMZ 183041a. 24-26. Gastrocopta cristata (Pilsbry and Vanatta), X 15 (p. 69). 24. Sanders local fauna. UMMZ 182183b. 25. Sanders local fauna. UMMZ 182183c. 26. Sanders local fauna. UMMZ 182183a. 27. Gastrocopta procera (Gould), X 15 (p. 67). Figured specimen. Dixon local fauna. UMMZ 182217. 28. Gastrocopta pellucida hordeacella (Pilsbry), X 15 (p. 71). Figured specimen. Rexroad local fauna. UMMZ 183037a. 29. Gastrocopta franzenae Taylor, n. sp., X 15 (p. 67). Figured specimen. Saw Rock Canyon local fauna. Specimen with bifid anguloparietal lamella and strong labial tubercle. UMMZ 177563a. 30. Gastrocopta rexroadensis Franzen and Leonard, X 15 (p. 73). Red Corral local fauna. UMMZ 183004a. 31, 32. Gastrocopta paracn'stata Franzen and Leonard, X 15 (p. 69). 31. Figured specimen. Bender local fauna. UMMZ 177507b. 32. Figured specimen. Bender local fauna. Basal fold absent. UMMZ 177507a. 33, 34. Gastrocopta scaevoscala Taylor, n. sp., X 15 (p. 70). 33. Holotype. UMMZ 184320. 34. Paratype. Bender local fauna. Lateral view of dissected specimen showing ascending columellar lamella. Dotted line marks shell axis. UMMZ 183017a. 35. Pupoides inornatus Vanatta, X 7/£ (p. 74). Rexroad local fauna. UMMZ 183074a. 36. 37. Pupoides aUiilabris (Adams), X 7H (p. 74). 36. Sanders local fauna. UMMZ 182158b. 37. Sanders local fauna. UMMZ 182158a. GEOLOGICAL SURVEY PROFESSIONAL PAPER 337 PLATE 1
37 LATE PLIOCENE AND PLEISTOCENE MOLLUSKS FROM THE HIGH PLAINS GEOLOGICAL SURVEY PROFESSIONAL PAPER 337 PLATE 2
10 11
"ll 12 13 A .I**- t 16 A15 - »* . 17 f V 20 W A t21
23 24 25 PLATE 2 FIGURES 1-3, 7, 8. Marstonia crybetes (Leonard), X 10 (p. 50). Topotypes, Saw Rock Canyon local fauna. 1. UMMZ 177554e. " 2. UMMZ 177554b. 3. UMMZ 177554a. 7. UMMZ 177554c. 8. UMMZ 177554d. 4, 5. Polygyra rexroadensis Taylor, X 3 (p. 82). Holotype, Rexroad local fauna. 4. Apical view. 5. Basal view. 6, 12-14, 17, 18. Fossaria dalli (Baker), X 10 (p. 54). Topotypes of Lymnaea turritella Leonard, Saw Rock Canyon local fauna. 6. UMMZ 177555a. 12. UMMZ 177555b. 13. UMMZ 177555f. 14. UMMZ 177555d. 17. UMMZ 17755oe. 18. UMMZ 177555c. 9. Marstonia derepta (Baker), X 10 (p. 51). Figured specimen, Saw Rock Canyon local fauna, UMMZ 186155a. 10, 11, 15, 16. Fossaria dalli Baker, X 10 (p. 54). Figured specimens, Jinglebob local fauna. 10. UMMZ 181151a. 11. UMMZ 181151d. 15. UMMZ 18ll51b. 16. UMMZ 181151c. 19-27. Fossaria dalli Baker, X 10 (p. 54). Figured specimens, Rexroad local fauna. Population showing wide range of variation and approaching F. parva. 19. UMMZ 177598d. 20. UMMZ 177598b. 21. UMMZ 177598a. 22. UMMZ 177598c. 23. UMMZ 177598i. 24. UMMZ 177598h. 25. UMMZ 177598g. 26. UMMZ 177598f. 27. UMMZ 177598e. PLATE 3 FIGUKES 1-4. Physa hordacea Lea, X 5 (p. 64). Paratypes, Recent, Vancouver, Washington. USNM 170764. 5, 6. Physa skinneri Taylor, X 5 (p. 63). Recent, Uinta Mountains, Utah. USNM 533484. 9-11. Physa skinneri Taylor, X 5 (p. 63). Paratypes, Berends local fauna. 9. UMMZ 177533c. 10. UMMZ 177533a. 11. UMMZ 177533b. 7, 8, 12-20, 23, 24, 27, 28. Promenetus kansasensis (Baker), X 20 (p. 59). Sand. Draw local fauna loc. 6. Specimens from an unusually variable population showing variation in sculpture and shape. 7,12,16. UMMZ 177430a. 8,13,17. UMMZ 177430b. 14, 15, 18. UMMZ 177430d. 19. 23, 27. UMMZ 177430e. 20. 24, 28. UMMZ 177430c. 21, 22, 25, 26, 29, 30. Promenetus kansasensis (Baker), X 20 (p. 59). Sand. Draw local fauna loc. 2. Unusually large specimens with expanded body whorl, analogous to P. exacuous (Say) form megas (Dall). 21. 25, 29. USNM uncat. 22. 26, 30. USNM uncat. GEOLOGICAL SURVEY PROFESSIONAL PAPER 337 PLATE 3
27 28 29 " '30
LATE PLIOCENE AND PLEISTOCENE MOLLUSKS FROM THE HIGH PLAINS GEOLOGICAL SURVEY PROFESSIONAL PAPER 337 PLATE 4
LATE PLIOCENE AND PLEISTOCENE MOLLUSKS FROM THE HIGH PLAINS PLATE 4 FIGURES 1-3, 5-7, 10-12. Gyraulus parvus (Say), X 20 (p. 58). Dixon local fauna loc. 1. 1. 5, 10. UMMZ 182200b. 2. 6, 11. UMMZ 182200a. 3. 7, 12. UMMZ 182200c. 4, 8, 9, 13, 17, 18 Gyraulus parvus (Say), X 5 (p. 58) Topotypes of G. enaulus Leonard, Saw Rock Canyon local fauna. 4,8,9. UMMZ 177556a. 13. 17, 18. UMMZ 177556b. 14-16, 19-26. Deroceras aenigma Leonard, X 10 (p. 80). Topotypes, Rexroad local fauna loc. 3. 14. UMMZ 183047d Ventral view. 15. UMMZ 183047k Dorsal view. 16. UMMZ 183047h Dorsal view, sinistral specimen. 19. UMMZ 183047J Ventral view. 20. UMMZ 183047e Dorsal view. 21. UMMZ 1830471 Dorsal view. 22. UMMZ 183047b Dorsal view. 23. UMMZ 183047c Dorsal view. 24. UMMZ 183047f Dorsal view. 25. UMMZ 183047g Dorsal view. 26. UMMZ 183047a Dorsal view.
U.S. GOVERNMENT PRINTING OFFICE: 1960 O -508945