Five Illinoian Molluscan Faunas from the Southern Great Plains

Home , Anisus, Carychium exiguum, Illinoian (stage), Physa skinneri, Pupilla blandi, Succineoidea

FIVE ILLINOIAN MOLLUSCAN FAUNAS FROM THE SOUTHERN GREAT PLAINS

Barry B. Miller

Department of Geology, Kent State University Kent, Ohio, U. S. A.

MALA COLOGIA Vol. 4, No. 1, p 173-260

Reprinted

MALACOLOGIA, 1966, 4(1): 173-260

FIVE ILLINOLAN MOLLUSCAN FAUNAS FROM THE SOUTKERN GREAT PLAINS^,^

Barry B. Miller

Department of Geology, Kent State University Kent, Ohio, U. S. A.

ABSTRACT

The mollusks from 5 late Cenozoic (Illinoian) assemblages in the southwestern Kansas-northwestern Oklahoma area: the Berends local fauna, of Beaver County, Oklahoma; the Doby Springs local fauna, of Harper County, Oklahoma; and the Adams , Butler Spring and Mount Scott local faunas, of Meade County, Kansas, are listed. A summary of information consisting of geologic range, distribution and ecology is given for all of the mollusks in the Mount Scott and Doby Springs local faunas, and for previously unreported species from the Berends, Adams and Butler Spring local faunas. Taxonomic changes include assignment of Promenetus exacuous and Pro- menetus kansasensis to a single species complex. There is no satisfactory way of separating these 2 uspecies" in Recent and fossil populations that contain individuals with both riblet and non-riblet type surface ornamentation. The riblet type is designated Promenetus exacuous form kansasensis and the non-riblet form Promenetus exacuous form exacuous. Interpretation of these molluscan assemblages is predicated on the assumptions that climatic factors are the primary agents controlling the distribution of non- marine mollusks; that fossil shells similar to those of living species represent the same species and were subject to the same limiting factors; and that the areas in the Great Plains presently containing associations of mollusks most similar to those found in these fossil assemblages, probably closely simulate the climatic conditions that existed at the time these fossils lived. Recon- struction of local habitat and climatic conditions under which these faunas lived has been based on a synthesis of information obtained from the fossil mollusks, pollen and vertebrates. In no case have interpretations based on the mollusks conflicted with conclusions reached independently from other fossil data. The 5 Illinoian assemblages show compositional differences that are probably related to climatic changes during the nlinoian. The relatively large number of species with predominantly northern distributions, together with the low repre- sentation of exclusively southern faunal elements, indicate that the Doby Springs local fauna represents the coolest of the assemblages studied, and probably lived during maximum nlinoian glaciation. The Mount Scott local fauna contains a greater number of species with southern distributions than any of the other faunas studied, suggesting that it represents the warmest of these Illinoian faunas. There is no evidence inthis area indicatingthat these faunas lived during more than one substage. Assuming that the transition from maximum glacial conditions, represented by the Doby Springs local fauna, towards warmer inter-

'~ontrl~utionNo. 2, Department of Geology, Kent State university. Research supported (in part) by National Science Foundation grant G- 5635. 2~daptedfrom a dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy at the University of Michigan. I74 B. B. MILLER

glacial conditions, represented by the Mount Scott local fauna, was unidirectional, then the climatic conditions implied by the Adams and Butler Spring assemblages must have occurred sometime between the time represented by the Doby Springs and Mount Scott local faunas. The first and last appearances of molluscan species and climatic implications of the Berends local fauna suggests that this assemblage is probably the oldest of these 5 Lllinoian faunas. The Berends assemblage contains 45 species of mollusks, 15 of which are new to this local fauna. One species, Sphaerium securis, represents the first fossil occurrence of this species from the southern Plains. This fauna lived in and around the margin of a small, shallow lake where the vegetation cover probably consisted mainly of conifers, grasses and composites, with only scattered patches of deciduous trees. The climate at the time this assemblage lived was probably similar to that presently occurring in the northeastern Neb- raska-northwestern Iowa region of the Great Plains. The Doby Springs local fauna contains 59 species of mollusks, one of which, Veertigo elatior is recorded for the first time as a fossil in the southern Plains. The mollusks indicate the existence of a cool, upstream lake, bordered by a marsh containing cattails, sedges and scattered deciduous and coniferous trees, which graded into a better drained shrub and tree community further back from the lake. The inferred former local climate combined shorter, cooler summers, with winters that were slightly cooler and longer than those presently prevailing in this region. The Adams molluscan assemblage contains 22 species, one of which, Lasmi- gona complanata is first recorded as fossil from this area. The medium to coarse, cross-bedded sands from which the fauna was recovered, together with the occurrence of molluscan species requiring continuity of aquatic habitat, indicate the existence of a stream. The presence of only 1 woodland species among the terrestrial gastropods suggests that there was only a thin covering of trees and shrubs. The climate probably combined winters perhaps no more severe than those presently occuring in northeastern Nebraska, with cooler, more moist summers, similar to those in the Dakotas and eastern Montana. There are 54 species of mollusks in the Butler Spring local fauna, one of which, Ligumia cf recta, is new to the southern Great Plains. This fauna lived in or near a medium sized river, with a slow to moderate current and a probable depth of several feet. There were scattered groups of trees along the river, but the predominant vegetation consisted of grasses and shrubs. The climate suggested by the assemblage consisted of cooler summers, similar to those now occurring in the eastern North Dakota region. Winters were probably no more severe than those in northcentral Nebraska. The Mount Scott molluscan fauna, containing 63 species, is the largest such assemblage known from the southern Plains. It contains the only fossil record of Zonitoides nitidus from the Great Plains. Sedimentary and paleontologic evidence indicate that 5 major habitat associations are represented. The climate at that time probably combined summers similar to those now occurring in the Lake Okoboji region of northwest Iowa, with winters no more severe than those in the northeast Kansas-southeast Nebraska area.

CONTENTS

Page Page

INTRODUCTION ...... I75 Introduction ...... I76 Berends Local Fauna ...... I77 FAUNAL DISCUSSION ...... I76 Doby Springs Local Fauna ...... I81 FIVE ILLINOIAN FAUNAS 175

Contents (cont.)

Page Page

Adams Local Fauna ...... I86 Family Lymnaeidae...... ,225 Butler Spring Local Fauna ...... I88 Superfamily Ancyloidea...... 227 Mount Scott Local Fauna ...... I89 Family Planorbidae...... 227 Summary of Illinoian molluscan Family Ancylidae ...... 236 I - faunas ...... 212 Family Physidae ...... 237 Order Stylommatophora ...... 240 SYSTEMATIC DISCUSSION ...... 216 Superfamily Cionelloidea ...... 240 e Class Pelecypoda ...... 217 Family Cionellidae ...... 240 Order Prionodesmacea ...... 217 Superfamily Pupilloidea ...... 240 Family Unionidae ...... 217 Family Strobilopsidae ...... 240 Order Teleodesmacea ...... 220 Family Pupillidae ...... 240 Family Sphaeriidae ...... 220 Family Valloniidae ...... 246 Class Gastropoda . . . , ...... 220 Superfamily Succineoidea ...... 247 Subclass Prosobranchia ...... 224 Family Succineidae ...... 247 Order Mesogastropoda ...... 224 Superfamily Endodontoidea ...... 247 Superfamily Valvatoidea ...... 224 Family Endodontidae ...... 247 Family Valvatidae ...... 224 Superfamily Zonitoidea ...... 249 Superfamily Rissoidea...... 225 Family Limacidae ...... 249 Family Hydrobiidae ...... 225 Family Zonitidae ...... 249 Subclass Euthyneura ...... 225 Superfamily Polygyroidea ...... 252 Order Basommatophora ...... 225 Family Polygyridae ...... 252 Superfamily Ellobioidea ...... 225 ACKNOWLEDGEMENTS ...... 252 Family Ellobiidae...... 225 . Superfamily Lymnaeoidea...... 225 REFERENCES ...... 253

INTRODUCTION intervals have been recognized in the Meade County area, which are believed Under supervision of Claude W. to correlate withthe Nebraskan-Kansan- Hibbard, field parties from the Uni- Illinoian-Wisconsin glacial and Aftonian- versity of Kansas (1936-1946) and Uq- Yarmouth-Sangamon interglacial stages versity of Michigan (from 1947 onwards) in the classical Mississippi Valley have been engaged in an intensive study sequence. of the late Cenozoic fossils and strati- The washing technique developed by graphic sequence of the southwest Hibbard (1949b) has made possible the Kansas-northwest Oklahoma area. recovery of extensive collections of During the 27-year period from 1936- fossils which have included ostracods, 1963, 13 Pleistocene faunas ranging in mollusks, fishes, amphibians, reptiles, age from Aftonian to Wisconsin, have birds and mammals. Detailed study of been studied (see Fig. 1, and summaries the pollen associated with these faunas in Taylor, 1960; Hibbard & Taylor, has recently been undertaken by Kapp 1960). Correlation of the Pleistocene (1965). Some of these groups have sequence in this area with that of the barely been studied, while others have glaciated region to the north and north- been fairly intensively described and east, is based on the inferred climatic illustrated. In this respect the mollusks changes and stratigraphic position of from these faunas are comparatively these faunas. Four cooler-climate in- well known. With the exception of the tervals separated by 3 warmer-climate Deer Park and Borchers local faunas, 176 B. B. MILLER

AGE ROCK UNIT FAUNA

WISCONSIN VANHEM FM. JONES L. F.

JINGLEBOB L. F. SANGAMON CRAGIN QUARRY L. F. MOUNT SCOTT L. F. KINGSDOWN FM. BUTLER SPRING L. F ADAMS L. F. w z DOBY SPRINGS L. F. u 0 o ILLINOIAN BERENDS L.F. I- -V) W YARMOUTH ATWATER MEMBER J0. , BORCHERS L. F. WY ZWw ..PEARLETTE ASH) 0 [LZ CUDAHY F. a0 0 KANSAN STUMP ARROYO MEMBER SEGER GRAVEL PIT L. F.

AFTONIAN o SANDERS L.F. [L MISSLER MEMBER Q-jt- DEER PARK L. F. a ANGELL MEMBER NEBRASKAN m w BENDER L. F. 2 W o UPPER REXROAD FM. REXROAD L. F. 5! J SAW ROCK CANYON L. F. P

FIG. 1. Late Cenozoic stratigraphic sequence and faunal succession in southwestern Kansas- northwestern Oklahoma.

mollusks have usually been among the surficial geology of the portion of the most abundant fossils recovered from Big Springs Ranch, Meade County, Kan- these assemblages. The shells of fossil sas, from which the Mt. Scott local mollusks often provide the character- fauna was collected, is also included. istics upon which specific diagnoses are New materials from the other Illinoian based. Interpretations of the Illinoian assemblages of this area, the Berends, assemblages considered in this report Adams and Butler Spring local faunas, are predicated upon the uniformitarian which have become available since these assumption that fossil shells similar to local faunas were reported on by Hibbard those of living species represent the & Taylor (1960), have requiredcertain same species and were subject to the emendations which are presented herein. same limiting factors. Conclusions The interpretation of the paleoecology based on the molluscan elements of the of these faunas is based on a synthesis faunas in all cases are consistent with of evidencefromfossilpollen, vertebrate the interpretations reached from study and invertebrate faunas and sedimentary of the pollen and other animal groups. characters. This paper is concerned primarily with the description and interpretationof FAUNAL DISCUSSION the hitherto incompletely reported late Introduction Illinoian molluscan assemblages from the Mt. Scott and Doby Springs local The climatic and stratigraphic signifi- faunas. A map and description of the cance of the Mount Scott andDoby Springs )LAN FAUNAS 177

local faunas is perhaps most readily cene sink-hole deposits, after collapse shown by comparison with other Illinoian of the underlying Crooked Creek for- faunas from the southwest-Kansas north- mation. The fauna. collected from these west-Oklahoma area. This section of the sediments has been summarized by report, therefore, also includes dis- Hibbard & Taylor (1960: 55-57) from cussion of the Berends, Adams and Butler previous reports (Herrington & Taylor, Spring local faunas. These five assem- 1958; Hibbard, 1956b; Mengel, 1952; blages show compositional changes which Rinker & Hibbard, 1952; Smith, 1954, seem best explained by assuming 1958; Starrett, 1956; Taylor, 1954; climatic change. Evidence presented Taylor & Hibbard, 1955). Kapp (1965) below suggests that the Doby Springs and has recently analysed the pollen from Mount Scott local faunas represent the the Berends local fauna locality. Four- coolest and warmest climatic conditions teen species of mollusks, hitherto un- in this sequence of faunas, respectively. reported from this fauna, were re- Although several stadial-interstadial in- covered from matrix collected during the tervals probably occurred during the summer of 1959 by University of Illinoian, there is no evidence in this Michigan Museum of Paleontology field area indicating that these assemblages parties. lived during more than one substage. The Doby Springs local fauna probably Faunal List, Berends Local Fauna: lived earlier in this interval, sometime near maximum glacial conditions, while CLASS PELECYPODA the Mount Scott local fauna lived later Order Teleodesmacea in the Illinoian after the climate had started to change towards warmer inter- Sphaerium lacustre (Miiller) glacial conditions. If this transition from *S. securis Prime maximum glacial towards warmer inter- S. sulcatum (Lamarck) glacial conditions was unidirectional, *S. transversurn (Say) then the climatic conditions implied by Pisidium casertanum (Poli) the Adams and Butler Spring local faunas P. compressurn Prime probably occurred sometime after the P. nitidum Jenyns Doby Springs local fauna lived, but be- P. obtusale (Lamarck) fore the time represented by the Mount *P. variabile Prime Scott local fauna. This faunal sequence is also suggested by the first and last CLASS GASTROPODA appearances of molluscan species. The Order Mesogastropoda mollusks of the Berends local fauna suggest slightly warmer climatic con- Valvata tricarinata (Say) ditions than those represented by the glacial maximum Doby Springs assem- Order Basommatophora blage. At the present time it is not * Carychium exiguum (Say) certain whether this warmer interval Stagnicola caperata (Say) occurred before or after maximum S. exilis (Lea) z Illinoian glaciation. *S. reflexa (Say) Berends Local Fauna Fossaria dalli (Baker) F. obrussa (Say) Previous Work. The Berends local Gyraulus circumstriatzcs (Tryon) fauna is known from secs 5 and 6, T 5 N, G. parvus (Say) R 28 E, Beaver County, Oklahoma. It Armiger m'sta (Linnaeus) was recovered from a dark silty inter- *Helisoma anceps (Menke) val in a sequence of sediments that H. trivolvis (Say) accumulated as a series of late Pleisto- Planorbula amtigera (Say) 178 B. B. MILLER

Promenetus exacuous form kansasensis Pelicanus erythrorhynchw Gmelin, white (Baker) pelican *Physa anatina Lea P. gyrina Say CLASS MAMMALIA P. skinneri Taylor Order lnsectivora Aplexa hypnorum (Linnaeus) Sorex cf S. cinereus Kerr, masked shrew Order Stylommatophora Blarina cf B. brevicauda (Say), short-tailed shrew *Strobilops lubyrinthica (Say) Gastrocopta armifera (Say) Order Rodentia *G. cristata (Pilsbry & Vanatta) G. procera (Gould) Castoroides sp. , giant beaver G. tappaniuna (Adams) Paradipoides stovalli Rinker & Hibbard, *G. holzingeri (Sterki) Stovall's beaver Pupoides albilabris (Adams) Geomys sp. , eastern pocket gopher *Vertigo milium (Gould) Perognathus cf P. hispidus Baird, hispid V. ovata (Say) pocket mouse *Vallonia gracilicosta Reinhardt Peromyscus berendsensis Starrett, deer *V. parvula Sterki mouse cf Succinea Ondatra triradicatus Starrett , muskrat, Oxyloma sp. Microtus pennsy lvanicus (Ord) , meadow vole Helicodiscus parallelus (Say) Pedomys cf P. ochrogaster (Wagner), prairie *Deroceras aenigma (Leonard) vole * Euconulus fulvus (Miiller) Hawaiia minuscula (Binney) Order Carnivora Stenotrema leai (Binney) Canis lutrans Say, coyote CLASS OSTEICHTHYES Order Proboscidea. Lepisosteus sp. , gar Esox masquinongy Mitchill, muskellunge Mammuthus cf M. columbi (Falconer), Catostomus commersoni (~ace'p'ede), white Columbian mammoth sucker Notemigonus crysoleucus (Mitchill), golden Order Lagomorpha shiner Semotilus cf S. atromaculatus (Mitchill), Leporidae , indeterminate, rabbit creek chub Semotilus sp. , minnow Order Perissodactyla Cyprinidae, indeterminate Ictalurus melas (Rafinesque), black bullhead Equus sp. , horse I. punctatus (Rafinesque) , channel catfish Lepomis cf L. cyanellus Rafinesque, green Rejected records sunfish Perca f7avescens (Mitchill), yellow perch Deroceras laeve = Deroceras aenigma; Fundulus sp. Aplodinotus grunniens = Fundulus sp.

CLASS REPTILIA Order Chelonia Age and Correlation. The sediments Unidentifiable turtle remains containing the Berends local fauna over- lay the Pearlette ash, and are therefore CLASS AVES post-Kansas in age. Climatic implica- Order Pelicaniformes tions of the pollen (Kapp, 1965) and fauna (Hibbard & Taylor, 1960; Miller, this re- *Species not previously reported. port), indicate that the assemblage lived FIVE ILLINOIAN FAUNAS 179

TABLE 1. Summary of local habitats represented by the mollusks of the Berends Local Fauna. *

Habitat Species Individuals I Of Semiaquatic: among vegetation and debris I Oxyloma sp. 1 241 near water's edge.

I Hygrophilic: moist situations in leaf mold, Carychium exiguum under sticks and debris; shaded areas not Vertigo ovata far from water. V. milium Gastrocopta tappaniana Deroceras aenigma

Woodland: moist areas under leaf mold litter, Stenotrema leai 8 4 down timber, among tall marsh grass. Euconulus fulvus 1 Strobilops labyrinthica 1 Helicodiscus parallelus 8

Sheltered areas: among rocks, shrubs, grass Gastrocopta armifera 30 or in timbered situations. G. holzingeri 29

Sheltered situations: these species are not Gastrocopta cristata restricted to a woodland habitat and can G. procera tolerate drier conditions. Pupoides albilabris Hawaiia minuscula Vallonia parvula V. gvacilicosta

Marginal situations: wet mud, sticks, stones Fossaria obmssa 62 or any other debris along water's edge: F. dalli 97 shallow ponds and protected spots.

Shallow quiet water: small stream, pond, 1 Pisidium casertanum 1 4+4/2** slough or marsh that may become dry Stagnicola reflexa during part of the year. S. caperata S. exilis Armiger crista Gyraulus circumstriatus Physa gyrina Aplexa hypnomm

Shallow quiet water: small pond, stream, Sphuerium securis slough, or marsh with no current or areas S. sulcatum of rooted vegetation with little current; S. transversum soft sand or mud bottom; not subjected Pisidium o btusale to significant seasonal drying. Gyraulus parvus Helisoma trivolvis Planorbula armigera (cont . )

*Based on material published in Taylor, 1954; Herrington & Taylor, 1958; Hibbard & Taylor, 1960; this report. **Fractions refer to the number of isolated single valves. 180 B. B. MILLER

Table 1 (cont. )

Number of Habitat Species 1 Individuals

(cont. ) Promenetus exacuous form kansasensis 513 Physa amtinu 58 LI P. skinneri 208

Perennial water: stream or lake with slow Sphaerium lacustre to moderate current; areas of still water; Pisidium compressurn shallow spots with soft sand or mud P. nitidum substrate not affected by seasonal drying. P. variabile Valvata tricarinuta Helisoma anceps during a cool, moist, glacial interval. shallow permanent and temporary water The relatively large number of extinct near the shore of a lake, with little or species of rodents suggests that this no rough water. Connection of this lake interval was pre-Wisconsin (probably to a permanent stream at some time is Illinoian). indicated by Valvata, which must have Stephens (1960) correlated the Berends continuity in aquatic conditions.') This and Doby Springs local faunas on the interpretation is supported by the fishes basis of the mammals shared in common. (Smith, 1954), the sphaeriids (Herrington Kapp (1965) has interpreted the pollen & Taylor, 1958), and by the additional spectrum associated with the Berends aquatic mollusks reported herein. fauna as representing maximum Illinoian The pollen spectrum (Kapp, 1965) glaciation and considered it equivalent shows large per centages of conifers, both to locality 5 of the Doby Springs grasses and composites, with only small (p 181) and to the Adams (p 186) local quantities of deciduous tree pollen. The faunas. The mollusks, however, suggest relative paucity of terrestrial snails that the Berends lived under slightly usually associated with deciduous wood- warmer climatic conditions than those lands can probably be accounted for by represented by the Doby Springs assem- assuming a vegetation cover such as is blage. At the present time it is not indicated by the pollen. The better certain whether this warmer interval represented of the terrestrial species occurred before or after maximum (Cavychium exiguum, Vevtigo ovata, V. Illinoian glaciation. milium, Gastvocopta tappaniana and Habitat. A summary of the local Hawaiia minuscula) probably lived among habitats represented by the mollusks is the moss-covered down timber near the given in Table 1. This arrangement is lake margin. The remainder of the somewhat arbitrary since many species terrestrial mollusks were probably dis- c- are not necessarily restricted to the one tributed among scattered patches of habitat to which they may have been deciduous trees further back from the assigned. However, it is believed that lake, under accumulations of leaf mold these habitat groups probably reflect the and rotting timber. fact that each species does have a certain climate. The area of sympatry for "preferred') habitat in which it is most most of the species of mammals. fishes abundant and where it most commonly and mollusks-found in the ~erendslocal occurs. Taylor (1954: 7) has interpreted fauna is the northeastern Nebraska- the aquatic gastropods of this fauna as northwestern Iowa region of the Great representing ". . . a habitat which had Plains. This area is now characterized FIVE ILLINOIAN FAUNAS

by a climate with normal annual temper- CLASS PELECYPODA atures of about 45O F; normal summer Order Prionodesmacea temperatures averages of about 700 F; normal winter temperatures of 20' F; Uniomerus tetralasmus (Say) and a precipitation rate of between 20- Anodonta grandis Say 30 in. a year, (Visher, 1954). Most of this precipitation comes during the Order Teleodesmacea hot part of the year when it is most Sphuerium occidentale Prime needed. The Berends local fauna appears S. rhomboideum (Say) to have lived under climatic conditions S. striatinum (Lamarck) similar to those now essting in the area S. sulcatum (Lamarck) of sympatry. Pisidium casertanum (Poli) Doby Springs Local Fauna P. compressum Prime P. nitidum Jenyns Previous Work. Smith (1958) proposed P. obtusale (Lamarck) the name Doby Springs local fauna for P. variabile Prime the fishes and associated organic remains occurring in late Pleistocene de- CLASS GASTROPODA posits in the N 1/2 SW 1/2 sec 10, T 27 Order Mesogastropoda N, R 24 W, Harper County, Oklahoma. Meyers (1959) tentatively assigned these Valvata tricarinata (Say) beds to the Illinoian stage on the basis *Probythinella lacustris (Baker) of sedimentary and molluscan faunal similarities with other sink hole deposits Order Basommatophora of this age in the area. ~ibbard& Taylor (1960) recorded one species of Carychium exiguum (Say) Lymnaea stagnalis jugularis Say mollusk (Pvobythinella lacustris) from Stagnicola caperata (Say) the Doby Springs fauna. Stephens (1960) S. reflexa (Say) mapped and studied the geology of the Fossaria dalli (Baker) area and described the mammalian fauna. F. obmssa (Say) More recently, additional materials from Armiger crista (Linnaeus) this fauna have been reported in papers Gyraulus circumstriatus (Tryon) by Mingener (1963, on the jumping G. deflectus (Say) mouse, Zapus); Etheridge (1960; 1961, G. parvus (Say) on the lizard Ophisaum); Tihen (1962, Helisoma anceps (Menke) on the bufonid toads); and Gutentag & H. trivolvis (Say) Benson (1962, on the ostvacods). Kapp promenetus exacuous form kansasensis (1965) has recorded the pollen found (Baker) associated with the Doby Springs local p. umbilicatellus (Cockerell) fauna. The Doby Springs local fauna Fernssia fragilis pryon) is known from 5 localities, only 4 of Laevapex fusms (Adams) which contain mollusks. The locality Physa anatina Lea numbers referred to below are the same P. gyrina Say P. skinneri Taylor as those used by Stephens (1960). Aplexa hypnorum (Linnaeus) Age and Correlation. The Doby Springs Order Stylommatophora

Faunal List, Doby Springs Local Fauna: Cionella lubrica (Miiller)

CLASS OSTRACODA *Species reported previously in Hibbard & Candona nyensis Gutentag & Benson Taylor (1960). 182 B. B. MILLER

Strobilops labyrinthica (Say) CLASS MAMMALIA Gastrocopta armifera (Say) Order Insectivora G. contracts (Say) G. cristata (Pilsbry & Vanatta) Sorex arcticus Kerr , arctic shrew G. holzingeri (Sterki) S. cinereus Kerr, masked shrew G. procera (Gould) Sorex palustris Richardson, northern water G, tappaniana (Adams) shrew Pupoides albilabris (Adams) Blurina brevicauda brevicauda (Say), short- Pupilla blundi Morse tail shrew 4 P. muscorum (Linnaeus) Vertigo elatior (Sterki) Order Rodentia V. milium (Gould) V. ovata (Say) Citellus cf C. richardsonii (Sabfne), Richard- * Vallonia gracilicosta Reinhardt son ground squirrel V. parvula Sterki Citellus tridecemlineatus (Mitchill), thirteen cf Succinea lined ground squirrel Oxylom sp. Thomomys sp. , western pocket gopher Discus cronkhitei (Newcomb) Geomys bursarius (Shaw), Plains pocket Helicodiscus parallelus (Say) gopher Deroceras aenigma Leonard Castorid, genus and species indeterminate, Euconulus fulvus (Miiller) large beaver Nesovitrea electrina (Gould) Peromyscus cf P. cochrani Hibbard, mouse Hawaiia minuscula (Binney) P. oklahomensis Stephens, mouse Zonitoides arboreus (Say) Onychomys cf 0. leucogaster (Wied-Neuwied) , Stenotrem leai (Binney) northern grasshopper mouse Ondatra zibethica (Linnaeus) , muskrat Microtus pennsylvanicus (Ord) , meadow vole CLASS OSTEICHTHYES Zapus hudsonius transitionalis Klingener , meadow jumping mouse Catostomus commersoni (~ac6p&de),white sucker Order Carnivora Semtilus cf S. atrornaculatus (Mitchill), creek chub a large form, family indeterminate cf Hybopsis pacilis (Richardson), flathead a small form, family indeterminate chub Pimephales promelus Rafinesque, flathead Order Proboscidea minnow Cyprinidae sp. Mammuthus sp. , mammoth Ictalurus melas (Rafinesque), black bullhead Lepomis cf L. cyanellus Rafinesque, green Order Lagomorpha sunfish Perca flavescens (Mitchill), yellow perch Lepus sp. , hare

Order Artiodactyla CLASS AMPHIBZA Order Salienta Camelops sp. , camel Antilocaprid sp. , pronghorn Bufo 'Americanus groupn, toad Bison cf B. latifrons (Harlan), giant bison

CLASS REPTILIA Order Perissodactylsr Order Squamata Equus cf E. niobmrensis Hay, horse Ophisaums attenuatus Baird, glass lizard Equus sp. , horse FIVE ILLINOIAN FAUNAS 183

TABLE 2. Summary of local habitats represented by the mollusks of the Doby Springs Local Fauna.

I I Locality and Habitat Species

Semiaquatic: among vegetation and Oxyloma sp. 4 debris near water's edge.

Hygrophilic: moist situations in leaf Cayichium exiguum

C mold, under sticks and debris; shaded Vertigo ovata areas, not far from water. V. milium V. elutior Gastrocopta tappaniana Deroceras aenigma

Woodland: moist areas under leaf litter, I Cionellu lubrica down timber, among tall marsh grass. I Stenotrema leai Discus cronkhitei Zonitoides arboreus Nesovitrea electrina Euconulus fulvus Strobilops lubyrinthica Helicodiscus parallelus

Sheltered areas: among rocks, shrubs, Gastrocopta armifera grass, or in timbered situations. G. contracts G. holzingeri

Sheltered situations: These species are Gastrocopta cristata not restricted to a woodland habitat G. procera and can tolerate drier conditions. Pu$illu blandi P. muscorum Pupoides albilubris Hawaiia minuscula Vallonia pamula V. gracilicosta

Marginal situations : wet mud, sticks, Fossariu obrussa stones, or any other debris along F. dalli water's edge; shallow pools and-other protected spots.

Shallow quiet water: small streams, Pisidium casertanum

Y ponds, sloughs or marsh, that may Sphaerium occidentale become dry during part of the year. Stagnicola reflexa S. caperata Armiger crista Gyraulus circumstriatus G. deflectus Promenetus umbilicatellus Physa gyrina Aplexu hypnorum Ferrissia fragilis 184 B. B. MILLER

Table 2 (cont. )

Locality and Habitat Species Abundance* 1245

Shallow quiet water: small ponds, Uniomerus tetralasmus R- streams, sloughs, or marsh, with no Lymnaea stagnalis jugularis R - R R current or areas of rooted vegetation Sphaerium rhomboideum R-R- with little current; soft sand or mud S. sulcatum S-AR bottoms; not subject of significant Pisidium obtzlsale R-RR seasonal drying. Gyraulus parvus ACAA Helisoma trivolvis - -RR Promenetus exacuous form kansasensis A-AA Phy sa anatina R-A- P. skinneri R-A- Laeva9ex fuscus --s-

Perennial water: Stream or lake with Sphaerium striatinum --c- slow to moderate current; areas of Pisidium compressum ARAA still water; shallow spots with soft P. nitidum C AA sand or mud substrate; not affected P. variabile A-CA by seasonal drying. Valvata tricarinata A SAA Helisoma anceps SRAA Probythinella lacustds A- Anodonta cf A. grandis R-

*R=<25; S= 26-49; C= 50-100; A=)100 shells. **These specimens were questionably referred to species.

local fauna is considered Illinoian inage Habitat. Stephens (1960) recognized on the basis of both stratigraphic position 4 habitat associations in the vertebrate and vertebrate fossils and has been fauna: a lake and marsh border com- correlated with the Berends and Butler munity; a lowland meadow community; Spring local faunas (Smith, 1958; a shrub and tree community; and an Stephens, 1960; Hibbard & Taylor, upland prairie community. 1960). However, studies of the pollen, Table 2 summarizes the habitat as- collected from samples of sediments sociations indicated by the Doby Springs associated with these assemblages molluscan fauna. The abundant repre- (Kapp, 1965), indicate that only locality sentation of Probythiwlla lacustris is 5 of the =~obySprings local fauna is interpreted as indicating that the lake equivalent to the Berends and the lower in which the sediments containing the part of the section from whichthe Butler Doby Springs local fauna were deposited Spring local fauna was recovered. This probably had a depth of at least five ft. P lower portion of the Butler Spring local The carbonaceous dark-gray to black fauna (localities 2, 3, 4 and 5 of Hibbard silty clay and silt from which P. lacus- & Taylor, 1960), has been designated tris (Hibbard & Taylor, 1960: 81) was the Adams local fauna by Kapp (1965). recovered, considered together with the The predominantly northern and north- occurrence of Pisidium variabile which eastern aspect of the Doby Spring mol- prefers a bottom on which soft sediments luscan fauna indicates cool, moist, are accumulating, suggests that the sub- glacial conditions, and is compatible strate, in part, consisted of a soft organic with the interpretations based on the mud. Firmer patches of bottom, com- pollen, and vertebrates. posed of sand and silt are indicated by FIVE ILLIN()IAN FAUNAS 185

Sphaerium striatinum. lived at a time when summer conditions The shallower portions of the lake were no hotter or drier than those near shore contained dense patches of presently occuring in northern North immersed aquatic vegetation which Dakota and Minnesota, near the southern probably included such species asSagit- limit of range for Pupilla muscomm. taria, Myriophyllum and Pediastmm The winters were most likely similar (Kapp, 1965). These plants provided to those in northeasternNebraska-north- suitable habitats for Sphaerium sul- western Iowa, the area in which the catum, S. rhomboideum, Valvata tri- majority of the Doby Springs molluscan carinata, G yraulus parvus, Promenetus species presently occur. Some of the exacuous form kansasensis and Physa more easterly (e.g., Laevapex ficscus), anutina. and southerly (e.g., Gastrocopta cris- k Temporary bodies of water in the tata, G. procera) distributed species, marsh area along the margin of the lake are at or near the periphery of their contained Pisidium casertanum, Sphaer- ranges in this region. Summers in the ium occidentale, Stagnicola caperata, S. northeastern portion of North Dakota are reflexa, Lymnaea stagnulis jugularis, characterized by normal temperatures Armigev crista, Gyraulus circum- of between 65' - 700 F (Vischer, 1954: stm'atus, G. deflectus, Promenetus um- map 5). The normal annual precipitation bilicatellus, Physa gyrina, Aplexa hyp- in this area is about 18-20 in. (Visher, nomm and Fevrissia fragilis. Rest 1954: map 492), slightly less than in marks on some of the P. casertanum Harper County, Oklahoma (Stephens, and the presence of some septate shells 1960). The greater part of this pre- amongst the Ferrissia, indicate that cipitation in North Dakota, however, some of the water situations were sub- occurs during the summer months, when jected to periodic drying. it is most needed. Cooler summers The terrestrial element in the mol- and lower evaporation rates, in the North luscan fauna is dominated by hygrophilic Dakota region, make the smaller amount species, which also lived in the marsh of rainfall more effective and have made near the lake margin. The vegetational it possible for a relatively large mol- cover postulated for this area by Kapp luscan fauna to maintain itself. The (1965), would have provided suitable normal winter temperature in the north- habitats for most of the terrestrialmol- eastern Nebraska-northwestern Iowa lusks listed. area, is between 15O-20° F (Visher, Gastrocopta cristata, G. procera, 1954: map 4). Pupilla blandi, P. muscomcm, Pupoides The inferred climate inHarper County, a Zbilabris, Hawaiia minuscula, Vallonia Oklahoma, at the time the Doby Springs pavvula and V. gracilicosta are terres- local fauna lived, combined shorter, trial species that can tolerate drier cooler summers, with winters that were situations and probably in part lived slightly cooler and longer than those amonst the better drained shrub and tree presently prevailing in this area. The community further back from the lake. normal summer temperature was Climate. Fifty-six of the 59 molluscan probably between 65' - 70' F. The forms present in the Doby Springs local normal annual precipitation was about '' are identifiable to species that are now 18-20 in., 8-10 in. of which fell during extant. Fourty-six of these species the summer months. The normal evapo- have been reported from the Lake Okobo- ration during the warmer part of the ji region of Iowa. There is, however, no year was between 30-35 in., compared one geographic locality known to me at with today's 50-60 in. in Harper County which all the species associated in the (Visher, 1954: map 483). The normal Doby Springs assemblage occur. winter temperatures were probably The Doby Springs local fauna probably between 15O - 20° F. 186 B. B. MILLER

Adams Local Fauna Order Basommatophora

Previous work. The term Adams local Stagnicola caperata (Say) fauna was proposed by Kapp (1965) for Fossaria dalli (Baker) the fossils recovered from the sequence Gyraulus parvus (Say) of medium to coarse, limonitic, cross- Promenetus exacuous form kansasensis bedded sands which occur stratigraph- (Baker) ically below the Butler Spring local Physa anatina Lea fauna (Hibbard & Taylor, 1960), in the Order Stylommatophora SE 1/4 sec 32, T 34 S, R 29 W, Meade County, Kansas. Kapp found that there Gastrocopta cristata (Pilsbry & Vanatta) was a sharp change in the frequency of G. procera (Gould) pine, grass and composite pollen, G. tappaniuna (Adams) between the top and bottom of unit 1, Pupilla blandi Morse measured section 2 of Hibbard & Taylor P. muscovum (Linnaeus) (1960: 43). Within this interval a trend P. sinistra Franzen towards increasing pine and decreasing Vevtigo milium (Gould) grass and composites is reversed, and V. ovata (Say) there is a sharp rise in the frequency Vallonia cyclophorella Sterki of the grass pollen with a concomitant V. gracilicosta Reinhardt decline in pine pollen. Kapp suggests cf Succinea that this striking reversal may repre- Discus cronkhitei (Newcomb) sent an unconformity between a cool, Helicodiscus singleyanus (Pilsbry) moist and a drier and/or warmer inter- Hawaiia minuscula (Binney) val. This sharp transformation in the composition of the pollen spectrum CLASS AMPHIBIA occurs at a point in the section that is Order Caudata stratigraphically between locality 1 and localities 2, 3, 4 and 5 of the Butler Ambystoma tigrinum (Green), tiger Spring local fauna (as used in Hibbard salamander & Taylor, 1960), and forms the basis for the separation of the Adams and CLASS REPTILIA Butler Spring local faunas. In this Order Chelonia report the Butler Spring local fauna is restricted to locality 1 and the Adams Emydoidea twentei (Taylor), Plains semibox local fauna to localities 2, 3, 4 and 5. turtle

CLASS MAMMALIA Faunal List; Adams Local Fauna: Order Edentata

C=LASS PELECYPODA Megalonyx sp. , ground sloth Order Prionodesmacea Order Rodentia *Lasmigona complanata (Barnes) Quadrula quadvula (Rafinesque) Citellus sp. , ground squirrel cf Geomys , eastern pocket gopher CLASS GASTROPODA Castoroides cf C. ohioensis Foster, Ohio Order Mesogastropoda giant beaver Microtus pennsylvanicus (Ord), meadow vole Valvata tricarinata (Say) Order Proboscidea

*Species not previously reported Mammuthus sp. , mammoth FIVE ILLINOIAN FAUNAS 187

Order Artiodactyla represented by Doby Springs local fauna locality 5. Camelops sp. , camel Habitat. The medium to coarse, cross- Order Perissodactyla bedded sands from which the fauna was recovered (Hibbard & Taylor, 1960), Equus scotti Gidley, horse together with the occurrence of Valvata Equus sp. , horse tricarinata, Gyraulus paruus, Quudmla quudmla and Lasmigona complanuta, Age and Correlation. Kapp (1965)ten- species requiring continuity of aquatic tatively correlated the Adams local fauna habitat, are interpreted as indicatingthe with locality 5 of the Doby Springs and existence of a stream. The presence of with the Berends local faunas on the only one wood-land species among the . occurrence of spruce pollen. The occur- terrestrial gastropods suggests that rences of spruce at these 3 sites are the there was only a thin covering of trees only unequivocal records of this pollen and shrubs, rather than a continuous entity in the Meade County area. The forest belt along the stream. Adams and Butler Spring molluscanfau- Climate. The Adams local fauna con- nas, however, are more closely related to tains many species presently living in each other, than is either one to the Doby the northern Great Plains. Valvata tri- Springs. All of the species found in the carinata, Pupilla blandi, P. muscorum, Adams local fauna, with the exception of P. sinistra and Vallmia cyclophorella, the unionid clams, also occur inthe But- occur in the Great Plains only in the ler Spring assemblage. Three species, northernmost states, in the Rocky Vallonia cyclophorella, Pupilla sinistra Mountains, or in the more humid Central and He licodiscus single yanus are present Lowland where they may range farther in the Adams and Butler Spring local south, but are still of obviously northern faunas, but are absent from the Doby distribution (Hibbard & Taylor, 1960). Springs local fauna. This has beeninter- A relatively southern element repre- preted to mean that the Doby Springs sented by Gastrocopta procera, G. cris- mollusks probably lived in the area tata, Physa anutina and Helicodiscus earlier in the Illinoian, before these singleyanus, reaches as far north as speciesreachedthis part of the Plains. South Dakota or Nebraska in the Great The mammals (Hibbard, 1963), the Plains, but is more common farther pollen (Kapp, 1965) and the mollusks south. Many of these northern and south- (Miller, this report) indicate a trend ern species are not now found together towards a warmer climate between the in the same area and it is assumed time the Doby Springs fauna and the that they are presently being kept apart Mount Scott fauna lived. This trend was primarily by climatic extremes. The characterized by a decrease in spruce climate during that part of the Illinoian and pine pollen and an increase in the when the Adams local fauna livedproba- - number of southern species in the faunas bly combined winters which were perhaps of the area. The first Illinoian ap- no more severe than those presently pearance in this region of Helicodiscus occurring innortheasternNebraska, with singleyanus, a species with a relatively cooler, more moist summers, similar T southern distribution, is in the Adams to those in eastern Montana and the local fauna and suggests that this assem- Dakotas. blage lived in the area sometime after The normal winter temperatures were this warming trend began. The strati- probably between 15O - 20' F, with graphic position of the Adams local fauna normal summer temperatures of 65O - indicates that it lived before the Butler 70° F (Visher, 1954). The precipitation Spring local fauna, probably sometime was perhaps similar to that now after the Illinoian glacial maximum occurring in Meade County, about 18.43 B. B. MILLER in. per year (Frye, 1942), but because of Probythinella lacustris (Baker) the lower evapotranspiration rates during the warm months (Visher, 1954), Order Basommatophora it was more effective. Cary chium exiguum (Say) Butler Spring Local Fauna Lymnaea stagnalis jugularzs Say Previous Work. The name Butler Stagnicola caperata (Say) Spring local fauna was first used by S. rej7exa (Say) Fossaria obrussa (Say) Herrington & Taylor (1958: 4) for the F. dalli (Baker) fossils- recovered from one locality in Anisus patte~soni(Baker) the SE 1/4 sec 32, T 34 S, R 29 W, Gyraulus circumstriatus (Tryon) Meade County, Kansas. Smith (1958) G. parvus (Say) reported on the fishes fromthis locality. Helisoma anceps (Menke) Hibbard & Taylor (1960) discussed the H. trivolvis (Say) stratigraphy of the area and described Promenetus exacuous form kansasensis additional mollusks and vertebrates from (Baker) this locality and from 4 additional sites p. umbilicatellus (Cockerell) which were included as part of the Butler Ferrissia fYagilis (Tryon) Spring local fauna. Analysis of the pollen Laevapex fuscus (Adams) from the sequence of sediments con- Physa anatina Lea taining the Butler Spring assemblage P. gyrina Say (Kapp, 1965) indicates that there are P. skinneri Taylor significant differences between locality Aplexa hypnovum (Linnaeus) 1 and localities 2, 3, 4 and 5 of Hibbard & Taylor (1960). In this report the Order .Stylommatophora Butler Spring local fauna is restricted to include only the fossils reportedfrom Gastrocopta armifera (Say) G. contracts (Say) locality 1 (U. S. GeologicalSurvey Ceno- G. cristata (Pilsbry & Vanatta) zoic locality 21042) of Hibbard & Taylor G. holzingeri (Sterki) (1960). G. procera (Gould) G. tappaniana (Adams) Faunal List, Butler Spring Local Fauna: Pupoides albilabyis (Adams) P. inornatus Vanatta CLASS PELECYPODA Pupilla muscorum (Linnaeus) Order Prionodesmacea P. blandi (Morse) P. sinistra Franzen Anodonta grandis (Say) *Vertigo elatior (Sterki) *Ligumia cf L. recta (Lamarck) V. milium (Gould) V. ovata (Say) Order Teleodesmacea Vallonia cyclophorelh Sterki V. gracilicosta Reinhardt (Lamarck) Sphaerium striatinum V. pamula Sterki S, transversum Say cf Succinea Sphuerium, indeterminate Oxyloma sp. (Poli) Pisidium casersfanum Discus cronkhitei (Newcomb) Prime P. compressum Helicodiscus singleyanus (Pilsbry) P. nitidum Jenyns Hawaiia minuscula (Binney) P. mlkeri Sterki ? Zonitoides arboreus (Say) Stenotrema leai (Binney) CLASS GASTROPODA Order Mesogastropoda

Valvata tricarinata (Say) "Species not previously reported FIVE ILLINOIAN FAUNAS 189

CLASS OSTEICHTHYES which the Mount Scott local fauna lived. These data suggest a trend toward a Lepisosteus sp. , gar slightly warmer climate between the time Catostomus commersoni (Lac6p&de), white the Doby Springs and Mount Scott local sucker faunas lived. The climatic change pro- Ictalums cf I. punctatus (Rafinesque), duced a diminution in the abundance of channel catfish certain of the dominant northern ele- I. melus (Rafinesque) , black bullhead ments present in.the Doby Springs local Perca flavescens (Mitchill) , yellow perch , fauna and permitted Helicodiscus sing- leyalvus to enter the area by the time CLASS REPTILIA Order Chelonia the Adams and Butler Spring local faunas . lived. The increase in southern faunal Chelydra serpentina (Linnaeus) , snapping elements within the Mount Scott local turtle fauna is interpreted as a continuation of this climatic change toward slightly CLASS MAMMALIA warmer conditions. In this context the Order Rodentia Butler Spring and Adams assemblages appear to have lived after the Doby Microtus pennsylvanicus (Ord) , meadow vole Springs and before the Mount Scott local faunas. Rejected Records Habitat. The Butler Spring fauna lived Vertigo elutior = Vertigo gouldi; Oxyloma in or near a medium sized river, proba- sp. = Oxyloma retusa. bly the ancestral Cimarron River (Hibbard & Taylor, 1960). This river Age and Correlation. Examination of had a slow to moderate current, with a the mollusks from the Mount Scott (p 189 probable depth of several f eet. Protected et seq.), Butler Spring and Doby Springs areas of the river, with less current, local faunas, suggests that the Butler contained dense beds of submergent Spring is intermediate between the other aquatic vegetation. The terrestrial 2 in terms of first and last occurrences gastropods lived among scattered groups of species. The first Illinoian occur- of trees along the river, but the pre- rences of the 4 northern species Pro- dominant vegetation consisted mostly of bythinella lacustvis, Lymnaea stagnulis grasses and shrubs. jugulavis, Vevtigo elatior and Pupilla muscorum, in this region, is in the Doby Climate. The climate suggested by Springs local fauna. These species con- the Butler Spring local fauna (Hibbard tinue in the area through the time repre- & Taylor, 1960) consisted of cooler sented by the Butler Spring local fauna summers than now occur in Meade but in greatly reduced relative numbers. County. These conditions are similar The absence of the first 3 of these to those now occuring in the eastern species from the Adams local fauna is North Dakota region. Winters were possibly due to the smaller size of the probably no more severe than those in sample from which this assemblage was north-central Nebraska. ' recovered. (The Adams local fauna was sorted from approximately 10 pounds of Mount Scott Local Fauna of matrix, whereas the Butler Spring Previous work. In the summer of local fauna was picked from about 750 1953, Claude W. Hibbard and members pounds of matrix [Hibbard & Taylor, of the University of Michigan Museum of 19601). The Adams assemblage also in- Paleontology field party opened aquarry cludes the first Illinoian occurrence of (UM-K4-53) on the north side of Spring Helicodiscus singleyanus, a relatively Creek, a little over 2 miles upstream southern species which persists locally from Mount Scott, Big Springs Ranch, through that portion of the Illinoian during Meade County, Kansas. A total of 5 190 B. B. MILLER

tons of matrix was removed from this Order Teleodesmacea quarry during the summers of 1953, 1957 and 1960, and has yielded a large Sphuerium lacustre (Miiller) biota containing pollen, ostracods, mol- S. occidentale Prime lusks, fishes, amphibians, reptiles, birds S. partumeium (Say) and mammals. A second locality (UM- S. rhomboideum (Say) K2-59) opened near the base of Mount S. striatinurn (Lamarck) Scott, was worked during the 1959 and S. sulcaturn (Lamarck) 1960 field seasons. Two additional S. transversum (Say) localities (UM-K1-60 and UM-K3-60), Pisidium. casertanum (Poli) were discovered along Spring Creek and P. compressurn Prime Hart Draw, during the summer of 1960. P. nitidum Jenyns The ostracods from localities UM-K1- P. obtusale (Lamarck) P. subtruncatum Malm 60 and UM-K2-59 (Gutentag, written P. variabile Prime communication); the mollusks from P. walkeri Sterki locality UM-K4-53 (Miller, 1961); some of the reptiles from locality UM-K2-59 CLASS GASTROPODA (Etheridge, 1961); the fishes (Smith, Order Mesogastropoda 1963) and mammals (Klingener, 1963; Hibbard, 1963) from localities UM-K4- Valvata tvicarinata (Say) 53, UM-K1-60 and UM-K2-59; and the pollen (Kapp, 1965) from UM-K4-53, Order Basommatophora UM-K1-60, UM-K2-59 and UM-K3-60, represent all the fossil remains re- Carychium exisurn (Say) covered from these collections that have Stagnicola caperata (Say) thus far been studied. The name Mount S. exilis (Lea) Scott local fauna was first used by S. refilexa (Say) Etheridge (1961: 181) for the fossil Fossaria dalli (Baker) material recovered from locality UM- F. obrussa (Say) K2-59, near the base of Mount Scott. Armiger crista (Linnaeus) Gyraulus circumstriatus (Trj The following faunal list summarizes G. parvus (Say) the above and includes all the mollusks Helisoma anceps (Menke) from localities UM-K1-60, UM-K2-59 H. trivolvis (Say) and UM-K3-60, none of which have been Promenetus exacuous form previously reported. exacuous (Say) P. umbilicatellus (Cockerell) Faunal List; Mount Scott Local Fauna: Fervissia fragilis (Tryon) Laevapex fuscus (Adams) Locality Physa anatina Lea mom0 mwmw P. gyrina Say Species &A&& P. skinnevi Taylor YYY? Aplexa hypnorum (Linnaeus) zzzz DDDD Order Stylommatophora CLASS OSTRACODA Cionella lubrica (Miiller) Candona reonensis Gutentag & Strobilops labyrinthica (Say) Benson x x Gastrocopta amifera (Say) Cypridopsis vidua (0. F. Miiller) x x G. contracts (Say) Candona sp. Yav x G. cristata (Pilsbry & Vanatta) G, holzingeri (Sterki) CLASS PELECYPODA G. pellucida hordeacella (Pilsbry) Order Prionodesmacea G. procera (Gould) G. tappaniana (Adams) Uniomerus tetralasmus (Say) x*x x Pupoides albilabris (Adams) FIVE ILLINOLAN FAUNAS

Locality CLASS AMPHIBIA m 0 a 0 Order Salienta L? w L9 w 1111 Species *3NC3 % g g Bufo sp. , toad 6 ?Acris sp., frog a a a Rana pipiens Schreber , leopard Pupilla blandi Morse x x frog vertigo milium (Gould) xxx . V. ovata (Say) xxxx CLASS REPTILJA Vallonia gracilicosta Reinhardt x x x x V. parvula Sterki Order Chelonia cf Succinea xxxx Oxyloma sp. Terrapae llunensis Oelrich, Mscus cronkhitei (Newcomb) xxxx plains box turtle Helicodiscus parallelus (Say) xxxx H. singleyanus ,(Pilsbry) x Order Squamata Punctum minutissimusn Lea x Deroceras aenigma Leonard Ophisaurus attenuatus Baird, Euconulus fulvus (Miiller) xxx glass lizard Nesovitrea electrina (Gould) x x Hawaiia minuscula (Binney) xxxx CLASS AVES Zonitoides arbo?*eus (Say) xxxx 2. nitidus (Miiller) Unstudied Steaotrema leai (Binney) x x CLASS MAMMALIA CLASS OSTEICHTHYES Order Insectivora Lepisosteus platostomus Rafines- que, Shortnose gar x x Sorex cinereus Kerr , masked shre~% Esox masqz~inongyMitchill , S. arcticus Kerr , arctic shrew muskellunge x S. palustris Richardson, Hy bognathus hankinsoni Hubbs , northern water shrew brassy minnow x Blarina brevicauda carolinensis Campostoma anomalunz (Rafines- (Bachman), southern shorttail que) , stoneroller x x shrew Semotilus atromaculatus (Mitchill), Cqptotis parva (Say), least shrew creek chub xxx Dionda nubila (Forbes), ozark Order Chiroptera minnow x Cyprinidae , indeterminate x x Lasiuns cinereus (Beauvois) , Ictiobus, sp. Rafinesque, hoary bat buffalo fish x Catostonzus commersoni Order Rodentia (Lac kp'ede) , white sucker x x iMoxostoma duquesnei (LeSueur) , Cynomys sp. , prairie dog black redhorse x Citellus cf C. tridecemli~zeatus Ictalurus punctatus (Rafinesque) , (Mitchill), ground squirrel channel catfish x Citellus sp. , ground squirrel I. melas (Rafinesque) , black Geomys sp. , pocket gopher bullhead x x Paradipoides or Castor sp. , Microptens salmoides beaver (Lackp'ede) , largemouth bass x x Reithrodontom3ls sp. , harvest Lepomis cyanellus Rafinesque, mouse green sunfish x peyomyscus besendseais L. humilis? (Giarard), orange- Starrett , deer mouse spotted sunfish x P. prog7

Locality centrate from these collections has been sorted for mollusks. SW 1/4 sec 13, Species Locality UM-K1-60. T 32 S, R 29 W, approximately 528 ft north and 396 ft east of the southwestern section corner, about 1/2 mile down- Neotoma cf A'. floridana (Ord) , stream from locality UM-K4-53. The packrat x faunule from this locality was removed Oryzomys fossilis Hibbard, fossil from the sand, silt and marl, repre- -. rice rat x x sented by units 1-3, in Measuredsection Synaptonzys australis Simpson, 4 (see p 202). Seven tons of matrix Simpson's bog lemming were washed from this site during the Ondatra zibethica Linnaeus , /x/XIXI 1 1960-1961 field seasons. Only about muskrat, 12% of this concentrate has been sorted Micvotus pennsylvanicus (Ord), for mollusks. meadow vole Locality UM-K2-59. SE 1/4 sec 18, Pedomys ochrogastev (Wagner), T 32 S, R 28 W, 396 ft north and 1386 prairie vole Zapus hudsonius tvansitionalis ft west of the southeastern section Klingener , meadow jumping corner, a little over 2 miles downstream mouse on Spring Creek, from locality UM-K4- 53. The fossils were collected from a Order Carnivora coarse, greenish brown silt (unit 8 of IIIII Measured Section 3, see p 202), ex- Mustela vison Schreber , mink posed in a quarry excavated on the lower slope of Mount Scott, approxi- Order Lagomorpha mately 23 ft above the bed of Spring Sylvilagus sp. , cottontail rabbit Creek. Twenty-seven tons of matrix Lepus sp. , hare have been removed from this locality, less than 5% of which has been sorted Order Artiodactyla for mollusks. Locality UM-K3-60. NE 1/4 SW 1/4 Bison cf B. latif~ons(Harlan), sec 17, T 32 S., R 28 W, about 1980 ft giant bison north and 924 ft east of the southwestern section corner. The fossils occur in dark brown, silty clay, near the base of Localities. The Mount Scott local Hart Draw, downstream from and fauna is known from the following 4 approximately 40 ft below locality 1 of localities, situated on the Big Springs the Cragin Quarry local fauna (Hibbard ranch, Meade County, Kansas (Figs. 2 & Taylor, 1960). The mollusks repre- and 3): sent all the fossils recovered from Locality UM-K4-53. SE 1/4 sec 14, about 10 pounds of matrix. T 32 S, R 29 W, approximately 1914 ft Local Stratigraphy. A sequence of north and 1254 ft west, of the south- sediments ranging in age from upper eastern section corner. The fossils Pliocene to Recent is exposed in the -r were recovered from a greenish-brown Big Springs Ranch area (Fig. 1). The silty clay (unit 3 in. Measured Section Rexroad, Ballard and Crooked Creek 1, see p 200), 2-3 ft above the stream formations represent cycles of wide- bed of Spring Creek (Miller, 1961: 104). spread sheet deposition, in which Five tons of matrix have been removed aggrading streams filled their valleys, from this locality since it was first spreading sands, silts and clays across worked in 1953. About 10% of the con- the interstream divides. The Kingsdown FIVE ILLINOIAN FAUNAS

- - /'.. , \.:j .I ". ., \,.. P.../" T .I 30 S

I US 54-160 T 3 2 S

T 33 S KAN 96

T 3 4 S

L I

FIG. 2. Map of Meade County, Kansas showing location of Big Springs Ranch area (diagonal lines).

and Vanhem formations in this area, Rexroad Formation. The upper Plio- by contrast, occur as valley fills or cene in this area is represented by the terrace deposits that are usually situated Rexroad Formation, exposed along the topographically below the older beds. south bank of Spring Creek and the north At many localities, the beds are ob- and south sides of Hart Draw, inportions scured by slopewash and wind blown sand, of sections 13,14,24, T 32 S, R 29 W, and and good exposures are usually re- sections 7, 18, 19, T 32 S, R 28 W. Good stricted to the relatively few steepbluffs outcrops of the Rexroad are com- - occurring on the outside of meanders. paratively rare, and are usually re- Where the bedding of the sediments could stricted to the base of bluffs along the be observed, it showed a very low main stream valleys where erosion has apparent dip, the greatest angle been deep enough to expose the upper measured not exceeding 4O. All these portion of the formation, There are difficulties made mapping difficult and 2 exceptions to this usual mode of accurate measurement of strike and dip occurrence: the first is an outcrop, impossible. composed of approximately 3 1/2 ft of B. B. MILLER

FIG. 3. Geologic map of the Big Springs Ranch area, Meade County, Kansas. FIVE ILLINOIAN FAUNAS 196 B. B. MILLER

orange, yellow and green sands and Creek, in SE 1/4 NW 1/4 sec 19, T 32 S, clays, exposed in a low bank along a R 28 W. At this locality, the Ballard tributary of Spring Creek in the SW 1/4 is 42.5 ft thick and has an apparent dip NW 1/4 sec 24, T 32 S, R 29 W; the of about 4O east (see Measured Section second is an exposure in the floor of 5, p 203). the Sanders Gravel pit, SE 1/4 NW 1/4 Hibbard (1958) introduced the name sec 7, T 32 S, R 28 W. Angell member, for the basal portion of The thickest section of Rexroad ex- the Ballard Formation, designating the amined in the area outcrops on the 13.7 ft of cross-bedded, partly cemented, south side of Hart Draw, in the NW 1/4 sand, gravel and cobbles, exposed near NE 1/4 sec 18, T 32 S, R 28 W. It is the base of the Sanders Gravel pit in composed of 23 ft of reddish-gray to SE 1/4 NW 1/4 sec 7, T 32S, R 28 W, as gray sandy silt and fine sand, capped the type section. . by one foot of caliche. This exposure Differences in amount of cementation yielded the mollusks reported by Taylor have produced some lateral variation in (1960), from locality 1 of the Rexroad the appearances of the Angell member. local fauna. A second occurrence of Three different modes of occurrence mollusks in the Rexroad has been re- have been recognized. In its most com- corded by Taylor (1960) from deposits mon form, it is a well indurated, cross- exposed along the south side of Spring bedded, dark gray sand and gravel, Creek in the NW 1/4 NE 1/4 sec 24, T stained with manganese and iron, best 32 S, R 29 W. displayed in outcrops along Spring Creek Ballard Formation. The name Ballard and its tributaries, in sections 11, 12 Formation was proposed by Hibbard and 24, T 32 S, R 29 W, and in sections (1958) as a repkcement for Meade 7 and 8, T 32 S, R 28 W. Less fre- Formation, which had undergone a quently, the Angell occurs as a partially variety of usages (see summaries in cemented, cross-bedded, yellow-buff Hibbard, 1955, Chart I; and Hibbard sand and gravel, exemplified by ex- & Taylor, 1960, fig. 2). The type posures in the NE 1/4 SE 1/4 sec 14, locality of the Ballard Formation is T 32 S, R 29 W, and NW 1/4 SE 1/4 situated on the Big Springs Ranch, onthe sec 19, T 32 S, R 28 W. A third type south side of Hart Draw, in Sections 7 of occurrence, in which the deposit is a and 18, T 32 S, R 28 W (Hibbard, 1958). yellowish-tan, unconsolidated sand and At the type locality, the Ballard rests gravel, was observed in the SW 1/4 unconformably on the Rexroad. It is NW 1/4 sec 24, T 32 S, R 29 W. relatively thin, consisting of about 8 ft Fossils are not abundant in the Angell of cemented basal sand and gravel, over- member although some vertebrate re- lain by 12 ft of buff-gray silt and clay. mains have been collected from ex- The upper 3 ft of the Ballard at this posures at the Sanders Gravel pit exposure yielded the mollusks and mam- (Hibbard, 1956a). mals comprising locality 1 (UM-K1-53) The Missler member of the Ballard of the Sanders local fauna (Hibbard, Formation (Hibbard, 1949a) is the 1956a; Taylor, 1960). reddish-buff sand and silt overlying the The Ballard Formation occurs along Angell sand and gravel sections in 6 and the valleys of Spring Creek and Hart 7, T 32 S, R 28 W. Good outcrops, - Draw in portions of all the mapped where the member is as much as 35 ft sections of the Big Springs area. thick, occur along Spring Creek and its Wherever the base of the Ballard is tributaries in sections 12, 14 and 23, clearly exposed, it rests unconformably T 32 S, R 29 W, and in sections 7, on the Rexroad. The maximum thickness 8 and 19, T 32 S, R 28 W. The Missler, of the Ballard measured in the area is where it is well exposed because of the in a bluff along the south side of Spring steepness of slope, usually appears as FIVE ILLINOIAN FAUNAS 197

reddish-brown and green sands, silts Vestiges of what seem to be the channel and clays, interbedded with zones of or channels, in which the Stump Arroyo caliche nodules and stringers. A well member was deposited, crop out along indurated, dense, white caliche, about the tributaries of Spring Creek in 6 in. thick, often occurs near the top of sections 10, 11, 14 and 15, T 32 S, R 29 this member. W. Some of the exposures reveal Crooked Creek Formation. The channels at least 12 ft deep, as evi- Crooked Creek Formation as defined bv denced by the relief on the Stump Arroyo- / Hibbard (1949a), includes those sedi: Missler contact. ments which were laid down during the The only fossils collected from the cycle of deposition which followed the Stump Arroyo member inthe Bigsprings Ballard Formation. It is by far the Ranch area are a lower molar (UMMP most widespread formation exposed in 3 1352*) and astragalus (UMMP 31353) the study area, and reaches a thickness of Nannippus sp., from the N 1/2 sec 14, locally of at least 61.5 ft. T 32 S, R 29 W. The basal Stump Arroyo sand and Exposures of the Atwater member of gravel member of the Crooked Creek the Crooked Creek Formation occur in Formation is extensively exposed in all sections of the mapped area. With portions of all sections of the map area, the exception of the Kingsdown For- reaching a thickness of up to 31.5 ft. mation exposed in the S 1/2 sec 17, T It occurs most frequently as an uncon- 32 S, R 28 W, this member forms most solidated, reddish-tan sand and pebble of the upland surface. The thickest rubble. The gravel contains a dis- section of Atwater measured consists of tinctive milky white quartz component, 3 1.25 ft of reddish-brown sand and silt, which in overall aspect makes the Stump intermixed with stringers and nodules of Arroyo appear lighter in color than the caliche and ash. It is exposed in the Angell sand and gravel. The sands and E 1/2 of sec 14, T 32 S, R 29 W (see gravels are generally not well sorted Measured Section 2, p 200). The Pearl- and cross-bedding is poorly preserved. ette ash, the most distinctive unit in Locally the Stump Arroyo occurs as a the Atwater when present, is well ex- lime cemented, cross-bedded, sand and posed in the NW 1/4 sec 18; SW 1/4 and gravel which grades upward to a NW 1/4 SE 1/4 sec 7, T 32 S, R 29 W, massive greenish-gray sand. The and in the NE 1/4 sec 14, T 32 S, R surface of the entire unit usually is 28 W. The greatest thickness of ash stained a dark gray. measured was a 160 in. section exposed The gray color of the Stump Arroyo in the NW 1/4 SE 1/4 sec 7, T 32 S, is possibly imparted by finely dissemi- R 28 W. The fossiliferous base of this nated particles of manganese and iron sequence is listed as locality 11, of the which have been reworked from Cudahy fauna, in Hibbard (1949a, map 2). weathered portions of the Angell mem- Getz (1960) has reported an astragalus ber of the Ballard. No pieces of re- of Canis sp. (UMMP 38406) from below worked Angell were observed at the a lentil of Pearlette ash exposed in the localities examined, although such an NW 1/4 sec 18, T 32 S, R 28 W. occurrence has been reported by Hibbard Kingsdown Formation. The Kingsdown (1951), in unconsolidated Stump Arroyo is represented in the vicinity of the in the Seger Gravel pit. The sand Big Springs Ranch by scattered, dis- presently being deposited in the beds of continuous remnants of stream channel Hart Draw and Spring Creek contains and floodplain deposits. It reaches its local concentrations of this black manganese and iron stain; here, too, the color is apparently derived from re- *University of Michigan Museum of Paleon- worked older beds. tology catalogue number. greatest development in the SE 1/4 sec caliche, are exposed along the valleys 18, T 32 S, R 28 W, at Mount Scott, on of the north-south tributaries of Spring the north side of Spring Creek, where Creek, in the S 1/2 sec 18. Lithology a thick sequence of at least 61 ft of and stratigraphic and topographicposi- gravel, sand, silt, clay and caliche are tion suggest that these deposits are exposed (see Measured Section 3, p 202). equivalent to the upper part of the Kings- Other outcrops of the Kingsdown occur down exposed at Mount Scott (see Meas- in the SW 1/4 sec 17, T 32S, R 28 W, on ured Section 3, p 202). the south side of Hart Draw; in the SW The lower portions of the Kingsdown, c. 1/4 sec 13, the SE 1/4 sec 14, T 32 S, exposed along the north side of Spring R 29 W, and the S 1/2 sec 18, T 32 S, Creek and the southwest side of Hart R 28 W, on the north side of Spring Draw, include sediments which contain Creek. the Mount Scott local fauna and its The absence of the Kingsdown For- stratigraphic equivalents. These sedi- mation from the south side of Spring ments consist primarily of sands, silts, Creek in section 19, to the southwest of silty clays and marls, deposited on the Mount Scott, may in part be the result floodplain of ancestral Spring Creek. of local sinkhole collapse. There is The Kingsdown exposed in a quarry at some evidence for at least one episode locality UM-K2-59, near the base of of late Pleistocene collapse that in- Mount Scott, contains at its base a 2 ft volved the Rexroad, Ballard and Crooked fossiliferous, greenish-brown to green Creek formations in this area. Unit 11 silt, representing a floodplain phase of of the Ballard Formation (see Measured deposition. Overlying these sediments, Section 5, p 203) is exposed some 42 ft is a 6.5 ft channel phase sequence of above the base of a high bluff on the cut and fill structures, composed of south side of Spring Creek, in the NW lenses of sand and gravel containing 1/4 sec 19. A green clay which appears cobbles up to 10 in. in length. The to be the same as unit 11, is exposed gravels appear to be composed of re- about 1/8 mile downstream near the base worked Stump Arroyo gravel and pieces of the stream. Lf the bed at these 2 ex- of Ballard caliche. The close proximity posures is the same there must have of the Kingsdown at this locality to known been approximately 42 ft of displacement. exposures of the Stump Arroyo-Ballard The Kingsdown and Crooked Creek contact (see Measured Section 3, p 202), formations in section 18, are both capped together with the size and high degree of by a caliche, which underlies the up- angularity of the caliche cobbles, all in- land surface. The contact betweenthese dicate a local origin for these coarser 2 formations is uncertain probably be- elastics, and by implication, that an- cause of the continued caliche accumu- cestral Spring Creek had become en- lation on the Crooked Creek during the trenched in the Crooked Creek For- Sangamon. The boundary has been mation, and had cut downward, at least questionably mapped alongthe outermost locally, to the Ballard caliche. occurrence of several caliche-capped A similar succession occurs on the hummocks which rise a short distance southwest side of Hart Draw (Hibbard above the adjoining Crooked Creek & Taylor, 1960: 29). A dark brown silt caliche. The tops of these hummocks, containing mollusks and pollen (Kapp, ). approximately at the same elevation as 1965) is exposed at locality UM-K3-60, the caprock at Mount Scott, have been near the base of Hart Draw, several interpreted as erosional remnants of hundred feet downstream from the what was formerly a more extensive section measured by Taylor (Hibbard Sangamon caliche bed. & Taylor, 1960: 29-30). These sedi- Kingsdown sediments, composed of ments are considered correlatives of red-brown silts capped in places by those at UM-K2-59 on the basis of FIVE ILLINOIAN FAUNAS 199

lithology and stratigraphic position, even Ballard and Rexroad formations to at though this relationship has not clearly least the present stream floor. These been established by paleontologic evi- channel fills are believed to be equi- dence. valent to the Vanhem exposed on the north side of Spring Creek, andprobably The remaining outcrops of Kingsdown represent the position of ancestral Spring exposed along Spring Creek, upstream Creek during a portion of the Wisconsin. from UM-K2-59, are floodplain deposits 1 similar to those at the base of Mount A younger cycle of post-Kingsdown Scott. The fossil-bearing units at locality UM-K1-60, however, are marls and cutting is represented by the present stream system. This valley cutting * sands which apparently represent a small lake or impoundment of the stream episode was discontinuous, as evidenced (Hibbard, 1963). locally by at least one terrace level approximately 6 ft above the present stream floor of Spring Creek and Hart Post-Kingsdown Sediments. At least 2 cycles of post-Kingsdown incision and Draw and by the series of terrace de- deposition are recognizeable in the Big posits of Late Wisconsin age located along the north side of the Cimarron Springs Ranch area. Evidence for the older channel cutting phase is recorded River (Hibbard & Taylor, 1960: 22, along the north bank of Spring Creek, in McLaughlin, 1946: 36; Byrne & Mc- Laughlin, 1948: 84). sections 13 and 14, T 32 S, R 29 W. Here the Kingsdown Formation has been channeled into, at places, to a depth of The entrenchment of the Recent at least 12 ft, and filled with fine gravel streams has not proceeded to the depth and light brown-buff sand and silt (Fig. of the earlier Illinoian and Wisconsin 4). These sediments were previously erosional phases, for the base of the interpreted as Wisconsin in age (Miller, Kingsdown and Vanhem formations are 1961: 105), and placed in the Vanhem not exposed. The valley walls along most Formation, even though, at the time, of the major stream valleys are usually there was no way of excluding the possi- not steep, and the light brown sands, bility of their Recent origin. Fossil silts and gravels of Recent alluvium, mollusks since recovered from the upper exposed in the beds of these streams, part of these deposits (UM-K2-60 located often grade imperceptibly into the slope in the SE 1/4 sec 14, T 32 S, R 29 W), wash and wind blown sand obscuring the include 6 species no longer living in older deposits. These sediments mask the Meade County area, thus supporting the Vanhem Formation over much of the the original assignment of the beds to area. The relatively few good exposures the Wisconsin Vanhem Formation. of the Vanhem occur in steep bluffs and are not frequent enough to permit deter- A thicker succession of sediments, mination of the aerial extent of this for- with similar lithology, is exposed in mation. These difficulties made it im- bluffs along the south side of Spring practical to separate the Vanhem from Creek and along some of its tributaries. * the younger Recent sediments, and the At some localities these deposits reach entire sequence of post-Kingsdown 51 ft in thickness, filling channels that deposits has consequently been mapped have been cut throughthe Crooked Creek, as one unit. 200 B. B. MILLER

Measured Sections

Measured Section 1. Section measured in SW 1/4 NE 1/4 SE 1/4 sec 14, T 32 S, R 29 W, on north bank of Spring Creek. The section is a composite, with the upper 7.15 ft measured about 85 ft upstream from the lower 6. 33 ft. Thickness Total of unit thickness Top of bank (in feet) (in feet) Vanhem Formation 8. Gravel, sand and silt, light brown-buff; channeled into gray clay below...... 2.25 2.25 Kingsdown Formation 7. Clay, light gray, with joints apparently dipping 430 W, 75O N and80°~...... 2.30 4. 55 6. Clay, "varved", with light and dark gray laminae...... 2.60 7.15 Measured section continued 8 5 ft upstream. 5. Clay, gray-black, with sandy limonitic nodules...... 2.20 9.35 4. Clay black, with caliche nodules...... 0. 75 10.10 3. Clay, silty, greenish-brown, grading upward into black clay (loc. UM-K4-53)...... 1.33 11.43 2. Clay, black, streaked with white flecks...... 0.25 11.68 1. Silty clay, greenish-brown on fresh exposures, gray- black when dry; limonite stains; shell zone in upper 15" (base not exposed)...... 1.80 13.48 Floor of Spring Creek

Measured Section 2. Started in the floor of Spring Creek, NW 1/4 SE 1/4 sec 14, T 32 S, R 29 W, Meade County, Kansas and measured along a northward traverse that ascends to the High Plains surface. Thickness Total of unit thickness Crooked Creek Formation (in feet) (in feet) Atwater member 11. Silts and caliche, gray-brown, containing caliche nodules and stringers...... 28.75 28.75 10. Volcanic ash (Pearlette ash), white, impure, containing reworked fragments of jasper, feldspar; contact above and below very irregular ...... 2.50 31. 25 9. Sandy silt, thin-bedded, thin clay layer on top, olive color with orange mottling...... 1.00 32.25 Stump Arroyo member 8. Gravel, with milky white quartz most distinctive component, gravel imbedded in gray- green sandy silt matrix, bed cemented in places...... 29. 50

Ballard Formation Missler member 7. Silt, brown...... 1.50 6. Clay, green...... 0. 80 5. Caliche, fine grained, well indurated...... 0. 50 FIVE ILLINOZAN FAUNAS

Thickness Total of unit thickness (in feet) (in feet) 4. Clay, thinly laminated, light brown, high in organic content ... 1.20 65.75 3. Silts, red, interbedded with caliche nodules...... 3.20 68.95 2. Sand, yellow-brown, with lime cement, grades upward into red-brown to red-buff silt, with zones of caliche nodules. Sand stained orange with limonite streaks...... 6.00 74.95 .4 Angel1 member 1. Sand and gravel, medium to coarse yellow-buff, indurated in places. Light buff color on dry surface. Top 18" cross bedded (base not exposed)...... 4.00 78.95

Measured Section 3. Section measured on the north side of Spring Creek, in the SE 1/4sec 18, T 32 S, R 28 W, Meade County, Kansas. The section was started at a willow grove in the bed of Spring Creek and measured along a north northwest traverse which ascends to the top of Mount Scott. The description of sediments is based on fresh exposures. The stratigraphic section is approximately 3 ft greater than the usual topographic thickness because of Kingsdown deposits in channels cut into the Crooked Creek and Ballard Formations. Units 15and 18 contain respectively, localities 2 and 3 of the Cragin Quarry local fauna (Hibbard & Taylor, 1960) of Sangamon age.

Thickness Total of unit thickness Topsoil (in feet) (in feet) Kingsdown Formation 19. Caliche, white, indurated, porous, with irregular cavities. Surface covered with a black lichen ...... 2.00 18. Sand and silt, red-brown, with zones of lime enrichment; some caliche nodules; orange-yellow limonite stringers; coarser at base, grading upward into finer sand and silt (Cragin Quarry local fauna, locality 3, comes from red- brown silt near top of unit)...... 11.40 17. Silt, gray-green, with limonite blotches and stringers ; shell at base...... 0.80 16. Sand, medium to fine, light brown, grading upward to fine sand with orange limonite blotches and stringers...... 7.40 15. Silt, brown, with some limonite stringers. Contains Sphaerium and other mollusks. (Cragin Quarry local fauna, locality 2)...... 6.70 14. Clay, gray-green, with some fine silt; silt component increases towards top; with manganese dioxide and limonite strains; contains shells...... 7. 30 13. Clay, brown, with some silt; grades upward to light brown-buff clay...... 11.00 12. Silty clay, light brown. ..; ...... 1.60 11. Sandy clay, red-brown...... 5.00 Top of west wall of quarry 10. Sand and gravel, yellow and buff, occur in cut and fill type deposition, with lenses of sand and gravel that pinch out laterally; sands containing some shells and clay balls...... 5.50 9. Gravel, over 2 in. in diameter (reworked Stump Arroyo gravel) in yellow sand; cobbles of caliche (reworked Ballard caliche) up to 10 in. long; base irregular, channeled into lower unit...... 0.80 B. B. MILLER

Thickness Total of unit thickness (in feet) (in feet) 8. Silt, coarse, greenish-brown, with mollusks (Mount Scott local fauna, locality UM-K2-59) grading upward into green silt and olive green clay with orange limonite clay...... Quarry floor. Base of Kingsdown is not exposed

Crooked Creek Formation Stump Arroyo member (exposed approximately 75 ft east southeast of quarry). 7. Gravel and sand; sand colored yellow-orange...... 2.50 Ballard Formation Missler member 6. Silt, green; indurated, dense, white caliche at top forms small ridge...... 1.90 65.90 5. Sand, fine, orange-brown...... 0.90 66.80 4. Clay, green, with lime enriched zone near top...... 1. 20 68.00 3. Silt, red-brown...... 1.20 69.20 2. Clay, silty, green, with caliche nodules...... 2.30 71.50 1. Covered interval...... 11.30 82.80 Floor of Spring Creek

Measured Section 4. Section measured in the SW 1/4 SW 1/4 sec 13, T 32 S, R 29 W, on the north side of Spring Creek. The Vanhem formation at this locality is channeled into the Kings- down and is topographically lower than the upper portion of that formation.

Thickness Total of unit thickness Topsoil (in feet) (in feet) Kingsdown Formation 7. Clay, gray-green, with red-brown limonite stringers; base not exposed. Top 2 ft contain mollusks...... 5. 66 5. 66 Vanhem Formation 6. Covered interval...... 5. 66 11.32 Top of bank 5. Sands and silts, reddish-brown, with small caliche nodules; coarser at base, grading upward into finer sands and silts; contains lenses up to 6 1/2 in.thick of light gray clay. These sediments are channeled into the black clay of unit 4...... 7. 00 Kingsdown Formation 4. Clay, black on fresh exposures; gray when dry; with nodules and streaks of calcium carbonate 3/4 of an inch in diameter, and red-orange limonite spots; unit containing many unionid fragments...... 0. 70 3. Silt, light gray, with layers 1/2 in. thick stained with limonite; white calcareous stringers interspersed throughout the unit; unit containing many mollusks (Mt. Scottlocal fauna, locality UM-K1-60)......

MILLER

Summary of the Late Cenozoic Geo- on the upper portion of the Crooked Creek lodc- Historv of the Bie:-* S~rinrrs u Ranch Formation. area. The late Cenozoic history of Sometime during the late Yarmouth- southwestern Kansas and northwestern early Illinoian interval, ancestral Spring Oklahoma has been reviewed in papers Creek, flowing as a northwest-southeast by Hibbard (1949a; 1950; 1954; 1955), stream, started to entrench its course on and Hibbard & Taylor (1960). The dis- the Yarmouth surface formed on top of cussion below concerns primarily that the Crooked Creek Formation, and cut portion of this history pertaining to the downward to at least the Angel1 member e Big Springs Ranch area. of the Ballard Formation. The topo- There are no known exposures of pre- graphic occurrence of the Kingsdown, Rexroad Pliocene sediments in the Big below the top of the older CrookedCreek Springs Ranch area. The Rexroad For- Formation in the Big Springs Ranch area mation represents a local accumulation of Meade County, and in northwest Clark of sediments from a through-flowing County, along the upper part of Bluff stream, which were trapped in a basin Creek (Hibbard & Taylor, 1960: 20), produced by subsidence along the west over 30 miles to the northeast, suggests side of the Crooked Creek Fault some- that this cycle of valley cutting was time during the early part of the upper related to regional, rather than local Pliocene (Smith, 1940: 98). A climatic events. This erosional cycle was change toward conditions of increased followed by one of deposition, which aridity near the end of Rexroad time is started later in the early Illinoian and implied by the caliche occurring at the continued through middle Sangamon time, top of the Rexroad Formation (Hibbard, nearly filling the valley to the Yarmouth 1954: 235). surface. Sediments representing the Increased runoff and regional warping lower portion of the Kingsdown, exposed rejuvenated the through-flowing streams at Mount Scott, are the result of stream at the close of Rexroad time, and in- aggradation; some of the units in the augurated a new cycle of erosion and upper portion of this sequence, however, deposition, during which the sediments are probably eolian in origin. The of the Ballard Formation (Nebraskan- Kingsdown sediments occuring along Aftonian) were laid down (Hibbard, 1948: Spring Creek record only the late Illi- 593). The decrease in particle size noian-early Sangomon part of this towards the top of the formation and sequence. An older portion of the Kings- the presence of a buried caliche seem down, representing sediments equivalent to indicate aperiod of increasing drought. to those containing the earlier Illinoian The climatic implications of the Sanders Butler Spring, Adams, Doby Springs and local fauna, which occurs above the -Berends local faunas, was probably de- buried caliche, suggests that this cycle posited in the deeper main channel phase terminated in a mesothermal, subhumid somewhere to the south of Mount Scott. climate (Taylor, 1960). A late Sangamon-early Wisconsin ero- The sediments represented by the sion interval is indicated by the thick Crooked Creek Formation (Kansan-Yar- series of sediments filling the channels mouth) are the product of the episode cut into the Kingsdown, Crooked Creek, of downcutting and deposition which Ballard and Rexroad formations. During followed the Ballard. Increased pre- this phase, ancestral Spring Creek, cipitation and uplift are implied by the probably aided by local sinkhole collapse channels cut into the Ballard and coarser in the area to the south and southwest clastics represented by the Stump Arroyo of Mount Scott, migrated to the south sands and gravels. During late Yarmouth of the present valley as it became en- time, climatic conditions were again trenched in the older formations. The conducive to the formation of a caliche apparent dip of 40 E on the Ballard For- FIVE ILLIFOLAX FAUSAS

FIG. 4. Photozrxph taken near locnlity VJI-K4-.53 show in^ sediments of the l'nnhem Formntion fillinq chnnnrl. cut into thc Kinqsclo~inFormation. The white rinshed line sho\vc; the npnrosimntc contact. In thc mirldlc force7.ounrl this conhct is covereci 11:' slump. mation (see Measured Section 5). is & Taylor, 1960: 24). Later in the Wis- believed to be related to ths event. consin. aggrading streams filled the Those portions of the Kinqsdown de- vallevs cut durinq the previous erosional posited on the south side of Spring Creek cycle with sediments belonqing to the were either buried during the episode of Vanhem Formation. sinkhole collapse or vere removed The present stream s:i.stem represents during the subsequent cycle of stream en- the most recent cycle of valley cuttine;. trenchment. This new7 cycle of valley The entrenchment of the Recent streams cutting was perhaps in part related to a has been discontinuous, as evidenced by climatic chanq~involving an increase the series of terraces alonq the Cimarron in preciptiation and runoff, such as is River (3kLaughlin, 1947: Byrne ). llc- implied by the late Sangamon Jinglebob Lauqhlin. 1948: Hibbard Taylor. 1950) local fauna (Hibbard, 1955: 204: Hibbard and at least one terrace level alone; 206 B. B. MILLER

Spring Creek and Hart Draw. the probability that these sediments are Age and Correlation. The Mount Scott of Recent origin. The fossils are inter- local fauna is considered Late Illinoian preted asindicating aprobable Wisconsin in age on the basis of the following age for these sediments. The bed con- stratigraphic and paleontologic evi- taining the UM-K4-53 and UM-K1-60 dence. faunules is thus post-Yarmouth (Crooked 1) It occurs in sediments that are Creek Formation) and pre-Wisconsin topographically lower, but strati- (Vanhem Formation) in age. graphically younger, than the Crooked 3) The presence of many species with b. Creek Formation of Kansan and Yar- predominantly northern and/or north- mouth age. At locality UM-K2-59, the eastern distributions implies a glacial fossils occur in a coarse, greenish- fauna, which, on the basis of the strati- brown silt of the Kingsdown Formation graphic position of the containing beds, which is channeled into the Stump Arroyo must be Illinoian. Miller (1961: 123) af- member of the Crooked Creek For- ter studying the mollusks from UM-K4- mation. The section at this locality may 53, concluded "If future work can defi- be followed up the slope of Mount Scott, nitely establish the age of the fossils as to locality 3 of the Cragin Quarry local Illinoian, the term 'Butler Spring local fauna of Sangamon age (Hibbard & Taylor, fauna' should be used for this local fau- 1960). The only gap in sedimentation in nule. " Subsequent study however, has this interval, occurs at the base of unit suggested that the Mount Scott is slightly 9 (Measured Section 3); it is believed younger than the Butler Spring local fau- to represent only a small local uncon- na. The Mount Scott local fauna appears formity produced by the migration of the to be the youngest of the 5 Illinoianmol- main stream channel of ancestralspring luscan assemblages (Mount Scott, Butler Creek northward toward Mount Scott Spring, Adams, Doby Springs and Be- soon after deposition of the sediments rends local faunas) considered in this containing the UM-K2-59 faunule on the report . floodplain of this same stream. A climatic change from a cooler glacial 2) The Kingsdown Formation at locality maximum, represented by the Doby UM-K4-53, again is an alluvial deposit, Springs local fauna, to slightly warmer occupying the valley of ancestral Spring conditions during the time the Mount Creek. This stream had become en- Scott local fauna lived, is indicated by trenched inthe Crooked Creek Formation the mammals (Hibbard, 1963) and the early in the Illinoian and cut downward pollen (Kapp, 1965). The mollusks imply to at least the Angel1 member of the a similar trend with the number of Ballard Formation. The fossils at species with southern distributions in- localities UM-K4-53 and UM-K1-60 are .creasing from 4, in the Doby Springs from the same stratigraphic interval of local fauna (Uniomems tetvalasmus, the Kingsdown Formation, which can be Physa anatina, Gastvocopta pyoceva and traced downstream about 1/2 mile (see G. cvistata), to 6 in Mount Scott local Fig. 3) as an almost continuous bed. fauna (which moreover includes Gastvo- The stratigraphic position of the Kings- copta pellucida hovdeacella and Helico- down in this area indicates that it is discus singleyanus). post-Crooked Creek in age. Habitat. Table 3 listsby habitat group, A post-Kingsdown episode of channel the species of mollusks occuring in the cutting is recorded by gravel and light Mount Scott local fauna. brown-buff sands and siltsfilling valleys Miller (1961: 123), concluded that the which have been cut into the Kingsdown molluscan assemblage at locality UM- Formation (Fig. 4). Mollusks recovered K4-53 represented 3 major habitats: from this interval (locality UM-K2-60), "(1) a small permanent stream of the include 6 species no longer living in same approximate size as the present Meade County area, thus eliminating Spring Creek; (2) a temporary water FIVE ILLINOM FAUNAS 207

TABLE 3. Summary of the local habitats represented by the mollusks of the Mount Scott Local Fauna.

1 1 Locality and 1 Abundance* momo Habitat LOWL~W Species Ill I .

Semiaquatic: among vegetation and Oxylom sp. A-A - debris near water's edge.

Hygrophilic: moist situations in leaf Cary chium exiguum mold, under sticks and debris; shaded Velertigo ovata areas, not far from water. V. milium Gastrocopta tappaniana Deroceras aenZgma

Woodland: moist areas under leaf litter, Cionella lubrica down timber, among tall marsh grass, Stenotrema Zeai on the floodplain. Discus cronkhitei Zonitoides arboreus Z. nitidus Nesovitrea electrina Euconulus fulvus Strobilops labyrinthica Helicodiscus parallelzls Punctum minutissimum

Sheltered areas: among rocks, shrubs, Gastrocopta armifera ACAR grass, or in timbered situations. G. contracts AAA- G. holzingeri ARAR

Sheltered situations: these species are G. cristata CAR not restricted to a woodland habitat G. procera RCR and can tolerate drier conditions. G. pellucida hordeacella -R - Pupilla blandi -R - Pupoides albilabris SAR Hawaiia minuscula CA S Vallonia parvula RAR V. pacilicosta RA S Helicodiscus singleyanus R - -

Marginal situations: wet mud, sticks, Fossaria dalli A R stones, or any other debris along F. obmssa C R water's edge; shallow pools and protected spots.

Shallow, quiet water: small streams, Pisidium case~tanum A A ponds, or sloughs on the floodplain, Sphaerium occidentale R - that may become dry during part of Stagnicola exilis - - the year. S. caperata A R S. reflexa R R (c ont .) 208 B. B. MILLER

Table 3 (cont. )

------I I LLality and /

Armiger crista Gyraulus circumstriatus Promenetus umbilicatellus Physa gyrina Aplexa hypnomm Ferrissia fragilis

Shallow, quiet water: small pond, Uniomems tetralusmus streams or sloughs with no current or Sphaerium partumeium areas of rooted vegetation with little S. rhomboideum current; soft sand or mud bottoms; S. sulcatum not subject to significant seasonal S. transversum drying. Pisidium obtusale Gyraulus parvus Helisoma trivolvis Promenetus exacuous form exacuous Phy sa anatina P. skinneri Laevapex fuscus

Perennial water: stream or lake, with Sphaerium lucustre slow to moderate current; areas of S. stktinum still water; shallow spots with soft Pisidium compressurn sand or mud substrate; not affected P. nitidum by seasonal drying. P. subtruncatum P. variabile P. walkeri Valvata tricarinata Helisoma anceps

*~=<25;S= 26-49; C= 50-100; A=>100 shells. situation, perhaps a slough or temporary Sedimentary and paleontologic evi - pond, situated on the floodplain of the dence indicate that the Mount Scott stream; and (3) a woodland habitat, local fauna contains the following major situated near the stream." New infor- habitat associations: (1) a small lake mation, based on subsequent studies of or permanent pond; (2) a perennial the pollen (Kapp, 1965), the fishes (Smith, stream; (3) a temporary water situation; 1963), the mammals (Hibbard, 1963), the (4) a marsh-woodland; and (5) anupland remainder of the molluscan fauna and meadow. field observations, have confirmed some The thin bed of marl (unit 2, Measured of these earlier conclusions and filled Section 4) at locality UM-K1-60 implies in certain details of the environment. slack water deposition at this site, and FIVE ILLINOW FAUNAS 209

is interpreted as indicatingthe existence These cut and fill structures are be- of a small oxbow lake or permanent lieved to represent a younger depo- pond on the floodplain of ancestral Spring sitional phase when the main stream Creek. Faunal evidence, based pri- channel of ancestralspring Creek shifted marily on the occurrence of Esox mas- north over the older floodplain. (3) The quinongy, Ictioh sp., Perca flaves- large fish assemblage reported from cens and Micropterms salmoides, (Smith, the Mount Scott localfauna (Smith, 1963), 1963), an association of fish most fre- as well as some of the mollusks (Miller, quently found in a lacustrine habitat 1961; this paper), suggest permanently (Smith, oral communication), supports flowing water. This stream was per- this interpretation. Shallow areas near haps several feet deep and in places had - the shore of this lake probably con- a slow to moderate current, a sub- tained dense growths of aquatic vege- strate of mud, soft sand and some gravel, tation which provided suitable habitat and shallow spots with dense rooted conditions for Promenetus exacuous vegetation that greatly reduced the form exacuous, Valvata tricarimta, Gy- effects of the current. raulus parvus, Physa anutinu, Sphaerium The presence of ephemeral bodies of sulcatum, Pisidium variabile, P. walk- water on the floodplain of ancestral evi, P. nitidum and P. compressum. Spring Creek is evidenced by the occur- These species, which constitute the rence of prominent Urest marks" on largest portion of the total number of many of the Pisidium casevtanum and individual mollsusks at this locality, Sphaevium occidentale, a condition gen- are believed for the most part to have erally considered indicative of a fluctu- been autochthonous and to have lived at ating water level, and by the abundance or near the site of deposition. Some of of aquatic mollusks frequently associated the shells in this group, especially those with temporary water situations. The of the Valvata, show signs of abrasion overwhelming predominance among the suggesting that they have been trans- aquatic mollusks of locality UM-K2-59 ported. These individuals, together with of temporary water species, together the thin layers of interbedded sand with the relatively large terrestrial associated with the marl and more faunal element, suggests that it was obvious allochthonous faunal elements, probably part of a backwater swamp or were probably washed into the lake during marsh. The mollusks requiring perma- periods when the nearby stream was in nent water, as well as the fish, were floodstage. probably carried to this site from the The efistence of a small permanent nearby stream during floods. stream near the site at which the Mount A marshy -woodland community repre- Scott assemblage lived is indicated by senting animals which probably lived the following: (1) the intermittent nature on the floodplain of the stream among of the outcrop is suggestive of a me- patches of tall grass, sedges, shrubs andering stream that followed a course and some scattered stands of trees in similar to that of the present Spring which moist leaf mold accumulated, is Creek. Similarities in lithology and implied by the pollen spectra (Kapp, fauna of localities UM-K4-53 and UM- 1965) and the abundance of Strobilops K2-59 are interpreted as meaning that labyrinthica, Gastrocopta contracts, G. they were essentially contemporaneous tappaniana, Vevtigo ovata, V. milium, and were once part of the floodplain of Oxyloma sp,, Helicodiscus par~llelus, ancestral Spring Creek. (2) The sedi- Deroceras aenigma, Carychium exi- ments directly overlying the Mount Scott guum and Zonitoides nitidus. Some of local fauna at locality UM-K2-59 (units the mammals represented by Swex cin- 9 and 10, Measured Section 3) consist ereus, S. arcticus, Ovyzomys fossilis, of lentils of cobbles, gravel and sand. Synaptomys australis, Blurina brevi- MILLER

cauda carolinensis, Micvotus pennsyl- and fossil molluscan faunas, signifies a vanicus and Zapus hudsonius tvansition- climate markedly different from that alis (Hibbard, 1963), support this con- which characterizes the area today. clusion. The majority of the fossil species are A better drained upland, containing aquatic, requiring permanent or semi- some trees, herbaceous composites and permanent water, and for the most part grasses, is indicated by the pollen (Kapp, are presently distributed well to the 1965). This community included Cvypto- north and/or northeast of the Mount Scott tis pavva, Pedomys ochvogastev, Lepus fauna locality. The variety of habitats * and Citellus (Hibbard, 1963; Stephens, indicated by the fauna requires a supply 1960), and some of the less numerous of water not available in the area at the xerophilic terrestrial gastropods repre- present time. Hibbard (1963) suggested . sented by Gastvocopta pellucida hovdea- that incision by ancestral Spring Creek cella, G. cristata, Helicodiscus singley- of the highly charged aquifers repre- anus and Pupoides albila bvis. sented by the Stump Arroyo and Angel1 Climate. The climatic conditions sands and gravels would have produced under which the Mount Scott local fauna a series of springs that provided much lived may be inferred from the elements of the water for the aquatic habitats. of the fauna. Interpretation of the Mount The amount of downcutting necessary to Scott molluscan assemblage is predi- expose these aquifers, together with the cated on 2 assumptions: (1) that cli- occurrence of land snails that probably matic factors are the primary agents would not have been influenced by these controlling the distribution of non- springs, imply greater runoff due to an marine mollusks (Taylor, 1960; Boycott, increase in rainfall. 1921; 1934; 1936). The rationale for Although the Mount Scott molluscan believing that the north-south and east- assemblage contains some associations west distributions of some groups of of species that are no longer known to mollusks appear to be controlled occur together (e.g., Pupilla blandi and essentially by temperature extremes and Stenotvema leai) it most resembles the available moisture has been presented fauna living in the tri-state area of by Taylor (1960). (2) The area in the Nebraska, South Dakota and Iowa, far Great Plains containing a molluscan to the north. The mollusks of this area fauna most similar to the Mount Scott have not been intensively studied, but assemblage will probably most closely published records (Shimek, 1915; Over, simulate the climatic conditions repre- 1915, 1928; Roscoe, 1954, 1955; Jones, sented by the Mount Scott mollusks. 1932; and Taylor, 1960), together with Additions to the Mount Scott molluscan materials examined in the University fauna and new distribution data for some of Michigan Museum of Zoology and the of the species in the Great Plainsregion United States National Museum col- has necessitated a slightly different lections indicate that 50 of the 63 species climatic interpretation from that given included in the Mount Scott local fauna by Miller (1961). Perhaps the most occur in this general region. conspicuous and significant aspect of Forty-seven of the species repre- the Mount Scott molluscan fauna is the sented in the Mount Scott local fauna relatively large number of species it have been recorded from the area contains as compared with the de- surrounding the Lake Okoboji region of pauperate Recent fauna of the Meade northwest Iowa. The great similarities County area. Hibbard & Taylor (1960) between the molluscan fauna of this rela- listed only 29 species living in the county tively restricted area and the Mount today, whereas the Mount Scott local Scott assemblage, together with the fact fauna contains some 63 species. The that many species with predominantly contrast in size alone, between the Recent northern, northeastern and southern dis- FIVE ILLINOIAN FAUNAS

TABLE 4. Comparison of climatic data for Meade County, Kansas, and the Lake Okoboji region of Northwest Iowa*

Temperature Meade County Lake Okoboji

Annual temperature 55'-60° F 40'-45' F Normal winter temperature 1 250-35'~ / 15O-20' F Normal summer temperature 1 75O-800 F 1 65'-70' F Normal annual numbers of days with maximum temperature of 90° or higher 60- 80 I 10-20 Normal annual number of days with daily maximum temperatures above 85O 80-100 10-30 Normal daily maximum temperature in July / goo-9s0 F 1 84O-87OF Normal annual number of days of very hot 1 weather (day and night averaging 750 or higher) ! 0 Evaporation

and shallow lakes 50-70 inches 30-40 inches Normal annual evaporation from pans 90 inches 1 50-60 inches Normal annual excess of precipitation over evaporation 1 0 inches 0-10 inches Precipitation

Normal annual precipitation 20-30 inches 20-30 inches Percent of precipitation that occurs in summer 1 30-40% 1 40-5090 Normal summer precipitation / 6-8 inches 1 10-12 inches *Based on Visher . 1954

tributions, are close to or at the peri- first 2 species in this group are pres- phery of their respective ranges in the ently known to occur in South Dakota Great Plains region in this area, are and eastern Iowa, while P. nitidum has interpreted asindicating that the climatic been recorded from eastern Iowa and conditions prevaling in the Lake Okoboji Douglas County, Nebraska (UMM Z region represent fairly closely those 110376)*. P. subtruncatum and P, ob- that existed in southwestern Kansas at the tusale reach their southern limits in time the Mount Scott local fauna lived. the Great Plains, in South Dakota Twelve of the extant species not (Herrington, 1962). Two species are occurring in the Lake Okoboji areaform included in the northeastern category. part of the 3 major distribution groups Ferrissia fragilis and Laevapex fuscus mentioned above. The northern element includes Pisidium variabile, P. walkeri, P. nitidum, P. subtruncatum, Sphuerium *University of Michigan Museum of Zoology occidentale and Pupilla blandi. The catalogue number. MILLER occur in Iowa to the south and east of deacella is the most southerly mollusk Lake Okoboji (Leonard, 1959; Basch, in the fauna, with respect to its range, 1963). in the Great Plains; it presently ex- Three species with predominantly tends as far north as the southeastern southern distributions, Physa anatina, Nebraska-northeastern Kansas area. Gastrocopta cristata and Helicodiscus The association in the Mount Scott local singleyanus, occur in northeastern Neb- fauna of G. p. hordeacella and other raska and southeasternSouthDakota, and southern elements, with more northerly * might be expected in the Okoboji region species seems best explained by assum- of Iowa (Taylor, 1960; Roscoe, 1954). ing that the winter temperatures were A fourth member of this group, Gastro- not quite as cold as those now occurring copta pellucida hordeacella, reaches the in the vicinity of Lake Okoboji. northern limit of its range in the Plains, A winter temperature between 25O and in the northeastern Kansas-southeastern 350 F, similar to that existing in the Nebraska region. area of the Great Plains in which G. p. The occurrence in the Mount Scott hordeacella reaches its northern limits, local fauna of species that are now is believed to have prevailed during the separated by several hundred miles time the Mount Scott local fauna lived. seems most reasonably explained in The summers during this interval were terms of climatic change in which sea- probably like those of northwesternIowa sonal extremes were not of the same at the present, with temperatures magnitude as those which presently between 65O and 700 F, and withbetween characterize the Meade County or over- 20-30 in. of rainfall, 10-12 in. of which lap areas. falls during the warmer part of the year The climatic differences between (see Table 4). Meade County, Kansas and the Lake Summary of Illinoian Okoboji region of Iowa are most pro- Molluscan Faunas nounced in terms of summer temperatures and moisture extremes (Table 4). The 5 molluscan assemblages con- The normal summer temperatures in the sidered in this report show compositional Okoboji area are about 5' cooler than in differences which seembest explainedby Meade County. The latter moreover has assuming climatic change. The Doby higher maximum summer temperatures Springs and Mount Scott local faunas which persist for longer periods of time. appear to represent the climatic end Both areas are shown by Visher (1954) points in a sequence of Illinoian biotic as having an annual rate of precipi- assemblages. Vertebrate and pollen evi- tation of between 20-30 in., but the dence show that the Doby Springs local Okoboji region receives a larger portion fauna represents the coolest and the of this precipitation during the summer Mount Scott local fauna the warmest of months when it is most needed. The these faunas. The mollusks show a greater summer rainfall, lower similar trend with the Doby Springs temperature, and smaller evapotran- containing fewer southern elements than spiration rates, produce cooler, more does the Mount Scott local fauna. The moist summers in the Lake Okoboji Doby Springs local fauna has beeninter- area. preted by Kapp (1965) andHibbard (1963) The low frequency of some of the as having probably lived during maxi- southern faunal elements in the Mount mum Illinoian glaciation, while the Mount Scott local fauna, represented by Heli- Scott local fauna has been considered codism singleyanus and Gastrocopta as transitional between the Illinoian and pellucida hordeacella, suggests that Sangamon (Kapp, 1965). Although several these species may have been close to stadial-interstadial intervals may have the limits of their range. G. p. hor- occurred, there is no evidence in this FIVE ILLINOIAN FAUNAS 213

area indicating that these faunas lived with summers that were no more severe during more than one substage of the than those now occurring in northern Illinoian. This interval was charac- North Dakota and Minnesota. The in- terized by a trend towards increasingly ferred climate for the Harper County, warmer climatic conditions which Oklahoma area at the time the Doby culminated in the Sangamon interglacial. Springs local fauna lived probably con- A tentative chronology based on the sisted of summer temperatures of be- climatic interpretation of the faunas, tween 65O-70° F, with winter tem- first and last occurrences of molluscan peratures of between 15O-20° F, and species, and pollen analytical data, an annual precipitation of about 18- suggests that the Doby Springs probably 20 in., 8-10 in. of which fell during the represents the oldest, followed by the summer months. Adams, Butler Spring and Mount Scott The overlap area which encompasses local faunas. most of the species present in the Adams The proper placement of the Berends and Butler Spring local faunas is not local fauna in this sequence is not cer- very different from that of the Doby tain. The overlap area for most of the Springs local fauna. The climate at mollusks of the Berends local fauna is the time the Adams and Butler Spring the northeasternNebraska-northwestern local faunas lived combined winters Iowa region. This has been interpreted which were similar to those now occur- to mean that the Berends assemblage ring in northeastern Nebraska with sum- lived at a time when the climate in the mers like those that presently prevail panhandle of Oklahoma was similar to in eastern Montana and the Dakotas. that now occuring in the Plains farther Winter temperatures were probably be- to the north. The winters probably tween 15O-20° F, with normal summer average about 20° F, with average sum- temperatures of 650-70° F. The annual mer temperatures of 700 F. The pre- precipitation was perhaps between 18- cipitation was between 20-30 in. (Visher, 19 in. 1954). This indicates a slightly warmer, The Adams and Butler Spring local more moist climate than that of the time faunas appear to have lived during a of the Doby Springs and Adams local slightly warmer phase of the Illinoian faunas. The last appearance in this than did the Doby Springs local fauna. area of the Plains of Planorbula armi- Some of the same northern species gera and Sphaerium sews occurs in present in the Doby Springs local fauna the Berends local fauna. The absence continued to live in the area through the of these 2 species from the Doby Springs, time represented by the Butler Spring Adams, Butler Spring and Mount Scott local fauna, but they were greatly re- local faunas has been interpreted to duced in numbers. In addition, the Adams mean that the Berends assemblageprob- and Butler Spring local faunas contain ably lived earlier in the Illinoian, before the earliest Illinoian occurrences in the - these species had become locally ex- M eade County area of ~e'licodiscussing- tinct. leyanus, a relatively southern species. The Doby Springs mollusks represent These 2 lines of evidence suggest that the coolest of the 5 Illinoian assemblages the climate was changing towards slightly considered in this report. The distri- warmer conditions. This warming trend bution pattern of the Doby Springs mol- evidently brought about a reduction in lusks suggests that this assemblage many of the northern species whichwere probably lived in a climate that com- present in the Doby Springs local fauna, bined winters similar to those in north- and permitted more southerly elements eastern Nebraska-northwestern Iowa to enter this area. This influx of (the area in which the majority of the southern species continued through at Doby Springs species currently occur), least that portion of the Illinoian repre- B. B. MILLER

PLATE I

FIGS. 1, 2. Sphaerium sulcatum (Lamarck) , Mount Scott Locality UM-K2- 59. Inner and outer view of right valve. X4, UMMZ 208885a. FIG. 3. Anodonta grandis Say, Doby Springs locality 4. Outer view of right valve. X0. 8, UMMZ 213832. FIGS. 4, 5. Pisidium obtusale (Lamarck), Mount Scott locality UM-K2- 59. Inner and outer views of left valve. X8, UMMZ 208872a. FIGS. 6, 7. Pisidium subtruncatum Malm. Fig. 6, Mount Scott locality UM-K4-53. Outer view of right valve. X8, UMMZ 200632. Fig. 7, Jones Sink, lower level. Inner view of right valve. X8, UMMZ 195840a. FIG. 8. Cionella lubrica (Miiller), Doby Springs locality 4. Apertural view. X8, UMMZ 208994. FIGS. 9,lO. Sphaerium occidentale Prime. Mount Scott locality UM-K2-59. Inner and outer views of left valve. X8, UMMZ 208880a. FIG. 11. Vwtigo elatiov Sterki, Doby Springs locality 5. Apertural view. X8, UMMZ 213770a. FIG. 12. Oxyloma sp. , Mount Scott locality UM-K2-59. Apertural view. X8, UMMZ 213697a.

216 B. B. MILLER

sented by the Mount Scott local fauna. at locality UM-K2-60 are also included The Adams and Butler Spring local here because of their relevance to the faunas, thus, appear to be intermediate dating of the Mount Scott local fauna. between the Doby Springs and Mount Scott Only previously unreported species of local faunas. mollusks appearing in the faunal lists The Mount Scott local fauna isbelieved of the Berends, Adams and Butler Spring to be the youngest of the 5 Illinoian local faunas are described. A summary assemblages considered in this report. of information is given for each species, Locality UM-K2-59 of the Mount Scott consisting of geologic range, distri- local fauna occurs stratigraphically be- bution, ecology and such taxonomic low the Cragin Quarry local fauna of changes as are considered necessary. Sangamon age (Hibbard & Taylor, 1960). References listed under the synonymies There is no evidence in the intervening for some of these species are not sequence of sediments of an unconfor- necessarily complete andusually include mity or of any faunas equivalent to the only some of the more important reports. Butler Spring, Adams, Doby Springs or The treatment of the various species Berends local faunas. It is assumed has not been uniform, since many of the that the correlatives of these assem- mollusks have been quite thoroughly dis- blages occur somewhere below the in- cussed in recent papers by Taylor (1960) terval from which the Mount Scott local and Hibbard & Taylor (1960). However, fauna was recovered. species not covered in these papers, as Several different lines of evidence, well as species requiring taxonomic the overlap area of the mollusks, the changes have generally received detailed larger number of species withrelatively attention in this report. The super- southern distributions, the mammals familial and familial taxa used for the (Hibbard, 1963), and the pollen (Kapp, gastropods follow the arrangement pro- 1965), all indicate that the Mount Scott posed by Taylor & Soh1 (1962). The local fauna is the warmest of the 5 terminations for the superfamily names, Illinoian faunas from this region. The however, follows recommendation 29aof majority of the molluscan species occur- the International Commission on Zoo- ring in the Mount Scott local fauna are logical Nomenclature (Stoll, 1961: 29), now found living in northwest Iowa in and end in "-oidea". the area of Lake Okoboji. Some of the The ecologic and distributional data more southern species, however, only included for each species are in large reach as far north in the Plains as the part based on published information. southeast Nebraska-northeast Kansas The more important references include: area. These distribution patterns Pilsbry (1946; 1948), Baker (1939), Leon- suggest that the Mount Scott local fauna ard (1959), Taylor (1960) and Hibbard & lived at a time when local winter Taylor (1960), for the terrestrial gastro- temperatures were between 25O-35' F, pods; Baker (1911; 1928a), Leonard summers between 65'-70' I?, with be- (1959), Taylor (1960) and Hibbard & tween 20-30 in.of rainfall per year, 10- Taylor (1960), for the aquatic gastropods; 12 in. of which fell during the warmer and Baker (1928b), Herrington & Taylor part of the year. (1958), Herrington (1962), Simpson (1914), Ortmann (1919) and Murray & SYSTEMATIC DISCUSSION Leonard (1962), for the pelecypods. Additional information obtained from The systematic discussion in this por- personal collections and study of materi- tion of the paper includes all the mol- als in the United States National Museum lusks from the Mount Scott and Doby and University of Michigan Museum of Springs local faunas. Materials col- Zoology, has also been included. The lected from post-Kingsdown sediments geologic range of each species is based FIVE ILLINOLAN FAUNAS 217

primarily on the information published tem; upper St. Lawrence drainage; Red by Taylor (1960) and Hibbard & Taylor River of the North; Lake Winnepeg, (1960), and in general only includes Manitoba; southwest to Texas; south- records for the southwest Kansas-northeast to Pennsylvania. It is in the Tennes- west Oklahoma region. see and Cumberland drainages (Ortmann, In the discussion that follows, the 1919: 147). University of Michigan Museum of Ecology. In the southwestern part of Zoology (UMMZ) catalogue number, and its range it is usually a pond or lake the number of specimens counted (in form. The typical form is an inhabitant parenthesis), are enteredafter "Materi- of smaller creeks, where it prefers -al" for each species. Numbersin excess quiet pools and eddies, and often lives of 200 have usually been estimatedvolu- in fine sand and mud (Ortmann, 1919: metrically. Integers are used to indicate 145). specimens of pelecypods that have both Material. Doby Springs local fauna, valves intact. Fractions indicate locality 4, 213832 (1/2). Butler Spring separate pelecypod valves. local fauna, locality 1, 208898 (1/2). Remarks. An early Pleistocene Class Pelecypoda record, from the Sand Draw local fauna Order Prionodesmacea of Brown County, Nebraska (Taylor, Family Unionidae 1960: 45), is based on fragmentary material, and was questionably ref erred Uniomems tetralasrnus (Say) to genus and species. An identifiable Plate 11, Figs. 3-5 portion of a left valve from locality 1, Geological Range. Late Pleistocene of the Butler Spring local fauna, con- (Illinoian) to Recent. firms the presence in this assemblage Distribution. "Throughout the Missis- of Anodonta grandis, which had pre- sippi Valley as far northas forty degrees viously been queried (Hibbard & Taylor, latitude and southwest to Mexico. It i's 1960: 74). also found in the Ohio and Alabama Lasmigom complamta (Barnes) river systems" (Scammon, 1906: 337). Ecology. It is found living in tem- Geologic Range. Late Pleistocene porary ponds, tanks, sloughs and inter- (Illinoian) to Recent. mittent streams, in situations where no ' ~istribution. Upper Mississippi other mussels are likely to be found. drainage as far south as Arkansas in This species is capable of withstanding the west; Ohio River system; upper periods of drought by burrowing into the St. Lawrence and its tributaries; north substrate and hibernating over the dry in MacKenzie River; Alabama drainage; period (Isely, 1925; Baker, 1928b). 'In it is found in northern Louisiana and Kansas it occurs in habitats of soft or extends westward to Iowa, Kansas, hard mud bottoms in shallow water" Arkansas and Oklahoma (Ortmann, 1919: . (Murray & Leonard, 1962: 84). 136). The record of Recent occurrence Material. Doby Springs local fauna, of this species nearest to the Meade locality 4, 208937 (1 + 4/2). Mount County area is from the Little Arkansas Scott local fauna, locality UM-K4-53, River, Wichita, Sedgwick County, Kan- 200649 (2/2); locality UM-K1-60,213802 sas (Murray & Leonard, 1962). (1/2); locality UM-K3-60, 214173 (4/2). Ecology. Prefers deep andquiet water with muddy bottoms; but is found oc- Anodonta grandis Say casionally in riffles and in gravel (Ort- Plate I, Fig. 3 mann, 1919: 135). Geologic Range. Late Pleistocene Material. Adams local fauna, locality (Illinoian) to Recent. 3, 208899 (1/2), 214072 (2/2). Distribution. Entire Mississippi sys- Remarks. Parts of the umbone region B. B. MILLER

PLATE I1

FIGS. 1, 2. Sphaerium rhomboideum (Say), Doby Springs locality 1. Inner and outer views of right valve. X4, UMMZ 213739a. FIGS. 3-5. Uniomerus tetralasmus (Say), Doby Springs locality 4. Figs. 3, 4, Inner and outer views of right valve. X0.8, UMMZ 208937a. Fig. 5, Outer view of left valve showing umbonal sculpture. X4, UMPIZ 2089 37b. FIGS. 6-8. Punctum minutissimum (Lea), Mount Scott locality UM-K4- 53. Apical, apertural and basal views. X16, UMMZ 200636a. FIGS. 9, 14. Pisidium walkeri Sterki, Mount Scott locality UM-Kl-60. Inner and outer views of right valve. X8, UMMZ 208873a. FIGS. 10, 11. Pisidium vanabile Prime, Mount Scott locality UM-K2-59. Inner and outer views of left valve. X8, UMMZ 208887a. FIGS. 12, 13. Sphaem'um secuvis Prime, Berends local fauna. Inner and outer views of left valve. X8, UMMZ 208840a. FIG. 15. Sphaenum lacustve (Mdler), Mount Scott locality UM-K4-53. Outer view of right valve. X4, UMMZ 200642a.

220 B. B. MILLER of 2 valves (UMMZ 213072) are only and S. lacustre form jayense from the identifiable to genus, but it seems prob- mud of Rice Lake, Peterborough Co., able that they belong to the same species Ontario, and in Hay Bay, Lennox and as the more complete specimen found at Addington Co., Ontario. In the latter the same locality. This is the first place most specimens were obtained record of this species from the Meade from black ooze a quarter to a half County region. mile from shore, at a depth of 6-8 meters" (Herrington, 1962: 20). Ligumia cf L . re cta (Lamar ck) Material. Mount Scott local fauna, Geologic Range. Late Pleistocene locality UM-K4-53, 200642 (12/2). (Illinoian) to Recent. Remarks. Sphaerium lacustre form Distribution. "Entire Mississippi ryckholti has been reported from the drainage; Alabama River Drainage; Dixon local fauna, Kingrnan County, Red River of the North; St. Lawrence Kansas, which is believed to be either drainage" (Simpson, 1914: 95). The latest Nebraskan or earliest Aftonian in river form of this species L. r, latis- age (Herrington & Taylor, 1958). The sima has been reported from the Marais earliest record of this forminthe Meade des Cygnes and Elk Rivers in south- County area is from the Cudahy fauna eastern Kansas (Murray & Leonard, of Kansan age (Herrington & Taylor, 1962). 1958). The one valve from the UM-K4- Ecology. It prefers lakes and large 5 3 material whichis complete, possesses rivers, where it may be found living a straighter hinge, truncated ends, and under a variety of conditions (Ortmann, a more gently rounded central margin 1919). than S. lacustre form ryckholti, giving Material. Butler Spring local fauna, the shell a rectangular outline, charac- locality 1, 214070 (1/2). teristic of the ecologic form jayense. Remarks. The material at hand is a The Mount Scott shells represent the fragmentary left valve, in which the only known fossil occurrence of this beak sculpture is somewhat worn. The form from the southwest Kansas-north- double loop sculpture is characteristic west Oklahoma area; all of the older of Ligumia and is probably Ligumia records belong to S. lacustve form vyck- recta. The specimen represents the first holti. fossil record of this species from the Sphaerium occidentale Prime southwest Kansas-northwest Oklahoma area. Plate I, Figs. 9-10 Geologic Range. Late Pleistocene Order Teleodesmacea (Illinoian) to Recent. Family Sphaeriidae Distribution. Northern North Ameri- ca: New Brunswick, Quebec, Ontario; Sphaerium lacustre (Miiller) Eastern United States from Vermont, Plate 11, Fig. 15 south to Georgia and Alabama; west to Geologic Range. Middle Pleistocene Great Lakes region; Western United (Kansan) to Recent. States from Montana south to Utah and Distribution. Throughout most of Colorado, west to Oregon and Washington

North America, north of Mexico; Japan; (Herrington,-. 1962). Hawaii; Europe (Herrington, 1962). Ecology. Ustill waters of swamps, Ecology. "Most plentiful in small ditches, and ponds; among grass and lakes and ponds, but also found in large leaves. This species has a preference lakes, rivers and creeks. It appears to for, or requires, a habitat that dries have preference for a muddy bottom. I up for part of the year" (Herrington, have collected the S. lacustre form 1962: 22). rychkholti from ponds and bog-ponds, Material. Doby Springs local fauna, FIVE ILLINOIAN FAUNAS 221

locality 4, 213833 (1/2). Mount Scott Remarks. The Doby Springs record local fauna, locality UM-K4-53, 200643 represents the oldest fossil occurrence (1/2); locality UM-K2-59,208877 (85/2), of this species from the southern Plains 208880 (9/2). region. Remarks. The Doby Springs local Sphaerium securis Prime fauna record represents the oldest known 11, occurrence of this species in the south- Plate Figs. 12-13 ern Great Plains. Geologic Range. Late Pleistocene (Illinoian) to Recent. Sphaerium partumeium (Say) Distribution. Newfoundland, Maritime Geologic Range. Late Pliocene (Rex- Provinces, Quebec, Ontario (southern road Formation) to Recent. part), British Columbia; most of states Distribution. Southern Canada from of Mississippi River in United States; New Brunswick west to Saskatchewan. Minnesota, Nebraska, Iowa, Arkansas, Most of the United States east of Missis- Texas, Montana and Washington (Her- sippi River; Minnesota, Nebraska, Iowa, rington, 1962). Missouri, Kansas, Oklahoma, Louisiana, Ecology. '(Ponds, lakes and rivers. Texas, Montana, California and Nevada Frequently found in fine sand" (Herring- (Herrington, 1962). ton, 1962: 26). Taylor (1960: 46) col- Ecology. [(Ponds, swamps, small lected it from a small permanent or lakes, and slow-moving streams. It has nearly permanent pond in sandy bottom a preference for a muddy bottom, and is among dense vegetationin shallow water. fairly common" (Herrington, 1962: 24). Material. Berends local fauna, 208840

Material. Mount Scott local fauna, (7/2).. . locality UM-K2-59, 208878 (5/2). Remarks. This is the first known Remarks. The late Pliocene record fossil record of this species from the is based on fragmentary material from southern Great Plains. locality UM-K1-59, of the Bender local Sphaerium striatinurn (Lamarck) fauna, of Meade County, which, because of its condition was only questionably Geologic Range. Late Pliocene (Rex- referred to this species (Miller, 1964). road Formation) to Recent. Distribution. Most of North America Sphaerium ~homboideum(Say) from Panama to the Great Slave Lake, Plate 11, Figs. 1-2 Northwest Territories (Hibbard & Geologic Range. Late Pleistocene Taylor, 1960). (Illinoian) to Recent. Ecology. The typical form of this Distribution. Ontario (as far north as species is restricted to perennial bodies James Bay and Manitoba); New England of water with some current action. It and New York west to Minnesota and has been found living in sand, sandy Iowa; Montana and Idaho. Questionably gravel and sandy mud substrates, in - recorded from Alaska, British Columbia water from 2-3 in. down to a depth of and Nevada (Herrington, 1962). 13.5 meters. It has never been collected Ecology. In still water of enlarge- from ponds, lagoons, bog-ponds, swamps ments in creeks and rivers; sheltered or anywhere in stagnant water (Herring- places in small lakes. It has a prefer- ton, 1962). ence for a soft muddy bottom, with con- Material. Doby Springs local fauna, siderable vegetal growth of weeds and locality 4, 213826 (52/2). Mount Scott algae (Herrington, 1962). local fauna, locality UM-K4-53, 200645 Material. Doby Springs local fauna, (14/2); locality UM-K2-59, 208875 (3/2). locality 1, 213739 (9/2); locality 4, Remarks. The first occurrence of 213834 (3/2). Mount Scott local fauna, this species in the Meade County area locality UM-K4-53, 200644 (1/2). is in the Saw Rock Canyon local fauna, 222 B. B. MILLER

Seward County (Taylor, 1960). local faunas (Herrington & Taylor, 1958; Taylor, 1960), but the material from Sphaevium sulcatum (Lamarck) the Berends local fauna represents the Plate I, Figs. 1-2 oldest record from the Meade County Geologic Range. Late Pliocene (Rex- area. road Formation, Bender local fauna) to Sphaevium sp. Recent. Distribution. Northeastern North Included here are shells, which, be- America from Quebec south to Virginia, cause of their fragmentary condition or . west to South Dakota, Wyoming, Montana immaturity, could not be identified to and Washington (?). It does not apparently species. occur south of those areas that were Material. Doby Springs localfauna, glaciated (Herrington, 1962). locality 2, 185789 (2/2); locality 4, ' Ecology. S. sulcatum lives on the soft 213828 (112); locality 5, 213752 (2/2). bottom of small lakes, creeks or rivers, Mount Scott local fauna, locality UM-K2- that have areas of still water (Herring- 59, 208893 (5/2), 208894 (112); UM-K3- ton & Taylor, 1958). It prefers soft 60, 214167 (2/2). sand, with vegetation, and is never found Pisidium casevtanum (Poli) in swamps or ponds (Herrington, 1962). Material. Berends local fauna, 208829 Geologic Range. Early Pliocene (La- (6sbySprings local fauna locality verne Formation, Laverne local fauna, 1, 213736 (42/2); locality 4, 213826 Beaver County, Oklahoma) to Recent (116/2); locality 5, 213753 (1/2). Mount (Herrington & Taylor, 1958). Scott local fauna, locality UM-K4-53, Distribution. Almost cosmopolitan. 200646 (54/2); locality UM-K1-60, In the Western Hemisphere from Pata- 208882 (26/2); locality UM-K2-59, gonia to the Arctic Circle. It has been 208885 (23/2). recorded from all of the states in the United States, with the exception of Sphaevium tvansvevsum (Say) Hawaii, Kentucky and North Dakota (Her- Geologic Range. Early Pleistocene rington, 1962). (Nebraskan) to Recent. Ecology. This widespread species is Distribution. North America east of capable of adapting itself to a diverse the Rocky Mountains, north to the North- range of habitats and has a correspond- west Territories and south to Mexico. ingly variable shell. It has been found Europe (Herrington, 1962). living in bog ponds, swamps that dry up Ecology. aLarge lakes, rivers, and for several months of the year, swamp- sloughs" (Herrington, 1962: 30). Hibbard creeks, creeks with considerable cur- & Taylor (1960: 77) state that: NInMeade rent, rivers and lakes, including the County, Kansas, it occurs commonly in Great Lakes (Herrington, 1962). "The Crooked Creek, and lives in both sand form vopevi Sterki, as well as some- and mud in slow current or quiet water. what similar shells with thin walls and Empty shells were abundant in a dry smooth outlines, comes from ponds, metal stock tank, usually kept full by a swamps, bog ponds, and small lakes windmill pump. Before the water had that are filling up with marl. The dried up the habitat was a soft mud heavier -shelled, typical casevtanum bottom, with a thin cover of submerged lives in rivers or fairly large creeks" vegetation." (Herrington & Taylor, 1958: 14). Material. Berends local fauna, 208947 Material. Berends localfauna, 208835 (172)unt Scott local fauna, locality (29/2).by Springs local fauna, locality UM-K2-59, 208892 (1/2). 1, 213738 (95/2); locality 4, 213829 Remarks. This species has been re- (87/2); locality 5, 213748 (200/2). Mount ported from the Sand Draw and Dixon Scott local fauna, locality UM-K4-53, FIVE ILLINOIAN FAUNAS 223

200629 (600/2); locality UM-K1-60, with marl, rivers and large creeks, 208870 (200/2); locality UM-K2-59, where it thrives in soft sand. Unlike 208896 (60/2), 208876 (3/2), 208879 the typical form, P. n. pauperculum (200/2), 208891 (33/2); locality UM-K3- does not apparently inhabit ponds (Her- 60, 214183 (1/2). Recent in Harper rington & Taylor, 1958). County, Oklahoma. Material. P. n. form nitidum Jenyns: Remarks. The variability of this Berends local fauna, 208828 (64/2). Doby species has evidently been greatly under - Springs local fauna, locality 1, 213737 estimated, for in a recently published (74/2); locality 4, 213831b (6/2); local- revision of the North American Sphaeri- ity 5, 213750b (700/2). MountScottlocal idae, 41 names are listed under the fauna, locality UM-K4-53, 200630b synonymy of Pisidium casertanum (Her- (91/2); locality UM-K1-60, 208868 (2 + rington, 1962: 33). 200/2). P. n. form pauperculum Sterki: Doby Pisidium compressum Prime Springs local fauna, locality 4, 213831a Geologic Range. Middle Pleistocene (425/2); locality 5, 213750a (200/2). (Kansan, Cudahy Fauna) to Recent. Mount Scott local fauna, locality UM-K4- Distribution. North America, from 53, 200630a (234/2); locality UM-K2-59, Great Slave Lake, Northwest territories, 208881 (2/2), 208884 (1/2). south to Mexico (Herrington, 1962). Pisidium obtusale (Lamarck) Ecology. It is restricted to per- Plate I, Figs. 4-5 manent water situations, such as creeks, rivers and lakes, with some current, and Geologic Range. Middle Pleistocene prefers a sandy bottom with some vegetal (Kansan, Cudahy fauna) to Recent. growth. It never occurs in ponds, Distribution. Holarctic; in North swamps, lagoons or bog-ponds (Hibbard America from New Jersey and New York & Taylor, 1960; Herrington & Taylor, west through Ohio, Illinois, Minnesota 1958). and the Dakotas, to the west coast; north- Material. Berends local fauna, 208839 ward to Northwest Territories; a single (313/2). Doby Springs local fauna, record from Mexico (Herrington, 1962). locality 1, 213740 (160/2); locality 2, Ecology. The form rotundatum pre- 185788 (9/2); locality 4, 213830 (4960/2); fers shallow water and sheltered spots locality 5, 213746 (620/2). Mount Scott in lakes, creeks, ponds, lagoons and local fauna, locality UM-K4-53, 200629 flood plains, where there are con- (1095/2); locality UM-K1-60, 208869 siderable accumulations of grassy vege- (500/2); locality UM-K2-59, 208886 tation or fallen leaves from trees (Her- (5/2), 208890 (4/2); locality UM-K3-60, rington, 1962). 214194 (1/2). Material. Berends local fauna, 208946 (1/2). Doby Springs local fauna, locality 1, 214160 (2/2); locality 4, 213835 Pisidium nitidum Jenyns (21/2); locality 5, 213751 (3/2). Mount Geologic Range. Early Pliocene (La- Scott local fauna, locality UM-K4-53, verne Formation, Laverne local fauna) 200631 (63/2); locality UM-K1-60, to Recent (~errington& Taylor, 1958): 208874 (1/2); locality UM-K2-59,208872 Distribution. Holarctic; Eurasia and (11/2), 208889 (450/2). Locality UM-K2- North Africa; in North America from 60, 214186 (112). Mexico to Northwest Territories, but Remarks. Only the ecologic form absent from the southeast United States. rotundatum Prime is represented iq Ecology. ULarge ponds, bog ponds, the fossil record. lakes, creeks, and rivers. Seems to Pisidium subtruncatum Malm prefer shallow water" (Herrington, 1962: Plate I, Figs. 6-7 4 6). Pisidium nitidum form pauperculum Sterki lives in lakes, now filling Geologic Range. Late Pleistocene 224 B. B. MILLER

(Illinoian) to Recent. Pisidium walkeri Sterki Distribution. Eurasia; in North Plate 11, Figs. 9, 14 America it is found from the Great Slave Lake, Northwest Territories, south Geologic Range. Late Pleistocene to the northern tier of states in the (Illinoian) to Recent. United States; New York, the Great Distribution. "Northern NorthAmeri- Lakes, Colorado, Wyoming and Cali- ca: Canada from Ontario to Alberta, fornia (Herrington, 1962). northward to James Bay and Northwest Ecology. 'Streams, bays, and lakes, Territories: in the United States, Maine including the Great Lakes" (Herrington, to Virginia, and Michigan, Ohio, Mis- 1962: 49). Baker (1928b) reports it souri, Iowa, South Dakota, Minnesota, from mud, sand and gravel bottom, at Montana and Arizona" (Herrington & depths ranging from 0.8-5.6 meters. Taylor, 1958: 17). Material. Mount Scott local fauna, Ecology. Pisidium walkevi lives 9n lbcality UM-K4-53, 200632 (1/2). rivers, creeks, and small lakes" (Her- Remarks. Herrington & Taylor rington & Taylor, 1958: 17). (1958) suggest that the northern distri- Material. Mount Scott local fauna, bution of this species may reflect its locality UM-K4-53, 200634 (1/2); inability to survive the warmer summers locality UM-K1-60, 208873 (6/2). that prevail further to the south. Remarks. The Mount Scott material represents the oldest confirmed fossil Pisidium variabile Prime record of P. walkevi. Plate 11, Figs. 10-11 Geologic Range. Late Pleistocene Class Gastropoda (Illinoian) to Recent. Subclass Prosobranchia Distribution. Prince Edward Island Order Mesogastropoda west to British Columbia; New England Superfamily Valvatoidea states south to District of Columbia; Family Valvatidae Tennessee, Alabama, Ohio, Michigan, Valvata tricavinata (Say) Indiana, Wisconsin, Illinois, South Dakota, Iowa, Montana, Washington, Geologic Range. Early Pliocene (La- Idaho, California, Utah and Colorado verne Formation, Laverne local fauna) (Herrington, 1962). to Recent (Hibbard & Taylor, 1960). Ecology. P. variabile has been taken ~istributidn. "Eastern united States from both lake and river situations. It west to Iowa. Great Slave Lake south lives on a variety of substrates (marl, to Virginia and the Ohio River" (Baker, clay, mud, silt, sand and gravel), at 1928a: 14). Cherry County, Nebraska depths down to 12 meters (Baker, 1928b). (Taylor, 1960). "Creeks, rivers, and lakes; usually in Ecology. "Valvata tricarinuta is a still water where soft sediments accumu- species of rivers, lakes, and permanent late" (Herrington, 1962: 50). ponds, particularly where there is abun- Material. Berends local fauna, 208838 dant vegetation. It is found on all (4mYSprings local fauna, locality varieties of bottoms and in all depths 1, 213735 (200/2); locality 4, 208888 down to 18 ft. It is usually associated (69/2); locality 5, 21 3747 (150/2). Mount with the filamentous algae Oedogonium Scott local fauna, locality UM-K4-53, and Cladophora, and has been observed 200633 (9/2); locality UM-K1-60,208871 on Vaucheria upon which it was ap- (79/2); locality UM-K2-59, 208883 (4/2), parently feeding . . . it prefers habitats 208887 (3/2). where the bottom sediments are more or Remarks. These records represent less firm and will enable the individuals the only known fossil occurrence of to move freely about" (Reynolds, 1959: Pisidium variabile from the High Plains. 160). FIVE ILLINOIAN FAUNAS 225

The single Nebraska occurrence of wider distribution for this species during this snail was from a spring-fed pond the Late Pleistocene. with a water temperature of 15O C, Ecology. The geographic and bathy- where it was found living in association metric range of this species indicates with Ceratophyllum, Elodea and algae that it occurs almost exclusively in (Taylor, 1960). rivers and lakes at depths of about 5 Material. Berends local fauna, 208821 ft or more. It has been found living on (319). Doby Springs local fauna, locality vegetation, as well as a variety of sub- 1, 213705 (1900); locality 2, 185775 (26); strates, including fine sand, coarse sand, locality 4, 208900 (8800); locality 5, gravel, marl and soft mud (Hibbard & 213744 (1600). Mount Scott local fauna, Taylor, 1960). locality UM-K4-53, 200652 (3600); local- Material. Doby Springs local fauna, . ity UM-K1-60, 21.3788 (2600); locality locality 4, 213789 (10,750), 185789 (3) UM-K2-59, 213700 (102). Remarks. The variations in number and arrangements of the carinae in V. Subclass Euthyneura tvicarinuta have long been recognized Order Basommatophora (Robertson, 1915; Baker, 1928a; La Superfamily Ellobioidea Roque, 1956), and given sub-specific Family Ellobiidae designations. However none of these Carychium exiguum (Say) variations have been definitely related to any known fixed set of conditions, Geologic Range. Late Pliocene (Rex- either geographic, stratigraphic or eco- road Formation, Rexroad local fauna, logic, and probably do not deserve sub- Meade County, Kansas) to Recent specific rank. In this work they are all (Taylor, 1960). treated as V. tvicarinuta. Distribution. "Newfoundland to Colo- rado, south to Mobile Bay, Alabama, and Superfamily Rissoidea near Deming, southwesternNew Mexico" Family Hydrobiidae (Pilsbry, 1948: 1052). Ecology. "C. exiguum lives in the Probythinella lacustris (Baker) crevices of rotten logs or ondeadleaves Geologic Range. Late Pleistocene in moist places, or sometimes it is found (Illinoian) to Recent. in very wet places, such as Pomatiopsis Distribution. #NorthAmerica between lapidaria frequents" (Pilsbry, 1948: the Rocky Mountains and the Appalach- 1054). In northern Nebraska it was ians, from Great Slave Lake to Arkansas, found living on wet leaves in seepage Alabama, and New York. The species areas and along a spring-fed brook is known from the MacKenzie River, (Taylor, 1960). Hudson Bay, St. Lawrence River and Material. Berends local fauna, 208837 Mississippi River drainages, but not (188). Doby Springs local fauna, locality from the shorter streams of the Atlantic 2, 195781 (1); locality 4, 208941 (160); or Gulf Coastal plains" (Hibbard & locality 5, 213745 (570). Mount Scott Taylor, 1960: 80). local fauna, locality UM-K4-53, 200603 Branson (1959) records this species (1500); locality UM-K2-59, 213659 (350). from Hughes and Payne counties, Okla- homa. Fossil occurrences of P. lacus- Superfamily Lymnaeoidea tris in McPherson County, Kansas Family Lymnaeidae (UMMZ uncat.), the Butler Spring local Lymnaea stagnalis &gularis Say fauna, Meade County, Kansas (Hibbard & Taylor, 1960), the Doby Springs local Geologic Range. Late Pleistocene fauna, Harper County, Oklahoma, and (Illinoian) to Recent. The oldest definite Lake Thatcher beds, Caribou County, occurrence of this species is from the Idaho (Miller, 1963), indicate a much Illinoian Doby Springs local fauna, of MILLER

Harper County, Oklahoma. (Ogallala Formation, Buis Ranch local Distribution. "Found over most of fauna, Beaver County, Oklahoma, to northern North America north of about Recent (Hibbard & Taylor, 1960). 41 degrees; southward in the Western Distribution. "From Quebec and United States to northern California, Massachusetts west to California; Yukon Nevada, and southern Colorado; un- Territory and James Bay south to Mary- known in the northern Great Plains be- land, Indiana, Colorado, and California" tween the Rocky Mountains and the (Baker, 1928a: 263). eastern Dakotas and northwestern Ecology. "In the Mississippi Valley . Nebraska" (Hibbard & Taylor, 1960: 85). this species seems to almost invariably Ecology. "The American form of occupy intermittent streams or small stagnalis is usually foundinmore or less pools, ponds and ditches which dry up stagnant parts of ponds or lakes and in summer. In Illinois it is usually rivers about vegetation. It may fre- found in association with Aplexa hyp- quently be seen floating among pond norum and Sphaevzum occidentale, either weeds and algae, the foot applied to in small streams, pools or sloughs, or the surface film of the water, the shell in spring pools in the woods which be- hanging downward" (Baker, 1928a: 202). come completely dry in the late spring Material. Doby Springs local fauna, and summer. The species hibernates locality 1, 213742 (8); locality 4, 208902 to a greater degree than any of the (21); locality 5, 213756 (2). other Lymnaeas, a fact attested to be Remarks. All the material from the the many rest varices observed on the Doby Springs local fauna consists of shell of large individuals. In these fragments of spires. These compare dry ponds living specimens may fre- well with the more complete material quently be found by digging into the mud, from the Bar M. local fauna of Harper leaves and other debris. In Wisconsin, County, Oklahoma (Taylor & Hibbard, capevata has been found almost invari- 1955). ably in small woodland pools which be- Hibbard & Taylor (1960: 86) have come dry in summer and fall, or in suggested the possible presence of small streams which become wholly or Lymnaea stagnalis jugularis in the partially dry" (Baker, 1928a: 263). Hib- Cudahy fauna of Lincoln County, Kansas. bard & Taylor (1960) have found S. They surmised that the record of Acella capevata in association with Stagnicola haldemani from that fauna reported by palustvis, Promenetus umbilicatellus Frye, Leonard & Hibbard (1943: 41) and Aplexa hypnomm in temporary might have been based on a misidenti- bodies of water in northern Nebraska. fication of immature spires of L. s. Material. Berends local fauna, 208830 jugularis. Neither of these species has (25). Doby Springs local fauna, locality been recorded from that locality in sub- 1, 213721 (16); locality 4, 208901 (350); sequent faunal lists for the Cudahy locality 5, 213755 (27). Mount Scott local fauna (Leonard, 1950: Frye & Leonard, fauna, locality UM-K4-53, 200647 (1200); 1952). Acella haldemani, however, has locality UM-K1-60, 213790 (6); locality been recently collected by a University UM-K2-59, 213703 (4000). Locality of Michigan Museum of Paleontology field UM-K2-60, 214197 (1). party, along with other undescribed mol- Stagnicola exilis (Lea) lusks, from younger beds in nearby McPhearson County, Kansas, thus posing Geologic Range. Late Pliocene (Rex- the question of whether the original road Formation) to Recent. The oldest record from Lincoln County was indeed reported occurrence of S. exilis in the based on Lymnaea stagnalis jugularis. Meade County area is from the Rexroad local fauna (Hibbard & Taylor, 1960). Stagnicola caperata (Say) Distribution. "Ohio to Kansas, north- Geologic Range. Middle Pliocene ward to northern Minnesota and north- FIVE ILLINOIAN FAUNAS 227

ern Michigan" (Baker, 1928a: 227). situations, near water, on the mud, moist Ecology. "Exilis is an inhabitant of leaves and other debris. In Meade sloughs, ponds and streams which dry County, Leonard (1943) reports it to up more or less during a portion of the prefer living on wet ground among sedge year" (Baker, 1928a: 227). and grass. I have collected it from Material. Berends local fauna, 208814 leaf litter in a seepage area near the (21). Mount Scott local fauna, locality Doby Springs local fauna locality in UM-K2-59, 213696 (8). Harper County, Oklahoma, where it was associated with Succinea, Veertigo mil- Stagnicola rejlexa (Say) ium, Gastrocopta tappaniam and Fos- Geologic Range. Late Pliocene (Rex- saria obmsa. , road Formation, Bender local Fauna) to Material. Berends localfauna, 208832 Recent (Miller, 1964). (60). Doby Springs local fauna, locality Distribution. Eastern Quebec from 1, 213712 (137); locality 2, 185779 (4); approximately 65' longitude, west to locality 4, 208903 (286); locality 5, Nebraska to about 100' longitude, north 213785 (27). Mount Scott local fauna, to Manitoba to near 50' latitude, south locality UM-K4-53, 200608 (159); local- to southern Illinois and southern Kansas, ity UM-K1-60, 213792 (8); locality 37' latitude (Baker, 1911; 1928a). UM-K2-59, 213660 (110). Recent, Harper Ecology. S. rejlexa is found in creeks, County, Oklahoma. ponds, lakes and rivers, in sheltered Fossaria obmssa (Say) localities, attached to floating debris, submerged vegetation, stones, etc. Many Geologic Range. Late Pliocene (Rex- of these habitats become dry in the road Formation) to Recent. The earliest summer. In northern Illinois and Wis- tertiary record of this 3pecies in the consin it has been collected from small Meade County area is from the Bender pools or ponds and in swales located in local fauna (Miller, 1964). woods or open field; none have been Distribution. "From the Atlantic to found in large streams or lakes (Baker, the Pacific Oceans, and from the Mac- 1911; 1928a). Kenzie Territory, Canada, south to Ari- Material. Berends localfauna, 208830 zona and northern Mexico" (Baker, (25). Doby Springs local fauna, locality 1928a: 296). 1, 213743 (1); locality 4, 208942 (133); Ecology. "The normal habitat of this locality 5, 213754 (2). Mount Scott local species is in small bodies of water, as fauna, locality UM-K4-53, 200648 (12); creeks, ponds, sloughs, bays, and marshy locality UM-K1-60, 213791 (8); locality spots along river banks. It is at home UM-K2-59, 213661 (700). on sticks, stones, and any other debris that may be in the water or along its (Baker) Fossaria dalli edge" (Baker, 1928a: 296). Geologic Range. Early Pliocene (La- Material. Berends local fauna, 208807 - verne Formation) to Recent. The oldest (39). Doby Springs local fauna, locality Cenozoic record of this species in the 4, 208904 (50). Mount Scott local fauna, Meade County area is from the Laverne locality UM-K4-53, 200609 (90); locality local fauna (Hibbard & Taylor, 1960). UM-K1-60, 213793 (11); locality UM-K2- Distribution. "Ohio to northernMichi- 59, 213662 (2). Recent, Harper County, gan and Montana, south to Kansas and Oklahoma. Arizona" (Baker, 1928a: 288). It has been collected in Meade County, Kansas at Superfamily Ancyloidea the State Park (Leonard, 1943). Family Planorbidae Ecology. This species is semi-aquatic Armiger crista (Linnaeus) in its habits and is only rarely found submerged in water. It lives inmarginal Geologic Range. Late Pleistocene to B. B. MILLER

PLATE 111

FIGS. 1-3. Zonitoides nitidus (Muller) , Mount Scott locality UM-K4- 53. Apertural , basal and apical views. X8, UMMZ 213789a. FIG. 4. Laevapex fuscus (Adams), Doby Springs locality 4. Apical view. X8, UMMZ 208916a. FIGS. 5-7. Gyraulus deflectus (Say), Doby Springs locality 1. Apertural, basal and apical views. X8, UMMZ 213706a. FIGS. 8-10. Strobilo@s labyrinthica (Say), Mount Scott locality UM-K4-53. Apertural, apical and basal views. X8, UMMZ 200640a. FIVE ILLISOIAS FAUSAS 2 MILLER

Recent. In the southwest Kansas-north- 1960: 57). west Oklahoma area, the oldest record Material. Berends local fauna, 208818 is from the Berends local fauna, of (99). Doby Springs local fauna, locality Beaver County, Oklahoma. 1, 213708 (75); locality 2, 185788 (2); Distribution. &Holarctic; in North locality 4, 208907 (114); locality 5, America, north of about 41 degrees, but 213759 (35). Mount Scott local fauna, south of this latitude on the Pacific locality UM-K4-53, 200616 (900), Coast to San Mateo County, California, 200617a (3800); locality UM-K1-60, and southward in central UtahJ' (Hibbard 213795 (200); locality UM-K2-59,213664 & Taylor, 1960: 101). (7300). Ecology. Armiger crista is a species of ponds and marshes (Hibbard &Taylor, Gyraulus deflectus (Say) 1960). Goodrich (1932: 65) states that Plate 111, Figs. 5-7 this snail "lives upon water-logged Planorbis deflectus Say, 1824: 261, PI. sticks and rotting leaves in stagnant XV, Fig. 8 water." In temporary ponds in southern Planorbis hirsutus Gould, 1840: 196 Michigan, U. S. A., A. crista is believed Planorbis obliquus DeKay , 1843: 62, PI. to hibernate after burrowing into the mud IV, Fig. 57a,b (Kenk, 1949). Beetle (1965) collecting Gyraulus deflectus (Say), F. C. Baker, from ponds in Grand Teton National Park, 1928a: 370, PI. XXIII, Figs. 15-18, 20- Wyoming, U. S. A., found this species 21 to be restricted to one pond. This pond, Gyraulus deflectus obliquus (DeKay) , F. which was subjected to periodic drying, C. Baker, 1928a: 372, PI. XXIII, Figs. was the only one that posessed a bottom 22-26 composed of colloidal muck, inter- Gyraulus hirsutus (Gould) , F. C. Baker, spersed with a mat of decaying vege- 1928a, PI. XXIII, Figs. 8-14 tation. The surface of the pond was Gyraulus deflectus (Say), Taylor, 1960: 5 7 completely covered with a marsh trefoil. Material. Berends local fauna, 208811 Geologic Range. Middle Pleistocene (19). Doby Springs local fauna, locality (Kansan) to Recent. The earliest known 1, 213709 (2); locality 4, 208905 (32); occurrence in the Meade County area is locality 5, 213763 (146). Mount Scott from the Cudahy fauna (UMMZ 177244). local fauna, locality UM-K4-53, 200602 Distribution. - ~aritimeProvinces in (19); locality UM-K1-60, 213794 (I), Canada, south to Potomac River, Vir- locality UM-K2-59, 208663 (77). ginia; northwest to Montana, Idaho, British Columbia, Northwest Terri- Gyraulus circumstviatus (Tryon) tories and Alaska. Geologic Range. Middle Pleistocene The distribution map (Fig. 5) is based (Kansan, Cudahy Fauna) to Recent. on published records and materials ex- Distribution. 'North America approxi- amined in the U. S. National Museum mately between 40° and 50' N. latitude: (USNM) and University of Michigan south in Rocky Mountains to central New Museum of Zoology (UMMZ) collections. Mexico" (Taylor, 1954: 3). OIn the The following is a list of localities Great Plains it ranges south to northern along the southern periphery of its dis- Nebraska" (Taylor & Hibbard, 1955: 14). tribution: Ecology. "Gymlus civcumstriatus is Virginia: RoachesRun (UMMZ 87959); characteristic of small, seasonal water Belvoir, Ft. Humphrey's, drift of Poto- bodies, such as wood pools, marshes, mac River (USNM 466527); Fairfax ponds on flood plains or prairie ponds. County, Potomac River at Mt. Vernon In northern Nebraska it was found in (USNM 572015); Potomac River at seepages beside streams, and in Alexandria (UMMZ 156782). temporary Sand Hills ponds'' (Taylor, District of Columbia: Anacostia, FIVE ILLINOIAN FAUNAS 231

FIG. 5. Distribution of Gyraulus deflectus. 232 B. B. MILLER

Poplar Point (USNM uncat.). include intermediates which grade into Ohio: Hamilton County, Cincinnati the G. deflectus form, Baker (1928a: 372) (USNM 121165); Franklin County, has tried to preserve the G. deflectus Columbus (USNM 30228a). obliquus, even though he admitted that it Illinois: Mason County, Quiver Creek; was not always possible to separate it Quiver and Dogfish Lakes (Baker, 1906). from G. hivsutus. Champaign County, Crystal Lake, Ur- Gyraulus hivsutus has been separated bana (Zetek, 1918). from G. deflectus by its (1) rounded, sub- Iowa: Muscatine County, Muscatine angular periphery (2) deeper umbilicus, (USNM 476433); Linn County, Cedar and (3) more hirsute condition. None Rapids (USNM 476528); Cerro Gordo of these characters are consistent, with County, Mason City (USNM 506111); the hivsutus, deflectus and intermediate Dickenson County, Robinson Lake (USNM forms frequently occurring in the same 476519); Palo Alto County, Ruthven lot. (UMMZ 66256). The material in the National Museum Nebraska: Cherry County, Fort Nio- and University of Michigan Museum of brara National Wildlife Refuge, 5 miles Zoology collections, deposited under the east of Valentine. Pond and seepage name Gyvaulus borealis Westerlund, area at old fort reservoir (Taylor, 1960). falls within the range of variation of Montana: Flathead County, sec 1, T 28 G, deflectus as understood here, andhas N, R 21 W, 8 miles east of Kalispell been included in the distribution map. (UMMZ 181060). Some authors (Binney, 1865; Dall, Idaho: Coeur dlAlene County, Coeur 1905; Baker, 1928a) have suggested that d'Alene (USNM 474743). Gyraulus deflectus might be a synonym British Columbia: Kamloops (USNM of the European G. albus (Miiller). Com- 382125). parison of G. deflectus with 8 lots of Ecology. "In northern Nebraska this G. albus from Europe, in the University species was found in a spring-fed pond of Michigan Museum of Zoology, in- with a temperature of 150 C. The snails dicates that they differ in the type of were crawling on aquatic plants, largely sculpture. G. albus has more regularly Cevatophyllum, Elodea, and algae, near spaced, raised, spiral and axial orna- the edge of the pond" (Taylor, 1960: mentation. The spiral sculpture is 5 7) better developed and appears as fine Material. Doby Springs local fauna, lirae resting on the more subdued axial locality 1, 213705 (200); locality 2, sculpture. In G. deflectus the sprial 185787 (12); locality 4, 208908 (140). sculpture, when present, is produced by Remarks. Examination of material in the alignment of impressed pits, which the collections of University of Michigan accommodate the hair-like projections Museum of Zoology and the U. S. National in the fresh shell. These pits are lower Museum indicates that the name com- than the axial growth lines. binations Gyvaulus obliquus, G. deflectus The differences in sculpture appear " obliquus and G, hivsutus, are all junior distinct enough to at least justify their synonyms of Gyvaulus deflectus. specific separation, but sufficient The differentia used by DeKay (1843: European material has not been examined - 62) to distinguish G. obliquus from G. to determine the consistency of this deflectus are the obliquity of the character. aperture and the absence of the medial Gyraulus pavvus (Say) peripheral angulation. Neither of these characters are considered valid. There Geologic Range. Late Pliocene (Rex- are many lots with individuals in which road Formation) to Recent. The first both of these characteristics attain occurrence of this species in the Meade different degrees of development, and County area is in the Saw Rock Canyon FIVE ILLINOIAN FAUNAS 233

local fauna, Seward County, Kansas (Hib- souri. The southern distribution is not bard & Taylor, 1960). clear owing to mixing with related Distribution. UEasternNorth America species" (Baker, 1928a: 332). east to the Rocky Mountains from Florida Ecology. This species is primarily northward to Alaska and northern British an inhabitant of quiet, stagnating water. Columbia" (Baker, 1928a: 377). In the Meade County area, it has been Ecology. LfUsuallyin quiet bodies of found living in temporary ponds and water; often of small size . . . . This more permanent bodies of water (Leon- . species is more often found in vegetation ard, 1943). than in any other situation" (Baker, Material. Berends local fauna, 208836 1928a: 376-377). In Meade County, Kan- (43). Doby Springs local fauna, locality sas, and Harper County, Oklahoma, this 4, 208910 (12); locality 5, 213757 (2). species was collected from rooted Mount Scott local fauna, locality UM-K4- vegetation growing in premanent water 53, 200622 (10); locality UM-K2-59, situations. 213701 (49). Recent, Harper County, Material. Berends localfauna, 208801 Oklahoma. (45). Doby Springs local fauna, locality Planorbula avmigera (Say) 1, 213707 (700); locality 2, 185782 (50); locality 4, 208906 (1700); locality 5, Geologic Range. Middle Pleistocene 213760 (1900). Mount Scott local fauna, (Kansan, Cudahy fauna) to Recent. locality UM-K4-53, 200618 (1340), Distribution. "New England west to 200617b (5660); locality UM-K1-60, Nebraska, south to Georgia and Louisi- 213797 (1650); locality UM-K2-59, ana, north to Great Slave Lake" (Baker, 213665 (226). Recent, Harper County, 1928a: 359). Oklahoma. Ecology. "Planorbula armigera is largely a species of swales or of small Helisoma anceps (Menke) stagnant bodies of water" (Baker, 1928a: Geologic Range. Late Pliocene (Rex- 358). road Formation. Saw Rock canvon local Material. Berends local fauna, 208823 fauna) to ~ecent'(Taylor, 1960)." (700). Distribution. "Oregon to Maine, south- Promenetus exacuous-kansasensis westward to western Mexico and Ala- Complex bama" (Hibbard & Taylor, 1960: 103). Pvomenetus exacums form exacuous Ecology. #. . . primarily a river and creek species, not living in large lakes" (Say) (Baker, 1928a: 319). Geologic Range. Late Pleistocene Material. Berends local fauna, 208805 (~llinoian)to Recent. (17). Doby Springs local fauna, locality Distribution. NNorth America north 1, 213714 (32); locality 2, 185785 (3); to about 39'; south in the Colorado locality 4, 208909 (126); locality 5, Plateau and Rocky Mountains to New 213758 (600). Mount Scott local fauna, Mexico; south along the Atlantic Coast- locality UM-K4-53, 200621 (161); al Plain to North Carolina; an isolated locality UM-K1-60, 213798 (175). occurrence in cool springs, Meade County, Kansas" (Hibbard & Taylor, Helisoma trivolvis (Say) 1960: 107). Geologic Range. Early Pleistocene Ecology. "Promenetus exacuous is (Nebraskan-Aftonian, Dixon local fauna) found in shallow, perennial or subperma- to Recent (Taylor, 1960). nent, quiet-water bodies, such as ponds, Distribution. "Atlantic Coast and oxbow lakes, marshes, and sloughs or Mississippi River drainages, northward back waters along streams. It is usually to Arctic British America and Alaska on the submerged vegetation in such and southward to Tennessee and Mis- habitatsn (Hibbard & Taylor, 1960: 108). 234 B. B. MILLER

Material. Mount Scott local fauna, Draw local faunas) through the late locality UM-K4-53, 200635 (500); local- Illinoian (Mount Scott local faunas), seem ity UM-K1-60, 213799 (200); locality to indicate a trend toward a decrease UM-K2-59, 213667 (190). in the proportion of riblet-bearing to Remarks. Hibbard & Taylor (1960: non-riblet-bearing individuals. This 106) recognize 2 distinct species of trend appears to be reversed in the P~omenetusfrom the late Cenozoic of younger Cragin Quarry local fauna, in the southern Great Plains, P. kansas- which most of the shells possess well ensis, which occurs in faunas from Plio- developed riblet sculpture. The younger ' cene to late Pleistocene, and P. exacuous, Jinglebob and Jones local faunas con- that has a geologic range of late San- tain shells that are predominantly with- gamon to Recent. They state that "Plio- out riblets. In most of the Recent lots - cene to late Pleistocene series of P. examined the sculpture consists almost kansasensis differ consistantly from P. entirely of growth lines, with only an exacuous in that all or most shells have occasional individual possessing riblets a sculpture of riblets instead of growth- in the later whorls. However, at least lines. " Evolutionary replacement of 5 lots of Recent shells examined in the P. kansasensis by P. exacuous some- University of Michigan Museum of time in the Sangamon between the time Zoology collections (UMMZ 47019; the Cragin Quarry and Jinglebob local 36844; 179041; 179040; 179038) con- faunas lived, is suggested by these tain individuals in which all or most authors. The shells from the Mount possess well developed axial riblets in Scott local fauna do not contain the the adults. characteristic riblet sculpture of P. The distribution of riblet andnon-rib- kansasensis, and must be considered P. let-bearing shells through time seems exacuous. The occurrence of P. exac- susceptible of several possible inter- uous in the Illinoian, makes untenable pretations: the suggested placement of the nomen- (1) The 2 types of sculpture may clateral change from P. kansasensis to represent distinct sympatric species P. exacuous, sometime in the Sangamon. which have existed together since at Examination of the P. kansasensis-P. least the Pliocene Saw Rock Canyon exacuous complex from llfaunasoccur- fauna. The fluctuating proportions of ring in beds that range in age from Upper riblet to non-riblet-bearing individuals Pliocene (Saw Rock Canyon local fauna) in the different faunas may be due to through Wisconsin (Jones local Fauna), sampling error and/or selective en- confirms Hibbard & Taylor's obser- vironmental pressures. vation that there are no consistent differ- (2) There may be 2 species involved, ences in size, compression of whorls, or with P. kansasensis evolving into P. strenth of carina through time. However, exacuous sometime in the late Illinoian, the strength of the axial sculpture, its between the time represented by the persistence to the adult stage, and the Butler Spring and Mount Scott faunas. proportion of riblet to non-riblet-bearing The coarse riblet shells in the Cragin individuals in a given population also Quarry local fauna might be a pheno- appear to be inconsistent. typic response to the effects of in- There is a higher proportion of non- creasing aridity, a condition suggested riblet to riblet-bearing individuals in by some of the other faunal elements the Pliocene Saw Rock Canyon local (Hibbard & Taylor, 1960). fauna than in the younger Bender local (3) The 2 types of sculpture may fauna population from locality UM-K1- represent the extremes of variation with- 59. The Pvomenetus examined from the in one very variable species, with the sequence of Pleistocene faunas from changes in the proportions of riblet to Nebraskan-Aftonian (Dixon and Sand non-riblet-bearing individuals afunction FIVE ILLINC)IAN FAUNAS

of selection pressures by different en- populations of P. kansasensis-P. exacu- vironmental conditions. ow complex are the result of environ- The first of these explanations is mental or genetic differences. It seems rejected on the grounds that there appear best at this stage to recognize the 2 to be no conchologic characteristics that types of populations, i.e., those com- can be used to separate individuals of posed predominantly of shells bearing these 2 species if they occur together riblets, and those in which the non- in the same fauna. Hibbard & Taylor riblet-bearing shells are in the majority, (1960: 106) in a discussion of the vari- with the names Promenetus exacuacs ability of P. kansasensis state that form kansasensis and P. exacuous form "Axial riblets, the most conspicuous ele- exacuous, respectively. ment of sculpture, vary from strong Promenetus exacLLous and widely spaced . through weaker and . . form kansasensis (Baker) closely spaced .. . to a point at which the sculpture may be said to consist of Geologic Range. Late Pliocene (Rex- growth lines rather than riblets . . . ." road Formation) to Late Pleistocene At this point the sculpture is indis- (Kingsdown Formation). The oldest tinguishable from P. exacuous. record of this form in the Meade County The proportion of riblet to non-riblet- area is from the Saw Rock Canyon local bearing individuals in a population can- fauna (Hibbard & Taylor, 1960). not be used to separate these 2 forms Distribution. Pliocene in southern since they show no consistent tendency Idaho, northern Texas and southwest- with respect to these ratios through ern Kansas; early Pleistocene in the time. Thus, while the Mount Scott Great Plains, from Iowa to Texas; late population is composed primarily of the Pleistocene southwestern Kansas, north- non-riblet-bearing form, the younger western Oklahoma (Hibbard & Taylor, Cragin Quarry local fauna, which is 1960). located approximately 50 ft higher up Ecology. P. e. form kansasensis is the slope of the same hill, contains assumed to have had habitat require- shells in which individuals with riblets ments similar to those of the form exacu- predominate. In the still younger Jingle- OUS . bob and Jones local faunas the non- Material. Berends localfauna, 208808 riblet-bearing form is dominant. (469). Doby Springs local fauna, locality There is the possibility that the shells 1, 213715 (257); locality 4, 208911 (850); possessing strong riblet sculpture in the locality 5, 213761 (350). Cragin Quarry local fauna might be the Promenetus umbilicatellus (Cockerell) result of increased aridity, with a concomitant rise in the concentrations of Geologic Range. Late Pliocene (Rex- alkaline salts. This explanation receives road Formation, Saw Rock Canyon some support from the xeric character local fauna) to ~ecent(Hibbard & ~a~ior, of the lizard fauna and the 2 mammals, 1960). Notiosorex crawfordi and Dasypterus Distribution. North America north of golliheri (Hibbard & Taylor, 1960). How- about the 41st parallel from western ever, the absence of riblets in the New York, northwest to Alaska, south 7 associated aquatic mollusks would seem to Washington and Oregon; in Rocky to argue against this interpretation. Mountains south to Colorado; sporadic At the present time there appears to in eastern Kansas and Oklahoma (Hibbard be no satisfactory way of separating the & Taylor, 1960). predominantly riblet and non-riblet- Ecology. This species is an inhabit- bearing populations. More faunas must ant of temporary bodies of water, where be studied before it can be determined it frequently is found associated with whether the observed variations in the Stagnicola caperata, S. palustris and MILLER

Aplexa hypnorum (Hibbard & Taylor, UM-K4-53, 200607 (4); locality UM-K1- 1960). 60, 213803 (7); locality UM-K2-59, Material. Doby Springs local fauna, 213698 (7). locality 1, 213715 (1); locality 2, 185791 Remarks. Basch (1963) in a recent (1); locality 5, 213762 (3). Mount Scott review of the North American fresh- local fauna, locality UM-K2-59,213666 water limpets found that the significant (5200). Locality UM-K2-60, 214187 (2). characteristics of Fevvissia meekianu overlap those of F. fvagilis in such a Family Ancylidae manner as to make equivocal their Fevvissia fvagilis (Tryon) specific distinction, and placed F,meek- iana in the synonymy of F.fvagilis. Many Ferrissia meekiuna (Stimpson), Hibbard of the collections of F. fvagilis examined & Taylor, 1960: 113; Miller, 1961: 108 by Basch contained both septate and non- Fe?rissia fragilis (Tryon), Basch, 1963: septate individuals. Specimens with 435 septa, however, were found to be indis- tinguishable from those in which there Geologic Range. Late Pliocene (Rex- was no septum formation, and he con- road Formation) to Recent. In the concluded "that there is no reasonable southern Plains it is first known from basis for placing the North American the Bender local fauna, locality UM-K1- septate forms in a separate genus (Gund- 59, UMMZ 208848. lachia)" (Basch, 1963: 438). Distribution. "Fevvissia fvagilis In this report the writer follows the appears to be among the most widely specific dispositions of the freshwater distributed of North Americafreshwater limpets proposed by Basch. snails" (Basch, 1963: 436). It probably occurs in suitable habitats throughout Laevapex fuscus (Adams) the United States. It has been recorded Plate 111, Fig. 4 from Alabama, North Carolina, Mis- Laevapex kirklandi (Walker), Hibbard & souri, Virginia, District of Columbia, Taylor, 1960: 114 Texas, California, Oregon, Washington, Laevapex fuscus (C. B. Adams), Basch, Nebraska, Kansas, Oklahoma and from 1963: 419-420, Fig. 7 Iowa eastward to Massachusetts (Hib- bard & Taylor, 1960; Basch, 1963; Geologic Range. Late Pleistocene Branson & Peters, 1964). (Illinoian) to Recent. The earliest record Ecology. "These snails often may be is from the Doby Springs local fauna, found on the bottom inch of stems of in the Meade County area. Typha and other rooted flowering plants Distribution. "United States and Can- growing in ditches and swamps . . . . The ada, generally east of the Great Plains. water in which they live is often dark Great Lakes area, Florida and south- and foul-smelling, and the habitats eastern states. Generally absent from usually have a mud bottom. One often mountainous areas" (Basch, 1963: 420). finds leeches, sphaeriid clams, and small Ecology. "Commonly in impoundments planorbid snails such as Gy~aulusand or cut-off stagnant backwaters of rivers. Menetus associated with these lim- The species is also found in lakes and pets .... Septate forms are known from occasionally in slow-flowing rivers on 2 principal types of habitat, temporary dense vegetation near the shores .. . L. woods pools such as those found in the fuscus is often found on the under sides northern states, and roadside ditches of lily pads, on cat-tails, sedges, and from whichwater presumably disappears other emergent rooted vegetation. * for part of the year" (Basch, 1963: 436- (Basch, 1963: 420). 4 37). Material. Doby Springs local fauna, Material. Doby Springs local fauna, locality 4, 208916 (27), 213825 (8). Mount locality 4, 208915 (6); locality 5, 213784 Scott local fauna, locality UM-K4-53, (2). Mount Scott local fauna, locality 200623 (17); locality UM-K1-60, 213804 FIVE ILLINOIAN FAUNAS 237

(75), 213805 (68); locality UM-K2-59, living in a stock tank on the Big Springs 213703 (6); locality UM-K3-60, 214174 Ranch, Meade County, Kansas. (5). Recent, Harper County, Oklahoma. Ecology. "Hildvethiana is character - Remarks. Basch (1963) found suf- istic of swales and summer-dry ponds, ficient overlap in the characters of L. where it is deprived of moisture for a fusw and L. kirklandi which led him large part of the year. The variety in to believe that they are conspecific. The such localities rarely reaches maturity, classification proposed by Basch is fol- the form being small, short, shell with lowed herein, and L. kivklandi is placed dome-shaped spire, greatly resembling in the synonymy of L. fuscus. oleacea and elliptica. These small forms usually show no rest marks, living only Family Phy sidae as long as the water remains inthe pool. Physa anatim Lea Mature shells may often be found by digging in the mud or in the cracks in Geologic Range. Late Pliocene (Rex- the pool where the mollusk has descended road Formation, Saw Rock Canyon local for moisture. Large specimens may fauna) to Recent (Hibbard & Taylor, have as many as four rest marks on 1960). the shell. In some sloughs the water Distribution. Some of the problems may become very low in summer but does which have resulted in the confused taxo- not completely disappear. In such nomic state of the genus Physa have localities hildrethiana attains its finest recently been summarized by Branson development . . ..I' (Baker, 1928a: 454). (1961). Until such time as the entire Material. Berends local fauna, 208817 family is revised, the geographic dis- (97). Doby Springs local fauna, locality tribution, ecology, life history and fossil 4, 208913 (300). Mount Scott local fauna, record of members of this group will locality UM-K4-53, 200626 (270); remain unclear. locality UM-K1-60, 213801 (45); locality Ecology. "Physa anatina is an in- UM-K2-59, 213669 (135). habitant of quiet and stagnant water; it Remarks. Branson (1961) reports that thrives in small ponds, or even in metal Physa g-yvina occurs inall the counties of stock tanks" (Leonard, 1950: 21). In Oklahoma. The Big Springs Ranch re- Meade County, Kansas, and Harper cord is the first reported occurrence of County, Oklahoma, this snail has been this species from Meade County, Kansas. collected in small spring fed streams. Physa skinneri Taylor Material. Berends local fauna, 208816 (58).- Springs local fauna, locality Physa elli$tica 7ninor Crandall, 1901, 1, 213723 (20); locality 4, 208912 (300). Nautilus, 15: 55 (non Physa fontinalis mi- Mount Scott local fauna, locality UM-K4- nor Moquin-Tandon, 1855: 451, 452) 53, 200625 (320); locality UM-K1-60, Physa skinneri Taylor, 1954. Taylor & 213800 (125); locality UM-K2-59,213670 Hibbard, 1955; Hibbard & Taylor 1960; (170). Taylor, 1960; Miller, 1961 Physa gyrina Say, form hildrethiana Lea Geologic Range. Lower Pleistocene Geologic Range. Middle Pleistocene (Nebraskan) to Recent. Earliest record - (Kansan, Cudahy fauna) to Recent. from the southern Great Plains is from Distribution. Probably found through- the Nebraskan Dixon local fauna (UMMZ out most of North America north of 182208, 191518) (Hibbard & Taylor, Mexico. Baker (1928a: 454) states 1960). "hildrethiana is common east of the Distribution. From Alaska southward Mississippi River from western New to northern Utah; eastward through York and Pennsylvania to Illinois and South Dakota, Iowa, Michigan to south- south to Alabama. Occurs also in Iowa eastern Ontario. The following is a list and possibly in other states west of the of additional Recent localities, which Mississippi River. " It has been found supplements the one published in Hib- B. B. MILLER FIVE ILLINOIAN FAUNAS 239

bard gs Taylor (1960: 119-120). The Physa were common" (Hibbard & distribution map (Fig. 6) is based on all Taylor, 1960: 120-121). The collections known Recent occurrences of this made from Hays, Alberta, contain Physa species. skinneri associated with Stagnicola cap- British Columbia: Stoney Creek, near erata, Physa gyrinu, Gyraulus parvus, Vanderhoof (USNM 601 340). Helisoma trivolvis and Promenetus um- Alberta: Beaver Hills Lake, Tofield bilicatellus. (USNM 471229). Taber (UMMZ 214557, Material. Berends localfauna, 208817 . 214558). Hays (UMMZ 214554, 214555). (97). Doby Springs local fauna locality Rolling Hills (UMMZ 214556). Shore, 1, 213722 (1); locality 4, 208914 (2i). Hastings Lake, 30 miles east of Ed- Mount Scott local fauna, locality UM- monton (UMMZ 214159). K4-53, 200627 (3); locality UM-K2-59, Manitoba: Pothole 9, near Minnedosa 213671 (40). (UMMZ uncat.). Pike Lake, The Pass Remarks. There is every reason to (UMMZ 214154). believe that the lot catalogued under Ontario: Cochrane District; Smoky UMMZ 118095 are the specimens on Falls, Kapuskasing (UMMZ 214157). which Crandall based Physa elliptica Hastings Co.; Bay of Quinte, 3 miles minor. Enclosed with the shells is a east of Belleville (UMMZ 214158); Pond note stating "These little shells were 2 miles south of Thomasburg, Hungerford sent to me by Strengfrom GrandRapids." Township (UMMZ 214155, 214156). This agrees with the statement made Wyoming: Teton County (Beetle, 1962: by Crandall (1901: 55) "Physa elliptica 74). minor n.v. was sent to me from Grand Iowa, Palo Alto County: Elbow Lake, Rapids, Mich., by Mr. Streng." This Ruthven (UMMZ 157548); Trurnbell material is part of the Crandall col- Lake (UMMZ 157549); Lost Island Lake lection acquired by Bryant Walker. (UMMZ 157547). The measurements in millimeters of South Dakota: Roberts County, Big 11 individuals from this lot (UMMZ Stone Lake (UMMZ 157551). 118095) are given below: Michigan: Wayne County, Grosse Isle (UMM Z 28800). Kent County, Grand Height of Rapids (UMMZ 118096, 118095). Bay Height Width Aperture Number of County, Saginaw Bay, Union Beach mm mm mm Whorls (UMMZ 28795). 3.5 2.75 2.5 3. 25 Ecology. The Recent records of this 3.5 2. 25 2. 5 2. 75 species suggest that it lives in shallow 3.6 2. 3 2. 5 2.75 bodies of water, either perennial or 3.6 2. 25 2. 3 2.5 temporary, ponds, sloughs or backwaters 3.75 2. 3 2. 55 2.5 along streams. The presence of P. 3. 12 2. 0 2. 3 2. 25 skinneri in the Bay of Quinte on Lake 2. 66 1. 87 1.75 2. 25 Ontario, may indicate a new type of 3. 0 2. 25 2. 6 2. 25 habitat not previously recorded, but there 3.12 2. 12 2. 5 2. 25 is no detailed information on the label 3. 5 2. 12 2. 6 2. 25 to indicate the local environmental 3. 5 2. 25 2. 5 2.5 - conditions. lnLincoln County, Wyoming, Taylor collected it from a shallow per- Examination of this series of syntypes ennial pond on the floodplain of the Bear of P. e. minov and the types of P. skin- River. This pond contained a rich neri, indicates that the latter is a junior growth of submergent aquatic plants synonym of the former. The name com- growing in a soft organic mud bottom, bination Physa minor, however, is pre- marginal cattails, and many mollusks. occupied by a European species, Physa Valvata, Lymnaea, Promenetus and fontimlis minor Moquin-Tandon, 1855. MILLER

The first available name for this species places; under wood, such as old board is Physa skinnevi Taylor. sidewalks; in chinks of stone walls and under stonesJ' (Pilsbry, 1948: 1049). Aplexa hypnomm (Linnaeus) Material. Doby Springs local fauna, Geologic Range. Early Pleistocene locality 4, 208944 (1); locality 5, (Ballard Formation) to Recent. The 213781 (1). Mount Scott local fauna, oldest record of this species inthe Meade locality UM-K3-60, 214171 (1). County area is from the Sanders local fauna, Meade County, Kansas (Hibbard Superfamily Pupilloidea & Taylor, 1960). Family Strobilopsidae Distribution. Circumpolar: in North Stvobilops labyvinthica (Say) America it occurs from the Cascade Plate 111, Figs. 8-10 Mountains east to the Atlantic and from Alaska and Hudson Bay south to Nevada, Geologic Range. Late Pliocene (Rex- Colorado, the Platte River; the Ohio road Formation, Saw Rock Canyon local River and the District of Columbia. fauna) to Recent (Hibbard & Taylor, Ecology. According to Baker (1928a: 1960). 474) Aplexa hypnomm is a species of Distribution. Eastern North America, swales and intermittent streams or from Quebec to northernAlabama, west- stagnant pools in Wisconsin. It is re- ward to Manitoba, Black Hills, South ported as especially abundant in wood- Dakota, eastern Kansas and Oklahoma. land pools which become dry in sum- Ecology. "This snail inhabits forested mer, in company with Stagnicola capev- areas, and is found in and among the ata, Physa hildvethiana and Sphaev- leaf litter, decaying logs, and beneath ium occidentale. started bark and other debris of the Material. Berends local fauna, 208831 forest floor" (Leonard, 1959: 167). (25). Doby Springs local fauna locality Material. Berends local fauna, 208799 4, 208943 (4); locality 5, 213783 (1). (1). Doby Springs local fauna, locality Mount Scott local fauna, locality UM-K4- 1, 213741 (4); locality 4, 208917 (4). 53, 200601 (142); locality UM-K2-59, Mount Scott local fauna, locality UM- 213672 (500). K4-53, 200640 (338); locality UM-K1-60, 213806 (90); locality UM-K2-59, 213673 Order Stylommatophora (55). Superfamily Cionelloidea Remarks. Branson (1961), in one of Family Cionellidae his series of papers summarizing the Recent gastropod fauna of Oklahoma, Cionella lubvica (Miiller) listed the occurrence of Stvobilops lab- Plate I, Fig. 8 rinthica in that state without distinguish- Geologic Range. Late Pliocene (Rex- ing between Recent and fossil material. road Formation, Rexroad local fauna) The Recent records of S, texasianu to Recent (~a~lor,1960: 77). (which he considers a synonym of S. Distribution. Holarctic. NPointBar- labyvinthica) cited from Harper County, row, Alaska, and Queen Charlotte Islands may be based on fossil shells. to Labrador and Newfoundland, south in the East to Washington, D. C. and south- Family Pupillidae ern Missouri; in all the Western and Gastvocopta avmifera (Say) Mountain States except California; to the Mexican boundary in Arizona; inthe Geologic Range. Early Pliocene (La- Sierra Madre of western Chihuahua" verne Formation. Laverne local fauna) (Pilsbry, 1948: 1048). to Recent (~ibbard& Taylor, 1960). Ecology. "C. lubvica lives among Distribution. Eastern North America the damp under-leaves indensely shaded from the Maritime Provinces in Canada, FIVE ILLINOIAN FAUNAS 241

south to northern Florida; west to UM-K2-59, 213674 (2500). Locality Rocky Mountains. UM-K2-60, 214188 (1). Recent, Beaver Ecology. "Gastrocopta armifera is and Harper counties, Oklahoma. a gregarious species occurring com- Gastrocopta procera (Gould) monly on wooded slopes, near or removed from a stream. It is to be found Geologic Range. Middle Pleistocene under dead wood, limestone rocks, or (Kansan, Cudahy fauna) to Recent. light cover of leaf mold or other debris. Distribution. Maryland to South Caro- G, armifera frequently occurs under lina and Alabama; west to Black Hills, boards or rocks in gardensn (Franzen South Dakota, southeastern Wyoming, & Leonard, 1947: 329). I have collected and Arizona (Hibbard & Taylor, 1960; this species in large numbers in tall Reigle, 1963). - grass, under boards and beneathdead Ecology. "Gastrocopta procera is wood in cottonwood groves, in Meade typically an inhabitant of timbered slopes County State Park. At drier stations near streams, where it lives in leaf it was often associated with Gastro- mold, beneath fallen logs or loosened copta cristata and Pupoides albilabvis. bark, or beneath stones, but sometimes Material. Ber ends local fauna, 208824 it is found living in meadows in dead (29). Doby Springs local fauna, locality grass" (Leonard, 1950: 32). Its dis- 1, 213717 (3); locality 2, 185783 (1); tribution in Kansas is indicative of its locality 4, 208918 (27); locality 5, ability to withstand periods of high 213766 (97). Mount Scott local fauna, temperature and drought (Leonard & locality UM-K4-53, 200610 (1500); Goble, 1952). locality UM-K1-60, 213807 (50); locality Material. Berends local fauna, 208825 UM-K2-59, 213678 (1150); locality UM- (5). Doby Springs local fauna, locality 2, K3-60, 214178 (17). Locality UM-K2-60, 195796 (1); locality 4, 208921 (110); 214189 (87). Recent, Harper andBeaver locality 5, 213764 (15). Mount Scott counties, Oklahoma. local fauna, locality UM-K4-53, 200615 (2100); locality UM-K1-60, 213871 (19); Gastrocopta tappaniana (Adams) locality UM-K2-59, 213680 (83); locality Geologic Range. Late Pliocene (Rex- UM-K3-60, 214175 (7). Recent, Beaver road Formation, Saw Rock Canyon local and Harper counties, Oklahoma. fauna) to Recent (Hibbard & Taylor, Gastrocopta cvistata (Pilsbry & Vanatta) 1960). Distribution. "Ontario and Maine to Geologic Range. Late Pliocene (Rex- Virginia and Alabama, west to South road Formation, Bender local fauna) ~akotaand Kansas, southwest to Ari- to Recent (~ibbard& Taylor, 1960). zona, but not known from the south east- Distribution. Arizona and New Mexi- ern Atlantic States, Virginia to Florida" co, east to Arkansas and Louisiana; (Pilsbry, 1948: 889). north to Nebraska, Indiana. Ecology. 'This snail is found be- Ecology. This snail can be found in 4 neath wood, logs and similar debris in timbered areas, both upland and flood- moist places, especially on floodplains plain situations, as well as in grass- and in moist upland forested areas" lands (Leonard, 1943). In Meade County - (Leonard, 1959: 177). State Park, I have collected it in large Material. Berends localfauna, 208802 numbers from under pieces of scrap (696). Doby Springs local fauna, locality wood lying in a dry upland meadow. At 1, 213728 (64); locality 2, 185794 (6); this locality it was frequently an asso- locality 4, 208922 (800); locality 5, ciate of Pupoides albilabris. Both of 213765 (560). Mount Scott local fauna, these species are apparently xerophilic. locality UM-K4-53, 200615 (2100); local- Material. Berends local fauna, 208826 ity UM-K1-60, 213812 (96); locality (67). Doby Springs local fauna, locality 1, MILLER

213731 (1); locality 4, 219920 (350); water courses, under dead wood, leaf locality 5, 213767 (8). Mount Scott local mold and grass" (Franzen & Leonard, fauna, locality UM-K4-53, 200612 (710); 1947: 331). locality UM-K1-60, 213809 (88); locality Material. Doby Springs local fauna, UM-K2-59, 213677 (850); locality UM- locality 4, 208919 (1). Mount Scott K3-60, 214180 (2). Recent, Beaver and local fauna, locality UM-K4-53, 200611 Harper counties, Oklahoma. (830); locality UM-K1-60, 213808 (118); Gastvocopta pellucida locality UM-K2-59, 213676 (735). hovdeacella (Pilsbry) Gastvocopta holzilzgevi (Sterki) Geologic Range. Late Pliocene (Rex- Geologic Range. Late Pliocene (Rex- road Formation, Rexroad local fauna). to road Formation, Saw Rock Canyon local Recent (~ibbard& Taylor, 1960). fauna) to Recent (Hibbard & Taylor, - Distribution. "Southern and Baja, 1960). California (locally) through southeastern Distribution. "Ontario and western Colorado and Kansas to Florida; south New York to Helena, Montana, south to to Sinaloa and Tampico, Mexico; north- Illinois, Kansas, and Albuquerque and ward on the Atlantic Coastal Plain to Mesilla, New Mexico" (Pilsbry, 1948: southern New Jersey" (Hibbard & Taylor, 883). 1960: 123-124). "The northward ex- Ecology. "Gastvocopta holzingevi is tension along the Atlantic Coast is very found on timbered slopes under leaf narrow, discontinuous so far as known, mold . . . . In order to remain established and perhaps mainly confined to the in northwestern Kansas, as it is doingto coastal islands" (Pilsbry, 1948: 914). some extent, it must be tolerant of the Ecology. "Uncertain. From the geo- hot, dry climatic conditions that obtain graphic distribution, frequent occur- in the summer time. G. holzingevi is rence in stream drift in the southern not one of the predominating species Great Plains, and occasional occurrence of the fauna of Kansas. According to in samples of fossils washed from bulk its general distribution in North Ameri- collections of matrix, it appears this ca, it prefers decidedly cooler climates" species may live among the grass roots, (Franzen & Leonard, 1947: 334-335). perhaps sometimes even on exposed Material. Berends local fauna, 208809 slopes. So far as known, the snail has (29). Doby Springs local fauna, locality 4, not been collected alive in the Plains" 208940 (1); locality 5, 213749 (1). Mount (Hibbard & Taylor, 1960: 124). Scott local fauna, locality UM-K4-53, Material. Mount Scott local fauna, 200613 (980); locality UM-K1-60, 213810 locality UM-K2-59, 213675 (2). Recent, (1); locality UM-K2-59, 213679 (180); Harper County, Oklahoma. locality UM-K3-60, 214169 (1). Gastvocopta contvacta (Say) Vevtigo ovata (Say) Geologic Range. Early Pliocene (La- Geologic Range. Early Pliocene (La- verne Formation, Laverne local Fauna) Verne Formation, Laverne local fauna) to Recent (Hibbard & Taylor, 1960). to Recent. Distribution. Eastern North America Distribution. Labrador south to the from Maine, westward to Manitoba, South West Indies, west to Mexico, Arizona, .. Dakota, central Kansas, Oklahoma and Utah, Oregon and Alaska. western Texas, south to the states of Ecology. "Vertigo ovata ... lives only Morelos and Vera Cruz, in Mexico. The in moist environs afforded by shaded typical form is replaced by a different slopes near streams and shores of ponds. subspecies along the Gulf Coastal Plain Its range in Kansas extends westward (Hibbard & Taylor, 1960). to the generally dry regions of the High Ecology. "On shaded slopes along the Plains. In these regions are local FIT= ILLINOUN FAUNAS 243

ponds and streams, many of which are Geologic Range. Middle Pleistocene fed by the artesian springs, along whose (Kansan) to Recent. The earliest re- shaded slopes V. ouata is.found, though cords from the Meade County area are not in great numbers" (Franzen & Leon- from the Cudahy fauna, Sunbrite Ash ard, 1947: 355). Pit, UMMZ 177255a, and the Cudahy Material. Berends local fauna, 208803 Mine UMM Z uncatalogued. (1070). Doby Springs local fauna, local- Distribution. The distribution map ity 1, 213718 (438); locality 2, 185795 (Fig. 7) is based on material examined - (7); locality 4, 208925 (166); locality 5, in the collections of the University of 213768 (300). Mount Scott local fauna, Michigan Museum of Zoology, the U.S. locality UM-K4-53, 200654 (1280); local- National Museum, and published records. ity UM-K1-60, 213814 (85); locality The following is a list of peripheral UM-K2-59, 213682 (5500); locality localities: UM-K3-60, 214176 (2). Recent, Beaver British Columbia: Field; Pilsbry, and Harper counties, Oklahoma. 1948: 956 Alberta: Laggan; Pilsbry, 1948: 956. Vertigo milium (Gould) Manitoba: Ilford (UMMZ 180135). Geologic Range. Late Pliocene (Rex- Montana: Darby and White's Spring road Formation, Saw Rock Canyon local (West of Ward); Vanatta, 1914: 370). fauna) to ~ecent(Hibbard & -~a~lor, Wyoming: Teton County, Moran 1960). Junction; Beetle, 1962: 74. Distribution. uMaine and Quebec to Colorado: Lake George, Henderson, the Florida Keys, west to South Dakota, 1924: 136. Grand County, Colorado, and southeast- New Mexico: Socorro County, Oscoro ern Arizona; Tampico, Meldco; Mountains; Pilsbry, 1948: 956. Jamaica; Santo Domingo" (Pilsbry, Ontario: Moose Factory (UMMZ 1948: 945). 55957); Cochrane District, Smokey Falls Ecology. "Found in numbers under (UMMZ 180148); Patricia District, sticks, stones and leaves in moist situ- Borthwick Lake, (UMMZ 180150). ations along flood-plains and wooded Quebec: Hull County, Hull, Fairy stream borders'' (Leonard, 1959: 185). Lake (UMMZ 180136). It has been collected from beneathpieces Newfoundland: Boone Bay, Brooks of moss covered wood, in a seepage & Brooks, 1940: 72. area at Doby Springs Park, Harper Maine: Aroostook County, Caribou County, Oklahoma. (USNM 160160). Material. Berends local fauna, 208820 New York: Staten Island (USNM (21). Doby Springs local fauna, locality 4, 47497a). 208924 (92); locality 5, 213769 (255). West Virginia: Jefferson County; Mount Scott .local fauna, locality UM-K4- Brooks & Kutchka, 1938: 77; Preston 53, 200653 (245); locality UM-K1-60, County, Camp Dawson (USNM 533530). 213820 (2); locality UM-K2-59, 213681 Virginia: Montgomery County, Rad- (350). Recent, Harper County, Okla- for (USNM 523439); Fredrick County, homa. 1 mile east of Hayfield (USNM 533414). Ohio: Summit County, Hudson (UMMZ Vertigo elatior Sterki - 112178); Tuscarawas County, vicinity of Plate I, Fig. 11 New Philadelphia; Sterki, 1894: 5. Vertigo elutior, Pilsbry, 1948: 956, Indiana: Hendricks County, Danville Figs, 514; 515, number 6 (USNM 160140b). Vertigo elutior, Franzen & Leonard, Illinois: Winnebago County; Baker, 1947: 357, PI. XX, Fig. 2 1939: 106. Vertigo gouldi, Hibbard & Taylor, 1960: Iowa: Cerrogordo County, Mason 134-135, PI. XI, Fig. 10 (non Binney) City, dredged in Big Slough (USNM B. B. MILLER FIVE ILLIN()IAN FAUNAS 245

508637); West Lake Okoboji; drift on basal fold can occur as a single high north shore of Millers Bay (USNM tubercle, alone or in close proximity 476262). to an infrapalatal fold, that is also small, North Dakota: Polette County, Upsilon low and tuberculate; or as a high Lake; Winslow, 1921: 9. tubercle, with a closely associated infra- Ecology. "Vertigo elatior prefers a palatal tubercle. The palatal callus is cool humid climate characteristic of also variable, and is completely lacking northern United States or mountainous or poorly developed in some individuals. areas of southern states .. . . Prolonged Hibbard & Taylor (1960: 134) referred drought and high temperatures, charac- one shell (UMMZ 197569) from locality teristic of Kansas' summers, are factors 1 of the Butler Spring local fauna to ,, excluding V. elatior from Kansas today" Vertigo gouldi. However, this speci- (Franzen & Leonard, 1947: 357). men lacks the well developed axial Material. Doby Springs local fauna, sculpture characteristic of that species. locality 1, 213730 (30); locality 4, It does, though, compare favorably with 208938 (165); locality 5, 213770 (450). the large series of fossil Vertigo elatior Butler Spring local fauna, 197569 (1). from the Doby Springs local fauna, and Remarks. Vertigo elatior, as under- with the lots of Recent V. elatior shells stood in this report, may be recognized in the U. S. National Museum and Uni- by (1) its elongate-oval to conic-ovate versity of Michigan Museum of Zoology shell; (2) a strongly biarcuate aperture, collections. produced by a fold, which leaves an The description and illustration of V. elongate groove on the exterior of the gouldi in Franzen & Leonard (1947: palatal wall about halfway fromitsadap- 358, PI. 11, fig. 3) suggests that their ical end; (3) the outer margin of the material is probably V. elatior. The aperture, below this groove in the palatal presence of V. elatior in a collection of wall, which is slightly reflected; (4) the shells made in 1958, by a University of base of the shell which tends to be con- Michigan Museum of Paleontology field stricted and appea-rs pinched; (5) a party from the same locality listed by crest, parallel to the outer marginof the Franzen & Leonard for Vertigo gouldi aperture, which is usually present; (6) (Cudahy Mine, sec 2, T 31 S, R 28 W, the number of lamellae andfolds, charac- Meade County, Kansas), tends to support teristically 5 in number: one high, short this conclusion. angular lamella, located at the center of Pupoides albilabris (Adams) the parietal wall, whichis directed downward and towards the outer margin of Geologic Range. Early Pliocene (La- the aperture; 2 palatal folds, an upper, verne Formation, Laverne local fauna) which merges outwardly toward the to Recent (Hibbard & Taylor, 1960). mouth of the aperture with the inden- Distribution. iiEasternNorthAmerica tation in the palatal wall; and a more from southern Canada to the Gulf of ' deeply immersed lower palatal fold; a Mexico, west to the Dakotas, Colorado, low basal fold, located at the base of and Western Arizona; northern Mexico, the collurnelar wall; and a subvertical, Cuba, Haiti, Puerto Rico, Bermuda" collumelar fold, which ascends rapidly (Leonard, 1959: 181). inwardly. The palatal and basal folds Ecology. "This species, tolerant of are usually situated on and joined by a high summer temperatures and drought, callus. is found in woodlands, in deep grass, The chief variations observed occur or even among the roots of short grass in the number and size of the palatal in unshaded areas" (Franzen & Leonard, folds. A small, weak tubercle, in the 1947: 371). In Meade County, I have position of the suprapalatal fold, may observed this species crawling about on be developed in some individuals. The a barren outcrop during a bright sunny 24 6 B. B. MILLER

day after a good rain. In this area, from 2.5-3.5mm and 1.60-1.75mm re- the animal must seek shelter infractures spectively. and joints in the dried ground during Pupilla mwcovum (Linnaeus) the hotter part of the day, coming out only at night or after a rain. Geologic Range. Middle Pleistocene Material. Berends localfauna, 208819 (Kansan, Cudahy fauna) to Recent (Frye & (11). Doby Springs local fauna, locality Leonard, 1952: 158-159). 4, 208923 (15); locality 5, 213771 (14). Distribution. Holarctic. In North Mount Scott local fauna, locality UM-K4- America, from Newfoundland, U... Anti- 53, 200637 (620); locality UM-K1-60, costi Island south to Atlantic City, New 213813 (36); locality UM-K2-59, 213685 Jersey, westward in Canada and in the (275), locality UM-K3-60, 214181 (13). northern tier of states to Milton, Oregon; Locality UM-K2-60, 214191 (1). Recent, south in the Rocky Mountain region Beaver and Harper counties, Oklahoma. through Colorado to Socorro County, New Mexico, and northern Arizona; north to Pupilla blandi Morse Anuk, Alaska* (Pilsbry, 1948: 934). Geologic Range. Middle Pleistocene Ecology. "Lives in regions having a (Kansan, Cudahy fauna) to Recent. cool, humid climate. Is found living on Distribution. URocky Mountains, from the ground, under wood, stones, and New Mexico (at high elevations) to leaves* (Franzen & Leonard, 1947: 376). southern Alberta; northern Great Plains Material. Doby Springs local fauna, east to western North Dakota; Black locality 1, 213727 (105); locality 4, Hills, South Dakota" (Hibbard & Taylor, 208939 (58); locality 5, 213773 (190). 1960: 130). One lot in the U. S. National Museum collection (USNM 598457) from Family Valloniidae river drift of the Little Missouri River, Vallonia gracilicosta Reinhardt Dunn County, North Dakota, contains many shells that still have the perio- Geologic Range. Late Pliocene (Rex- stracum and indicate a more easterly road Formation, Rexroad local fauna) occurrence than any reported by Taylor to Recent, (Hibbard & Taylor, 1960). (Hibbard & Taylor, 1960: 131). Distribution. Rocky Mountains from Ecology. "Inthe higher, forested parts southern Arizona and New Mexico (at of the Rocky Mountains Pupilla blandi high elevations) northward to Alberta is widespread. It occurs in forest, or and Fort Norman, Northwest Terri- in meadows and parks where cover and tories; eastward to Manitoba, Nebraska, some moisture are available. Together Minnesota and Iowa. The eastern limits with Vallonia, it may often be found in of its distribution are uncertain because protected situations somewhat less damp of the difficulty of distinguishing V. than those required by snails such as gracilicosta from V. albula, which ex- Vitvina, Vertigo or Discus" (Hibbard & tends further east. Taylor, 1960: 131). Ecology. In northern Nebraska, onthe Material. Doby Springs local fauna, Fort Niobrara Wildlife Refuge, I have locality 5, 213772 (21). Mount Scott local collected V. g~acilicostafrom amongst fauna, locality UM-K4-53, 200638 (1); moist leaf mold, bark and fallen timber locality UM-K2-59, 213683 (5). Locality in cottonwood groves. UM-K2-60, 214185 (405). Material. Berends local fauna, 208806 Remarks. The large series of Pupilla (7). Doby Springs local fauna, locality 1, blandi from locality UM-K2-60 is rela- 213725 (1650); locality 2, 185797 (12); tively consistent in the number and de- locality 4, 208926 (620); locality 5, velopment of the denticles. The most 213774 (1120). Mount Scott local fauna, variable characters in this series are locality UM-K4-53, 200650 (800); local- the height and diameter, which range ity UM-K1-60, 213815 (20); locality FIVE ILLINOW FAUNAS 247

UM-K2-59, 213687 (110); locality UM- Remarks. Recent studies of Oxyloma K3-60, 214166 (32). Locality UM-K2-60, retusa and 0. haydeni by Franzen (1963, 214193 (500). 1964) have shown that it is not possible to separate these 2 species on shell Vallonia parvula Sterki characteristics. Fossil shells belonging Geologic Range. Late Pliocene (Rex- to the genus Oxyloma may, therefore, road Formation, Bender local fauna) to have beenincorrectly assigned to species Recent (Hibbard & Taylor, 1960). in earlier reports (Leonard, 1950; Frye Distribution. "Great Plains from & Leonard, 1952; Taylor, 1954, 1960; South Dakota to north-central Oklahoma Taylor & Hibbard, 1955; Hibbard & eastward to northern Missouri, northern Taylor, 1960; Miller, 1961). Ohio, southern Ontario, and western cf Succinea " New York" (Hibbard & Taylor, 1960: 138). Material. Berends local fauna, 208834 Ecology. " Vallonia parvula inhabits (6). Doby Springs local fauna, locality 1, wooded areas in both upland and flood- 213719 (160); locality 2, 185793 (13); plain situations. It has been found in locality 4, 208928 (163); locality 5, and about logs, sticks, rocks and similar 213780 (5). Mount Scott local fauna, niches, and often buries itself in the locality UM-K4-53, 200641 (66); locality ground to a depth of several times its UM-K1-60, 213817 (9); locality UM-K2- diameter* (Leonard, 1959: 189). In 5 9, 213693 (71); locality UM-K3-60, Meade County, it has been collected 214179 (1). Locality UM-K2-60, 214196 from leaf litter, in a cottonwood grove (150). near the west end of Lake Larrabee. Remarks. Most succineids cannot be Material. Berends local fauna, 208800 satisfactorily placed to genus or species, (23). Doby Springs local fauna, locality on the basis of shell characteristics 4, 208927 (4). Mount Scott local fauna, (Miles, 1958; Taylor, 1960). In the locality UM-K4-53, 200651 (561); local- present study, it has only been possible ity UM-K1-60, 213816 (8); locality to identify shells belonging to the genus UM-K2-59, 213686 (145); locality Oxyloma with reasonable confidence. All UM-K3-60, 214168 (4). Recent, Harper the remaining succineid material has County, Oklahoma. been referred to cf Succinea.

Superfamily Succineoidea Superfamily Endodontoidea Family Succineidae Family Endodontidae Oxyloma sp. Discus cronkhitei (Newcomb)

Oxyloma naverrei Leonard, 1950: 23, P1. Geologic Range. Middle Pleistocene 4, Fig. H. Frye & Leonard, 1952: 158, (Kansan, Cudahy fauna) to Recent (Frye P1. 15, Fig. ee & Leonard, 1952: 158-159). Oxyloma haydeni (Binney), Taylor, 1954: Distribution. Alaska south to the 5. Taylor & Hibbard, 1955: 8 mountains of southern California; south Oxylorna retzrsa (Lea), Taylor, 1960: in Rocky Mountains to Colorado, Ari- 78. Hibbard & Taylor, 1960: 141. Miller, zona and New Mexico; eastward across 1961: 108 northern Canada from British Columbia to Labrador and Newfoundland. East of Material. Berends localfauna, 208810 the Rocky Mountains the southern limits (188). Doby Springs local fauna, locality of its range are reached in south-central 1, 213720 (8); locality 4, 208929 (12). Nebraska, Missouri, Illinois, Indiana, Mount Scott local fauna, locality UM-K4- Ohio, West Virginia, Kentucky, Mary- 53, 200624, (143); locality UM-K2-59, land. A collection made from the flood- 213697 (200). plain of the Republican River, in Red 248 B. B. MILLER

Willow County, Nebraska, represents the Helicodiscus singleyanus (Pilsbry) southernmost record for this species in the High Plains. Geologic Range. Late Pliocene (Rex- Ecology. "In the east it lives in road Formation, Saw Rock Canyon local humid forest, under dead wood, and fauna) to Recent (Hibbard & Taylor, among rotting leaves or grass in rather 1960). wet situations. It is a common snail Distribution. "United States from New in the Canadian and Transition faunas, Jersey and Florida west to South Dakota, and occurs sporadically in the Caro- southwest to Arizona" (Leonard, 1959: linian" (Pilsbry, 1948: 604). In Red 133). Willow County, Nebraska, it was found Ecology. This species has apparently under sticks, logs and forest litter, never been taken alive in the High Plains on the flood plain of the Republican (Hibbard & Taylor, 1960). The occur- River, in association with Zonitoides rence of fresh shells among the fossil avboveus, Gastvocopta avmifeva, Devo- bearing matrix that has been washed cevas laeve and Gastvocopta tappanianu. from various localities in the southwest Material. Doby Springs local fauna, Kansan-northwest Oklahoma area, locality 1, 213724 (132); locality 2, seems to indicate that Helicodiscus 185776 (14); locality 4, 208930 (104). singleyanus can survive the hot dry Mount Scott local fauna, locality UM-K4- summers that characterize this region. 53, 200605 (2); locality UM-K1-60, The animal probably lives among the 213818 (10); locality UM-K2-59, 213691 grass roots during the hot, dry part of (1); locality UM-K3-60, 214182 (1). the day, and seeks shelter inthe fractures and cracks that have formed in the more He licodiscus pavallelus (Say) exposed surfaces. Geologic Range. Late Pliocene (Rex- Material. Mount Scott local fauna, road Formation, Bender local fauna) locality UM-K1-60, 213821 (2). Recent, to Recent (Miller, 1964). Beaver and Harper Counties, Oklahoma. Distribution. "Eastern North Ameri- Punctum minutissimum Lea ca, from Newfoundland south to Georgia Plate 11, Figs. 6-8 and Alabama; westward to South Dakota and Oklahoma" (Hibbard & Taylor, 1960: Geologic Range. Late Pleistocene 144). (Illinoian) to Recent. Ecology. "It lives on decaying wood ~istribution. Newfoundland south to in shady or humid places, also on damp Florida; west to southern Ontario, the leaves" (Pilsbry, 1948: 627). "Ineastern Lake States, South Dakota, Colorado, Kansas the species may occur in grassy New Mexico, Oregon and Mexico, (Pils- fields, on sparsely timbered slopes, and bry, 1948). The nearest Recent occur- on rocky ledges as well as in more rence to the Meade County area is from moist places. In the more arid parts of Phillips County Kansas (Leonard, 1959). the state, however, H. pavallelus is Taylor (1960) has collected Punctum limited to woodland cover, and usually minutissimum from Brown County, occurs around decaying timber" (Leon- Nebraska. ard & Goble, 1952: 1039). Ecology. "This dwarf among pygmies Material. Berends localfauna, 208812 lives on damp leaves, around decaying (7). Doby Springs local fauna, locality 1, logs, and is chiefly obtained by sifting 213733 (45); locality 4, 208931 (1425); leaves . .. Morse states that 'dense, hard- locality 5, 213776 (195). Mount Scott wood growths appear to be their favorite local fauna, locality UM-K4-53, 200620 position. They prefer the rotten bark (280); locality UM-K1-60, 213819 (20); of beech trees, and frequently are found locality UM-K2-59, 213692 (350); local- in the large forms of fungi, such as ity UM-K3-60, 214170 (52). Polypom and Boletus' " (Pilsbry, 1948: FrVE ILLINOIAN FAUNAS 249

645). usually be obtained by leaf sifting where Material. Mount Scott local fauna, its presence would otherwise be un- locality UM-K4-53, 200636. (43). suspected" (Pilsbry, 1946: 236). I have Remarks. The only previously re- collected it in Washtenaw County, Michi- ported fossil occurrence of P. minutis- gan, from a marsh on the floodplain of simum in the southern Plains region, has the Huron River among high grass and been from the Sangamon, Jinglebob local scattered shrubs. fauna (van der Schalie, 1953). Its Material. Berends localfauna, 208815 presence in the Mount Scott local fauna (1). Doby Springs local fauna, locality extends its known range to the Illinoian. 1, 213713 (10); locality 4, 208933 (23); locality 5, 213775 (37). Mount Scott Superfamily Zonitoidea local fauna, locality UM-K4-53, 200606 Family Limacidae (84); locality UM-K1-60, 213823 (3); locality UM-K2-59, 213684 (6). Deroceras aenigma Leonard Nesovitrea electrinu (Gould) Geologic Range. Late Pliocene (Rex- road Formation. Saw Rock Canyon local Geologic Range. Late Pliocene (Rex- fauna) to Late '~leistocene(sangamon, road Formation, Rexroad local fauna) Cragin Quarry local fauna) (Hibbard & to Recent (Hibbard & Taylor, 1960). Taylor, 1960). Distribution. Vorth America north Ecology. Unknown. It probably had of about latitude 38O, southward in the habitat requirements similar to those Rocky Mountains to Arizona and New of living species of Deroceras. Mexico" (Hibbard & Taylor, 1960: Material. Berends local fauna, 208827 148). (24). Doby Springs local fauna, locality Ecology. Nesovitrea electvina is pri- 1, 213710 (8); locality 4, 208932 (42), marily a wood snail, living among dead 213779 (51). Mount Scott local fauna, leaves, under bark and decaying logs. locality UM-K4-53, 200604 (200); local- It is often found in association with ity UM-K1-60, 213822 (17); locality Zonitoides arboreus, Strobilops laby- UM-K2-59, 213690 (250). rinthica and Retinella indentata (Leon- Remarks. Several lots contain speci- ard & Goble, 1952). Leonard (1950: mens that are thinner and smaller than 37) reports that N. electrim is a com- the typical D. aenigma; they may per- mon snail in the forested areas of east- haps be D. laeve. At present, however, ern Kansas, "where the annual rainfall it is not possible to separate these 2 is generally more than 35 in.but it de- species when they occur together, for clines in frequency of occurrence to- the smaller, thinner shells may repre- wards the more arid' Plains Border sent either D. laeve or immature D. Province and is unknown in the Plains aenigma (see discussionin Taylor, 1954). Province even where timber is locally available." Although this species evi- Family Zonitidae dently prefers a woodland habitat, it has been collected from more open Euconulus fulvus (Miiller) situations " in marsh grass; in ferns Geologic Range. Middle Pleistocene in meadows; . . . in herbacious tall weeds, (Kansan, Cudahy fauna) to Recent. (Frye in grass and under burdocks in open & Leonard, 1952: 158-159). fields ..." (Archer, 1939: 19). Distribution. "Almost throughout the Material. Doby Springs local fauna, Holarctic realm, but wanting in the Gulf locality 1, 213734 (15); locality 4, and South Atlantic States from Texas to 208934 (5); locality 5, 213777 (31). Mount North Carolina" (Pilsbry, 1946: 236). Scott local fauna, locality UM-K4-53, Ecology. "E. fulvus lives among 200639 (107); locality UM-K2-59, 213689 leaves in well-shaded places, and may (22). 250 B. B. MILLER

Hawaiia minuscula (Binney) on or under the bark of logs, under boards, bricks or stones in the grass, Geologic Range. Late Pliocene (Rex- or in any like situation offering pro- road Formation, Saw Rock Canyon local tection from the sun and a reasonable fauna) to ~ecent(Hibbard &- Taylor, degree of moisture. In the southern 1960). Alleghanies I have found it up to about Distribution. nNorth America from 5800 feet, and in the Colorado Rockiesit Alaska and Maine to Costa Rica . . . . It has been taken at 10,000 feet" (Pilsbry, is generally spread over every eastern 1946: 482). and midwestern state, and in Florida Material. Doby Springs local fauna, as far south as Miami and Cape Sable, locality 1, 213726 (25); locality 2, though not seen from the Keys. It be- 185786 (2); locality 4, 208935 (146). comes rather local in Rocky Mountain Mount Scott local fauna, locality UM- States, and has not been seen from K4-53, 200655 (150); locality UM-K1-60, Washington, Oregon, Idaho, Nevada and 213796 (17); locality UM-K2-59, 213695 Utah, and only towards the south in (131), 213699 (63); locality UM-K3-60, California, where some records are 214177 (7). Locality UM-K2-60, 214192 probably owing to importation with (1). plants7' (Pilsbry, 1946: 421-423). Zonitoides nitidus (MUller) Ecology. "Hawaiia minuscula found is 111, under logs, sticks, stones and in clumps Plate Figs. 1,3 of grass in both floodplains and upland Geologic Range. Late Pleistocene situations" (Leonard, 1959: 120). I have (Illinoian, Mount Scott local fauna) to collected this species from sticks buried Recent. in moist leaf litter that filled a small Distribution. Holarctic. The ac- gulley, in a stand of cottonwood trees, companying distribution map (Fig. 8) is at the west end of Lake Larrabee, in based on published records, andmaterial Meade County State Park. examined in the University of Michigan Material. Berends local fauna, 208822 Museum of Zoology (UMMZ), and the (543). Doby Springs local fauna, locality U. S. National Museum (USNM) col- 1, 213732 (25); locality 4, 208936 (1300); lections. locality 5, 213778 (108). Mount Scott The following is a list of peripheral local fauna, locality UM-K4-53, 200619 occurrences along the southern margin (2300); locality UM-K1-60, 213824 (54); of its distribution: locality UM-K2-59, 213688 (650); local- Newfoundland: Bonne Bay; Brooks ity UM-K3-60, 213172 (30). Locality & Brooks, 1940: 71. UM-K2-60, 214184 (3). Recent, Harper Quebec: Anticosti Island; Pilsbry, County, Oklahoma. 1948: 477. Nova Scotia: Victoria County; Mac- Zonitoides avboveus (Say) Millan, 1953: 399. Geologic Range. Early Pliocene (La- Virginia: Orange County, Orange, verne Formation, Laverne local fauna) UMMZ 115097. to Recent (Hibbard & Taylor, 1960). Kentucky: Warren County, USNM Distribution. North America from 568444. northern Canada south to Banana River, Tennessee: Knox County, Knoxville, Costa Rica (Pilsbry, 1946). Its north- USNM 38881. ern limit is placed by Dall (1905) at Alabama: Jackson County, USNM 61' north latitude. 116596. Ecology. the eastern states and Iowa: Linn County, Cedar Rapids, Mississippi valley this snail is every- USNM 507625; Dubuque County, Dubuque, where abundant, to be found wherever USNM 507744; Bremer County, Waverly, there are trees or shelter of any kind; USNM 507820; Cerro Gordo County, FIVE ILLINOIAN FAUNAS

FIG. 8. Distribution of Zonitoides ~zitidzis. 252 B. B. MILLER

Clear Lake, USNM 507724; Dickinson Superfamily Polygyroidea County, West Lake Okoboji, USNM Family Polygyridae 525186. Nebraska: Cuming County, Beemer, Stenotvema leai (Binney) USNM 506141. Geologic Range. Middle Pleistocene Utah: Morgan County, 3 miles east (Kansan, Cudahy fauna) to Recent. of Morgan, UMMZ 167030; Cache County, Distribution. New York southto Mary- Logan, Blacksmith Fork of Bear River, land, west to Kansas; Nebraska and UMMZ 167029. South Dakota (Pilsbry, 1940: 678). Washington: King County, Seattle; Ecology. Stenotvema leai prefers Pilsbry, 1948: 477. moist wooded areas. Baker (1939: 60) Oregon: Clatsop County, Astoria; states that it is found "... in lowlands, Pilsbry, 1948: 477. usually on the floodplains of rivers and California: Alameda County, Berke- creeks or the margins of lakes, inwood- ley; Los Angeles County, San Diego lands of oak, hickory, elm and willow. " County, San Diego; all in Pilsbry, Taylor (1960) has reported it from the 1948: 477. floodplain of the Niobrara River in The few and scattered west coast Cherry County, Nebraska, were it was records probably represent relatively collected from beneath scattered, moist, recent introductions (Pilsbry, 1948). debris. I have collected it from a Ecology. ((InIllinois Zonitoides nitidus similar habitat on the floodplain of the lives in decaying logs and under forest Republican river, south of Franklin, debris on floodplains of streams. It Nebraska. shows a preference for muddy places. Material. Berends local fauna, 208804 Trees usually characteristic of its (73). Doby Springs local fauna, locality habitat are elm, maple or hickory" 4, 213837 (1); locality 5, 213782 (1). (Baker, 1939: 79). Goodrich (1932: 32) Mount Scott local fauna, locality UM-K1- states that 2. nitidus &lives in fairly 60, 213704 (2); locality UM-K2-59, large colonies close to the water's edge 213694 (32). of creeks, lakes and marshes." Remarks. One lot of S. leai (s.s.) Material. Mount Scott local fauna, (UMMZ 206549) collectedfrom theflood- locality UM-K4-53, 213789 (89); locality plain of the Republican River, south of UM-K2-59, 213786 (13). Franklin, Nebraska, represents the Remarks. Z. nitidus may be distin- closest confirmed record of this species guished from 2. avboveus by the more to the Meade County area. rounded base of its aperture and body whorl; its greater diameter and height ACKNOWLEDGMENTS of shell for any given number of whorls; and the relatively greater width of its Financial support for travel expenses umbilicus in relation to the total shell was made available from the Graduate diameter. Student Research Fund of the Horace In North America, 2. nitidus is rare H. Rackham School of Graduate Studies. as a fossil. To my knowledge, one lot A research grant from the National (U. S. Geological Survey Cenozoic local- Science Foundation (G-5635) to Claude W. ity 22414) from Weber County, Utah, Hibbard helped defray the field and represents the only other fossil occur- laboratory expenses, for the study of the rence of this species from west of the Mount Scott local fauna. Mississippi River. The scarcity of this I am most grateful to Arthur D. species as a fossil makes it highly pro- Sanders, Jack T. Sanders and J. D. bable that its occurrence at 2 localities Golliher for allowing me to map and in the Mount Scott local fauna were collect fossils on portions of their land, essentially contemporaneous. and to members of the University of )IAN FAUNAS 253

Michigan Museum of Paleontology field BEETLE, D. E., 1962, Additions to parties who collected most of the Teton County, Wyoming, Mollusca. materials studied in this report. Thanks Nautilus, 76(2): 74. are due to Henry van der Schalie of the , 1965, Molluscan Fauna of University of Michigan Museum of Zo- Some Small Ponds in Grand Teton ology and J. P. E. Morrison of the United National Park. Nautilus, 78(4): 125- States National Museum, for granting me 130. access to the collections in their care. BINNEY, W. G., 1865, Land and fresh-

d The sphaeriids reported herein were water shells of North America. Part checked and/or identified by H. B. Her- 11. Pulmonata, Limnophila and Thalas- rington, of Westbrook Heights, Ontario. sophila. Smithsonian rnisc. Coll., 143: The identifications of the unionid clams 1-161. ' were checked by Henry van der Schalie. BRANSON, B. A., 1959, Notes on I wish to acknowledge my indebtedness Oklahoma snails (Gastropoda) with new to Claude W. Hibbard of the University records. Southwestern Nat., 3: 224- of Michigan Museum of Paleontology, 226. for the help, advice and encouragement , 1961, Recent Gastropoda of he has so freely given throughout the Oklahoma. Part 11. Distribution, course of this study; and to Dwight W. ecology and taxonomy of fresh-water Taylor, University of Michigan Museum species, with description of Helisoma of Zoology, who critically read the travertina sp. nov. OMa. State Univ. manuscript and made many helpful com- Publ., Arts and Sci. Stud., Biol. Ser., ments and suggestions. 6: 1-72. BRANSON, B. A. & PETERS, L., 1964, REFERENCES Limax maximus and Fevvissia meek- iana in Oklahoma. Nautilus, 77 (3): 107- ARCHER, A. F., 1939, The ecology of 108. the Mollusca of the Edwin S. George BROOKS, S. T. & BROOKS, B. W., Reserve, Livingston County, Michigan. 1940, Geographical distribution of Occ. Pap., Univ. Mich. Mus. Zool., the Recent mollusca of Newfoundland. 398: 1-24, Ann. Carnegie Mus., 28: 53-75. BAKER, F. C., 1906, A catalogue of BROOKS, S. T. & KUTCHKA, G. M., the Mollusca of Illinois. Bull. Ill. 1938, Occurrence of the family Pupill- State Lab. nat. Hist., 7(6): 53-136. idae in West Virginia. Art. 111. Ann. , 19, The '~~mnaeidaeof Carnegie Mus., 27: 63-85. North and Middle America, recent BOYCOTT, A. E., 1921, Notes on the and fossil. Chicago Acad. Sci., Spec. distribution of British land andfresh- Pub., 3: 1-539, water Mollusca from the point of view , 1928a, The Fresh Water of habitat and climate. Proc. malac. Mollusca of Wisconsin, Part I. Gastro- Soc. Lond., 14: 163-167. , 1934, The habitats of land < poda. Bull. Wisc. Geol. nat. Hist. Survey,. . 70 (1): 1-507. Mollusca in Britain. J. Ecology, 22(1): , 1928b, Id., Part 11, Pelecy- 1-38. poda. Ibid., (2): 1-495. , 1936, The habitats of fresh- , 1939, Fieldbook of Illinois water Mollusca in Britain. J, animal land snails. Illinois nat. Hist. Survey Ecology, 5(1): 116-186. Manual, 2: 1-166. BYRNE, F. E. & McLAUGHLIN, T. G., BASCH, P. F., 1963, A review of the 1948, Geology and groundwater re- recent freshwater limpet snails of sources of Seward County, Kansas. North America (Mollusca: Pulmon- Kans. geol. Survey Bull., 69: 1-140. ata). Bull. Mus. comp. Zool., 129: CRANDALL, A. O., 1901, The American 399-461. Physae. Nautilus, 15: 25-30, 42-45, 2 54 B. B. MILLER

54-58, 69-71. tral High Plains. Kans, geol. Survey DALL, W. H., 1905, Land and fresh- Bull., 157(4): 1-60. water mollusks of Alaska and adjoining HENDERSON, J., 1924, Mollusca of regions. Srnithsonian Instit., Harri- Colorado, Utah, Montana, Idaho and man Alaska Expedition, 1899, Alaska, Wyoming. Univ. Colorado Studies, 13: 1-171. 13(2): 65-223. DeKAY, J., 1843, Zoology of New York HERRINGTON, H. B., 1962, A revision or the New York fauna. Part V. Mol- of the Sphaeriidae of North America lusca. Albany, New York, 271 p. (Mollusca: Pelecypoda). Misc. Publ. ,, ETHERIDGE, R., 1960, The slender Univ. Mich. Mus. Zool., 118: 1-74. glass lizard, Ophisaums attenuatus, HERRINGTON, H. B. & TAYLOR, D. W., from the Pleistocene (Illinoian glacial) 1958, Pliocene and Pleistocene of Oklahoma. Copeia, 1960(1): 46-47. Sphaeriidae (Pelecypoda) from the I , 1961, Late Cenozoic glass central United States. Occ. Pap. Univ. lizards (Ophisaums) from the south- Mich. Mus. Zool., 596: 1-28. ern Great Plains. Herpetologia, 17(3): HIBBARD, C. W., 1948, Late Cenozoic 179-186. climatic conditions in the High Plains FRANZEN, D. S., 1963, Variations in of western Kansas. In: Colbert, E.H. the Anatomy of the Succineid Gastro- (ed.), Pleistocene of the Great Plains. pod Oxy loma retusa. Nautilus, 76(3): Bull. Geol. Soc. Amer., 59: 592-597. 82-95. , 1949a, Pleistocene strati- , 1964, Anatomy of the Succin- graphy and Paleontology of Meade eid Gastropoda Oxyloma haydeni. County, Kansas. Contrib. Univ. Mich. Nautilus, 77(3): 73-81. Mus. Paleontol., 7(4): 63-90. FRANZEN, D. S. & LEONARD, A. B., , 1949b, Techniques of col- 1947, Fossil and living Pupillidae lecting microvertebrate fossils. Con- (Gastropoda-Pulmonata) in Kansas. trib. Univ. Mich. Mus. Paleontol., 8(2): Univ. Kans. Sci. Bull., 31: 311-411. 7-19. FRYE, J. C., 1942, Geology and ground- , 1950, Mammals from the water resources of Meade County, Rexroad Formation from Fox Canyon, Kansas. Kans. geol. Survey Bull., Meade County, Kansas. Contrib. Univ. 45: 1-152. Mich. Mus. Paleontol., 8(6): 113-192. FRYE, J. C. & LEONARD, A. B., 1952, , 1951, Vertebrate fossils from Pleistocene geology of Kansas. Kans. the Pleistocene Stump Arroyo mem- geol. Survey Bull., 99: 1-230. ber, Meade County, Kansas. Contrib. FRYE, J. C., LEONARD, A. B. & HIB- Univ. Mich. Mus. Paleontol., 9(7): 227- BARD, C. W., 1943, Westward ex- 245. tension of the Kansas UEquus beds". , 1954, Second contribution to J. Geol., 51: 33-47. the Rexroad fauna. Trans. Kans. Acad. GETZ, L. L., 1960, Middle Pleisto- Sci., 57: 221-237. cene Carnivores from southwestern , 1955, The Jinglebob inter- Kansas. J. Mammology, 41(3): 361- glacial (Sangamon?) fauna from Kan- 365. sas and its climatic significance. GOODRICH, C., 1932, Mollusca of Contrib. Univ. Mich. Mus. Paleontol., Michigan. Michigan Handbook Series 12(10): 179-228. No. 5. University Mus., Univ. Mich., , 1956a, Vertebrate fossils 120 p. from the Meade Formation of south- GOULD, A. A., 1840, Miscellanies. western Kansas. Pap. Mich. Acad. Amer. J. Sci., 38: 196. Sci., Arts and Letters, 41: 145-203. GUTENTAG, E. D. & BENSON, R. H., 1956b, Micvotus pennsylvani- 1962, Neogene (Plio-Pleistocene) mid)from the Hay Spring local fresh-water Ostracodes from the cen- fauna of Nebraska. J. Paleontol., 30: FIVE ILLIN()IAN FAUNAS 255

1263-1266. liminary survey of the land andfresh- , 1958, New Stratigraphic water Gastropoda, Cape Breton, Nova names for early Pleistocene deposits Scotia. Proc. Nova Scotian Inst. Sci. in southwestern Kansas. Amer. J. 1953-1954, 24(4): 389-408. Sci., 256: 54-59. MCLAUGHLIN, T. G., 1946, Geology , 1963, A late Illinoian fauna and ground-water resources of Grant, from Kansas and its climatic signifi- Haskell, and Stevens counties, Kansas. cance, Pap. Mich. Acad. Sci., Arts Kans. geol. Survey Bull., 61: 1-221. and Letters, 48: 187-221. , 1947, Accelerated channel HIBBARD, C. W. & TAYLOR, D. W., erosion in the Cimarron Valley in 1960, Two late Pleistocene faunas southwestern Kansas. J. Geol., 55: from southwestern Kansas. Contrib. 76-93. Univ. Mich. Mus. Paleontol., 16(1): MENGEL, R. M., 1952, White Pelican 1-223. from the Pleistocene of Oklahoma. ISELY, F. B., 1925, The fresh-water Auk, 69: 81-82. mussel fauna of eastern Oklahoma. MEYERS, A. J., 1959, Geology of Har- Proc. Oklahoma Acad. Sci., 4: 43-118. per County, Oklahoma. Oklahoma JONES, D. T., 1932, Mollusks in the geol. Survey Bull., 80: 1-108. vicinity of YanMon, South Dakota. MILES, C. D., 1958, The family Suc- Nautilus, 45(4): 115-118. cineidae (Gastropoda: Pulmonata). KAPP, R. O., 1965, Illinoian and San- Univ. Kans. Sci. Bull., 38(II) 24: 1499- gamon Vegetation in Southwestern 1543. Kansas and Adjacent Oklahoma. Con- MILLER, B. B., 1961, A late Pleisto- trib. Univ. Mich. Mus. Paleontol., 19 cene molluscan faunule from Meade (14): 167-255. County, Kansas. Pap. Mich. Acad. KENK, R., 1949, The animal life of Sci., Arts and Letters, 46: 103-125. temporary and permanent ponds in , 1963a, A fossil occurrence southern Michigan. Misc. Publ. Univ. of P~obythinellalacustris (Baker) Mich. Mus. Zool., 71: 1-61. (Gastropoda: Arnnicolidae) from west KLINGENER, D., 1963, Dental evo- of the continental divide. J. Paleontol., lution of Zapus. J. Mammalogy, 44(2): 37(3): 703-704. 248-260. , 1964, Additional mollusks La ROQUE, A., 1956, Variation of car- from the late Pliocene Bender local inae in Valvata tricarinata. Nautilus, fauna, Meade County, Kansas. J. 70(1): 13-14. Paleontol. 38(1): 113-117. LEONARD, A. B., 1950, A Yarmouth MOQUIN-TANDON, A. C. H. B., 1855, molluscan fauna in the midcontinent Histoire naturelle des mollusques region of the United States. Univ. terrestres et fluviatiles de France. Kans. Paleontol. Contrib., Mollusca, J.-B. ~aili&re,Paris, Vol. 11, 646 p. Art., 3: 1-48. MURRAY, H. D. & LEONARD, A. B., - , 1959, Handbook of Gastro- 1962, Handbook of Unionid mussels pods in Kansas. Univ. Kans. Mus. in Kansas. Univ. of Kans. mus. Nat. nat. Hist., Misc. Publ., 20: 1-224. Hist., Misc. Publ., 28: 1-184. LEONARD, A. B. & GOBLE, C. R., ORTMANN, A. E., 1919, A monograph .. 1952, Mollusca of the University of of the Najades of Pennsylvania. Pt. Kansas Natural History reservation. III. Systematic account of the genera Univ. Kans. Sci. Bull., 34 (II)16: 1013- and species. Memoirs Carnegie Mus., 1055. 8(1): 1-343. LEONARD, A. E., 1943, The Mollusca OVER, W. H., 1915, Mollusca of South of Meade and Clark counties. Trans. Dakota. Nautilus, 29(8): 90-95. Kans. Acad. Sci., 46: 226-240. , 1928, Additional records MACMILLAN, G. K., 1953, A pre- of South Dakota Mollusca. Nautilus, 256 B. B. MILLER

41(3): 92-93. SMITH, G. R., 1963, A late Illinoian PILSBRY, H. A., 1946, Land Mollusca fish fauna from southwestern Kansas. of North America (north of Mexico). Copeia, 1963(2): 278-285. Acad. Nat. Sci. Phila., Monogr. 3, SMITH, H. T. U., 1940, Geologic studies Vol. II(1):. , 1-520. in southwestern Kansas. Kans. geol. , 1948, Id., Ibid., Vol. II(2): Survey Bull., 34: 1-212. 521-1113. STARRETT, A,, 1956, Pleistocene mam- REIGLE, N. J., 1963, Northern records mals of the Berends fauna of Okla- of Gastvocopta pvoceva. Nautilus, homa. J. Paleontol., 30(5): 1187-1192. + 77(1): 16-18. STEPHENS, J. J., 1960, Stratigraphy and REYNOLDS, M. B., 1959, Pleistocene Paleontology of a late Pleistocene Molluscan faunas of theHumboldtDe - basin, Harper County, Oklahoma. Bull. posit, Ross County, Ohio. Ohio J. Geol. Soc. Amer., 71: 1675-1702. Sci., 59(3): 152-166. STERKI, V., 1894, The land and fresh- RINmR, G. C. & HIBBARD, C. W., water Mollusca in the vicinity of New 1952, A new beaver, and associated Philadelphia. A contribution to the vertebrates, from the Pleistocene of natural history of Tuscarawas County, Oklahoma. J. Mammology, 33: 98-101. Ohio. New Philadelphia, Ohio, 14 p. ROBERTSON, A. D., 1915, The Mollusca STOLL, N. R., ed., 1961, International of Georgian Bay. Contrib. Canadian Code of Zoological Nomenclature Biol., Supplement to the 47th Annual adopted by the XV International Con- Report Dept. Marine and Fish., Ses- gress of Zoology. International Trust sional Pap., 39b: 95-111. for Zoological Nomenclature. London. ROSCOE, E. J., 1954, Terrestrial 176 p. Gastropoda from the BlackHills, South TAYLOR, D. W., 1954, A new Pleisto- Dakota. Proc. Utah Acad. Sci., Arts cene fauna and new species of fossil and Letters, 31: 67-72. snails from the High Plains. Occ. , 1955, Additional South Dakota Pap. Univ. Mich. Mus. Zool., 557: 1- terrestrial Gastropod records. Amer. 16. Midl. Natur., 54(2): 511-512. , 1960, Late Cenozoic mollus- SAY, T., 1824, (In: KEATING, W. H.) can faunas from the High Plains. U.S. Narrative of an expedition to the Geol. Survey Prof. Pap., 337: 1-94. source of the St. Peter's River, Lake TAYLOR, D. W. & HIBBARD, C. W., Winnepeek, Lake of the Woods, etc. 1955, A new Pleistocene fauna from Philadelphia. H. C. Carey & I. Lea. Harper County, Oklahoma. Oklahoma Vol. 11, 459 p. geol. Survey Circular, 37: 1-23. SCAMMON, R. E., 1906, The Unionidae TAYLOR, D. W. & SOHL, N.F., 1962, An of Kansas, part I. Univ. Kans. Sci. outline of Gastropod classification. Bull., 3(9): 279-373. Malacologia, l(1): 7-32. SHIMEK, B., 1915, The Mollusca of the TIHEN, J. A., 1962, A review of the Okoboji region. Bull. fr. Labs of Nat. New World fossil bufonids. Amer. - Hist., State Univ. of Iowa, 7(2): 70-88. Midl. Natur. 68(1): 1-50. SIMPSON, C. T., 1914, A descriptive VAN DER SCHALIE, H., 1953, Mollusks catalogue of the Naiades or pearly From an interglacial deposit (San- fresh-water mussels. Bryant Walker, gamon? age) in Meade County, Kansas. A Detroit, 1540 p. Nautilus, 66: 80-90. SMITH, C. L., 1954, Pleistocene fishes VANATTA, E. G., 1914, Land and fresh of the Ber ends fauna of Beaver County, water shells from eastern Canada. Oklahoma. Copeia, 1954 (4): 282-289. Proc. Acad. Nat. Sci. Phila., 66: 209- , 1958, Additional fishes from 226. Kansas and Oklahoma. Copeia, 1958 VISHER, S. S., 1954, Climatic Atlas (3): 176-180. of the United States. Harvard Univ. FIVE ILLINOIAN FAUNAS 257

Press, Cambridge, 403 p. ZETEK, J., 1918, The Mollusca of WINSLOW, M. L., 1921, Mollusca of Piatt, Champaign and Vermilion North Dakota. Occ. Pap. Univ. Mich. counties, of Illinois. Trans. Ill. State Mus. Zool., 98: 1-18. Acad. Sci., 11: 151-182.

RESUMES CIKCO MALACOFAUNM DE EDAD ILLINOIANA DE LAS GRAXDES LLA~URASSURE~~AS

B. G. Miller Se enumeran 10s moluscos de cinco conjuntos del Cenozoico superior (Illinoian) en el sudoeste de Kansas- noroeste de Oklahoma: la fauna local de Berends, de Beaver County, Oklahoma: la fauna local de Doby Springs, Harper Co., Oklahoma; y las faunas locales de Adams, Butler Springs y Mount Scott, de Meade Co., Kansas. Un sumario informativo sobre la distribuci6n geolhgica, geograica y ecol6gica se da para todos 10s moluscos de Mount Scott y Doby Springs, y para las especies previamente seflaladas de 10s Berends, Adams y Butler Springs. Cambios taxon6micos incluyen la asignaci6n de Promenetus exacwnci y Promenetus knnsnsensis a un Gnico complejo especifico. No eldste manera satisfactoria para separar esas dos 'especies" enpoblacionesrecientes y f6siles que contienen individuos con ambos tipos de ornamentaci6n superficial, costulados y no costuladcs. El tipo costulado es designado P. exocuous form kansasemis y el liso P. exacuous form exncuous . En la interpretaci6n de estos conjuntos se asume que 10s factores climaticos son 10s agentes primarios que controlan la distribucibn de 10s moluscos no-marinos; que ejemplares f6siles similares a 10s vivientes representan la misma especie y estuvieron sujetas a 10s mismos factores limitadores; y que 10s ireas de las Grandes Llanuras del presente, conteniendo asociaciones de moluscos mis similares a aquellas de 10s conjuntos f6siles, simulan probablemente condiciones climaticas en que esas faunas vivieron. Reconstrucci6n de habitat local y condiciones climaticas de las mismas, has sido basadas sobre una sintesis de la informaci6n obtenida de 10s moluscos f6siles, polen y vertebrados, yen ninghcaso estas interpretaciones basadas en moluscos han contradicho conclusiones alcanzadas independientemente por datos de otros f6siles. Los cinco conjuntos Illinoianos muestran discrepancias de composici6n que se relacionan probablemente con 10s cambios climaticos durante aquella ipoca. El ncmero relativamente grande de especies con distribucibn predominante en el norte, junto con la poca representaci6n de elementos faunisticos exclusivamente surefios, indican que la fauna local de Doby Springs representa el conjunto de clima dsfrio estudiado y probablemente vivi6 durante el mama de la glaciaci6n de Illinois. La fauna de Mount Scott contiene un nGmero mayor de especies de distribuci6n en el sur, sugiriendo que es la fauna de clima mis cilido en esa ipoca. No hay evidencia de que esas faunas locales vivieran durante m6s de urn unidad de tiempo (substage). Asumiendo que la transici6n desde condiciones glaciales mamas, representada por la fauna local de Doby Springs, hacia condiciones interglaciales mis calidas, representada por la fauna de Mount Scott, fue unidireccional, entonces las condiciones climaticas implicadas por 10s conjuntos de Adams y Butler Springs, deben haber ocurrido en algun momento entre el tiempo representado por las fauias de Doby Springs y Mount Scott. Las primeras y atims apariciones de especies de moluscos, y las implicaciones climaticas de la fauna local de Berends, sugiere que este conjunto es probablemente el mis antiguo de 10s cinco niveles del Illinoian. El conjunto de Berends contiene 45 especies de moluscos, 15 de 10s cuales son nuevos para esta fauna local. Spluerium securis representa el primer f6sil de esta especie para las Llanuras del Sur. Esta fauna vivi6 en, y alrededor, de un pequefio y poco profundo lago, donde la vegetaci6n marginal consistia probablemente de co- 258 B. B. MILLER

niferas, gradneas y compuestas, con s6lo algunas manchas de irboles de hoja caduca. El clima en ese tiempo era probablemente similar a1 del presente en el norte de Nebraska y noroeste de Iowa de las Grandes Llanuras. La fauna local de Doby Springs contiene 59 especies y una de el1as;Vevtigo elatior, se registra por primera vez como f6sil de las Llanuras. Los moluscos indican la existencia de un lago frio bordeado por pantanos con totora, juncos, coniferas y otros irboles disperses, que engranaban conunazona de mejor desague conteniendo arbustos y irboles a distancia del lago. El clima local combinaba veranos mis cortos y mis frios, con inviernos que eran generalmente mis frios y largos que 10s ocurren en las condiciones presentes de la regi6n. El conjunto de Adams contiene 22 especies, con Lasmigonn complalucta que se registra pro primera vez en el irea. Las areniscas cruzadas de grano grueso y mediano, donde estos f6siles se obtuvieron, junto con la presencia de especies que requieren un habitat acuatico continuo, indican un cuerpo de agua corriente. Una sola especie de molusco terrestre de habitat boscoso, seAala que s61o existia una cubierta delgada de &boles y arbustos. El clima probablemente combinaba inviernos quiz6 no mis severos que 10s actuales en noreste Nebraska, con veranos mis hhmedos y frios, similar a aquellos de las Dakotas y oeste de Montana: Hay 54 especies de moluscos en la fauna local de Butler Springs, y entre ellas Ligumia cf recta es nueva para las Grandes Llanuras del Sur. Esta fauna vivi6 en, o cerca, de un rio de tamaflo mediano, de corriente moderada y probablemente con varios pi& de profundidad. Habia grupos esparcidos de irboles a lo largo del rio, per0 la vegetaci6n predominante consistia de gramineas y arbustos. El clima contaba con veranos mis frios, similares a 10s de hoy en el oeste de N. Dakota. Los inviernos eran probablemente no mgs severos que aquellos de 10s Nebraska norcentral. La malacofauna de Mount Scott contiene 63 especies y es el conjunto dsgrande conocido para las Llanuras del Sur. Contiene el hico f6sil de Zonitoides nitidus en las Grandes Llanuras. Evidencia paleontolhgica y estratigraicaindica que hay repre- sentadas cinco asociones mayores. El clima de aquel tiempo probablemente combinaba veranos similares a 10s que ocurren en el Lago Okoboji del noroeste de Iowa, con inviernos no mis severos que en el noreste de Kansas y sureste de Nebraska.

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