HARVARD UNIVERSITY Library of the Museum of Comparative Zoology lA 7^€MM^m Off y^isc. rtc^^Hi.'UK /^■: / I 4l/ If i I UNIONID MUSSELS ^ f4k Harold D. Murray A.Byron Leonard / 1 --."V/ / The University of Kansas Department of Zoology and State Biological Survey \ Cover: Blue-Point Mussel (Crenodonfa peruviana costata) on the bottom of a stream; a Black Crappie swims above, and a Muskrat sits on a log on the bank. HANDBOOK OF UNIONID MUSSELS IN KANSAS BY Harold D. Murray AND A. Byron Leonard Contribution from the Department of Zoology and State Biological Survey Museum of Natural History University of Kansas Lawrence, Kansas University of Kansas Museum OF Natural History editor: E. RAYMOND HALL Miscellaneous Publication No. 28, pp. 1-184, 45 plates, 42 figures Published Maij 10, 1962 University of Kansas Lawrence, Kansas MUS. COMP. ZOOL LIBRARV JAN -7 1963 NARlfARD UNIVERSITY PRINTED BY JEAN M. NEIBARGER. STATE PRINTER TOPEKA. KANSAS 1962 29-692 CONTENTS PAGE Introduction 9 Purpose 9 Economic Importance 9 Acknowledgments 10 Review of Literature 10 Classification of Mussels in Kansas 10 Morphology of Unionidae 14 Structure of Shell 14 Soft Anatomy 19 Muscles of Shell 19 Mantle 19 Gills 24 Foot 24 Digestive System 24 Kidney 25 Circulatory System 25 Nervous System 25 Reproductive System 25 Development and Growth of Fresh-Water Mussels 26 Development 26 Growth 27 Ecology of Fresh-Water Mussels 27 Condition of Water 28 Bottom Conditions 29 Depth of Water 29 Vegetation 30 Light 30 Current 30 Temperature 30 Predation and Parasitism 30 Collection and Preservation of Mussels 31 Key for Identification of Species and Subspecies of Mussels in Kansas 33 Introduction 33 Accounts of Species 37 Introduction 37 Maps and Illustrations 38 Phylum Mollusca 39 Class Pelecypoda 39 Order Eulamellibranchia 39 Family Unionidae (D'Orbigny) Ortmann 39 Subfamily Unioninae 39 Genus Fusconaia Simpson 39 Fusconaia flava ( Rafinesque ) 39 Genus Megolonaias Utterbach 42 Megalonaias gigantea ( Barnes ) 42 (3) / University of Kansas Publs., Mus. Nat. Hist. PAGE Genus Crenodonta Schliiter 45 Crenodonta peruviana peruviana (Lamarck) 47 Crenodonta peruviana costata (Rafinesque) 49 Genus Quadrula Rafinesque 50 Quadrula quadrula ( Rafinesque) 50 Quadrula pustulosa ( Lea ) 55 Quadrula nodulata Rafinesque 58 Quadrula metanevra Rafinesque 61 Quadrula cylindrica ( Say ) 64 Genus Tritogonia Agassiz 65 Tritogonia verrucosa ( Barnes ) 67 Genus Plethobasus Simpson 68 Plethobasus ctjphyus (Rafinesque) 68 Genus Pleurobema Rafinesque 71 Pleurobema cordatum coccineum (Conrad) 71 Pleurobema cordatum catillus ( Conrad ) 74 Pleurobema cordatum pyramidatum ( Lea ) 76 Genus Elliptio Rafinesque 78 Elliptio dilatatus ( Rafinesque ) 80 Genus Uniamerus Conrad 81 Uniomerus tetralasmus (Say) 83 Subfamily Anodontinae 84 Genus Lasmigona Rafinesque 84 Lasmigona costata ( Rafinesque ) 86 Lasmigona complanata ( Barnes) 87 Genus Anodonta Lamarck 90 A7wdonta grandis Say 92 Anodonta imbecilis Say 95 Genus Arcidens Simpson 96 Arciderxs confragosus (Say) 97 Genus Strophitus Rafinesque 99 Strophitus rugosus (Swainson) 99 Subfamily Lampsilinae 102 Genus Obliquaria Rafinesque 102 Obliquaria reflexa Rafinesque 103 Genus Cyprogenia Agassiz 105 Cyprogenia aberti (Conrad) 105 Genus Actinonaias Fischer and Crosse 108 Actinonaias carinata carinata (Barnes) 108 Genus Truncilla Rafinesque Ill Truncilla truncata Rafinesque Ill Truncilla donaciformis ( Lea) 114 Genus Plagiola Rafinesque 116 Plagiola lineolata (Rafinesque) 117 Genus Leptodea Rafinesque . • 119 Leptodea fragilis ( Rafinesque) 120 Leptodea laevissima ( Lea ) 123 Unionid Mussels in Kansas 5 PAGE Genus Proptera Rafinesque 126 Proptera data ( Say ) 126 Proptera purpurata (Lamarck) 129 Proptera capax ( Green ) 1"^2 Genus CaruncuJina (in Baker) Simpson 134 Carunrulina parva ( Barnes) 135 Genus Ligumia Swainson 137 Ligumia recta latissima (Rafinesque) 137 Ligumia stibrostrata (Say) 140 Genus Lnmpsilis Rafinesque 143 Lampsilis aiiodontoides anodontoides (Lea) 143 Lampsilis anodontoides jaUaciosa (Smith) Simpson 146 Lampsilis radiata siliquoidea (Barnes) 149 Lampsilis ovata ventricosa ( Barnes) 152 Genus Dysnomia Agassiz 155 Dysnomia triquetra (Rafinesque) 155 Species in Kansas That May Have Been Extirpated Since 1890 158 Allocation of Species Previously Reported From Kansas 165 Physiography of Kansas Drainage Systems 166 Systems Ancestral to Present Drainage 166 Present Drainage Systems 167 Origin of Kansas Unionid Fauna 168 Generalizations and Conclusions 170 References 171 Glossary 174 Index 180 ILLUSTRATIONS Plates Plate 1. Classes of Mollusca 13 Plate 2. Morphological features of the unionid shell 17 Plate 3. Morphological features of the soft anatomy of a unionid mussel, 21 Plate 4. Morphological features of the soft anatomy of a unionid mussel, 23 Plate 5. Fusconaia flava ( Rafinesque ) 40 Plate 6. Megalonaias gigantea ( Barnes) 43 Plate 7. Crenodonta peruviana costata (Rafinesque) and Crenodonta peru- viana peruviana ( Lamarck ) 46 Plate 8. Quadnda quadrula Rafinesque 51 Plate 9. Quadrula ptisttdosa ( Lea ) 54 Plate 10. Quadrula nodulata Rafinesque 57 Plate 11. Quadnda metanevra Rafinesque 60 Plate 12. Quadrula cylindrica (Say) 63 Plate 13. Tritogonia verrucosa ( Barnes) 66 Plate 14. Plethobasus cyphyus ( Rafinesque ) 69 Plate 15. Pleurobema cordatum coccineum ( Conrad ) 72 Plate 16. Pleurobema cordatum catillus (Conrad) 75 Plate 17. Pleurobema cordatum pyramidatum (Lea) 77 6 University of Kansas Publs., Mus. Nat. Hist. PAGE Plate 18. Elliptio dilatatus (Rafinesque) 79 Plate 19. Uniomenis tetralasmtis (Say) 82 Plate 20. Lasmigona costata ( Rafinesque ) 85 Plate 21. Lasmigona complanata (Barnes) 88 Plate 22. Anodonta grandis Say 91 Plate 23. Anodonta imbecilis Say 94 Plate 24. Arcidens confragosus (Say) 98 Plate 25. Strophitus rugosiis (Swainson) 100 Plate 26. Obliquaria reflexa Rafinesque 103 Plate 27. Cijfyrogenia aberti (Conrad) 106 Plate 28. Actinonaias carinata carinata ( Barnes ) 109 Plate 29. TrunciUa truncata Rafinesque 112 Plate 30. TrunciUa donaciformis (Lea) 115 Plate 31. Plagiola lineolata ( Rafinesque) 118 Plate 32. Leptodea fragilis ( Rafinesque) 121 Plate 33. Leptodea laevissima ( Lea ) 124 Plate 34. Proptera alata (Say) 127 Plate 35. Proptera purpurata (Lamarck) 130 Plate 36. Proptera capax ( Green ) 133 Plate 37. Carunculina parva ( Barnes) 135 Plate 38. Ligumia recta latissima (Rafinesque) 138 Plate 39. Ligumia subrostrata (Say) 141 Plate 40. Lampsilis anodontoides anodontoides (Lea) 144 Plate 41. Lampsilis anodontoides fallaciosa (Smith) Simpson 147 Plate 42. Lampsilis radiata siUquoidea (Barnes) 150 Plate 43. Lampsilis ovata ventricosa ( Barnes) 153 Plate 44. Dysnomia triquetra (Rafinesque) 156 Plate 45. Anodonta suborbiculata Say, Obovaria olivaria ( Rafinesque ) , and Ptychobranchus fasciolare (Rafinesque) 159 Text Figures 1. Distribution of Fusconaia flava in Kansas 41 2. Distribution of Megalonaias gigantea in Kansas 44 3. Distribution of Crenodonta peruviana peruviana and Crenodonta peruviana costata in Kansas 48 4. Distribution of Quadrula quadrula in Kansas 52 5. Distribution of Quadrula pustulosa in Kansas 55 6. Distribution of Quadrula nodulata in Kansas 58 7. Distribution of Quadnda metanevra in Kansas 61 8. Distribution of Quadrula cylindrica in Kansas 64 9. Distribution of Tritogonia verrucosa in Kansas 67 10. Distribution of Plethobasus cyphijus in Kansas 70 11. Distribution of Pleurobema cordatum coccineum in Kansas 73 12. Distribution of Pleurobema cordatum catillus in Kansas 76 13. Distribution of Pleurobema cordatum pyramidatum in Kansas 78 14. Distribution of Elliptio dilatatus in Kansas 80 15. Distribution of Uniomerus tetralasmus in Kansas 83 16. Distribution of Lasmigona costata in Kansas 86 17. Distribution of Lasmigona complanata in Kansas 89 Unionid Mussels in Kansas 7 PAGE 18. Distribution of Anodanta grandis in Kansas 92 19. Distribution of Anodonta UnhecUis in Kansas 95 20. Distribution of Arcidens confragosus in Kansas 97 21. Distribution of Strophitus rugosus in Kansas 101 22. Distribution of Ohliquarm reflexa in Kansas 104 23. Distribution of Cijprogcnia aberti in Kansas 107 24. Distribution of Actinonaias carinata carinata in Kansas 110 25. Distribution of Tnmcilla truncata in Kansas 113 26. Distribution of TruuciUa donaciformis in Kansas 115 27. Distribution of Plagiolu lincolata in Kansas 119 28. Distribution of Leptodea fragilis in Kansas 120 29. Distribution of Leptodea laevissima in Kansas 125 30. Distrilnition of Proptem (data in Kansas 128 31. Distribution of Proptera purpurata in Kansas 131 32. Distribution of Proptera capax in Kansas 134 33. Distribution of Caruncidina parva in Kansas 136 34. Distribution of Ligumia recta latissima in Kansas 139 35. Distril)ution of Ligumia mbrostrata in Kansas 142 36. Distribution of Lampsilis anodontoides anodontoides in Kansas 145 37. Distribution of Lampsilis anodontoides falhciosa in Kansas 148 38. Distribution of Lampsilis radiata siliquoidea in Kansas 151 39. Distribution of Lampsilis ovata ventricosa in Kansas 154 40. Distribution of Dtjsnomia triquetra in Kansas 157 41. Map of the State of Kansas showing major river drainages and major physiographic divisions 166 42. Map of Kansas showing counties {inside back cover) TABLES Table 1. List of species reported by Call and Scammon, invalidly re- corded for Kansas 164 Table 2. List of species reported by Call and Scammon, of doubtful oc- currence in the state of Kansas 164 Table 3. List of species reported by Call and Scammon, validly re- corded, but now absent from Kansas 164 Table 4. Total number of genera and species recorded by Call ( 1885- 1887), Scammon ( 1906), and in this report 165 Table 5. Occurrence of mussels in the four major river systems in Kansas, indicating mussels restricted to the Mississippi drainage and mussels occurring in that drainage south of the Missouri River 169 INTRODUCTION Purpose The objectives of the study here reported on are to list and de- scribe the unionid (fresh-water) mussels in Kansas, to show their distribution within the state and in North America, to provide the means for identifying unionids found in Kansas, to indicate the origin of the present populations in the state, and to appraise the ecological and economic importance of the unionid fauna. The report is designed to be useful to teachers and other non-specialists, as well as to malacologists having a professional interest in the unionids in Kansas. Economic Importance Because unionids are found in all bodies of permanent water — streams, ponds, and lakes — they represent an important part of the aquatic fauna of Kansas. Coker (1919) carefully considers the economics of the mussel industries that were once an important part of the economy of America. The manufacture of buttons from the shells of fresh-water mussels began about 1891. In 1912 the income from this industry amounted to approximately $6,173,486 yearly. With the advent of plastics, the shell industry decHned sharply; today, fresh-water mussel shells are used primarily for novelty work, such as charm bracelets, hat pins, belt buckles, and fancy buttons. In 1912, approximately 17,063 tons of shells were removed from the Neosho River in Kansas; this amounted to 17 per cent of the pearly products for the United States in that year. In the year 1960 in Kansas there was no commercial harvesting of unionid shells. In the past the meats of mussels have been used for hog and poultry food as well as for human consumption; however, unsanitary conditions prevailing in shell industries discouraged people from using meats of mussel for food. More recently farmers have not utilized meats of mussel for animal food. Fisherman frequently find the putrified meats of mussels to be excellent fish-bait. Mussels serve as food for raccoon, mink, otter, muskrat and several other mammals, and tlie immature mussels are frequently eaten by fishes. Baker ( 1918 ) considers the unionids as an important source of food for fishes. This subject is discussed in more detail under Predation and Parasitism (p. 30). Of less importance in large, fast-flowing rivers, but extremely significant in small ponds and stagnant pools is the consideration of the filtering eflFect on the water by the mussels. Allen (1914) (9) 10 University of Kansas Publs., Mus. Nat. Hist. estimated the siphoning rate of Lampsilis siliqiioidea of 200 grams weight at approximately 34,560 cc. (approximately 8.8 gallons) per day! ACKNOWLEDGMENTS We wish to thank Dr. Henry van der Schalie, University of Michi- gan, for his aid in identification of the species of the genus Creno- donta, and to thank Drs. Frank Cross, R. E. Beer, and E. Raymond Hall, University of Kansas, for criticism of the manuscript. Special thanks are extended to the State Biological Survey of Kansas for financial support in the preparation of illustrations. REVIEW OF LITERATURE In a series of publications, R. E. Call (1885-1887), Washburn College Laboratory of Natural History, listed unionids then known from Kansas. Localities were given by town and county, and in some instances by drainage systems. No description of the animals or illustrations accompanied Call's work; therefore, it is not possible to determine the validity of some of his records. In 1906 R. E. Scammon published "The Unionidae of Kansas," in which he described the various species occurring in Kansas and published illustrations of most of them. Unfortunately, those spe- cies presently thought to be invalid records for the state are those that were not illustrated in his study. Until now Scammon's pub- lication was the only comprehensive account of mussels in Kansas. A thorough study of mussels occurring in a tributary of the Kansas River was made by D. S. Franzen and A. B. Leonard and in 1943 resulted in their publication, "The Mollusca of the Wakarusa River Valley." That account added much to the knowledge of the unionid fauna of the northern part of Kansas. Clark and Gillette ( 1911 ), A. E. Leonard ( 1943), and A. B. Leonard and A. E. Leonard (1946) have published locality records for various areas of the state. CLASSIFICATION OF MUSSELS IN KANSAS Fresh-water mussels belong to that large group, a phylum, of animals, termed the Mollusca. Members of the phylum, inhabiting salt water, brackish water, fresh water, and land, have the following characteristics in common: bilateral symmetry; no segmentation (except Monoplachophora ) ; soft body covered by mantle that usually secretes calcareous shell; mouth having radula (except bi- valves); dorsal heart; respiration by gill-like ctenidia or "lung"; nervous system having paired ganglia, connectives, and nerves; sexes separated or united; trochophore, veliger or glochidial larva. Unionid Mussels in Kansas 11 The phylum Mollusca encompasses six classes (Plate 1): Class Monoplacophora: Nearly bilaterally symmetrical mollusks, exhibit- ing internal metamerism; dorsal, monovalve shell; posteromedian anus; well developed coelomic cavities; auricles paired, metamerically arranged; nephridia metameric; nervous system primitively orthoneurous; Neopilina, Tryblium ( figure 1 ) . Class Amphineura: Chitons. Body elongate; shell of eight plates or none; head reduced; nerve ring around mouth and two pairs of ventral nerve cords; no tentacles; having radula; marine; Chiton, Cryptochiton (fig- ure 2). Class Gastropoda: Snails, slugs. Shell usually spiral, reduced or absent; head distinct; foot large, flat for holdfast and creeping; visceral mass turned 180° counterclockwise (torsion) on head and foot; two or one ctenidia or replaced by secondary gills or "lung"; radula; marine, ter- restrial or fresh-water; Lijmnca, Phtjsa, Haliotis, Crepichila (figure 3). Class Scaphopoda: Tusk shells. Shell and mantle slenderly tubular and open at both ends; foot conical; delicate "tentacles" around mouth; no gills; having radula; marine; Dcntalium (figure 4). Class Pelecypoda: Clams, fresh- water mussels, oysters. Shell of two lateral halves, usually symmetrical having dorsal hinge ligament; mantle flattened having right and left lobes; head absent; fleshy labial palpi be- side mouth; no jaws or radula; mostly marine, some fresh-water; Pecten, Anodonto, Pholas, Sphacrium (figure 5). Class Cephalopoda: Squids and octopuses. Shell external, internal, or none; head large, well-developed eyes; mouth having homy jaws and radula surrounded by 8 to 10 tentacles; nerve ganglia grouped as a "brain" in cartilagelike covering; marine; Nautilus, Sepia, Octopus (fig- ure 6). The class Monoplacophora, recognized by some authors, was erected to include certain fossil mollusks and recently is considered to include the living genus 'Neopilina (Plate 1, figure 1), collected first in a deep trench off the west coast of Mexico in 1952 ( Lemche, 1957). The class is unusual in several ways, but especially in that 'Neopilina shows internal segmentation. It is now thought that some mollusks are segmented, and it may be that all primitive forms were. Mussels belong to the Class Pelecypoda, which includes four orders: Protobranchia, Filobranchia, Eulamellibranchia, and Septi- branchia. All of the fresh-water mussels are in the order Eula- mellibranchia; for information concerning the other orders of pelecypods, the reader is referred to Storer and Usinger ( 1957 ) or Borradaile, e^ a/. (1958). Eulamellibranchiates are characterized by having W-shaped gills, that have the lamellae of each half united by firm connections, and by having two adductor muscles of equal size. The order includes 12 University of Kansas Publs., Mus. Nat. Hist. PLATE 1 The Classes of Mollusca Fig. 1. Class Monoplacophora. Neopilina in dorsal view, showing dorsal shell and growth lines. Approximately X 1- Fig. 2. Class Amphineura. A chiton (Cryptochiton) in dorsal view, show- ing typical eight shell-plates exposed. Approximately X Vs- Fig. 3. Class Gastropoda. A snail ( Triodopsis ) seen in lateral view, showing the position in which the shell is carried. Approximately X Ws- Fig. 4. Class Scaphopoda. A tusk shell (Dentalium) in lateral view, showing the conical foot extended, and the peculiar tentacles, called captacula. Approximately X 1- Fig. 5. Class Pelecypoda. A mussel (Aiiodonta) in lateral view, showing ex- tended foot, and incurrent and excurrent siphons. Approximately XVs. Fig. 6. Class Cephalopoda. A squid ( Loligo ) in dorsal view. Approximately Xii. Figure 1 modified from Lemche, 1957; figures 2, 3, 4, 5, and 6 modified from Leonard, 1959. Unionid Mussels in Kansas 13 PLATE 1 14 University of Kansas Publs., Mus. Nat. Hist. marine and fresh-water species and numerous families, two of which occur in Kansas: Unionidae and Sphaeriidae. The Sphaeriidae are small, thin-shelled mussels, having a simple pallial line and no hinge plate. The right valve has four lateral teeth, two anterior and two posterior to the cardinal teeth; the left valve has two lateral teeth, one anterior and one posterior to the cardinal teeth. The Sphaeriidae possess other unique characters; but, because this bulletin is not concerned with that family, its characters will not be considered in further detail. The unionids are characterized as follows: shell nacreous having a thick epidermis; beaks usually sculptured, often showing remains of nuclear shell; ligament opisthodetic ( behind beak ) ; hinge having or lacking teeth, when teeth are present they are the pseudocardinals and laterals, but never having anterior laterals; pallial line simple. Three subfamilies of unionids are recognized in Kansas: Union- inae, Anodontinae, and Lampsilinae. Differentiation of the sub- families is based on characteristics listed below: Unioninae lack papillae on edge of mantle; branchial opening well devel- oped; marsupium formed by all four gills or by outer gills only; glochidia semi-elliptical or semicircular, lacking spine; hinge always complete, hav- ing rather strong teeth; generally no sexual differentiation of shell. Anodontinae have branchial opening well-defined, no flaps or papillae in front of opening on edge of mantle; marsupium formed by outer gills in entire length; glochidia semicircular to triangular having hook in middle of ventral margin of each valve; hinge rarely complete and if so reduced; secondary sexual differences of shell rarely present. Lampsilinae have edge of mantle in front of branchial opening smooth to crenulated or bearing papillae; marsupium rarely formed by whole outer gill, generally only by, or within, posterior part; glochidia semicircular or semielliptical, lacking spine, rarely cell-shaped and having two spines; hinge generally complete, having well-developed teeth, rarely reduced; sexual differences of shell usually noticeable, often strongly expressed. For a more complete discussion of the characteristics of the sub- families the reader is referred to Walker ( 1918 ) . MORPHOLOGY OF UNIONIDAE Structure of Shell All Unionidae secrete a bivalved shell by means of the mantle. The shell of unionids varies in shape, color, size, and thickness, but is consistently composed of three layers of secreted material. The outer layer, called the periostracum or epidermis, is a thin, horny, albuminoid material that protects the underlying portions of the shell from mechanical erosion and from dissolution by carbonic and other acids found in water. Damage to the periostracum, that is Unionid Mussels in Kansas 15 deposited only at the edges of the mantle, results in various degrees of erosion of the underlying layers that, incidentally, is characteristic of most unionid shells found in the streams of Kansas. The color of the periostracum of Kansas species varies from black or brown to yellow or green; in addition, the periostracum may be smooth or wrinkled. The middle or prismatic layer is relatively thin and is composed of vertical prisms of calcium carbonate in the form of calcite. The inner layer, called nacre or mother-of-pearl, is the thickest layer in all but the youngest shells and is composed of thin laminae or plates of calcite, aragonite or both. The nacreous and prismatic layers may be deposited by almost any part of the mantle. Among unionids in Kansas, the nacre varies in color from white, cream, or salmon to purple or pink. A fourth layer, the hypostracum, may occur as a secretion of the adductor, retractor, and protractor muscles; chemically, the hypostracum closely resembles the nacreous layer. The shell is divided into a right and left valve and is held together dorsally by a ligament (Plate 2, figures 2, 5). The ligament is a dark brown, proteinaceous substance; and varies in length and thickness, but in unionids the hgament functions by its elasticity and opens the two valves when the adductor muscles relax. Each shell has anterior, posterior, ventral, and dorsal margins ( Plate 2, fig- ure 1 ) . The anterior margin of the shell can be identified by the posi- tion of the pseudocardinal teeth, which in fresh-water mussels are always situated anteriorly. The anterior and posterior margins are curved, straight, or pointed whereas the dorsal and ventral margins are straight, convex, or concave. Anterior to the ligament is the inflated umbo (Plate 2, figure 1) that in young or well preserved shells usually shows the outlines of the embryonic or nepionic shell. Extending from the posterior margin of the umbo in a general posteroventral direction and developed to varying degrees is the posterior ridge ( Plate 2, figures 2, 3 ) . Some species ( Plate 2, figures 2, 3), have a lateral ridge anterior to the posterior ridge. The lateral ridge extends from the lateral area of the umbo over the lateral portion of the valve to its ventral margin. The area anterior to the umbo is termed the anterior slope; the posterior slope (Plate 2, figures 2, 3) is posterior to the umbo. These slopes differ according to species. In some species the pos- terior slope is extended dorsally and forms a posterior wing (Plate 2, figure 5), and in others the anterior slope extends dorsally and forms an anterior wing (Leptodea loevissima, Plate 33). The an- terior wing is rarely greatly developed. 16 University of Kansas Publs., Mus. Nat. Hist. MORPHOLOGICAL FEATURES OF THE UNIONID SHELL Fig. 1. Quadnila nodulata Rafinesque, catalogue no. 10502; exterior, right valve. Actual length, 1^ inches. Fig. 2. Quadnila quadnila Rafinesque, catalogue no. 10496; dorsal aspect of shell. Actual length, 2% inches. Fig. 3. Quadnila quadrula Rafinesque, catalogue no. 10496; exterior, right valve. Actual length, 2% inches. Fig. 4. Quadnila quadnila Rafinesque, catalogue no. 10496; interior, left valve. Actual length, 2% inches. Fig. 5. Proptera alata (Say), catalogue no. 10396; interior, left valve. Actual length, 4/2 inches. Unionid Mussels in Kansas 17 PLATE 2 umbo posterior morgin knob ventral margin' dorsol morgin -•— onterior morgin rest period growth line posterior ridge posterior slope loteral ridge posterior ridge posterior slope position of ligoment anterior slope lateral ridge pustule interdentum umbonal covity loterol tooth ^ onter ior slope pseudocordinol tooth posterior wing posterior retractor muscle scar ligoment posterior adductor muscle scar palliol line dorsal muscle scor anterior retractor muscle scar onterior adductor muscle scar onterior protractor muscle scar 18 University of Kansas Publs., Mus. Nat. Hist. The exterior of the valve is smooth (Ligumia suhrostrata, Plate 39, figure 3), marked by undulations {Crenodonta peruviana peru- viana, Plate 7, figures 3, 4), set with pustules (Quadrula quadrula, Plate 2, figure 3), or knobs (Quadrula nodulata, Plate 2, figure 1), or roughened by growth lines (Propfcra capax, Plate 36, figure 3). Internally the hinge line of most unionid shells is provided with hinge teeth, referred to as pseudocardinal and lateral teeth (Plate 2, figures 4, 5). These teeth are interlocking, elevated structures holding the two valves in position when open and preventing the valves from shearing apart when closed. The pseudocardinal tooth of one valve fits into a corresponding depression in the hinge of the opposing valve. Although variable in size, shape, thickness, and ornamentation, the pseudocardinals are always situated anterior to the lateral teeth and usually anterior to or directly below the umbo. The number of pseudocardinals varies from one to two in either the left or right valves. Accessory denticles may appear anterior and posterior to the main teeth, but are of little taxonomic significance. The pseudocardinal teeth vary from compressed, thin, and sharp to thick, wide, and heavily serrated. The lateral teeth are situated posterior to the umbo and posterior to the pseu- docardinals, and are high or low, straight or curved, serrated or smooth, long or short. There are usually two lateral teeth in the left valve and one in the right valve. Frequently an accessory lateral tooth develops ventral to the main lateral tooth in the right valve. On many kinds of fresh-water mussel shells, such as those of Quad- rula quadrula, there is an area between the lateral and the pseudo- cardinal teeth referred to as the hinge plate or interdentum (Plate 2, figure 4). The length of the interdentum is more or less parallel with the anteroposterior axis of the shell, and its width is approxi- mately parallel with the dorsoventi'al axis of the shell. The inter- dentum is absent in some species ( Propfera alafa, Plate 2, figure 5 ) and weak in others {Truncilla truncata, Plate 29, figures 1, 2). Both the pseudocardinals and laterals can be absent (Anodonta grandis, Plate 20) or reduced (Strophitus rugosus, Plate 15). In one genus (Lasmigona, Plates 20, 21) the lateral teeth are absent, and the pseudocardinal teeth are massive. The umbonal cavity (Plate 2, figure 4) extends into the umbo, but in some species (Ellipfio dUatatus, Plate 18) the umbonal cavity is shallow or nearly wanting. The inside of the shell has in addition to features already dis- cussed, at least seven scars or cicatrices produced by attachment of Unionid Mussels in Kansas 19 muscles. The position, size, and depth of each muscle scar is useful in the identification of the various species. The largest scars on the shell are the anterior and posterior adductor muscle scars ( Plate 2, figure 5). A posterior retractor muscle scar is situated dorsal to the posterior adductor muscle scar, and an interior retractor muscle scar is situated slightly dorsal and posterior to the anterior adductor muscle scar ( Plate 2, figure 5 ) . These may, however, fuse with or be incorporated into the adductor muscle scars and be diffi- cult to distinguish. Posterior to the anterior adductor muscle scar there is a single scar known as the anterior protiactor muscle scar ( Plate 2, figure 5 ) . An impression near the ventral edge of the shell, and extending from the anterior adductor muscle scar to the poste- rior adductor muscle scar, marks the point of attachment of the mantle to the shell and is refened to as the pallial line (Plate 2, figure 5). It is typically deeply impressed anteriorly and lightly impressed posteriorly. Within the cavity of the umbo there is typi- cally a series of scars for the attachment of the dorsal muscles from the mantle to the shell (Plate 2, figure 5). The dorsal muscle scars may be difficult to see, as in Actinonaias carinota carinata (Plate 28), or easily seen, as in Leptodea (Plates 22, 23). The function of the muscles is discussed beyond under the account of the soft anatomy. Soft Anatomy Muscles of shell (Plate 3, figure 3) The anterior and posterior adductor muscles close the valves and maintain their closure; if one or both of these muscles are cut the valves will open. The fibers of the adductor muscles extend from one valve directly across to the other; they have no direct connections to the body of the animal. The anterior and posterior retractor muscles insert on the shell and their fibers extend into the foot and body of the animal. These muscles retract the foot into the shell. Extension of the foot is accomplished with the aid of the anterior protractor muscle, most of the fibers of which extend over the viscera. Retraction of the mantle is accomplished by the pallial muscles at the margin of the shell and by the dorsal muscles in the umbonal cavity ( Plate 2, figure 5 ) . Mantle The mantle or pallium ( Plate 3, figure 1 ) secretes the shell, aids in respira- tion, serves as a sense organ, and aids in feeding. The mantle, which is open ventrally, anteriorly and posteriorly, but is closed dorsally, forms two lobes overlying the animal and immediately within the shell. The posterior portion of the mantle fomis the excurrent and incurrent siphons ( Plate 3, figure 1 ) for the intake and exit of water. There are numerous blood vessels ( Plate 4, figure 2) in the mantle and some exchange of gases occurs there. In addition, the epidermis of the mantle is ciliated and aids in feeding by trapping food in 20 University of Kansas Publs., Mus. Nat. Hist. MORPHOLOGICAL FEATURES OF THE SOFT ANATOMY OF A UNIONID MUSSEL Fig. I. Right valve and mantle removed exposing gills, foot, labial palpi, and incurrent and excurrent siphons. Length, approximately 5 inches. Fig. 2. Glochidium just before leaving the gills. Enlarged approximately X 20. Fig. 3. Right valve, mantle, and gills removed. Length, approx. 5 inches. Figures 1, 2, and 3, Aiwdonta fiuviatilis (Dillwyn) {^Anodonta cataracta Say) modified from Simpson, 1884 (pis. 4, 7, and 8) — courtesy of New York State Museum and Science Service. Unionid Mussels in Kansas 21 PLATE 3 suprabranchjal chamber outer gill inner gill excurrent siphon incurrent siphon mantle cavity visceral mass byssus thread adductor muscle hook valve posterior retroctor muscle posterior adductor muscle ntricle onterior retractor muscle onterior adductor muscle anterior protractor muscle 22 University of Kansas Publs., Mus. Nat. Hist. MORPHOLOGICAL FEATURES OF THE SOFT ANATOMY OF A UNIONID MUSSEL Fig. 1. Right valve removed showing the nervous and digestive systems. Length, approximately 5 inches. Fig. 2. Right valve removed showing circulatory and digestive systems. Length, approximately 5 inches. Figures 1 and 2, Anodonta ■fluviatilis ( Dillwyn ) ( = Anodonta cataracta Say ) modified from Simpson, 1884 (pis. 9 and 10) — Courtesy of New York State Museum and Science Service. Unionid Mussels in Kansas 23 PLATE 4 connecting nerves posterior mantle nerve posterior oorta venous sinus intestine stomach cerebral ganglion intestine connecting nerves pedal ganglion -onterior oorta 24 University of Kansas Publs., Mus. Nat. Hist. mucous and transporting the "mucous string" to the area of the labial palpi. The space between the mantle and the body of the animal is the mantle cavity (Plate 3, figure 1). Gills There is a pair of gills on either side of the foot; the gills next to the foot are the inner gills and those next to the mantle are the outer gills (Plate 3, figure 1 ) . The gills serve as organs of respiration, aid in feeding, and are used in reproduction. Water brought into the mantle cavity via the incurrent siphon enters the ostia of the gills by the action of cilia and passes into water tubes in the gills. Traveling dorsally and anteriorly in the tubes, the water eventually passes into the suprabranchial chamber ( Plate 3, figure 1 ) , and moves pos- teriorly to the excurrent siphon. As water passes over the gills gaseous exchanges occur and particulate food matter is trapped by mucous secreted from the gills. The food is trapped on all areas of the gills, but is concentrated ventrally by the cilia of the gills in a ventral food groove whence it is transferred anteriorly to the labial palpi (Plate 3, figure 1). At the anterior end, food is sorted from inorganic debris and passed on to the mouth by the elongate labial palpi. The unionids are unique among pelecypods in that they use all or some part of the gills as a marsupium for the developing eggs and glochidia, until they are discharged from the parent. Foot The class name Pelecypoda, or hatchet foot, indicates the shape of the foot ( Plate 3, figure 1 ) . The highly muscular foot is extended between the two valves and pushed into the substrate; therefore it serves as a holdfast organ but is primarily concerned with locomotion. Locomotion is accomplished in the following manner: with the aid of the anterior protractor muscles and blood sinuses of the foot, the foot is extended from between the two valves and inserted firmly into the substrate in the anteroventral direction. The animal contracts the retractor muscles, the muscles of the foot, and pulls itself forward. Locomotion of the unionid is slow and erratic. Digestive System The digestive system of the unionid is highly adapted for use of particulate matter, such as unicellular algae. Mouth, esophagus, digestive gland, intestine, rectum, and anus are present. The mouth ( Plate 3, figure 1 ) lacks masticatory structures, and the esophagus merely passes food on to the stomach. Digestion is accomplished in the stomach ( Plate 4, figure 1 ) with the aid of digestive juices from the digestive gland, which surrounds the stomach, and the crystalline style, which is specific in the digestion of carbohydrates. The crystalline style is an elongate, rodlike structure within a style sac, a diverticulum of the stomach. The intestine ( Plate 4, figure 1 ) extends from the stomach into the foot, makes several loops, and eventually passes dorsally where it is surrounded by the heart. Posterior to the heart the intestine enlarges and forms a rectum ( Plate 4, figure 1 ) , which empties via the anus ( Plate 4, figure 1 ) into the area near the excurrent siphon, where the waste materials are swept out of the mantle cavity and away from the animal. The part of the animal occupied by the digestive system, digestive glands, and gonads is referred to as the visceral mass (Plate 3, figure 1). Unionid Mussels in Kansas 25 Kidney Although physiologically the kidneys of all animals perform the same gen- eral function, that is to say, they are homeostatic organs, the kidney of the unionid is unique in its morphology and relationship to the heart. The kid- ney is situated immediately below the heart, and one end of the kidney opens into the pericardial chamber of the heart. Pericardial fluid, therefore, is filtered through the kidney and nitrogenous wastes and salts are removed or retained in the ventral glandular part of the kidney as is necessary for the internal stability of the animal. The nitrogenous wastes are eventually discharged from the more dorsally situated bladderlike structure of the kidney into the suprabranchial chamber of the gills and passed out the excurrent siphon. Circulatory System The circulatory system consists of a heart composed of a single ventricle ( Plate 3, figure 3 ) , two auricles ( Plate 3, figure 3 ) surrounded by a pericardial cavity, and numerous arteries and veins (Plate 4, figure 2). The almost transparent blood passes from the two auricles to the ventricle then out the anterior and posterior aortae ( Plate 4, figure 2 ) . The blood eventually reaches smaller vessels, which ramify over the body. Blood passing to the mantle is oxygenated and returned directly to the heart. Blood from the other parts of the body collects in a venous sinus (Plate 4, figure 2) and passes through the kidney, then is moved through the gills where it is oxygenated and is eventually returned to the auricles of the heart. The blood of the fresh-water mussels is an almost transparent, opalescent fluid, having no corpuscles for carrying oxygen as in the vertebrates. There are, nevertheless, small numbers of amoebocytes in the blood (Dundee, 1953). The specific gravity of the blood is variable depending on the species, but averages approximately 1.0026 (Ellis, Merrick, and Ellis, 1930). Nervous Syste7n ( Plate 4, figure 1 ) Two cerebral ganglia near the mouth on each side of the esophagus send connectives ( long nerves ) into the foot to the two pedal ganglia, and posteriorly to the two visceral ganglia. Small nerves extend from these ganglia into the tissues. There are no ears or eyes in the sense of vertebrate structures; there is an otocyst for balance, and various areas of the edge of the mantle are sensi- tive to light. Chemotactic sense receptors around the labial palpi and the mouth detect food. Reproductive System Fresh-water mussels are typically dioecious; that is to say, the organs of the male and female are in separate animals. Anodonta imbecilis has been re- ported as monoecious. The reproductive organs lie in a mass of tissue above the foot and their materials are discharged into the suprabranchial chambers of the gills. Sperm of the male pass to the outside, but eggs are discharged into the water tubes of the gills of the female where sperm fertilize the eggs and further development occurs. For a more thorough treatment of the soft anatomy of fresh-water mussels the reader is referred to Simpson (1884), Baker (1928), and Borradaile et al. (1958). 26 University of Kansas Publs., Mus. Nat. Hist. DEVELOPMENT AND GROWTH OF FRESH-WATER MUSSELS Development The development of the fresh-water mussel consists of five dis- tinct stages: 1) fertilized egg; 2) glochidium in the gill of the mussel; 3) parasitic stage on a fish or a salamander; 4) free-living Juvenal stage; and 5) adult stage. Eggs are fertilized by sperm that enter the gills with incurrent water and develop into small larvae called glochidia ( Plate 3, figure 2 ) . Glochidia range from % to % of a millimeter in height and from M to ?3 of a millimeter in length. They are typically rounded or oval and possess rudiments of a mouth, heart, foot, and intestines when discharged from the parent. The two valves, hinged dorsally, are held together by an adductor muscle ( Plate 3, figure 2 ) . There is usually an accessory sensory structure, the byssus thread (Plate 3, figure 2), which probably aids in the detection of a host fish. Some genera {Anodonta, Strophitus, and Lasmigona) have hook- type glochidia. The hook is at the ventral margin of the glochidial shell and is used to assure attachment of the glochidium to the fish. Hook-type glochidia are most frequently found on the fins and tail of the host fish. Hookless glochidia are most frequently found on the gills. Members of the genera Fusconaia, Crenodonta, Quadnda, Pleurobema, and EUiptio have this type of glochidium. A third type of glochidium, the axe-headed type related to the hook-type, is restiicted to the genus Propfera. Upon discharge from the parent the glochidia attach to, and embed in, the tissues of the proper fish host where further develop- ment occurs. During that parasitic stage, the glochidium completes the adult structures, but does not increase noticeably in size. The period of time that the larva is embedded in the host depends on the viability of the host, temperature of the water, and the place of attachment to the host. The fact that Aplodinottis gninniens (freshwater drum) and Ictalurus punctatus (channel catfish) are infected by several different kinds of glochidia probably is correlated with the bottom feeding habits of these fishes. In nature most infections are light, but heavy or massive infections may be induced artifically. The transformed glochidium breaks through the skin of the fish and falls to the stream bottom as a juvenal mussel. The adult shell forms under the glochidial shell, and the animal begins its growth to adulthood. Some glochidia of Strophitus edentulus, Anodonta imbecilis, and Unionid Mussels in Kansas 27 Obliquaria reflexa are known to develop without passing through the parasitic stage; it is thought, however, that other glochidia of those three genera pass through the parasitic phase. In the non- parasitic cycle the young are retained in the gills of the parent until mature, and discharged into the water where they subsequently fall to the stream bottom to become adults. Growth Increase in size of the shell is accomplished by secretions of the mantle at the margins of the valve. Increase in thickness of the valve is the result of secretions of all areas of the mantle. The periostracum of epidermis of the mussel shell typically shows dark, thin lines alternating with light, broader areas on the shell. The thin, dark lines are spoken of as "rest periods" or lines of arrested growth (Plate 2, figure 1). Less prominent lines of temporary arrested growth are called "growth lines" (Plate 2, figure 1). In some mussels a reliable way to estimate the age of the shell is to count the "rest periods," because these lines roughly correspond to the end of one year's growth. This is not necessarily true, because fluctuations in water level, pollution of water at various intervals, scarcity of food, and other adverse conditions may bring about a period of arrested growth. Growth of fresh-water mussels has been discussed by Lefevre and Curtis (1912); Isley (1914); and Coker, Shira, Clark, and Howard ( 1921 ) . ECOLOGY OF FRESH-WATER MUSSELS Mussels spend the entire juvenal and adult life partially or wholly buried in mud, sandy mud, or mud and rock bottoms of streams, ponds, lakes, canals, and swamps — any permanent body of water. Mussels are rarely found in stream bottoms of shifting sand. An animal generally orientates so that the posterior portion of the shell protrudes from the substrate and is directed upstream. In this way, materials brought in by the ventral incurrent siphon are in part forced into the mantle cavity by the force of the water current, and waste products that are eliminated from the dorsal excurrent siphon are quickly swept away from the animal. In Kansas there are two primary habitats for fresh-water mussels, impounded water (artificial lakes and ponds) and natural streams. The ecology of lakes, ponds, and streams in Kansas is poorly known. Permanent bodies of water in Kansas, even though artificial, un- doubtedly support large unionid populations. A one and one-half acre pond on the campus of The University of Kansas yielded 28 University of Kansas Publs., Mus. Nat. Hist. approximately 2,178 living unionids representing four different species Anodonta grandis, Crenodonta peruviana costata, Ligumia suhrostrata, and Uniomeriis tetralasmtis (Murray, 1960). Although fresh-water mussels in Kansas have a high tolerance to siltation, they are not tolerant to industrial pollution. Early records indicate that many streams in Kansas had relatively clear water; with the advent of agriculture and industry, both turbidity and industrial pollution have greatly increased. Although there is evi- dence that only three species have been extirpated from the state, the abundance of various species has been seriously affected. In areas where Scammon ( 1906 ) reported species as common or abundant, they are now rare, such as Actinonaias carinata carinata, Ligumia recta latissima and Plagiola lineolata, to mention but a few. In certain areas of the Mississippi Valley many species were har- vested to near extinction, and many beds were greatly depleted. Some harvesting occurred in streams in Kansas in the early 1900's, but not to the extent that any species were extirpated. Because the economic value of fresh-water mussels has declined in recent years, overharvesting does not appear to be likely in the near future. Continued industrial pollution and increased turbidity of the waters of Kansas are more likely to extirpate unionids, as well as other aquatic animals. Therefore, as well as for other reasons, Kansas should take the measures that are necessary to reduce and check soil erosion and to control industrial wastes by keeping them out of streams and by purifying those that do enter streams. Condition of Water The chemical and biological condition of the water greatly in- fluences the kinds and number of mussels present. Turbidity is probably the most important single negative factor and can result from either biological or physical conditions of the water. Biolog- ical turbidity generally benefits mussels because this is particulate food matter (plankton) in suspension, and mussels are dependent upon this for food. Turbidity resulting from mud and sand is generally detrimental to mussels, directly by clogging the feeding and respiratory mechanisms or indirectly by reducing the amount of food in the water. Turbidity of streams in Kansas measured from three to fifteen inches on a Secchi disk in July and August, 1956-1959. The amount of calcium carbonate available to the mussel and the amount of carbon dioxide (CO.,) and oxygen (O.) dissolved in the water are of primary importance. Other elements and Unionid Mussels in Kansas 29 compounds in solution are surely important but are of less obvious significance. Because calcium carbonate is necessary for the for- mation of the molluscan shell, a low calcium carbonate content in the water brings about a population of thin-shelled mussels. A high COo content in the water causes the formation of excessive amounts of carbonic acid, that in turn dissolves the calcium car- bonate deposited in the mussel shell. During drought years, when pondlike areas of a stream become stagnant, the lack of oxygen and the increase of carbon dioxide threaten mussel life in the stream. Bottom Conditions Although important to certain species, the type of stream bottom can vary considerably — within the limits of tolerance for any given species. Members of the genus Anodonta and Lcptodea frogiUs are often found in soft to hard mud, rarely in rocky areas. Mem- bers of the genera Crcnodonta, Tritogonia, and Qiiadnda are to be expected in rocky and gravelly areas as well as in mud bottoms. The most highly restrictive ecological factor for fresh-water mussels is the shifting sand bottom. Because most sti^eams of western and central Kansas have this type of bottom, few species of mussels exist in these streams, except as isolated populations in suitable areas. Depth of Water The depth of water inhabited by mussels depends upon tempera- ture, the presence of the proper amounts of dissolved oxygen, and suspended food material. In fast flowing streams mussels are found at depths of 15 feet or more. In Kansas, mussels are frequently found in shallow water (2 to 4 feet) during the summer and early autumn, but as the water cools in late autumn, there is a general movement to deeper pools in the stream. The depth at which mussels live in ponds varies with the time of the year, temperature of the water, presence of a thermocline, and dissolved gases present at various depths; in Kansas, this depth probably varies from 1 to 10 feet. Of the Kansas unionids, only Mcgalonaias gigantea re- mains in relatively deep water (5 feet or more) during even the warmer months. Some species, Caruncidina parva and Lampsilis subrostrata may be found in pondlike areas of a stream deep enough to cover only the siphons, at most an inch in depth. The water in these areas may be as warm as 40° to 45° C. 30 University of Kansas Publs., Mus. Nat. Hist. Vegetation Because plants covering the water cut the light incidence in the water and in turn lower the amount of plankton present at various depths, few mussels are found in heavily covered ponds. Because plant-cover in Kansas streams typically occurs only briefly during drought, the problem is not important for stream faunas. Light Although the matter has been poorly studied, it is evident that mussels react to variations in light intensity. On cloudy days they are less active and siphon water at a much slower rate than on bright sunny days. Mussels quickly close the shell if the light rays are suddenly interrupted, as by a shadow passing over the animal. Current The mixing action of moving water aids in the even distribution of dissolved gases, food, and minerals necessary to the mussel. In well mixed water, mussels tend to move into greater depths, and conversely into shallower water as the stream becomes quiet. Mussels react similarly in ponds and lakes as the water is circulated by the wind. Temperature Water maintains a fairly constant temperature in contrast to the air. The warmest temperature of a stream bottom recorded in this study was a temperature of 38° C. on the Delaware River, July 28, 1956. During the summer months and early autumn the bottom temperature of Kansas streams varies from 25° to 29° C, at least in areas where mussels are found. Winter temperatures on or near the bottom of Kansas streams vary from 5° to 20° C. Predation and Parasitism Mink, otter, raccoon, muskrat, fish, and some birds are reported as predators of mussels (Baker, 1928, p. 29). In Kansas, Stains ( 1956 ) reported that the following unionids were commonly eaten by raccoons: Crenodonta peruviana costata, Quadrula quadrula, Q. pustulosa, Lampsilis anodontoides anodontoides, Ligumia sub- rostrata, Lasmigona complanata, Tritogonia verrucosa, Anodonta grandis, and Proptera alata. It is not uncommon to find small piles of broken shells near or around a muskrat burrow, or on a small Unionid Mussels in Kansas 31 boulder near the stream. These piles usually are 10 to 20 feet from a muskrat burrow, but in several instances shells have been found up to and slightly inside the under-water entrance of the muskrat burrow. The most common species occurring in these small heaps of shells are: Anodonta grandis, Leptodca fragilis, Lampsilis ano- dontoides (young), Lasmigona complanata (young), and Uniomerus tetralasmus. These are the most common thin-shelled species in the streams of Kansas. Baker (1918) lists several species of Lamp- silis and Anodonta as fish food, as well as numerous species of sphaeriids. In Kansas, muskrats undoubtedly take the heaviest toll of adult mussels, and certain fishes take the heaviest toll of juvenal mussels. Aboriginal man used the meat of mussels for food, and the shells for tools and ornaments. Specimens recently identified in the collection of the Kansas Historical Society indicate that pre-Colum- bian Indians used Lampsilis ovata and Elliptio dilatattis, as well as other species, for food. European man has not acquired a taste for the meat of fresh-water mussels, but he has used the shells for buttons, and the meat as food for chickens and hogs. Three species of water mites are known to parasitize mussels in Kansas, Unionicola crassipes, U. wolcofti, and U. formosa (Mur- ray, unpublished data). The eggs and larvae are generally em- bedded in the gills, foot, and mantle, and the adults move around over the gills and body. In most instances they seem to cause little damage to the mussel. Several species of flukes parasitize mussels but have not been studied in the Kansas fauna. Baker (1928, p. 30) and Utterback (1916b, p. 520) report several species of flukes commonly found in the branchial and pericardial cavities, various muscular areas of the body, and in the ovaries. Discoloration of the nacre of Anodonta sp. is frequently the result of the presence of distomid flukes or larval mites. COLLECTION AND PRESERVATION OF MUSSELS Numerous methods have been devised for obtaining fresh-water mussels. The most successful method used in the tributaries of the Ohio and Mississippi rivers was a technique popular in the early 20th century, the bar and crowfoot hook method. Hooks composed of four prongs were suspended from 2 to 5 feet on a bar 6 to 10 feet long. These hooks were separated from each other by 4 to 6 inches. As the hooks were dragged along the bottom of the stream. 32 University of Kansas Publs., Mus. Nat. Hist. one prong of a hook might fall into the open valves of a mussel. When this happened the hook stimulated the mussel to close the valves, and the mussel could be pulled to the surface. Among other techniques used in the early 1900's were the dip-net drag, shoulder rake, shell tongs, fork, and the dredge. Each of these was ad- vantageous in particular situations depending on depth of water, bottom conditions, and the number of persons operating the equip- ment. A detailed account of the various techniques used is given by Lefevre and Curtis ( 1912 ) . One of our most effective methods was the hand method. By feeling along the stream bottom with the bare hands and placing the mussels in a sack or nearby boat, we could effectively sample and collect from an area. Shells of dead mussels are to be found along the sti^eam banks but these shells are frequently worn, and much of the original luster and beauty ai^e lost. Smaller specimens can be recovered by straining bottom-mud and water through a screen of }i inch mesh. In most years, the best time to collect mussels in Kansas is late summer or early autumn, inasmuch as 1 ) the streams then are usually lowest, allowing greatest accessibility to beds of mussels, and 2 ) most fresh-water mussels in Kansas reproduce at those times of year; consequently, the mussels move into shallower water and can be more easily obtained. The entire animal can be preserved in 10 per cent formalin or preferably in 70 per cent ethyl alcohol. In either case the animal must be washed several times in water and alcohol in order to remove the copious amounts of mucous. Most collectors save only the shell. By placing the live animal in boiling water or a steam oven, the adductor muscles can be relaxed to permit removal of the soft parts. Dead shells that are obtained from stream banks may frequently be restored to their original luster by soaking in a mild acid, such as oxalic acid, followed by light scrubbing with a wire brush. In our work, at the time of collection the collector records facts concerning water temperature, bottom temperature, sti'eam velocity, surrounding bank conditions, time spent collecting, time of day, date, and other important ecological information in a field notebook. The collector provides a label for the specimens taken at a par- ticular locality. He writes on this label the date, locality, collector's name, and field number, corresponding to the information in the collector's field notes. Once the specimens are properly identified Unionid Mussels in Kansas 33 a permanent label is placed in the container holding the specimens. All the data appearing on the field tag are transferred to this label as neatly as possible. Regardless of the size of the collection, a catalogue should be kept. This should be a book of good quality paper in which a catalogue number, name of shell (genus, species, authority), date, and collector's field number are recorded. The catalogue number is placed on the shell and on the permanent label accompanying the shell. The catalogue, specimens, and field notes should not be altered or destroyed. The same species from a different locality or from the same locality but obtained at a later date receives a different catalogue number. KEY FOR IDENTIFICATION OF SPECIES AND SUBSPECIES OF MUSSELS IN KANSAS Introduction The key is intended to aid in the identification of species and subspecies occurring in Kansas. The key should be used in con- junction with the descriptions of the several species and the plates accompanying each species. The key is arranged in couplets. Each statement of a couplet is in opposition to the other or at least distinctly different. It is nec- essary to read each statement within a couplet before making a choice. The choice may refer the reader to another couplet or to the name of a species; the page number following the name of a species refers the reader to the account of that species. If any part of the statement of the couplet does not seem to be true for a specimen, the other statement of the couplet should be true. 1. A. Surface of shell having knobs, pustules, or undulations ... 2 B. Surface of shell lacking knobs, pustules or undulations (only growth Hues interrupt smoothness of shell) 16 2. A. Surface of shell having undulations, lacking knobs or pustules 3 B. Surface of shell having or lacking undulations, having knobs, pustules or irregular W-shaped sculpture 7 3. A. Shell alate, highly compressed Lasmigona complanata p. 87 B. Shell not alate, not highly compressed 4 4. A. Undulations only on dorsoposterior slope never on lateral or posteroventral portions of shell Lasmigona costata p. 86 B. Undulations on dorsoposterior slope and on most of shell except anterior end 5 5. A. Shell longer than 165 mm Megalonaias gigantea p. 42 B. Shell shorter than 165 mm 6 2—692 7. A B. 8. A, B. 9. A, 34 University of Kansas Publs., Mus. Nat. Hist. 6. A. Shell compressed; shell having numerous small undulations over surface; umbo flat, not extending greatly above hinge line Crenodonta peruviana costata p. 49 B. Shell inflated; shell having fevi' large undulations over the surface; umbo inflated, extending well above hinge line. Crenodonta peruviana peruviana p. 47 Shell lacking distinct lateral teeth Arcidens confragostis p. 97 Shell having distinct lateral teeth 8 Umbonal region having small zigzag undulations, some forming irregular W's; lacking distinct pustules or knobs. Megalonaias gigantea p. 42 Umbonal region lacking small zigzag undulations, may have pustules or knobs 9 Length and height of shell nearly equal, shell quadrate, never elongate; posterior margin of shell perpendicular to dorsal and ventral margins 10 B. Length of shell more than height, shell elongate, never quadrate; posterior margin of shell usually not perpen- dicular to dorsal and ventral margins, if nearly perpen- dicular then height of shell less than 50% of total length of shell 14 10. A. Has 2-6 knobs extending over lateral portion of shell or on posterior ridge; usually lacking pustules; if pustules pres- ent shell having deep sinus above posterior ridge 11 B. May have knobs on shell; always having small pustules on shell; shell lacking sinus above posterior ridge 13 11. A. Knobs on posterior ridge of shell; posterior ridge high, having deep sinus above; epidermis sometimes marked by dark green chevrons Quadrula metanevra p. 61 B. Knobs on lateral areas of shell; posterior ridge not greatly enlarged, lacking sinus above; epidermis lacking dark green chevrons 12 12. A. Knobs on left valve opposite those on right valve. Quadnda nodulata p. 58 B. Knobs on left valve not opposite (alternating with) those on right valve Obliquaria reflexa p. 103 13. A. Pustules restricted to lateral portion of shell, remainder of shell smooth; shell inflated Quadnda piistulosa p. 55 B. Pustules frequently covering entire shell, restricted to posterior and lateral ridges, or any variation of the above, never restricted to lateral area; shell somewhat compressed Quadrula quadnda p. 50 14. A. No knobs on shell; small pustules usually over most of shell, except extreme anterior end; epidermis usually dark brown or black Tritogonia verrucosa p. 67 B. Knobs or pustules restricted to posterior or lateral ridges of shell or both; epidermis yellow, yellowish-green or light browTi 15 Unionid Mussels in Kansas 35 15. A. Lateral and posterior ridges prominent; epidermis lacking dark green chevrons Plethobasus cijphtjus p. 68 B. No lateral ridge, posterior ridge evident and represented by knobs extending to posteroventral margin; epidermis usually marked by dark green chevrons. Quadrula cylindrica p. 64 16. A. Pseudocardinal teeth absent or as nodulose swellings; lat- eral teeth absent 17 B. Pseudocardinal teeth present; lateral teeth usually present, if absent pseudocardinal teeth massive 19 17. A. Pseudocardinal teeth seemingly absent, but represented by small nodulous thickenings on hinge line. Strophitus rugosus p. 99 B. Pseudocardinal teeth absent, no indications of their pres- ence 18 18. A. Umbo not raised above hinge line Anodonta imbecilis p. 95 B. Umbo raised above hinge line Anodonta grandis p. 92 19. A. Lateral teeth absent or as small swellings, but sweUings of right valve do not interlock into swellings of left valve. Lasmigona complanata p. 87 B. Lateral teeth present, lateral tooth of right valve interlocks into lateral teeth of left valve 20 20. A. Interdentum extremely narrow or offering no bearing surface to opposing valve 21 B. Interdentum wide, may be long or short, offering obvious bearing surface to opposing valve 36 21. A. Nacre purple, pink, or salmon, if pink or salmon pseu- docardinal teeth thin, highly compressed, and sharp, slightly ragged 22 B. Nacre white, rarely pink, if pink pseudocardinal teeth thick, heavy, blunt, and extremely ragged 26 22. A. Pseudocardinal teeth thin, weak, not ragged 23 B. Pseudocardinal teeth thick, strong, slightly ragged 24 23. A. Epidermis entirely or partially glossy, usually yellowish- green; pseudocardinal teeth sharp in young and old specimens Leptodea laevissima p. 123 B. Epidermis dull, yellowish to yellowish-brown; pseudocardi- nal teeth sharp only in young individuals . Leptodea fragilis p. 120 24. A. Umbo cavity shallow, no more than one-third of the row of dorsal muscle scars situated within umbonal cavity. Proptera alata p. 126 B. Umbo cavity deep, more than one-third of the row of dorsal muscle scars situated within the umbonal cavity 25 25. A. Shell inflated, but not globose; dorsoposterior area having two to three indistinct ridges extending from umbo to posterior margin Proptera purpurata p. 129 B. Shell extremely globose; dorsoposterior area usually lacking indistinct ridges Proptera capax p. 132 26. A. Shell elongate, never truncate posteriorly 27 B. Shell globose or truncate posteriorly 33 36 University of Kansas Publs., Mus. Nat. Hist. 27. A. Umbo sculpture of 5-6 concentric loops curved sharply up- ward posteriorly 28 B. Umbo sculpture eroded or obsolete, if present of V-shaped or doubly looped parallel ridges 29 28. A. Adult shell never longer than 45 mm.; surface of shell smooth, growth lines not raised but marked by indistinct change of color Carunculina parva p. 135 B. Adult shell longer than 45 mm.; surface of shell not smooth, growth Unes raised and roughened, and usually marked by distinct change of color Uniomerus tetralasmus p. 83 29. A. Epidermis yellowish, yellowish-green, or yellowish-tan, usually glossy; nacre white to salmon, never purple; posterior end pointed 30 B. Epidermis light to dark brown, black, or yellowish-brown, usually dull; if yellowish-brown then posterior end rounded; nacre white, rarely pink 31 30. A. Epidermis having green rays. Lampsilis anodontoides fallaciosa p. 146 B. Epidermis lacking green rays. Lampsilis anodontoides anodontoides p. 143 31. A. Pallial hne lightly impressed anteriorly, rarely fonning crenulations; umbo of adult and young shell having distinct V-shaped parallel ridges Ligumia subrostrata p. 140 B. Pallial Hne deeply impressed anteriorly, usually forming crenulations; umbo of adult usually eroded, young shell rarely having indistinct V-shaped ridges 32 32. A. Posterior end of shell pointed; height of shell less than 50% of total length of shell; epidennis usually dark. Ligumia recta latissima p. 137 B. Posterior end of shell rounded; height of shell at least 50% of total length of shell; epidermis usually straw-colored or yellowish-brown Lampsilis radiata siliquoidea p. 149 33. A. Posterior ridge rounded; shell globose. Lampsilis ovata ventricosa p. 152 B. Posterior ridge angulate; shell truncate posteriorly 34 34. A. Epidermis having rays broken into squarish, triangular, or chevron shaped spots Dysnomia triquetra p. 155 B. Epidermis may have rays forming zigzag pattern resembling a series of W's, may have all rays continuous, or may have a mixture of continuous and zigzag rays 35 35. A. Posterior pseudocardinal tooth of left valve turned sharply dorsad and that portion of hinge line interrupted by tooth; surface of shell having numerous W-shaped green rays, few continuous green rays Truncilla donaciformis p. 114 B. Posterior pseudocardinal tooth of left valve usually straight, not turned sharply dorsad; surface of shell having few or no W-shaped green rays, having numerous thin green rays Truncilla truncata p. Ill 37. A B, 38. A. B. 39. A B. 40. A. B. 41. A. B. 42. A. Unionid Mussels in Kansas 37 36. A. Epidermis of shell having small dark brown or green rays extending from umbo to ventral margin and breaking to form small blocks or squares 37 B. Epidermis usually lacking rays from dorsal to ventral mar- gins; if present never broken into small blocks or squares. 38 Umbo greatly compressed Plugiola lineolata p. 117 Umbo moderately inflated Ctjprogcnia aherti p. 105 Umbo cavity shallow, barely extending under the inter- dentum Elliptio dilatatus p. 80 Umbo cavity deep, extending under the interdentum 39 Shell having lateral and posterior ridges . Plethohasus cyphtjus p. 68 Shell having posterior ridge, lacking lateral ridge 40 Shell quadrate; posterior margin of shell at nearly right angles to dorsal and to ventral margins. (nonpustulose) Quadrula pustulosa p. 55 Shell not quadrate, posterior margin of shell rounded or pointed 41 Posterior end of shell rounded, never pointed; umbones low, slightly inflated, not decidedly directed anteriorly. Actinonaias carinata carinata p. 108 Posterior end of shell pointed or bluntly pointed; umbones high, inflated, and directed anteriorly 42 Center of umbo posterior to posterior border of anterior adductor muscle scar; shape of shell forming a scalene triangle i^ Fusconaia flava p. 39 B. Center of umbo farther anterior, not posterior to posterior border of anterior adductor muscle scar; shape of shell forming an obtuse \^>^ or right [N^ triangle 43 43. A. Anterior margin of shell usually straight; anterior edge of umbo but slightly posterior to anterior margin of shell; center of umbo anterior to posterior border of anterior adductor muscle scar; umbo greatly inflated. Pleurobema coi datum pyramidatum p. 76 B. Anterior end of shell slightly or distinctly curved; anterior edge of umbo not close to anterior margin of shell; center of umbo not anterior to ( directly above ) posterior border of anterior adductor muscle scar; umbo moderately inflated 44 44. A. Shell inflated Pleurobema cordatum catillus p. 74 B. Shell flattened Pleurobema cordatum coccineum p. 71 ACCOUNTS OF SPECIES Introduction The abbreviated synonymy under each technical name consists of the fol- lowing: 1 ) original proposal of the name, 2) first usage of the current name-combination, 3 ) standard treatise in which a more nearly complete synonymy is given, 4) citations to previous records of the species or subspecies in Kansas. 38 University of Kansas Publs., Mus. Nat. Hist. When more than one usage is hsted under any one of the four categories mentioned above, the usages are arranged in chronological order. The type locality for each species is given as accurately as possible; unfor- tunately early workers did not always precisely indicate the locality from which the type specimen was obtained. No attempt has been made to redescribe each of the species of the Unionidae in Kansas; descriptions given are adapted from those of Walker (1918), Simp- son (1914), and Baker (1928), and are modified on the basis of personal observation. The ecology of each species is discussed, including the type of stream bottom inhabited. If habitats occupied in Kansas differ from those noted by other authors, this difference is indicated. Wherever possible the host fishes for the glochidia are given. This inade- quately studied but extremely important aspect of unionid life history needs additional and more careful study. Common and scientific names of fishes are, in so far as possible, those recommended by Chute (1948) and Bailey (1951, 1952, 1953, and 1956). Finally, remarks are made concerning the following aspects: 1 ) minor variations of the shell of species and subspecies in Kansas, as com- pared with the form in the Mississippi Valley, 2 ) variations of the shells of the species within the state, 3 ) economic value of the shell, appearance of the nacre, drying qualities of the shell and various facts concerning the species that might be of interest, 4) the largest specimens of each species recorded from Kansas. Species reported have been collected in Kansas within the past ten years ( except Cyprogcnia aherti, Scammon, 1906, and Dysnomia triquetra, Scammon, 1909). Catalogue numbers refer to, and all specimens are deposited in, the mollusk collections of the Museum of Natural History, The University of Kansas. Maps and Illustrations The distribution maps illustrate the presently known range of the species within Kansas. The insert map of North America on each distribution map indicates the approximate distribution of the species as recorded in the lit- erature. Black dots ( • ) indicate localities from which specimens have been studied by us; circles (o) represent localities of specimens cited in literature that have not been available for study, but which are thought to represent vaUd records. Any single symbol on the map may include one or more actual collecting sites. In the account of each species under "Occurrence in Kansas," the exact western- most locaHties for that species in Kansas have been given. Every effort has been made in Plates 5-44, to reproduce photographically the shells of each species in a manner most useful to those who attempt to identify shells without the aid of a reference collection. Interior views were illuminated at right angles to the shell with a diffusion ring between the fight source and the specimen. In most cases the muscle scars have been painted with water colors in order to denote the size, shape, and position of the scars. E.xternal views were photographed with two equidistant lights; a diffusion ring was placed between the light source and the specimen. Unionid Mussels in Kansas 39 PHYLUM MOLLUSCA Class Pelecypoda ORDER EULAMELLIBRANCHIA Family Unionidae Subfamily Unioninae ^ Genus Fusconaia Simpson 1900 *■ Shell round, rhomboid, triangular or short, elliptical, having mod- erate posterior ridge; beaks high, full, curved inward and forward; beak sculpture of few coarse, parallel ridges, curving upward be- hind; epidermis dark; surface not sculptured; hinge plate of mod- erate width; pseudocardinals strong; nacre white, salmon or purple (after Walker, 1918, p. 48). Type: L/nio trigomis Lea. Wabash Pig-Toed Mussel Fusconaia flava (Rafinesque) Plate 5, figures 1-4; text figure 1 Obliquaria flava Rafinesque, 1820, Ann. Gen. Sci. Phys., Bruxelles, .5:305, pi. 81, figs. 13, 14. Fusconaia flava, Ortmann, 1919, Mem. Carnegie Mus., 8:14; Franzen and Leonard, 1943, Univ. Kansas Sci. Bull., 29:399; Leonard and Leonard, 1946, Univ. Kansas Sci. Bull., 31:118. Quadrula ruhiginosa, Scammon, 1906, Univ. Kansas Sci. Bull., 3:359; Simp- son, 1914, A descriptive catalogue . . . mussels, p. 872. Unio ruhiginosus. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 96; Call, 1885, op. cit., p. 122; Call, 1886, op. cit., p. 182; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:22. Tijpe Locality: Small tiibutaries of Kentucky, Salt, and Green rivers. General Distrihiifion: Entire Mississippi River Drainage from western New York to eastern Kansas, Nebraska, South Dakota, and Texas; St. Lawrence River; Nelson River; Red River of the North; southern Ontario, Canada (Raker, 1928, p. 55; Strecker, 1931, p. 30; La Rocque, 1953, p. 91 ) . Occurrence in Katisas: Fusconaia fiava is common in the rivers of the southeastern drainages. It has been recovered from several localities in southern tributaries of the Kansas River. The westernmost records for F. flava in Kansas are: Kansas River, Topeka, Shawnee County (specimen not seen); 3/2 mi. W and 2/1 mi. N Americus, Lyon County ( sec. 20, T. 17 S, R. 10 E ) ; 4 mi. E and 2 mi. S Hamilton, Greenwood County ( sec. 21, T. 24 S, R. 12 E ) ; Fall River, Eureka, Greenwood County (specimen not seen); Elk River, VA mi. W Elk Falls, Elk County (sec. 34, T. 30 S, R. 11 E). 40 University of Kansas Publs., Mus. Nat. Hist. PLATE 5 Fusconaia flava (Rafinesque), no. 10611; from Neosho River, at a place VA mi. E and 1 mi. S St. Paul, Neosho Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, right valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length. SJ* inches. Unionid Mussels in Kansas 41 Museum of Natural History University of Kortsos 1959 Fig. 1. Distribution of Fuscoiwia flava in Kansas. Insert map shows approxi- mate distribution of this species in North America. Description of Shell: Shell elongate-quadrate, thin to solid, inequipartite to equipartite, compressed to rather inflated; beaks slightly elevated, somewhat inflated, turned forward; anterior end rounded, truncated above and meeting dorsal line at sharp angle; posterior end obliquely truncated; dorsal margin slightly curved or straight; basal margin straight, slighdy curved, or having slight emargination; posterior ridge conspicuous, roundly angled, having more or less distinct depression in front and usually ending in point at basal margin; surface marked only by growth lines that may form slight ridges; epidermis brownish or reddish, having tinge of green, faintly rayed in juvenile, dark and uniform in old individuals. Two elevated, triangular, elongated, serrated pseudocardinals in left valve; one large, biangular, elevated pseudocardinal in right valve; usually a denticle anterior to and one posterior to pseudo- cardinal of right valve; laterals long, straight, elevated, double in left, inclined to double in right valve; interdentum slight, narrow or absent; anterior muscle scar deeply impressed; posterior scar well marked; beak cavities deep, broad; nacre whitish to salmon, irides- cent posteriorly ( after Baker, 1928, p. 53. ) Although F. fiava is prob- ably the only representative of the genus in Kansas, we include a de- tailed description of the species in case F. ebeniis or F. undata occur in the state. 42 University of Kansas Publs., Mus. Nat. Hist. Ecology: In Kansas F. flava can be found in all habitats except those of shifting sand bottoms. It occurs in water no more than three inches deep, to pools from four to five feet deep. Host for Glochidia: Not recorded. Remarks: F. flava may be confused with but one other species in Kansas, Pleiirobema cordatum coccineum. In P. c. coccineum the posterior ridge is reduced to a rounded swelling or is lacking; the entire shell is more roundly oblique, and the pseudocardinals are more massive and less elevated. In addition to F. fiova, Scammon (1906, p. 360, 366) reported F. undaia and F. ehenus, which he did not illustrate. The F. undata in his Arkansas collection we identify as F. flava. F. ehenus and F. undata may still occur in Kansas, but no authentic records are known in the last 54 years, F. jiava was valued for buttons. The nacre is white in juveniles but in some older individuals has yellow blemishes or a salmon tint. Large specimens from Kansas are 4 to 4/2 inches long. Genus Megalonaias Utterback 1915 Shell large, heavy, obovate or rhomboid, alate posteriorly; dies obliquely folded; beak sculpture of coarse, doubly looped corruga- tions extending over upper surface of disc as nodulous plications; epidermis dark-brown or blackish; beak cavities narrow and deep; anterior muscle scars deep, filled by nacreous deposit; posterior scars large, indistinct (after Walker, 1918, p. 46). Type: Unio her OS Say. Giant Washboard Mussel Megalonaias gigantea (Barnes) Plate 6, figures 1-4; text figure 2 Unio gigantens Barnes, 1823, Amer. Jour. Sci., 6:119 [as variety of 17. crassus (Say)]. Megalonaias gigantea, Ortmann and Walker, 1922, Occas. Papers Mus. Zool., Univ. Michigan, no. 112, p. 7. Quadrula heros, Scammon, 1906, Univ. Kansas Sci. Bull., 3:346; Simpson, 1914, A descriptive catalogue . . . mussels, p. 825. Type Locality: Mississippi River, Prairie du Chein. General Distribution: Throughout the Mississippi River drainage, west to eastern Texas, Kansas, Iowa, and Wisconsin; Red River of the North; Manitoba, Canada; Tombigbee River, Alabama; Nuevo Leon, Northern Mexico. Unionid Mussels in Kansas 43 PLATE 6 Megalonaias gigantea (Barnes), no. 10595; from Neosho River, 2 mi. W St. Paul, Neosho Co., Kansas. Fig. 1, interior, right valve; fig. 2, interior, left valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 7% inches. 44 University of Kansas Publs., Mus. Nat, Hist. Occurrence in Kansas: Megalonaias gigantea is common in the lower Neosho River but rare in the Marais des Cygnes River. Al- though there are no valid records from the Verdigris River, it prob- ably occurs there. Unio undulatus reported by Call ( 1885-1887 ) in- cludes Crenodonta peruviana costata and M. gigantea; therefore, it is not possible to give accurate localities for Call's records for these two mentioned species. The westernmost records for M. gigantea in Kansas are: Marais des Cygnes River, 4 mi. S and 4 mi. E Admire, Lyon County; Neosho River, /2 mi. S Ottumwa, Coffey County. 39 38 - l^^s^ ■-E& i - l.._ Jo^f^\r^ ■ Scot* '0 0 M «) Miles Museum of Notural History University ol t r '^' Ctjprogenia aberti (Conrad), no. 3180; from Neosho River, Lyon Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, right valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 3 inches. Unionid Mussels in Kansas 107 39 38 Museum of Nolurol HiBtorj University ol Konsos I9i9 39 38 100 Fig. 23. Distribution of Cyprogenia aherti in Kansas. Insert map shows approximate distribution of this species in North America. Description of Shell: Shell solid, somewhat rhomboid, slightly inflated; beaks low, compressed, turned forward over kinule, having faint sculpture; posterior ridge rather sharp dorsally, gradually be- coming less distinct and slightly double in later growth; disc flat- tened above; posterior slope having wide, shallow, radial furrow behind posterior ridge; surface of shell having low, wide, concentric ridges; epidermis somewhat shiny or yellowish-green, having dots and flecks of dark green arranged in broken rays. Left valve having two low, radial pseudocardinal teeth, right valve having one pseudocardinal, sometimes an accessory denticle to each side; beak cavity shallow, compressed; interdentum wide, flat; muscle scars small, smooth, impressed; nacre white (after Simpson, 1914, p. 328). Ecology: There are no data on the ecology of this species in Kansas or Missouri. The bottom conditions of Kansas rivers from which C. aberti has been reported are rock, gravel, and soft mud. Host for Glochidia: Not recorded. Remarks: Because of the compressed shell, oval shape, and light green markings on the shell, C. aherti will not be confused with any other Kansas unionid. If C. aberti presently occurs in the southern drainages, it is rare. No specimens have been recovered since 1906. The largest C. aherti reported for Kansas measures approximately 3/8 inches long. 108 UNIVERSITi' OF KANSAS PuBLS., MuS. NaT. HiST. Genus Actinonaias Fischer and Crosse 1893 Shell ovate or subelliptical, distinctly longer than high, compressed or slightly inflated, lacking or having indistinct posterior ridge; disc not sculptured; beaks moderately anterior, never in middle, never at anterior end; beak sculpture of poorly developed, double-looped, faint bars obliterated in central parts; epidermis yellowish to green- ish, generally having distinct green rays; differences between male and female shells scarcely noticeable ( after Ortmann, 1912, p. 324 ) . Type: Unio sapotalensis Lea. Mucket Mussel Actinonaias carinata carinata ( Barnes ) Plate 28, figures 1-4; text figure 24 Unio carinatus Barnes, 1823, Amer. Jour. Sci., 6:259, pi. 13, fig. 10. Actinonaias carinata, Ortmann and Walker, 1922, Occas. Papers Mus. Zool., Univ. Michigan, no. 112, p. 47. Unio ligamentinus. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 120; Call, 1886, op. cit., p. 180; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:21. Lampsilis ligamentinus, Scammon, 1906, Univ. Kansas Sci. Bull., 3:289; Simpson, 1914, A descriptive catalogue . . . mussels, p. 79. Type Locality: Fox River, Wisconsin. General Distribution: Mississippi River drainage as far west as Kansas; St. Lawrence River drainage; southern Ontario and Mani- toba, Canada; Lake St. Clair and its drainage; Lake Erie drainage; Niagara River and its tributaries; Lake Ontario (Baker, 1928, p. 221; LaRocque, 1953, p. 85). Occurrence in Kansas: Actinonaias carinata carinata is now re- stricted to southeastern Kansas. Scammon ( 1906, p. 291 ) reported it as uncommon in the Kansas and Wakarusa rivers and common in the Marais des Cygnes River. In 1919 it occurred in the Neosho River as far north as Emporia (Coker, 1919, p. 28). Recent investi- gations in these same areas indicate that the "mucket mussel" is presently found only in the Neosho and Spring rivers. It probably ranges no farther north than the western edge of Allen County. The westernmost records for A. c. carinata in Kansas are: Neosho River, Emporia, Lyon County; Fall River, Eureka, Greenwood County ( specimen not seen ) . Description of Shell: Shell thick, heavy, subovate or subelliptical, inequipartite; anterior end rounded; posterior end pointed; ventral margin broadly rounded; dorsal margin almost straight; posterior ridge rounded, faint or absent; shell usually not much inflated, young Unionid Mussels in Kansas 109 PLATE 28 Actinonaias carinata carinata (Barnes), no. 10713; from Neosho River, VA mi. E Oswego, Labette Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, right valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 4/4 inches. 110 University of Kansas Publs., Mus. Nat. Hist. 100 39 39 38 Miislum of Natural Hielerf Unlversily of Kansas 1959 38 100 Fig. 24. Distribution of Actinoiuiias carinata carinata in Kansas. Insert map shows approximate distribution of this species in North America. individuals quite compressed; beaks somewhat swollen, not much elevated above dorsal margin; beak sculpture consisting of a few delicate, more or less distinct bars, somewhat double-looped; surface lacking sculpture, growth lines occasionally exaggerated at rest periods and dark bordered; epidermis yellowish, greenish, olive or brownish-yellow, having many broad green rays usually increasing in width from umbones; old individuals usually rayless becoming brown or blackish; concentric bands of color rarely present; female shell having post-basal region slightly more rounded than male shell. Two large, subtriangular, elevated, heavy, serrated, divergent pseudocardinal teeth in left valve, one in right valve, some indi- viduals having one small anterior and posterior tooth in right valve; laterals long, elevated, strong, heavy, somewhat roughened, two in left, one in right valve; interdentum long, narrow; beak cavity mod- erately deep; dorsal muscle scars in beak cavity and on hinge plate; anterior adductor muscle scar deeply excavated, sulcate; posterior scar distinct; nacre silvery white, often faintly cream colored, iri- descent posteriorly ( after Baker, 1928, p. 218 ) . Ecology: Typically this species is found in large rivers on riffles having sand and gravel bottoms. Occasionally it is found on hard, rocky bottoms wedged between large rocks (Baker, 1928, p. 220). In Kansas, specimens occur in sandy, muddy, rocky, and gravelly stream bottoms. Unionid Mussels in Kansas 111 Hosi for Glochiclia: The glochidia have been found naturally on the gills of: Lepomis cyonelhis, green sunfish; L. macrochirus, blue- gill; Micropteriis dolomieui, small-mouthed bass (Baker, 1928, p. 220). Infection by artificial means has occurred on several other game fishes. Remarks: The southern variety of this species, A. c. gibba Simp- son, would be expected to occur in Kansas. A. c. gibba is described as "Shell considerably shorter and more solid than the type, with a decided, curved, posterior ridge, the dorsal region much rounded or humped; surface having strong, low concentric ridges, muscle scars smaller than in the typical shell" ( Simpson, 1914, p. 82 ) . Be- cause specimens from Kansas are elongate and not humped or rounded, are unlike the gibba reported from Missouri, and agree with the description applied to the northern forms, we judge that specimens from Kansas are of the subspecies carinafa as previously reported by Scammon ( 1906, p. 290 ) . The even thickness of the shell, the smooth epidermis, and the beautiful nacre, made this one of the most highly valued shells for buttons. Older specimens show a definite bluing of the posterior area of the nacre, a darkening of the epidermis, and an absence of periostracum from the umbonal region. Large specimens from Kan- sas range from 5 to 5/2 inches in length. Genus Truncilla Rafinesque 1819 Shell rounded, oval or subtriangular, solid, inflated, generally smooth, rayed; beak sculpture delicate often obsolete, double- looped; shell of female differing from that of male, having post- basal region inflated, thinner than remainder of shell and of different texture; epidermis having broken, somewhat triangular rays; lacking a wide, radiate posterior furrow; pseudocardinals rather compressed, high, ragged; hinge plate narrow (after Walker, 1918, p. 80). Type: Truncilla truncafa Rafinesque. Deer Toe Mussel Truncilla truncata Rafinesque Plate 29, figures 1-4; text figure 25 Truncilla truncata Rafinesque, 1820, Ann. Gen. Sci. Phys., Bruxelles, 5:301. Pkigiola elegans, Scammon, 1906, Univ. Kansas Sci. Bull., 3:309; Simpson, A descriptive catalogue . . . mussels, p. 307. Unio elegans. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 96; Call, 1885, op. cit., p. 120; Call, 1886, op. cit., p. 180; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:21. 112 University of Kansas Publs., Mus. Nat. Hist. PLATE 29 Truncilla truncata Rafinesque, no. 10424; from Maiais des Cygncs River, at a place 4/2 mi. W and 2 mi. N Osawatomie, Miami Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, right valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 2 inches. Unionid Mussels in Kansas 113 Type Locality: Ohio River. General Distribution: Throughout the Mississippi River drainage from western Pennsylvania west to Iowa and eastern Kansas, north to Minnesota and Michigan, south to northern Ahibama, Tennessee and Texas (Baker, 1928, p. 226). Occurrence in Kansas: Triincilla truncata occurs in the eastern one-fourth of the state. It is common in the Marais des Cygnes River, but is rare in the Neosho River. Its occurrence north of the Kansas River is doubtful. The westernmost records for T. truncata in Kansas are: Wildcat Creek, Riley County (specimen not seen); Neosho River, 2/2 mi. W Americus, Lyon County ( sec. 9, T. 18 S, R. 10 E ) ; Little Arkansas River, Wichita, Sedgwick County ( specimen not seen ) . 39 38 Muieum of Nalural Hislory Un>»ersitif of Konsoi 1959 39 100 Fig. 25. Distribution ol TrunciUa truncata in Kansas. Insert map shows approximate distribution of this species in North America. Description of Shell: Shell subtrigonal, thick, solid, somewhat compressed; anterior end rounded, posterior end obliquely trun- cated, joining the short, slightly rounded dorsal margin at a sharp angle; ventral margin convex, curved upwards into posterior margin, producing sharp posterior end; posterior ridge sharp having exca- vated area behind and slight radial furrow in front; beaks elevated slightly, incurved; beak sculpture of 3-4 fine bars; epidermis green- ish, yellowish or yellowish-brown, usually distinctly rayed or having dark green spots, in some forming zigzag pattern; in older specimens 114 University of Kansas Publs., Mus. Nat. Hist. rays disappearing and shell brownish to black; secondary sexual differences slight. Pseudocardinals triangular, compressed, elevated, subparallel, strongly serrated, two in left valve, one in right valve; some shells having small denticle in front and above tooth of right valve, very old shells frequently having second denticle posteriorly; interdentum narrow; beak cavities shallow; anterior muscle scars deeply im- pressed; posterior muscle scars distinct; nacre silvery-white, occa- sionally pinkish, iridescent posteriorly ( after Baker, 1928, p. 224 ) . Ecology: In Kansas T. truncata is found in hard and soft mud, sand mixed with mud, rocky, and gravel bottoms of large and small streams. Host for Glochidia: Not recorded. Remarks: T. truncata may be confused with T. donaciformis. The differences between these are discussed under T. donaciformis. Because of the small size, T. truncata was not valued for buttons. The nacre is beautiful, and as in T. donaciformis it is rarely blem- ished. The drying qualities of the shell are excellent. Large speci- mens from Kansas range from 2/4 to 3 inches in length. Fawn's Foot Mussel Truncilla donaciformis (Lea) Plate 30, figures 1-4; text figure 26 Vnio donaciformis Lea, 1828, Trans. Amer. Phil. Soc, 3:267, pi. 4, fig. 3; Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 96; Call, 1886, op. cit., p. 180; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:22. Truncilla donaciformis, Ortmann and Walker, 1922, Occas. Papers Mus. Zool. Univ. Michigan, no. 112, p. 50; Franzen and Leonard, 1943, Univ. Kansas Sci. Bull., 29:378. Plagiola donaciformis, Scammon, 1906, Univ. Kansas Sci. Bull., 3:310; Simpson, 1914, A descriptive catalogue . . . mussels, p. 308. Unio zigzag. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:51; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:23. Type Locality: Ohio. General Distribution: Throughout the Mississippi River drainage from western Pennsylvania west to eastern Kansas and eastern Texas, from Louisiana north to Minnesota; Alabama (Baker, 1928, p. 230). Occurrence in Kansas: T. donaciformis occurs in the eastern portion of Kansas south of the Kansas River. It is common in the Neosho and Marais des Cygnes rivers. It has not been recorded Unionid Mussels in Kansas 115 from the Verdigris River; however it probably occurs in the lower stretches of that stream. PLATE 30 TrunciUa donaciformis (Lea), no. 10508; from Neosho River, 1 mi. S Ot- tumwa, CofFey Co., Kansas. Fig. 1, interior, right valve; fig. 2, interior, left valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 1% inches. 39 38 Scoi* '' 1 r r"'"' 100 97 Museum of Natural Hltrory Universil)! of Konso* 1959 39 36 Fig. 26. Distribution of TrunciUa donaciformis in Kansas. Insert map shows approximate distribution of this species in North America. 116 University of Kansas Publs., Mus. Nat. Hist. The westernmost records for T. donacijormis in Kansas are: Kan- sas River, Topeka, Shawnee County (specimen not seen); Neosho River, Strawn, Coffey County (sec. 33, T. 20 S, R. 14 E). Description of Shell: Shell ovate or elongate-ovate, inequipartite, strong, not solid; anterior end rounded; ventral margin strongly convex, curving up sharply posteriorly; dorsal margin slightly con- vex, curving with sloping posterior end to form more or less sharply biangulate point; beaks full, slightly elevated above dorsal margin; beak sculpture of 3-4 fine bars; posterior ridge well rounded, hav- ing slight posterior depression, no anterior radial furrow; epidermis greenish or yellowish, most shells marked by many dark green rays broken to form definite zigzag or arrow-head lines; growth lines roughened on posterior slope; shell gaps widely anteriorly; male shell usually more sharply pointed posteriorly than female shell. Hinge light; pseudocardinals compressed, slightly divergent, tri- angular, serrated, two in left valve, one in right valve; frequently vestigial tooth anterior to pseudocardinal of right valve; laterals elevated, straight, somewhat lamellar, slightly roughened; beak cavity shallow; muscle scars distinct; nacre silvery-white (after Baker, 1928, p. 229). Ecology: T. donociformis inhabits large rivers in sand or mud bottoms (Baker, 1928, p. 230). In Kansas it seems to occupy a wide variety of habitats from rocky mud, to sand and gravel bot- toms. It is not restricted to the larger streams of Kansas. Host for Glochidia: The glochidia are commonly found on fresh- water drum, Aplodinotus grunniens and rarely found on the sauger, Stizostedion canadense (Surber, 1913, p. 109). Remarks: In Kansas T. donaciformis is easily confused with T. truncata. When present, the definite zigzag marks on the shell of T. donaciformis distinguish it from T. truncata. T. donaciformis is more elongate and less trigonal, and has a rounded, angular pos- terior ridge. T. donaciformis may also be confused with Dysnomia triquctra; the differences between these two species are discussed in the account of D. triquctra. Because of its small size, T. donaciformis was not valued for but- tons. The drying qualities of the shell are excellent. Large speci- mens from Kansas range from 1% to 2 inches in length. Genus Plagiola Rafinesque 1819 Shell solid, surface irregularly, concentrically ridged; epidermis smoothish having larger and smaller scattered rays in irregular lu- nate or squarish blotches; hinge heavy and strong; hinge plate wide Unionid Mussels in Kansas 117 and flat; shell smaller in female than in male, more inflated and swollen at post-basal region (after Walker, 1918, p. 69). Type: Unio scciiris Lea (=: Ohliquaria lincolafa Raf. ). Butterfly Mussel Plagiola lineolata (Rafinesque) Plate 31, figures 1-4; text figure 27 Ohliquaria (Plagiola) lineolata Rafinesque, 1820, Ann. Gen. Sci. Phys., Bruxelles, 5:303. Plagiola lineolata, Agassiz, 1852, Arch, fiir Nat., 1:48. Plagiola securis, Scammon, 1906, Univ. Kansas Sci. Bull., 3:307; Simpson, 1914, A descriptive catalogue . . . mussels, p. 304. Unio securis. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:94; Call, 1886, op. cit., p. 178. Type Locality: Falls of the Ohio at Louisville, Kentucky. General Distribution: Mississippi River drainage from western Pennsylvania west to Iowa, north to Minnesota, south to Kansas and Oklahoma; Alabama ( Baker, 1928, p. 233 ) . Occurrence in Kansas: Plagiola lineolata occurs in the southern portion of the Neosho River drainage. Scammon ( 1906, p. 308 ) reported it from the Spring and Verdigris rivers. We have not seen specimens from these rivers, but there is no reason to doubt its oc- currence there. According to Utterback ( 1916a, p. 151 ) P. lineolata is common in the Osage River in Missouri, but there are no records of its occurrence in the same river ( Marais des Cygnes ) in Kansas. The westernmost records for P. lineolata in Kansas are: Neosho River, 2)2 mi. S and VA mi. W Tola, Allen County (sec. 9, T. 23 S, R. 18 E ) ; Fall River, Wilson County ( specimen not seen ) . Description of Shell: Because this is a monotypic genus, the generic description applies to this species. Ecology: The butterfly mussel is found in large rivers, where it usually inhabits areas of sand and gravel. In Kansas this species does not occur in a substrate of cobbles larger than 3 inches in diameter. Host for Glochidia: The glochidia are found on the gills of Ap- lodinotus grunniens, the freshwater drum (Baker, 1928, p. 232). Remarks: The dorsal ridge of the shell of the female is not so sharp as that of the male. The umbones of both sexes are greatly flattened, and the umbo cavities are correspondingly shallow; con- sidering the thickness of the shell, this is the most conspicuously compressed unionid in Kansas. 118 University of Kansas Publs., Mus. Nat. Hist. PLATE 31 Plagiola lineolata (Rafinesque) no. 10606; from Neosho River, at a place IJa mi. E and 1 mi. S St. Paul, Neosho Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, riglit valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, SJs inches. Unionid Mussels in Kansas 119 38 Scole K wuitrt I I Museum of Nafurol HItlorir univerfitf of Koneos 1959 39 38 97 Fig. 27. Distribution of Plagiola lineolata in Kansas. Insert map shows appro.ximate distribution of this species in North America. Shells of specimens from Kansas vary from a greenish-yellow to a light brown. The posterior, dorsal, and anterior areas of some are black; shells of young individuals are paler and lack the black areas. Although Scammon (1906, p. 308) reported this to be a common species, the authors have found it only rarely. The heavy siltation of the southern drainages in recent years may account for its decline in southeastern Kansas. The shell was valued for buttons. The drying qualities of the shell are good, but small cracks may appear in the ventral portion of the shell. The nacre is usually a beautiful white, but yellowing of the pseudocardinal and lateral teeth does occur. Large speci- mens from Kansas range from 3 to 4/2 inches in length. Genus Leptodea Rafinesque 1820 Shell large, thin, elliptical or slightly obovate, compressed, winged on dorsal margin; beaks low; epidermis rather smooth, often feebly rayed, dull colored, sometimes glossy; pseudocardinal teeth com- pressed, feebly or more often imperfectly developed; nacre purplish; shells of male and female nearly alike, that of female scarcely swollen at post-basal region (after Walker, 1918, p. 72). Type Uriio gracilis Barnes. 120 University of Kansas Publs., Mus. Nat. Hist. Fragile-Paper Mussel Leptodea fragilis (Rafinesque) Plate 32, figures 1-4; text figure 28 Unio (Leptodea) fragilis Rafinesque, 1820, Ann. Gen. Sci. Phys., Bruxelles, 5:295. Leptodea fragilis, Ortmann and Walker, 1922, Occas. Papers Mus. Zool. Univ. Michigan, no. 112, p. 53; Leonard and Leonard, 1946, Univ. Kansas Sci. Bull., 31:118; Franzen and Leonard, 1943, Univ. Kansas Sci. Bull., 29:386. Utiio gracilis. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 96; Call, 1885, op. cit., p. 120; Call, 1886, op. cit., p. 181; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:21. Latnpsilis gracilis, Scammon, 1906, Univ. Kansas Sci. Bull., 3:302. Leptodea gracilis, Simpson, 1914, A descriptive catalogue . . . mussels, p. 181. Ttjpe Locality: Ohio River. General Distribution: Throughout the Mississippi River drainage from eastern New York west to Kansas and Iowa, south to Texas, Mississippi and Alabama, and north to northern Wisconsin and Minnesota; St. Lawrence River drainage; Red River of the North; and Hudson River. Occurrence in Kansas: Leptodea fragilis is common in the Neosho, Verdigris and Marais des Cygnes rivers. It is not known north of the Kansas River except for the locality reported from Wild Cat Creek, Riley County, Kansas ( Call, 1885, p. 95 ) . This record could have been based on L. laevissima which generally replaces L. fragilis north of the Kansas River. 39 38 Mutcum of Noluror Hiilorr univeniijr ol Koniat 1959 36 100 Fig. 28. Distribution of Leptodea fragilis in Kansas. Insert map shows approximate distribution of this species in North America. Unionid Mussels in Kansas 121 PLATE 32 Leptodea fragilis (Rafinesque), no. 10484; from Neosho River, at a place /2 mi. N and 3 mi. W Neosho Rapids, Lyon Co., Kansas. Fig. 1, interior, right valve; fig. 2, interior, left valve; fig. 8, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 5% inches. 122 University of Kansas Publs., Mus. Nat. Hist. The westernmost records for L. fragilis in Kansas are: 2/2 mi. W Americus, Lyon County ( sec. 9, T. 18 S, R. 10 E ) ; Little Arkansas River, Wichita, Sedgwick County (specimen not seen); Elk River, 3 mi. N and 1 mi. W Elk Falls, Elk County (sec. 28, T. 30 S, R. HE); Silver Creek, Cowley County (specimen not seen). Description of Shell: Shell large, thin, brittle, subovate or sub- elliptical; anterior end rounded; ventral margin convex, curved upward into obliquely truncated posterior margin; dorsal margin straight, having slight wing posteriorly, wing obliterated in older shells; posterior ridge weak; valve compressed posteriorly; beaks only slightly elevated above dorsal line; beak sculpture of 3-4 faint, concentric, doubly looped bars, only posterior loop distinct; epi- dermis smooth, marked by concentric growth lines at rest periods; epidermis dull, yellowish, pale greenish-yellow or light brown; rayed or rayless, rays if present ill-defined, straight, wide or narrow; shell of female expanded at post-basal region. Pseudocardinals small, compressed, parallel to hinge line, more or less lamellar; two in left valve, one in right valve; laterals long, thin, compressed, lower in left valve feebly developed; no inter- dentum; beak cavity shallow; dorsal muscle scars form oblique row in beak cavity; anterior scars well impressed; posterior scars less so; nacre silvery-white having pinkish tint in posterior part, or all pinkish (after Baker, 1928, p. 234). Ecology: In Kansas L. fragilis inhabits deep and shallow streams. It occurs also in sand, mud, and rocky bottoms but usually in quiet pondlike areas of a stream. Host for Glochidia: Not recorded. Remarks: In Kansas L. fragilis may be confused with L. laevis- sima. The latter has a thinner shell, a richer and darker purple nacre, a shining greenish-yellow epidermis, and a more prominent dorsoposterior wing. L. fragilis is replaced by L. laevissima north of the Kansas River. The thin shell was never valued for buttons. Large cracks may appear in the shell, and the shell may break into several pieces. Because it lacks a solid hinge line and soHd teeth, there are fre- quently blemishes and mud in the dorsal area of the shell which may obliterate the hinge line area. Huge areas of the nacre may be pushed into folds in an attempt to restrict the mud. The nacre of young animals, of an iridescent pink or light purple, is strikingly attractive. Large specimens from Kansas range from 5)2 to 6 inches in length. Unionid Mussels in Kansas 123 Paper-Shell Mussel Leptodea laevissima (Lea) Plate 33, figures 1-4; text figure 29 Symphynota laevissima Lea, 1830, Trans. Amer. Phil. Soc, 3:444, pi. 13, fig. 23. Leptodea laevissima, Goodrich and van der Schalie, 1944, Anier. Midi. Nat., 32:316. Unio laeviss-imus, Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 120; Call, 1886, op. cit., p. 181; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:21. Lampsilis laevissima, Scanimon, 1906, Univ. Kansas Sci. Bull., 3:303; Simp- son, 1914, A descriptive catalogue . . . mussels, p. 183. Tijpe Locolity: Ohio. General Distribution: Throughout the Mississippi River drainage from western New York west to eastern Texas and central Kansas, from Minnesota south to Louisiana. Occurrence in Kansas: L. laevissima is common in the tributaries of the Kansas River; it occurs also in the Marais des Cygnes and Neosho rivers, but it is never common there. Its range includes the eastern half of Kansas. Records of localities of occurrence of L. laevissima indicate a more westerly distribution of this species than the authors have been able to confirm; westernmost localities include: Blue and Smoky Hill rivers (specimens not seen), Dickinson County; Little Arkansas River, Wichita, Sedgwick County (specimen not seen). Based on our studies, however, westernmost localities of occurrence of L. laevissima include: Delaware River, 4 mi. N Whiting, Jackson County (sec. 10, T. 5 S, R. 16 E); Neosho River, 2 mi. W and M mi. N Neosho Rapids, Lyon County (sec. 24, T. 19 S, R. 12 E). Description of Shell: Shell elliptical, thin, compressed ventrally; slightly inflated at umbones; high triangular, thin, small anterior wing; valves gape at each end; beaks compressed, flattened, not raised above hinge line; beak sculpture of several small, nodulous broken ridges; anterior and posterior ends rounded; ventral margin convex, dorsal margin straight, exclusive of wings; posterior ridge rounded; surface having irregular growth lines raised into ridges at rest period; epidermis olive-green or yellowish-green, yellowish near beaks, polished. Pseudocardinals feeble, thin, lamellar, diagonal to dorsal margin, slightly roughened, one or two in each valve; laterals well marked, long, thin, elevated; no interdentum; beak cavities shallow, having row of dorsal muscle scars; anterior and posterior muscle scars 124 University of Kansas Publs., Mus. Nat. Hist. PLATE 33 Leptodea laevissima (Lea), no. 10358; from Delaware River, at a place 2 mi. W and 2% mi. N Perry, Jefferson Co., Kansas. Fig. 1, interior, right valve; fig. 2, interior, left valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 6/4 inches. TJnionid Mussels in Kansas 125 39 38 39 luicum of Noturot Hlilory Univariitf of Konioi 1959 38 100 Fig. 29. Distribution of Leptodea laevissima in Kansas. Insert map shows the approximate distribution of this species in North America. lightly impressed; nacre rich mauve, lighter in young shells (after Baker, 1928, p. 247). Ecology: L. laevissima inhabits sand and mud bottoms of large rivers having good current (Baker, 1928, p. 248). In Kansas it is most common in areas of soft mud mixed with some sand, but occa- sionally occurs in rocky bottoms. It is not restricted to water with strong current, for the senior author has obtained numerous speci- mens from quiet pools. Host for Glocliidia: The glochidia of L. laevissima have been recovered from the gills of freshwater drum, Aplodinotiis grunniens and white crappie, Potnoxis annularis (Surber, 1913, p. 107). Remarks: In Kansas L. laevissima may be confused with L. fragilis. The differences between these two species are discussed under L. fragilis. L. laevissima from the northern drainages in Kansas have a more highly polished epidermis than those from the southern drainages. It is not clear why L. laevissima replaces L. fragilis north of the Kansas River; L. laevissima may have a higher tolerance to the larger amounts of sand present in the tributaries of the Kansas River. The highly polished, green epidermis, the beautiful mauve nacre, and the fragile nature of the shell make this one of the most beautiful of Kansas unionids. Unfortunately, because of its thinness, large 126 University of Kansas Publs., Mus. Nat. Hist. cracks usually appear in the shell when dried; consequendy shells were never valued for buttons. Large specimens from Kansas range from 6 to 6/2 inches in length. Genus Proptera Rafinesque 1819 Shell usually large, gaping at anterior end and at edge of dorsal slope; winged along dorsal region when young, often when adult; beak sculpture of an anterior and posterior loop, the former often wanting; epidermis generally brown, clothlike, rayless or feebly rayed; pseudocardinals frequently imperfect or nearly wanting; laterals remote (after Walker, 1918, p. 71). Type: Unio alatus Say. Pink Heel-Splitter Mussel Proptera alata (Say) Plate 34, figures 1-4; text figure 30 Unio alatus Say, 1817, Nich. Encyc, 2: (no pagination), pi. 4, fig. 2; Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 119; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:19. Proptera alata, Ortmann, 1912, Ann. Carnegie Mus., 8:333; Franzen and Leonard, 1943, Univ. Kansas Sci. Bull., 29:385. Lampsilis alata, Scammon, 1906, Univ. Kansas Sei. Bull., 3:299; Simpson, 1914, A descriptive catalogue . . . mussels, p. 162. Type Locolify: Lake Erie. General Distribution: Entire Mississippi River drainage from eastern New York west to eastern Kansas, north to Wisconsin, south to Tennessee and Arkansas; Red River of the North; Ottawa, Canada (LaRocque, 1953, p. 97). Occurrence in Kansas: Proptera alata occurs in the eastern one- half of Kansas as far north as the Kansas River. Specimens north of the Kansas River are unknown. Scammon (1906, p. 300) reported that P. alata occurred in Fall River; however, there have been no records from the Verdigris drainage since 1906. The westernmost records for P. alata in Kansas are: Kansas River, Topeka, Shawnee County ( specimen not seen ) ; Neosho River, % mi. N and 3 mi. W Neosho Rapids, Lyon County ( sec. 24, T. 19 S, R. 12 E). Description of Shell: Shell thick, heavy, subovate, roughly tri- angular; anterior end somewhat pointed; ventral margin slightly convex; dorsal margin straight, strongly alate posteriorly; posterior margin obliquely truncated, meeting dorsal wing at sharp angle; alate border slightly concave; posterior ridge rounded, having one or two elevated lines; valves more or less compressed; beaks slightly Unionid Mussels in Kansas 127 elevated above hinge line; beak sculptured by 3-4 fine bars; epi- dermis dark, brownish-green in young specimens to brownish or black in older specimens; young specimens frequently rayed; rest periods distinctly marked; shell in female slightly more broadly rounded in post-basal region than in male. PLATE 34 Proptera alata (Say), no. 10396, 9; from Marais des Cygnes River, at a place 2 mi. E and lli mi. S Pomona, Franklin Co., Kansas. Fig. 1, interior, right valve; fig. 2, interior, left valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 4/2 inches. 128 University of Kansas Publs., Mus. Nat. Hist. 39 39 38 Seolt 10 40Mir«i ■ I I Muiaum of Natural Hlttory Unlvartltjr of Konioi 1959 38 100 97 Fig. 30. Distribution of Proptera alata in Kansas. Insert map shows approxi- mate distribution of this species in North America. Pseudocardinals strong, flatly triangular, subparallel to hinge line, serrated, two in left valve, one in right valve, sometimes second tooth anterior in right valve; laterals long, almost straight, thin or lamellar; no interdentum; beak cavity shallow, having dorsal muscle scars in straight row; anterior and posterior muscle scars well marked; nacre light pink to purple, rose or salmon, iridescent posteriorly (after Baker, 1928, p. 242). Ecology: In Kansas P. alata is found in the larger streams in hard or soft mud, gravel, or rocky bottoms. It may be found in swift or still water but rarely in stagnant water. Host for GJochidia: Not recorded. Remarks: P. alata may be confused with P. ptirptirata. The former has a higher wing, lighter and less swollen shell, and is almost square behind; P. purptirata is more rounded. In P. alata no more than one-third of the row of dorsal muscle scars is situated in the umbonal cavity. In P. purpurata more than one-third of the row of dorsal muscle scars is situated within the umbonal cavity. P. alata may also be confused with Lasmigona complanata. The differences between P. alata and Lasmigona complanata are discussed in the account of the latter. Scammon ( 1906, p. 300 ) reported P. alata as common in the Kan- sas River drainages and scarce in the Marais des Cygnes and Neosho Unionid Mussels in Kansas 129 rivers. Recent studies indicate that it is extremely abundant in the Marais des Cygnes River, common in the Neosho River and ex- tremely rare in the Kansas River drainage. Because of the beautiful nacre and large size, this species is valued for novelty work. It was not valued for buttons. Large specimens from Kansas range from 7 to 7/2 inches in length. Purple Shell Mussel Proptera purpurata (Lamarck) Plate 35, figures 1-4; text figure 31 Unio purpurata Lamarck, 1819, An. sans Vert., 6:71. Proptera purpurata, Ortmann, 1912, Ann. Carnegie Mus., 8:334; Leonard and Leonard, 1946, Univ. Kansas Sci. Bull., 31:117. Lampsilis purpurata, Scammon, 1906, Univ. Kansas Sci. Bull., 3:300; Simp- son, 1914, A descriptive catalogue . . . mussels, p. 166. Vnio purpuratus. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 96; Call, 1885, op. cit., p. 121; Call, 1886, op. cit., p. 182; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:22. Tijpe Locality: "Africa!!" (Simpson, 1914, p. 167). General Distribution: Southern drainages of the Mississippi River from eastern Texas north to eastern Kansas through southern Mis- souri and western Tennessee to the Alabama River drainage. Occurrence in Kansas: Proptera purpurata is restricted to those streams draining the southeastern portion of Kansas. Call (1885- 1887) and Scammon (1906) reported it from the Kansas and Marais des Cygnes rivers. Either they confused P. alata and P. purpurata or else P. purpurata has been subsequently extirpated in those rivers. Recent studies show it to be absent from the Delaware, Marais des Cygnes, and Wakarusa rivers. It is abundant in the Neosho River. The westernmost records for P. purpitrata in Kansas are: Neosho River, 2/2 mi. W Americus, Lyon County (sec. 9, T. 18 S, R. 10 E); Little Arkansas River, Wichita, Sedgwick County (specimen not seen ) . Descriptioti of Shell: Shell large, somewhat obovate, inflated, solid, heavy; beaks full, high, having faint, corrugated sculpture; scarcely winged anteriorly; low, angular wing posteriorly, having 2-3 low, radiating ridges on posterior slope, surface nearly smooth, having shining, blackish epidermis; ligament large, long, exposed in adults; shell of female wide, rounded, marsupial swelling pos- teriorly, more truncated posteriorly than in male. 5—692 130 University of Kansas Publs., Mus. Nat. Hist. PLATE 35 Propfera purpiirata (Lamarck), no. 10571, 9; from Neosho River, }i mi. S of the dam at Humboldt, Allen Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, right valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 4/8 inches. Unionid Mussels in Kansas 131 Fig. 31. Distribution of Proptcra purpurata in Kansas. Insert map shows approximate distribution of this species in North America. Two, siibcompressed, ragged, elevated pseudocardinals in left and right valves, lower pseudocardinal of right valve the larger; laterals long, high, truncated in right valve; beak cavities deep having row of dorsal muscle scars; anterior muscle scar deep; posterior muscle scars scarcely impressed; nacre rich, dark, purple, iridescent pos- teriorly (after Simpson, 1914, p. 166). Ecology: P. purpurata inhabits streams of deep water having deep mud and fairly quiet pools. Host for Glochidia: Glochidia occur on the gills of Aplodinotus grunniens, the freshwater drum ( Surber, 1915, p. 6 ) . Remarks: In Kansas, P. purpurata can be confused with P. alata. The differences are discussed imder P. alata. P. purpurata replaces P. alata in the southern drainages where the former is extremely common. This is a beautiful shell and usually lacks blemishes on the nacre. Because of its purple color, it was not valued for buttons, but rather for novelties. In a novelty store in Kansas City, Missouri, in 1959, P. purpurata was being sold for 15 cents a half shell! Large Kansas specimens range from 6/2 to 6/4 inches in length. This is approxi- mately }2 inch longer than the largest specimen reported from Kansas by Call (1887, p. 22). 132 University of Kansas Publs., Mus. Nat. Hist. Pocketbook Mussel Proptera capax (Green) Plate 36, figures 1-4; text figure 32 Unio capax Green, 1832, Cab. Nat. Hist., 2:290. Proptera capax, Ortmann, 1914, Nautilus, 28:67. Larnpsilis capax, Simpson, 1914, A descriptive catalogue . . . mussels, p. 47. Type Locality: Falls of St. Anthony, Minnesota. General Distribution: The Mississippi River drainage from south- ern Wisconsin south to northern Arkansas and from eastern Tennes- see west to eastern Kansas. Occurrence in Kansas: One female of Proptera capax was re- covered by Harold Murray on 27 August, 1956 from the Neosho River, 7/2 mi. E and 1 mi. S Emporia, Lyon County, Kansas ( sec. 24, T. 19 S, R. 12 E ). This is the first record of P. capax in Kansas. The previous westernmost records were the St. Francis River, Arkansas, and two almost certainly invalid records from the Elkhorn and the Blue rivers, Nebraska ( Simpson, 1914, p. 47 ) . Description of Shell: Shell greatly inflated, obovate, subsolid, gaping at both ends; anterior end sharply rounded; dorsal margin curved; ventral margin straight, curving sharply into ends; umbones •elevated, greatly inflated; posterior ridge full, rounded, having exca- vated or flattened area posteriorly; shell alate anteriorly; ligament strong, passing forward under beaks forming a long lunule; surface smooth, more or less shining; epidermis yellowish or reddish-brown, more or less smoky, rayless; shell of female slightly more inflated in post-basal region than that of male. Pseudocardinals of left valve more or less double, thin, com- pressed, elevated, sulcate, before beaks; pseudocardinals of right valve double, erect, compressed; all pseudocardinals parallel to hinge line; laterals short, erect, curved, double in left, single in right valve; beak cavities deep, wide; anterior muscle scars well im- pressed; posterior muscle scars faintly impressed; nacre bluish white, pinkish, or salmon tinted (after Baker, 1928, p. 249). Ecology: The single specimen known in the state occurred in slowly moving water 3/2 feet deep in a substrate of cobble, sand, and very little silt. Host for Glochidia: Not recorded. Remarks: The specimen from Kansas has a somewhat darker Unionid Mussels in Kansas 133 PLATE 36 Proptera capax (Green), no. 10486, $; from Neosho River, at a place Ja mi. N and 3 mi. W Neosho Rapids, Lyon Co., Kansas. Fig. 1, interior, right valve; fig. 2, interior, left valve; fig. 3, exterior, left valve; fig. 4, dorsal aspect of shell. Actual length, G^Ae inches. 134 University of Kansas Publs., Mus. Nat. Hist, 39 38 39 Scale Uultum el Neturol Htlter| University of KenMS 1959 38 100 Fig. 32. Known occurrence of Proptera capax in Kansas. Insert map shows approximate distribution of this species in North America. purple nacre, and the shell is somewhat more elongate than typical specimens from the Mississippi River. P. capax is unknown in the area between the Neosho River and the Mississippi River. P. capax was never valued for buttons. The Kansas specimen measures 155 mm. (approximately 6^6 inches) in length, 88 mm. wide, and 101 mm. high. In length this specimen exceeds the pre- viously reported maximum length of P. capax by 12 mm. (Haas, 1941, p. 261). Genus Carunculina (in Baker) Simpson 1898 Shell small, inflated, obovate, rather solid, having thick, dark, ray- less or feebly rayed epidermis; beak sculpture of strong, concentric ridges forming a single rounded loop in front, strongly curved upward behind; female truncated obliquely in post-basal region; pseudocardinals compressed, smooth on inside, reflected upwards, somewhat ragged on edges (after Walker, 1918, p. 76). Type: Unio parvus Barnes. Unionid Mussels in Kansas 135 Lilliput Mussel Carunculina parva (Barnes) Plate 37, figures 1-4; text figure 33 Unio parvus Barnes, 1823, Amer. Jour. Sci., 6:274, pi. 13, fig. 18; Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 96; Call, 1885, op. cit., p. 120; Call, 1887, Bull. Washburn College Lab. Nat. Hist, 2:22. Carunculina parva, Utterback, 1916, Amer. Midi. Nat., 4:164; Franzen and Leonard, 1943, Univ. Kansas Sci. Bull., 29:384. Lampsilis parvus, Scammon, 1906, Univ. Kansas Sci. Bull., 3:298; Simpson, 1914, A descrij^tive catalogue . . . mussels, p. 151. Type Locality: Fox River, Wisconsin. General Distribution: Entire Mississippi River drainage from western New York, west to Iowa, eastern Kansas, and central Texas, north to southern Canada and Michigan, south to Louisiana (Baker, 1928, p. 253). Occurrence in Kansas: Cariinctdi7ia parva is common throughout the eastern one-fourth of Kansas. It has been reported from south- central Kansas where it is extremely rare. The westernmost records for C. parva in Kansas are: Delaware River, 8 mi. W and 3/2 mi. N Horton, Brown County; Big Creek, PLATE 37 Carunculina parva (Barnes), no. 10346, 9 ; from Delaware River, 8 mi. W and 3/2 mi. N Horton, Brown Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, right valve; fig. 3, dorsal aspect of shell; fig. 4, exterior, right valve. Actual length, IVi inches. 136 University of Kansas Publs., Mus. Nat. Hist. 38 luseum of Nolural Hlitory University ol Kontot 1959 38 Fig. 33. Distribution of Caruncilina parva in Kansas. Insert map shows ap- proximate distribution of this species in North America. Ellis County (specimen not seen); Turkey Creek, Barber County ( specimen not seen ) . Description of Shell: Shell small, rather solid, subelliptical, some- what inflated; anterior and posterior ends rounded; ventral margin straight or slightly convex; dorsal margin straight, almost parallel to ventral margin; posterior ridge absent; posterior slope somewhat flattened and compressed; beaks slightly elevated above hinge line, somewhat inflated; beak sculpture consisting of 5-6 distinct, sub- concentric bars, last 3-4 bars forming blunt angle posteriorly; epi- dermis unusually thick, usually dark brown or black, rayless, growth lines usually coarse and slightly elevated; shell somewhat more swollen in female than in male, having greatest height at posterior end of shell. Two rough, triangular, compressed, erect, pseudocardinals in left valve, one in right valve; laterals straight, rather strong, erect, rough; no interdentum; beak cavities deep, containing dorsal muscle scars in a diagonal row, pointed anteriorly; anterior adductor muscle scar deeply impressed, posterior muscle scar not distinct; nacre silvery white, more or less iridescent, some specimens having tinge of pinkish-yellow near beak cavities ( after Baker, 1928, p. 251 ) . Ecology: C. parva is found most often in small streams having sluggish current and mud bottom ( Baker, 1928, p. 253 ) . The species Unionid Mussels in Kansas 137 frequently occurs near the banks of large streams in water no more than 2 to 3 inches deep. Host for Glochidia: Not recorded. Remarks: In Kansas C. parva may be confused with small ex- amples of Ligumia siibrostrata. C. parva can be distinguished by its smaller size, its darker brown to black epidermis, its more rounded posterior end, and the presence of prominent half circled ridges on the beak. Sculpturing on the beak of L. siibrostrata consists of numerous inverted Vs. This is the smallest of the Kansas unionids. Because of its small size and fragile shell, it was never used for buttons. In some suitable habitat as many as 100 individuals occur within a radius of a few feet. Large specimens from Kansas range from 1 to l/s inches in length. Genus Ligumia Swainson 1840 Shell oval to oblong; surface smooth; beak sculpture delicate, double looped; shell of female more or less expanded or swollen in the post-basal region (after Walker, 1918, p. 76). Type: Unio recta Lamarck. Black Sand Mussel Ligumia recta latissima ( Rafinesque ) Plate 38, figures 1-4; text figure 34 Unio latissima Rafinesque, 1820, Ann. Gen. Sci. Phys., Bruxelles, 5:297, pi. 30, figs. 14, 15. Ligumia recta latissima, Ortmann and Walker, 1922, Occas. Papers Mus. Zool. Univ. Michigan, no. 112, p. 53. Unio rectus. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 96; Call, 1885, op. cit., p. 120; Call, 1886, op. cit., p. 182; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:22. Lampsilis recta, Scammon, 1906, Univ. Kansas Sci. Bull., 3:294; Simpson, 1914, A descriptive catalogue . . . mussels, p. 95. Type Locality: Ohio River. General Distribution: Throughout the Mississippi River drainage from western New York west to South Dakota and Kansas, north from Minnesota south to Louisiana, Alabama and Georgia; Mani- toba, Ontario, and Quebec, Canada. Occurrence in Kansas: Restricted to the drainages of southeastern Kansas. Although there are no valid records from the Neosho River, Ligumia recta latissima is thought to occur there. Inasmuch as L. r. latissima is extremely rare in Kansas, distribution records 138 University of Kansas Publs., Mus. Nat. Hist. PLATE 38 Ligumia recta latissima (Rafinesque), no. 10442, 9; from Marais des Cygnes River, /i mi. N and 2 mi. W La Cygne, Linn Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, right valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 5/4 inches. Unionid Mussels in Kansas 139 to date for the state probably do not show the actual geographic range. The westernmost valid records for L. r. latissima in Kansas are: Marais des Cygnes River, Franklin County; Elk River, 1 mi. S Howard, Elk County (sec. 12, T. 30 S, R. 10 E). 39 38 39 38 Mtittum of Nolurel History UnivtratiT ot K«fi908 1959 100 Fig. 34. Distribution of Ligumia recta latissima in Kansas. Insert map shows approximate distribution of this subspecies in North America. Description of Shell: Shell large, elongated, subelliptical, in some individuals lanceolate, longer than high, valves flattened; anterior end rounded; ventral margin straight or slightly convex; dorsal margin straight forming a pointed posterior end; posterior slope faintly indicated near beaks, rounded, low below; beaks low, scarcely raised above hinge line, sculptured by 3-5 faint doubly looped bars; epidermis more or less shining brown to black, some- times rayed; shell of female having inflated post-basal region. Pseudocardinals irregular, compressed, serrated, somewhat tri- angular, two in left, one in right valve; laterals long, straight, ele- vated, crenulated; no interdentum; beak cavity shallow, having dorsal muscle scars under hinge line; anterior muscle scars well impressed; posterior muscle scars faintly impressed; nacre usually dark purple, mauve, or white, iridescent posteriorly (after Baker, 1928, p. 255). Ecology: L. r. latissima is typically found in large rivers in gravelly bottoms in strong currents ( Baker, 1928, p. 259 ) . 140 University of Kansas Publs., Mus. Nat. Hist. Host for Glochidio: The glochidia occur on the gills of bluegill, Lepomis macrochirus and white crappie, Pomoxis annularis (Surber, 1913, p. 109; Coker, et al, 1921, p. 153). Remarks: In Kansas L. r. latissima may be confused with Elliptio dilatatus, but in the latter the hinge teeth are not pointed, there is an interdentum, the posterior end is less pointed, and the posterior ridge descends abruptly at the posterior margin. Scammon (1906, p. 295) reported that this species was common in the Marais des Cygnes River and rivers of more southern drain- age. Recent studies in these same areas indicate that it is now extremely rare. The increased turbidity of the water in recent years may account for the decline. The purple nacre of L. r. latissima is the most beautiful of that of all Kansas unionids. The drying qualities of the shell are excellent. The shell of L. r. latissima was rarely used for buttons; only rel- atively solid shells having white nacre were utilized. Large speci- mens from Kansas range from 5/2 to 6 inches in length. Common Pond-Mussel Ligumia subrostrata (Say) Plate 39, figures 1-4; text figure 35 Unio suhrostratus Say, 1831, New Harmony Disseminator, Jan. 15 (no pagination); Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:94; Call, 1885, op. cit., p. 122; Call, 1886, op. cit., p. 182; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:23. Ligumia subrostrata, Crier and Mueller, 1922, Nautilus, 36:100; Franzen and Leonard, 1943, Univ. Kansas Sci. Bull., 29:382; Leonard and Leonard, 1946, Univ. Kansas Sci. Bull., 31:117. Unio topekaensis. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50. Lampsilis subrostrata, Scammon, 1906, Univ. Kansas Sci. Bull., 3:295; Simp- son, 1914, A descriptive catalogue . . . mussels, p. 99. Type Locality: Wabash River. General Distribution: Entire Mississippi River drainage as far north as Fountain City, Wisconsin, and eastern South Dakota ( Simp- son, 1914, p. 99; Grier and Mueller, 1922, p. 100; Over, 1942, p. 5). Occurrence in Kansas: Ligtijnia subrostrata occurs in the eastern one-half of Kansas. The absence of records of occurrence from the Republican and Blue rivers in northern Kansas is probably related to inadequate collecting in those streams. The westernmost records for L. subrostrata in Kansas are: Dela- ware River, 8 mi. W and 3M mi. N Horton, Brown County (sec. 27, T. 17 S, R. 13 E); Marais des Cygnes River, 2'A mi. W and S'A mi. N Unionid Mussels in Kansas 141 PLATE 39 Ligumia suhrostrata (Say), no. 10652, i ; from Potter Lake, University of Kansas Campus, Lawrence, Douglas Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, right valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 2/4 inches. 142 University of Kansas Publs., Mus. Nat. Hist. Miller, Lyon County (sec. 1, T. 16 S, R. 12 E); Neosho River, 3M mi. W and 2'A mi. N Americus, Lyon County ( sec. 29, T. 17 S, R. 10 E ) ; Whitewater River, /2 mi. W Towanda, Butler County ( sec. 8, T. 26 S, R. 4 E); Little Arkansas River, Wichita, Sedgwick County (speci- men not seen ) . 38 39 lulfum of Natural Hiilorf University ot Kontat 1959 38 100 97 Fig. 35. Distribution of Ligumia subrostrata in Kansas. Insert map shows approximate distribution of this species in North America. Description of Shell: Shell elongate, irregularly elliptical, sub- solid, somewhat inflated; beaks full, having numerous delicate ridges drawn up in middle; slightly winged; rounded anteriorly; pointed posteriorly; posterior ridge moderately developed; surface dull, dirty greenish-yellow, generally having faint, wide rays on posterior por- tion, often having concentric bands of lighter and darker color; shell of female narrowed in front, having large, rounded post-basal swelling; base line incurved anterior to swelling; dorsal and ventral lines of shell of male nearly parallel forming sharp point posteriorly. Pseudocardinal teeth compressed, two in each valve, upper in right valve smaller; one, lamellar lateral in right valve, two in left valve; posterior and anterior muscle scars shallow; beak cavities moderate; nacre bluish-white, rarely thicker anteriorly (after Simp- son, 1914, p. 99). Ecology: L. subrostrata occurs abundantly in ponds and creeks in soft to hard mud. It is rarely found in rocky bottoms. In the larger streams it may be abundant locally, but always in shallow Unionid Mussels in Kansas 143 pondlike areas of the river. In June, 1958, the one-acre Potter Lake on the University of Kansas Campus, Lawrence, Douglas Co., Kan- sas, was drained. 2,178 living unionids representing 4 species were removed from the pond. L. suhrostrata accounted for 1,517 (ap- proximately 70 per cent) of the living specimens (Murray, 1960). Host for Glochidia: Not recorded. Remarks: L. suhrostrata may be confused with immature Unio- meriis tetralasmus, several species of the genus Lampsilis, and the adults of CariinciiUna parva. L. suhrostrata may be readily dis- tinguished from these species by the presence of numerous ridges on the umbones in the shape of V's having the open ends spread widely apart. Because of its small and fragile shell, L. suhrostrata was not valued for buttons. Large specimens from Kansas range from 3 to 3)1 inches in length. Genus Lampsilis Rafinesque 1820 Shell oval to elliptical, smooth or slightly, concentrically sculp- tured, usually without posterior ridge; epidermis generally smooth, shining, often rayed; beak sculpture of double-looped, parallel ridges, having posterior loop open behind or sculpture obsolete; hinge having one or two pseudocardinals; only lateral in right valve; two pseudocardinals, two laterals in left valve; female shell having strong inflation of shell, dilation in post-basal region, forming dis- tinct posterior truncation of shell ( after Walker, 1918, p. 78 ) . Type: Unio ovatus Say. Yellow Sand-Shell Mussel Lampsilis anodontoides anodontoides (Lea) Plate 40, figures 1-4; text figure 36 Unio anodontoides Lea, 1834, Trans. Amer. Phil. Soc, 4:81, pi. 8, fig 11- Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885', op. cit., p. 96; Call, 18S5, op. cit., p. 119; Call, 1886, op. cit., p. 180; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:19. Lampsilis anodontoides. Baker, 1898, Bull. Nat. Hist. Surv., Chicago Acad. Sci., Pt. 1, p. 100; Scammon, 1906, Univ. Kansas Sci. Bull., 3:291; Clark and Gillette, 1911, Proc. Biol. Soc. Washington, 24:68; Simpson,' 1914, A descriptive catalogue . . . mussels, p. 90; Franzen and Leonard, 1946, Univ. Kansas Sci. Bull, 31:117. Type Locality: Mississippi, Alabama, and Ohio rivers. General Distrihutiou: Entire Mississippi River drainage north as far as eastern South Dakota (Vermillion River), south as far as northern Mexico; all of the Gulf drainages from the Withlacoochee River, Florida, to the Rio Grande River ( Baker, 1928, p. 267; Over, 144 University of Kansas Publs., Mus. Nat. Hist, PLATE 40 Lampsilis anodontoides anodontoides (Lea), no. 10495, 9; from Neosho River, }» mi. N and 3 mi. W Neosho Rapids, Lyon Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, right valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 4% inches. Unionid Mussels in Kansas 145 1942, p. 5). Distribution includes Lampsilis anodontoides falla- ciosa. Occurrence in Kansas: Lampsilis anodontoides anodontoides in- habits primarily the streams draining southeastern Kansas. It is common in the Marais des Cygnes and Neosho rivers and rare in other southeastern Kansas streams. This species may occur as isolated populations in small streams in various parts of the state. It is occasionally found in the Wakarusa River, Douglas County. The westernmost records for L. a. anodontoides are: Neosho River, 2'A mi. W Americus, Lyon County (sec. 9, T. 18 S, R. 10 E); Ninnescah River, Reno County ( specimen not seen ) . 39 38 MuSSUffl of Nolurqr Hlslory Universllr o( Kansas 1959 39 Fig. 36. Distribution of Lampsilis anodontoides anodontoides in Kansas. Insert map shows approximate distribution of this subspecies in North America. Description of Shell: Shell large, thick, elongated, somewhat in- flated; gaping at both ends; anterior end rounded; posterior end pointed; dorsal and ventral margins straight, nearly parallel; pos- terior ridge rounded, low; beaks full, not greatly elevated above hinge line; epidermis smooth, shining, greenish-yellow or straw- colored, lacking rays; beak region frequendy reddish or brownish; female having conspicuous swelling behind center of venti-al bor- der. Two compressed pseudocardinals in each valve, anterior one elongate, posterior one elevated, serrated; laterals long, lamellar, nearly straight, roughened; no interdentum; beak cavities shallow having dorsal muscle scars; anterior and posterior muscle scars well 146 University of Kansas Publs., Mus. Nat. Hist. marked; nacre silvery-white, sometimes cream-colored or pinkish in beak cavity, iridescent posteriorly (after Baker, 1928, p. 266). Ecology: In Kansas L. a. anodontoides occurs in large and small streams, in slow and fast moving water, in rocky and muddy bot- toms, and in shallow and deep water, but it does not occur in shift- ing sand bottoms. Host for Glochidia: The following fish have been recorded as naturally infected: long-nosed gar, Lepisosteiis osseus; green sun- fish, Lepomis ctjanelhis; orange-spotted sunfish, L. humilis; large- mouthed bass, Micropteriis salmoides; white crappie, Pomoxis an- nularis; black crappie, P. nigromaculatus ( Baker, 1928, p. 267 ) . Remarks: It is not possible to distinguish between L. a. anodon- toides and L. a. fallaciosa in the records by Call ( 1885-1887 ) . Be- cause most of his localities are duplicated in later records at our museum, no attempt has been made to place his localities on the distribution map, figure 36. L. a. anodontoides may be distinguished from L. a. fallaciosa by its shell being somewhat larger, less cylindrical and, having a yellow- ish, usually rayless epidermis. As would be expected with inter- grades, there are examples in the eastern portion of the Marais des Cygnes River and southern portion of the Neosho River that are difficult to assign to one or the other subspecies. L. a. aiiodontoides may be confused with L. radiata siliqtioidea, but the latter is more rounded posteriorly and the shell is wider and heavier. The beautiful smooth nacre and even thickness of the shell make this the most prized of unionids for buttons. The drying qualities of the shell are excellent. Older specimens frequently have the periostracum eroded from the umbonal area. Large specimens from Kansas range from 4/2 to 5 inches in length. Slough Sand Mussel Lampsilis anodontoides fallaciosa Smith Plate 41, figures 1-2; text figure 37 Lampsilis fallaciosus Smith, 1899, Bull. U. S. Fish Comm., p. 291, pi. 79 (no description); Simpson, 1900, Proc. Acad. Nat. Sci., Philadelphia, p. 74, pi. 2, fig. .5; Scammon, 1906, Univ. Kansas Sci. Bull., 3:293; Simp- son, 1914, A descriptive catalogue . . . mussels, p. 92. Lampsilis anodontoides fallaciosa, Grier and Mueller, 1926, Bull. Wagner Free Inst. Sci. Philadelphia, 1(23): 18. Type Locality: Not given. General Distribution: Simpson (1914, p. 92) states that Lampsilis anodontoides fallaciosa occurs in the "upper Mississippi drainage, Unionid Mussels in Kansas 147 south to the Cumberland River, Tennessee and to Arkansas; Red River of the North?" Inasmuch as La Rocque (1953) does not record L. a. faUaciosa from the Red River of the North, it seems unhkely that the questionable locahty of Simpson is vaHd. Occurrence in Kansas: L. a. faUaciosa may have a wider range in Kansas than the distribution map (figure 37) indicates. Because L. a. faUaciosa is often confused with L. a. anodontoides, it is not always possible to distinguish one from the other in the literature. Validated occurrences indicate a range in the extreme eastern por- tion of the Kansas and Marais des Cygnes rivers and the extreme southern portion of the Neosho River, but L. a. faUaciosa probably occurs elsewhere in the state. PLATE 41 Lampsilis anodontoides faUaciosa Smith, no. 10647, 9 ; from Neosho River, Yi mi. E Chetopa, Labette Co., Kansas. Fig. 1, exterior, right valve; fig. 2, dorsal aspect of shell. Actual length, 3/4 inches. 148 University of Kansas Publs., Mus. Nat. Hist. The westernmost records for L. a. fallaciosa in Kansas are: Waka- rusa River, Douglas County; Marais des Cygnes River, SM mi. E Osawatomie, Miami County (sec. 10, T. 18 S, R. 23 E); Neosho River, % mi. E Chetopa, Labette County (sec. 35, T. 34 S, R. 21 E). 39- 38 39 Museum of Nolurol Hiitory Uni«*rsil)r of Kansas 1959 100 Fig. 37. Distribution of Lampsilis anodontoides fallaciosa in Kansas. Insert map shows approximate distribution of this subspecies in North America. Description of Shell: Shell as in L. a. anodontoides except more elongate and more cylindrical; epidermis of L. a. fallaciosa smooth- ish, greenish-yellow and having dark green rays covering greater part of shell; young specimens bright greenish-yellow having grass- green rays (rays absent in L. a. anodontoides) . Ecology: According to Baker (1928, p. 269) this subspecies typically is found in muddy sloughs or pondlike areas of streams. This is not true in Kansas, where the subspecies usually occurs in rocky areas in streams having a moderate current. Host for Glochidia: The glochidia have been observed on the gills of white crappie, Pomoxis annularis; sturgeon, Scaphirhynchtis platorhynchus; short-nosed gar, Lepisosteus platostomiis (Baker, 1928, p! 269). Remarks: Some authors (Baker, 1928; Simpson, 1914) have treated fallaciosa as a species, but recent authors (Goodrich and van der Schalie, 1944; van der Schalie and van der Schalie, 1950) have considered fallaciosa as a subspecies of anodontoides. In Kansas intergradation of fallaciosa and anodontoides occurs Unionid Mussels in Kansas 149 near the eastern and southern borders of the state. Specimens col- lected in those areas are not typical fallaciosa when compared to those of the upper Mississippi drainage. They are less distinctly rayed, and some examples are difficult to distinguish from anodon- toides, as might be expected. Utterback ( 1916a, p. 181 ) reported that anodontoides is supplanted by fallaciosa in the upper Osage River; therefore it is not clear why anodontoides should replace fallaciosa in the upper Marais des Cygnes drainage (Osage River in Missouri). Ecological stress factors may account for this distribu- tion; however, additional data are needed in order to clarify the situation. The distinguishing features and economic value of L. a. fallaciosa are discussed in the account of L. a. anodontoides. Fat Mucket Mussel Lampsilis radiata siliquoidea (Barnes) Plate 42, figures 1-4; text figure 38 Unio siliquoides Barnes, 1823, Amer. Jour. Sci., 6:269, pi. 13, fig. 15. Lampsilis radiata siliquoidea, Clarke and Berge, 1959, Memoir, New York State College Agriculture, no. 365, p. 60. La77}psilis siliquoidea, Franzen and Leonard, 1943, Univ. Kansas Sci. Bull., 29:379; Leonard and Leonard, 1946, Univ. Kansas Sci. Bull., 31:117. Lampsilis luteola, Scammon, 1906, Univ. Kansas Sci. Bull., 3:287; Simpson, 1914, A descriptive catalogue . . . mussels, p. 60. Unio luteolus, Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 96; Call, 1885, op. cit., p. 120; Call, 1886, op. cit., p. 181; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:21. Type Locality: Wisconsin River. General Distribution: Entire Mississippi Valley, from western New York west to North Dakota and Texas, from Louisiana north to Wisconsin and Minnesota; all of Canada east of the Rocky Moun- tains (Ortmann, 1919, p. 289; La Rocque, 1953, p. 93). Occurrence in Kansas: Lampsilis radiata siliquoidea occurs in the eastern one-half of Kansas, except north of the Kansas River. The absence of records north of the Kansas River may be a matter of inadequate sampling. It is common in smaller streams such as the Marais des Cygnes and Verdigris; it is rare in the Neosho River. The westernmost records for L. r. .siliquoidea in Kansas are: Kan- sas River, Manhattan, Riley County; Walnut River, Butler County (specimen not seen); Caney Creek, 1 mi. W Grenola, Elk County (sec. 7, T. 31S, R. 8E). Description of Shell: Shell heavy, large, solid, subelliptical, sub- ovate or ovate; somewhat compressed in young and in males, more 150 University of Kansas Publs., Mus. Nat. Hist. PLATE 42 Lampsilis radiata siliquoidea ( Barnes ) , no. 10373, 9 ; from Marais des Cygnes River, 1 mi. E and 9/4 mi. S Osage City, Osage Co., Kansas. Fig. 1, interior, right valve; fig. 2, interior, left valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 5^ inches. Unionid Mussels in Kansas 151 swollen in old individuals and in females; anterior end rounded; posterior end obliquely truncated, meeting straight dorsal and ven- tral margins in bluntly rounded point; posterior ridge indistinctly rounded; beaks slightly elevated above hinge line, flattened, sculp- tured by 6-10 fine, wavy bars; epidermis yellowish-green to brown- ish, smooth and shining, usually having dark green rays of varying width; growth lines indistinct; female having strong post-basal en- largement, swollen in older specimens. 39 38 -39 Wgsium of Nolurql Hiilorir UnivtrtilD el Kantai 1959 38 100 97 Fig. 38. Distribution of Lampsilis radiata siliquoidea in Kansas. Insert map shows approximate distribution of this subspecies in North America. Pseudocardinals variable, more or less triangular, erect, strong; laterally compressed, roughened, two in left valve, one large, some- times one small anterior tooth in right valve; laterals long, straight or slightly curved, thin; no interdentum; beak cavities shallow, hav- ing dorsal muscle scars; anterior muscle scar excavated; posterior muscle scars not well marked; nacre silvery or milky-white, iri- descent posteriorly (after Baker, 1928, p. 271). Ecology: According to Baker (1928, p. 273) L. r. siliquoidea is typically found in quiet water in mud bottoms. In Kansas it occurs in most types of stream bottoms except shifting sand. Host for Glochidia: Bluegill, Lepomis macrochirus; yellow perch, Perca favescens; and walleye, Stizostedion vitreum are natural hosts for the glochidia. Artificial infections have been successful on the gills of Microp- 152 University of Kansas Publs., Mus. Nat. Hist. tents dolomieui, M. salmoides, Pomoxis annularis, P. nigromaculatiis, StizostecJion canadense (Baker, 1928, p. 273). Remarks: In Kansas, L. r. siliquoidea is usually darker and less frequently rayed than typical specimens from the Mississippi River. Because of its even thickness and fine quality of nacre, L. r. sili- quoidea was highly valued for buttons. The epidermis of older specimens is usually greatly eroded. Large specimens from Kansas range from 5 to 5/2 inches in length. Plain Pocketbook Mussel Lampsilis ovata ventricosa (Barnes) Plate 43, figures 1-4; text figure 39 Unio ventricosus Barnes, 1823, Amer. Jour. Sci., 6:267, pi. 13, fig. 14 (out- line); Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:51. Lampsilis ovata ventricosa, Ortmann, 1919, Mem. Carnegie Mus., 8:301. Lampsilis ventricosa, Scammon, 1906, Univ. Kansas Sci. Bull., 3:285; Simp- son, 1914, A descriptive catalogue . . . mussels, p. 38; Leonard and Leonard, 1946, Univ. Kansas Sci. Bull., 31:117. Unio occidens. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:120; Call, 1887, Bull. Washburn College Lab. Nat. Hist., 2:22. Type Locality: Wisconsin and Mississippi rivers, Prairie du Chien, Wisconsin. General Distribution: Mississippi River drainage from northern Arkansas and Tennessee north to Minnesota and Wisconsin, from New York west to eastern Kansas; St. Lawrence River drainages, and southern drainages of the Hudson Bay ( La Rocque, 1953, p. 93 ) . Occurrence in Kansas: Lampsilis ovata ventricosa is found in the rivers draining the southern portion of Kansas. It is common in the Neosho River but rare in all other rivers in Kansas. The westernmost records for L. o. ventricosa in Kansas are: Kan- sas River, Topeka, Shawnee County (specimen not seen); Neosho River, 2 mi. W Americus, Lyon County; White Water River, To- wanda, Butler County ( specimen not seen ) ; Elk River, VA mi. N Elk Falls, Elk County (sec. 34, T. 30 S, R. 11 E). Description of Shell: Shell large, solid, obovate, inflated, having full, high beaks; beaks having few coarse, irregular corrugations; surface generally smooth, rest periods well marked; epidermis nor- mally shining green, greenish-yellow or brownish having bright green rays; hinge line incurved in front of beaks, outcurved behind; ligament narrow, prominent under beaks, narrow in front of beaks; female having post-basal region greatly swollen. Two, somewhat compressed pseudocardinals in left valve; two Unionid Mussels in Kansas 153 PLATE 43 Lampsilis ovata ventricosa (Barnes), no. 10548, 9; from Neosho River, % mi. E Neosho Falls, Woodson Co., Kansas. Fig. 1, interior, right valve; fig. 2, interior, left valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 4/8 inches. 154 University of Kansas Publs., Mus. Nat. Hist. 39- 3S Muieum of Natural Hiilory Universilf of Kdniai 1959 J9 Fig. 39. Distribution of Lampsilis ovata ventricosa in Kansas. Insert map shows approximate distribution of this subspecies in North America. compressed pseudocardinals in right valve; laterals short, slender, double in left valve, single in right valve; beak cavities deep, wide; muscle scars shallow, smooth, nacre brilliant, silvery, bluish-white, rarely pinkish (after Simpson, 1914, p. 38). Ecology: L. o. ventricosa is typically found in small rivers in sand and gravel bottoms in water a few inches to 4 feet deep (Baker, 1928, p. 284). Kansas specimens more often occur in gravel than in sand bottoms. Host for Glochidio: Glochidia of L. o. ventricosa have been ob- served on the gills of white crappie, Pomoxis annularis and sauger, Stizostedion canadense ( Baker, 1928, p. 284 ) . Remarks: Scammon (1906, p. 286) reported the variety L. ven- tricosa satura Lea, in Kansas. The variety satiira is "greatly in- flated, with livid or smoky-colored, sometimes blackish epidermis, and the marsupial swelling is remarkably developed" (Simpson, 1914, p. 41 ) . Scammon may have had examples of satura; however because he does not illustrate a specimen and because there are no specimens in his Kansas collection that are satura, we are not sure he had this variety from Kansas. The epidermis of a typical L. o. ventricosa is shiny, light to dark green, or greenish-yellow. There are usually bright green rays on the shell, but these fade in some older specimens. Unionid Mussels in Kansas 155 Because typical L. o. venfricosa are known from Missouri ( Utter- back, 1916a, p. 186) and because Kansas examples more closely resemble venfricosa than safura, we have referred Kansas speci- mens to L. o. venfricosa. A more thorough understanding of the trans-Mississippian forms of this species is necessary before the intergradation is adequately understood. Inasmuch as there are intergrades between venfricosa and ovafa (Ortmann, 1919, p. 301 ), it is not proper to elevate venfricosa to the species level as some recent authors have done. The value of the pocketbook mussel was considered low due to the frequent chalky consistency of the shell. The nacre is a beau- tiful white and rarely blemished. The drying qualities of the shell are good. Large specimens from Kansas range from 5/2 to 6 inches in length. Genus Dysnomia Agassiz 1852 Shell rounded, oval or subtriangular, solid, inflated, generally smooth, rayed; beak sculpture delicate, often obsolete, double- looped; female differs from male in having decided inflation of shell in post-basal region ( after Baker, 1928, p. 295 ) . Type: Unio foliatus Hildreth. Snuffbox Mussel Dysnomia triquetra (Rafinesque) Plate 44, figures 1-4; text figure 40 Truncilla triqueter Rafinesque, 1820, Ann. Gen. Sci. Phys., Bruxelles, 5:300, pi. 81, figs. 1-4. Dysnomia triquetra, Ortmann and Walker, 1922, Occas. Papers Mus. Zool. Univ. Michigan, no. 112, p. 65. Truncilla triquetra, Scammon, 1906, Univ. Kansas Sci. Bull., 3:283; Simpson, 1914, A descriptive catalogue . . . mussels, p. 5. Type Locolifij: Falls of the Ohio. General Disiribiifion: Mississippi River drainage from western New York west to Nebraska and eastern Kansas, north to Minnesota, and south to West Virginia, Tennessee, and northern Alabama (Baker, 1928, p. 298). Occurrence in Kansas: Scammon (1906, p. 284) reported Di/s- nomia friquetra from one locality in the Marais des Cygnes River, Ottawa, Franklin County, Kansas. This specimen is no longer available; however, there is one specimen obtained by Scammon on March 20, 1909, from the Wakarusa River. Description of Shell: Shell small, valves thickened, subovate, de- pressed, triangular or somewhat trapezoidal; anterior end rounded; 156 Unr^ersity of Kansas Publs., Mus. Nat. Hist. PLATE 44 Dysnomia triquetm ( Rafinesqiie), no. 3253; from Wakarusa River, near Law- rence, Douglas Co., Kansas. Fig. 1, interior, left valve; fig. 2, interior, right valve; fig. 3, exterior, right valve; fig. 4, dorsal aspect of shell. Actual length, 2 inches. Unionid Mussels in Kansas 157 39 38 tJ= Sc«lc Uuseym of Narural HIslorr university of Kdns09 1959 39 38 100 97 Fig. 40. Distribution of Dysnomia triqitetra in Kansas. Insert map shows approximate distribution of this species in North America. posterior end truncated; ventral margin straight or slightly curved, slightly emarginate posteriorly; dorsal margin short, slightly curved in young specimens, forming an angle with the sharply truncated posterior margin, in older specimens angle bluntly rounded; pos- terior ridge sharply defined; beaks swollen, somewhat elevated above hinge line; beak sculpture consisting of 3-4 faint, more or less double-looped bars; valves swollen but flattened on sides; epidermis yellowish or yellowish-green having many dark green rays covering all but flattened posterior slope behind posterior ridge; rays usually broken into squarish, triangular, or chevron-shaped spots of color; post-basal region somewhat swollen in females. Two strong, high, longer than wide, very ragged pseudocardinals in each valve, upper denticle of right valve smaller; laterals short, strong, serrated, elevated, roughened; no interdentum; cavity of beaks rather deep; adductor scars distinct; dorsal muscle scars in beak cavity; nacre silvery-white ( after Baker, 1928, p. 296 ) . Ecology: Baker ( 1928, p. 298 ) gives the habitat as gravel, stony, and sandy bottoms in shallow water having swift currents. Scam- mon (1906, p. 284) recovered a specimen from a hard mud and rocky bottom in shallow water. The current was not swift. The habitat of the specimen from the Wakarusa River is not known; however, this is typically a stream having a soft mud bottom. 158 University of Kansas Publs., Mus. Nat. Hist. Host for GJochidia: Not recorded. Remarks: Because only two specimens of D. triqtietra are known from Kansas, and because it has not been recovered from Kansas since 1909, it might well be absent from the state today. In Kansas, D. triquetra can be confused only with the closely related TrtinciUa truncata. T. triincata is more nearly triangular having a sharper posterior ridge, and the posterior part of the shell is excavated. The epidermis of D. triquetra is more highly polished and more heavily marked with rays. The only Kansas specimen available measures approximately 2 inches in length. SPECIES IN KANSAS THAT MAY HAVE BEEN EXTIRPATED SINCE 1890 On the bases of descriptions and illustrations by Scammon ( 1906), we think that Anodonta suborhiculata, Ohovaria olivaria, and Pty- chobranchiis fasciolare were validly reported for Kansas. For rea- sons unknown to us, the three species now seem to be absent from the unionid fauna of the state. So far as we know, no specimens of these three species are preserved from Kansas. If the tliree species should occur in Kansas, we hope that the following ac- counts taken in part from Scammon ( 1906 ) will aid in their identifi- cation. Subfamily Anodontinae Genus Anodonta Lamarck, 1799 Heel-Splitter Mussel Anodonta suborhiculata Say Plate 45, figure 3 Anodonta suborhiculata Say, 1831, New Harmony Disseminator (news- paper form, no pagination); Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:96; Call, 1885, op. cit., p. 123; Scammon, 1906, Univ. Kansas Sci. Bull., 3:325; Simpson, 1914, A descriptive catalogue . mussels, p. 400. Type Locality: Ponds near the Wabash River. General Distribution: Southern portion of Mississippi River sys- tem, Nebraska, Iowa, Illinois to Louisiana (Simpson, 1914, p. 400). Description of SJiell: "Shell very large, thin, slightly compressed, suborbiculate in outline. Anterior margin flatly curved; ventral margin almost circular; posterior margin rounded for the first fifth ventrally, and very slightly incurved for the remaining four-fifths; Unionid Mussels in Kansas 159 PLATE 45 Three unionid mussels probably extirpated in Kansas. The dates when last obtained in Kansas are unknown. Fig. 1, Ptychohranchus fasciolare (Rafinesque); fig. 2, Obovaria olivaria (Rafinesque); fig. 3, right valve of Anodonta suborhiculata Say. (Figs. 1, 2, and 3 from Scammon, 1906.) Ap- proximately /2 natural size. 160 University of Kansas Publs., Mus. Nat. Hist. dorsal margin straight or very slightly curved. Umboidal ratio variable, from 0.25 to 0.40. Umbones very low and flat, and orna- mented with four or five pairs of slight nodules arranged in a series which represent degenerate double-looped ridges. Anterior and lateral umboidal slopes flatly curved; posterior umboidal ridge low but distinct; posterior slope slightly excavated. Epidermis straw- yellow to dark brown in color, smooth and shining except for the posterior umboidal slope, which is slightly roughened; fine green rays are sometimes present. Lines of growth fine and continuous. Ligament dark and rather stout. "Interior: Hinge line very thin. Muscle scars large, very faint. Pallial line very faint. Cavity of shell rather large, of beaks very slight. Nacre white, varying to a light salmon, deepest in color near the umbones, very iridescent" ( Scammon, 1906, p. 325 ) . Ecology: Scammon (1906) reported the habitat of Anodonta suborhiculata in Kansas to be quiet water and muddy or somewhat sandy banks. This habitat is reproduced many times in eastern Kansas; however, intensive collecting in areas reported by Call (1885-1886) and Scammon (1906) have yielded no specimens of A. suborhiculata. As best as can be determined from the literature A. suborhiculata occurred in the eastern one-fourth of the state. Host for Glochidia: Not recorded. Remarks: Specimens of A. suborhiculata were obtained by one of us (Leonard) in 1949, but subsequently lost, from a pond east of the Neosho River, 4.5 mi. S Humboldt and 1.5 mi. W highways U. S. 169-59, Allen County, Kansas. These specimens were never catalogued. Murray carefully sampled the area in 1958 and 1959, but was unable to recover examples of A. suborhiculata. Leonard examined the exact location in 1959, and he was unable to recover examples; therefore, this species has probably been extirpated from Kansas. Unionid Mussels in Kansas 161 Subfamily Lampsilinae Genus Obovaria Rafinesque, 1819 Hickory-Nut Mussel Obovaria olivaria (Rafinesque) Plate 45, figure 2 Amblema olivaria Rafinesque, 1820, Ann. Gen. Sci. Phys., Bruxelles, 5:314. Obovaria olivaria, Ortmann and Walker, 1922, Occas. Papers Mus. Zool., Univ. Michigan, no. 112, p. 46. Unio ellipsis, Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50. Unio pealei. Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:50; Call, 1885, op. cit., p. 96; Call, 1886, op. cit., p. 181. Obovaria ellipsis, Scammon, 1906, Univ. Kansas Sci. Bull., 3:305; Simpson, 1914, A descriptive catalogue . . . mussels, p. 299. Type Locality: Ohio. General Distribution: "Upper Mississippi system as far south as the Tennessee and Arkansas River; St. Lawrence drainage area" (Simpson, 1914, p. 299). Description of Shell: Shell of moderate size, short, oval, quite heavy, especially anteriorly, somewhat inflated. Anterior margin rounded; ventral margin decidedly bowed; posterior margin rounded or roundly pointed dorsally; dorsal margin rather curved, and meet- ing the posterior at an angle of 130 to 160 degrees. Umboidal ratio generally small but highly variable; umbones full and swollen, de- curved and ornamented with approximately seven slightly double- looped ridges. Anterior umboidal slope abruptly rounded; posterior slope gradually rounded but flattened somewhat near the posterior margin in the female. Epidermis smooth, often clothlike, with occasional dark continuous lines of growth, in color from dark horn to honey-brown; young specimens a sage-green and marked with numerous narrow rays of dull green, which are obscurely present in shells of many adults. Ligament rather long, stout, chestnut brown. Interior: Pseudocardinals heavy, variable, generally high, erect, columnar or pyramidal, posterior left one often long and lamellar, double in left and single in right valve. Laterals of varying length and curvature, coarse and high. Anterior adductor cicatrix of mod- erate size, deeply excavated, set in front of pseudocardinals. Pos- terior scars of moderate size, well impressed, often confluent. Pallial line generally impressed its entire length. Dorsal cicatrices numer- ous round pits in cavity of umbones. Cavity of beaks and shell 6—692 162 University of Kansas Publs., Mus. Nat. Hist. moderate. Nacre white, slightly iridescent posteriorly ( After Scam- mon, 1906, pp. 305-306). Ecology: "It is a lover of water of moderate depth and of sandy riverbeds" (Scammon, 1906, p. 306). Although the above habitat for Ohovaria oUvoria is repeated many times in eastern Kansas, the species has not been recovered since approximately 1905. Host for Glochidia: The glochidia naturally occur on the gills of sturgeon, Scapliirhijnchus plotorlnjnchtis (Coker, et al., 1921). Remarks: Inasmuch as O. olivaria once occurred in large numbers in eastern Kansas and inasmuch as no examples have been recovered since approximately 1905, the species is assumed to be extirpated in Kansas. Subfamily Lampsilinae Genus Ptychobranchus Simpson 1900 Kidney-Shell Mussel Ptychobranchus fasciolare (Rafinesque) Plate 45, figure 1 Ohliquaria (Ellipsaria) fasciolaris Rafinesque, 1820, Ann. Gen. Sci. Phys., Bruxelles, 5:303. Ptijchobranchiis fasciolare, Ortniann and Walker, 1922, Occas. Papers Mus. Zool., Univ. Michigan, no. 112, p. 42. Unio phaseohts, Call, 1885, Bull. Washburn College Lab. Nat. Hist., 1:94; Call, 1885, op. cit., p. 121; Call, 1886, op. cit., p. 182. Ptychobranchus phaseohis, Scammon, 1906, Univ. Kansas Sci. Bull., 3:319; Simpson, 1914, A descriptive catalogue . . . mussels, p. 333. Ti/pe Locality: Muskingum River, Ohio. General Distribution: Ohio, Tennessee, and Cumberland river systems; Lower Peninsula of Michigan; Kansas; Arkansas; Okla- homa; Louisiana ( Simpson, 1914, p. 334 ) . Description of Shell: "Shell of moderate to large size, compressed, elongate ellipsoid in the outline, very heavy. Anterior margin very decidedly rounded; ventral margin gently bowed or straight; pos- terior margin roundly pointed, the point being directed ventrally; dorsal margin oblique and curved, generally passing gradually into the posterior margin. Umboidal ratio, about 0.23. Umbones low and compressed, marked with a number of very fine undulating ridges. Umboidal slopes flattened dorsally but decidedly curved marginally. Posterior umboidal ridge prominent but rounded. Epi- dermis light horn color (often with an olive-green cast) to dark chestnut-brown. Rays either fine, dark, and wavy, or broad and Unionid Mussels in Kansas 163 interrupted. Lines of growth numerous, coarse, and often imbri- cated. Lunule large and elongate. Ligament short and stout. "Interior: Pseudocardinals small, low serrate, bluntly pyramidal, single in the right and double in the left valve. Laterals short, heavy, slightly curved, oblique. Interdentum quite long, smooth, broad. Anterior adductor cicatrix deeply pitted, elongate, placed in front of the pseudocardinals. Posterior scars deeply impressed and distinct, the retractor almost hidden, and placed in on the tip of the lateral tooth; adductor scar elongate and pointed posteriorly. Dorsal muscle scar large and well impressed on the lower surface of the interdentum. Pallial line impressed its entire length. Branchial area well impressed, cavity of shell small. Beaks practically without a cavity. Nacre milk-white, slightly iridescent posteriorly" (Scam- mon, 1906, pp. 319-320). Remarks: Simpson ( 1914 ) expressed the opinion that P. fasciolare recorded from the areas of the southwest (Kansas included) was probably P. clintonense Simpson; conversely, Scammon (1906) ex- pressed the opinion that P. clintonense identified from Kansas was probably P. fasciolare. Inasmuch as no specimens of either species are available and inasmuch as the illustration of P. fasciolare by Scammon (1906) is insufficient to distinguish what Scammon had, we cannot determine whether or not Simpson was correct. We have listed P. fasciolare instead of P. clintonense for Kansas because we share Scammon's opinion concerning P. clintonense when he states "This species [P. clintonense] is described by Simpson from the Little Bed river near Clinton, Ark. I [Scammon] have a number of specimens from Spring river at Baxter Springs which will prob- ably fall under this species, but until there is more material in better condition at hand for identification I [Scammon] list this species for the state as doubtful. It is quite close to the preceding one." ( Scam- mon, 1906, p. 321. ) Agricultural practices, as well as lead and zinc mining, in the areas surrounding the Spring Biver in Kansas have polluted and increased the turbidity of this river. If examples of P. fasciolare or P. clintonense have been collected in Kansas since 1890, they are unknown to us. Inasmuch as specimens are unknown since 1890 and inasmuch as clear water streams do not exist in Kansas in 1961, it seems likely that P. fasciolare has been extirpated in Kansas. 164 University of Kansas Publs., Mus. Nat. Hist. c -d O y in > C G O s s o c U t; o a a; 'y y M n < c o a c o ^ a ^"rt 3- ~ y : J U3 M o 4: mi - 5j o 3 P ^ -ii 'T3 3 a 3 >~ O '^ 3 £■ ~ t: -^ '^ 3 5u ^, ^ 33:1^.^0,2."'^ — ^^'i)„"i;co"3o ■2 -2 -2 § 3 3 C 3 T3~'::s^~3~3~3~3 222o~'^^~ S553!33a3 ~53'^3333 kJ kJ -q a: OOOO c« Z -^ S < o Q Z < <: U n a Gi o 3 3 Zoo 00 y ni u OS y J ^ rt OJ 3 J '^ ^ 3 r- *•» ;^ ^ ^ .*.- a. •2 o o b f— 1 oi J-^ C o U •2 Co 3 O O 2 5j 3 CO Co O c o ^..» sr Co w "*« O ^ "+-, .C! a o o o CO M^ lO y a CO 03 J2 05 c ^-- 7 --a .2 ^ c ~, ^^ V 3 C Co K 3 3 Q y TO 3p:5 C8 C/3 3 a a 3 3 Co a a^ . -2 3 Co a 3 ~ a c ■2 8 2 2 5 3 n-i « ,— ^ a ^1^'^ ^3 t-H-I^.^ y C g '-' 3 03 J a « U -2 r^ J 2 J F^ 3 ■ CO 5u ^ 3 a — Co ~ 2 §2 3.2 11 a ti; 2 C V 2 -i ^ 2 a ^ a. 2.t- 53 •+^~3 a 'o Co ^ 3 3 a 3 3 3 C "t: ~3 "3: ~3 ~3 ^ ^ U o 2 o a a _;; _i; _~ _i; ^ ^ S •~ c- s *- ~ s,^ ^ s,»( ^ ^ *w ^ ^ ^ ^ ^ ^ ^ *2 c3 U OJ o a y "3 a CD CM u J CQ ■< 'n _c 2 c75 c >. o 03 ,— , a c« X r— CO o (32 U .2 "s ana Le Lea) Barnes (Lea) (Lea) ntonem X CD 0 I— ( ulata S Lea) a o a 02 sub orb ic ellipsis ( ^ 8 S S .Co 5^ 3 *^ 3.2.2 2 s 2.3 < c^ a i^ I^ ->) Co Qi < a 3 1 0 C/3 a. a 3 3 s e"^ c y 0 C/3 -ts a ^ 2 3 Q 13 3 3 3 a a 3) p 3-.a o O z -J ^ ^ -H(>i u 0 < U ^ > >■ 3 n •4-> 3 CQ O Q Q m Q a a 05 a a 3> CO Zoo 00 « s 03 y .03 J ^ 3 03 ^ ~ -> 3 '^ rt s ^ 3 1^ 3 I— I ^ 3 a h c 3 3 "^-a^il == -g § « 3 a § Si tjc CO 5 Q Cl) ». ■«- .3 o a a a a kD ". y > CO 'o ID a CO n < Zoo 3 00 '3^ y 1 -J 2 I a 2 a J ^(MC0'*''0CDt-0005O' (oq i<>icoTp»ocDi>ooa> I oi CO Unionid Mussels in Kansas 165 ALLOCATION OF SPECIES PREVIOUSLY REPORTED FROM KANSAS Call (1885-1887) and Scammon (1906) reported 12 species that are judged to be invalid records for Kansas (Table 1 ). Recognizing that the streams of the southern drainages pass through states hav- ing a typical southern fauna, early authors tended to identify vari- ants in Kansas species as members of the southern fauna. As an example, both Call and Scamman recorded Quadrula honstonensis typically found in Texas. We judge that this record was based upon a non-pustulose Quadrula pustulosa. It is possible that some of the species listed on Table 1 did at one time occur in Kansas, but on the basis of our present knowledge of the Kansas unionid fauna, the available specimens in Scammon's collections, and the illustra- tions as given by Scammon, it seems unlikely. In Table 2 are listed nine species of doubtful occurrence in Kansas. Because Call and Scammon gave few or no descriptions and no illustrations of these species, it is not possible to determine the validity of their records. Anodonta danielsii has the type locality of Topeka, Kansas; but because recent collections in that area indicated that only A. grandis occurs and because illustrations and descriptions of A. danielsii are judged to be of A. grandis, we have placed A. danielsii in synonymy with A. grandis. Three species are thought to have been extirpated in Kansas. They are listed in Table 3 and were discussed on pages 158-163. Table 4 summarizes the allocation of species reported by Call and Scammon, as well as those covered by this report. Table 4. Number of Genera and Species Recorded by Call (1885-1887), by Scammon (1906), and in This Report. Call Scammon This report (1885-1887) (1906) (1961) Total number of genera recorded .... 3 14 24 Total number of species recorded and presently recognized in Kansas. . . 36 39 41 Species validly recorded but now pre- sumably absent from Kansas 3 3 3 Number of species invalidly recorded 6 10 00 Number of named kinds now in synonymy 12 4 00 Number of species of doubtful occurrence 7 7 00 Total number of species recorded .... 64 63 44 166 University of Kansas Publs., Mus. Nat. Hist. PHYSIOGRAPHY OF KANSAS DRAINAGE SYSTEMS Systems Ancestral to Present Drainage Ancestral drainage systems cannot be deciphered in Kansas pre- vious to the beginning of the PHocene, inasmuch as no record of earHer Tertiary phenomena has been preserved in the western two- thirds of the state, and none but extremely fragmentary evidence exists east of the Flint Hills. West of the Flint Hills, Pliocene sediments were deposited on a mature topography underlain by Cretaceous rocks, in which relatively shallow valleys extended generally eastward or southeastward. East of the Flint Hills, high level chert gravels in a matrix of red clays, undoubtedly derived from prolonged weathering, lie above levels that can be fixed as Pleistocene in age. From the existing fragmentary evidence, it seems that drainage east of the Flint Hills extended in a generally east- ward or southeastward direction. Throughout Pliocene time and through early Pleistocene time, the Flint Hills continued to act as a drainage divide, there being no communication across them ( Frye and Leonard, 1959). The Flint Hills were breached in late Kansan time, and the Smoky Hill-Republican systems were captured by an ancestral Kansas River at or about the time the Kansan ice stood at the latitude of Riley County. If these drainage changes, occurring as they did in the relatively 39 38 -39 ,-38 Sealt ? 1 r r""" Ozork Ploteau' Muiauifi af Natvnl Hlilery unlvartlly ol ttaniot 1959 100 Fig. 41. Map ot the State of Kansas showing major river drainages and major physiographic divisions. Unionid Mussels in Kansas 167 recent geologic past, had any profound effect upon the unionid fauna, evidence is lacking to demonstrate the effects. All streams west of the Flint Hills left the present boundaries of the state flowing in a southeasterly or southerly direction prior to Late Kansan time, but lack of information from Oklahoma and other areas to the south makes it impossible to learn whether or not any of these streams flowed directly into the Gulf of Mexico (Frye and Leonard, 1952). Evidence on this point is further weakened by the absence of a satisfactory fossil record of unionids in Pliocene and early Pleisto- cene sediments. In summary, useful knowledge of ancestral drain- ages in Kansas insofar as they bear upon the unionid fauna, extends no farther into the past than late Kansan time, when, in general, the present-day drainage pattern became established. Present Drainage Systems (Fig. 41) The state of Kansas is typically a plains state that gently slopes from the western border (approximately 4,000 feet) to the eastern border (approximately 700 feet). The state is drained by four major river systems: Kansas-Smoky Hill-Republican, Marais des Cygnes, Neosho and Verdigris, and Arkansas river systems. Each of these systems varies in length and drains several types of soil over its length. Physiographically, Kansas may be divided into two major areas (Schoewe, 1949); the Great Plains and the Central Lowlands. The Marais des Cygnes as well as the Neosho and Verdigris systems are situated within the Central Lowlands. The Marais des Cygnes sys- tem drains the middle eastern portions of the state and has a rocky, mud bottom. This system empties into the lower Missouri River. The Neosho and Verdigris systems are here considered as one sys- tem because they join the Arkansas River immediately across the Kansas-Oklahoma border and because the mussel fauna of the two systems is essentially the same. The Neosho and Verdigris systems and their tributaries drain the southeastern portions of the state. The stream bottom of the lower portions of the systems is composed of mud and rock mixed with gravel. The source of the Verdigris does not drain areas having sandy soil; therefore the bottom is rarely sandy, but composed of silt and rock. The stream bottom of the upper Neosho is frequently sandy, but never shifting sand. The Neosho and Verdigris systems empty into the Arkansas River which eventually joins the lower part of the Mississippi River. The Arkansas River system drains the southern portion of the Great Plains and the extreme western portion of the Central Low- 168 University of Kansas Publs., Mus. Nat. Hist. lands. Because the majority of this system crosses the sandy soils of western Kansas, the stream bottom is primarily of shifting sand. The south-central portion does have some silt coming from the western edge of the Central Lowlands and is, therefore, suitable for some unionids. The Kansas system, the largest and longest system in Kansas, drains the entire northern /i to Yi of the state and flows, in an easterly direction to the Missouri River, over the Great Plains in western Kansas and the Central Lowlands in the eastern )i of Kansas. West of the Central Lowlands, the Kansas system has typically a shifting sand bottom; however within the Central Lowlands drainage, which also includes the Drift Plains of the Kansan ice advances, the stream bottoms become silty and do support several species of unionids. A small, poorly studied, but possibly highly significant drainage system is the Spring River. This will be given little attention be- cause the river traverses only the extreme southeastern portion of the extreme southeastern county of Kansas, Cherokee County, and because the fauna has been poorly sampled. Inasmuch as the stream originates as a clear water spring in southwestern Missouri and empties into the Neosho River in eastern Oklahoma, it might be expected to have an unusual fauna; however the scanty records from this system indicate that the fauna is essentially like that of the lower Neosho and Verdigris systems. In summary, that portion of the state lying west of the Central Lowlands is not suitable for most fresh-water mussels because of the shifting sand bottoms; nevertheless, populations of mussels do occur in isolated habitats. The Central Lowland drainages support large populations and at least 41 kinds of fresh-water mussels. ORIGIN OF KANSAS UNIONID FAUNA Bryant Walker ( 1917 ) proposed that the North American unionid fauna east of the Rocky Mountains originated in Cretaceous times in the swamps, streams, and bayous along the eastern shores of North America. Subsequent changes in the elevation of the con- tinent, and the various Pleistocene ice ages resulted in the periodic isolation of this fauna at different times and in different areas of the continent. These changes resulted in three major areas and three distinct faunal assemblages: 1) the fauna and area east of the Appalachian Mountains; 2) the fauna and area of the Mississippi Valley; 3) the fauna of the Alabama River and its tributaries. A fourth area and fauna occurs west of the Rocky Mountains and probably resulted from migration of prototypes from Siberia. Unionid Mussels in Kansas 169 \MAJOR RIVER SYSTEMS \\ IN KANSAS UNIONIDX \^ AND THE Sf^^C'" \v UNITED AND SUBSPECIES \STATES IN KANSAS \- a 3 O >< (0 «l z <» o to O > 3 a. Q. > o 3 (A o (A Restricted To Mississippi Drainage Restricted To South Of Missouri River Actinonaias carinota carinata Anodonto grandis • • Anodonta imbecilis • • Arcidens confragosus • Corunculina porva • Crenodonto peruviona costofa • Crenodonta peruviano peruviana • CyprogeniQ oberti • • Dysnomio triquetro • Elliptio dilatatus Fusconaia flavo Lompsilis onodontoides onodontoides • Lonrtpsilis anodontoides fallociosa • Lompsilis radlato siliquoideo Lampsllis ovoto ventricoso Losmigona complonato Losmigona costata Leptodeo frogilis Leptodeo loevissima • Ligumia recto latissimo Ligumio subrostroto • • Megaionoios gigonteo Obliquorio refiexa Plogiolo lineoloto • Plethobosus cyphyus • Pleurobemo cordotum catillus • PleurobennQ cordotum coccineum • Pleurobemo cordotum pyromidotum • Proptero oloto • • Proptero copax • Proptero purpuroto • • • Quodruio cylmdrico • Quodrulo metonevro • Quodruio nodulQto • Quodrulo pustuloso • • Quodruio quodrulo • Strophitus rugosus • Tritogonio verrucoso • Truncillo donociformis • Truncillo truncota • • Uniomerus tetralasmus • Total Number 41 22 29 39 20 19 2 Table 5. — Occurrence of Mussels in the Four Major River Systems in Kansas, Indicating Mussels Restricted to the Mississippi Drainage and Mussels Occurring in That Drainage South of the Missouri River. 170 University of Kansas Publs., Mus. Nat. Hist. Baker ( 1928 ) proposed that the North American unionid fauna originated in Texas during the Triassic period. If these early forms used fishes as hosts for the glochidia, migration to other areas of North America ( particularly the eastern and northeastern portions ) quickly ensued. The Mississippi embayment caused many groups of unionids to be isolated on either side of this inland sea. Changes in the earth's crust ultimately resulted in three areas of unionid development and distribution; these centers agree with those pro- posed by Walker ( see above ) , and are those areas recognized today as having distinctive faunal assemblages. On the basis of the relationships of the Kansas fauna to that of the Mississippi River (Table 5) and the known Pleistocene drain- ages, the Kansas unionid fauna is judged to have originated from the Mississippi Valley. Inasmuch as only two of the 41 species (Table 5) recognized in Kansas are restricted to drainages south of the Missouri River, and becavise all of the 41 species occur within the Mississippi River system, we judge that the present day fauna of unionids in Kansas is derived from the Mississippi Valley fauna and is composed of species having ubiquitous distribution in the valley. Although it is expected that some species restricted to southern areas should exist in Kansas, it is surprising that more southern species do not occur in the state; therefore it is assumed that the presence of but two southern species in Kansas reflects the peripheral position of the state with relation to the southern fauna. GENERALIZATIONS AND CONCLUSIONS 1) Twenty-four genera and 41 species and subspecies of mussels occur in Kansas streams. 2 ) All 41 species occur in the eastern % of Kansas, but probably no more than eight occur west of the division between the Central Lowlands and Great Plains. These eight are as follows: Ano- donta grandis, A. imbecilis, Cartinculina parva, Lampsilis anodontoides anodontoides, Lasmigona complanata, Leptodea laevissima, Tritogonia verrucosa, Uniomerus tetralasmus. 3) The kinds and numbers of mussels decrease from the south- eastern to the northwestern and southwestern parts of the state, and few or no mussels occur in the extreme western corners of the state. 4) The Neosho River has the greatest number and most kinds of mussels. Unionid Mussels in Kansas 171 5) The kinds and numbers of mussels present are greatly affected by the types of stream bottoms; the shifting sand bottoms of western Kansas exclude all species of mussels in Kansas. 6) The unionid fauna of Kansas is composed of species found pri- marily throughout the Mississippi Valley; only Cijprogenia aherti and Proptera purpurata are species of typically southern faunal assemblages. 7 ) The Kansas fauna of mussels is derived from the Mississippi Val- ley assemblage and is at the periphery of the western range; in addition, Kansas is at the periphery of the range of the southern fauna. The geographical location of Kansas is unique in relation to mussel distribution, and additional studies are necessary to clarify the numerous specific and subspecific relationships to the main fauna of the Mississippi Valley. 8 ) The decrease in abundance of several species and the extirpation of three species from Kansas since the studies by Call and Scam- mon indicate the necessity for immediate controls on industrial pollution and increasing siltation of Kansas streams. 9) Fresh-water mussels are an important source of food for many mammals and fishes, and are a valuable latent resource of the state of Kansas. REFERENCES Allen, W. R. 1914. The food and feeding habits of freshwater mussels. Biol. Bull., vol. 27, pp. 127-146, 3 pis. Bailey, R. M. 1951. Committee on names of fishes. Trans. Amer. Fish. Soc, vol. 80 (1950), pp. 356-357. 1952. Committee on names of fishes. Trans. Amer. Fish. Soc., vol. 81 (1951), pp. 324-327. 1953. Committee on names of fishes. Trans. Amer. Fish. Soc., vol. 82 (1952), pp. 326-328. 1956. A revised list of the fishes of Iowa, with keys for identification. In Iowa Fish and Fishing, by J. R. Harlan and E. D. Speaker, pp. 327-377, figs. 1-10. Baker, F. C. 1918. The productivity of invertebrate fish food on the bottom of Oneida Lake, with special reference to mollusks. New York State College of Forestry, vol. 18, no. 2, tech. publ. no. 9, 264 pp., 44 figs. 1928. Fresh water Mollusca of Wisconsin, Pt. II, Pelecypoda. Wisconsin Geol. and Nat. Hist. Survey, Bull. 70, 495 pp., 26 pis., 96 figs. Borradaile, L. a., Potts, F. A., Eastham, L. E. S., and Saunders, J. T. 1958. The Invertebrata. Cambridge Univ. Press, 3rd ed., 795 pp., 523 figs. Call. R. E. 1885a. Contributions to a knowledge of the fresh-water Mollusca of Kan- sas, I, fresh-water bivalves. Bull. Washburn College Lab. Nat. Hist., vol. 1, no. 2, pp. 48-51. 172 University of Kansas Publs., Mus. Nat. Hist. 1885b. Contributions to a knowledge of the fresh-water Mollusca of Kan- sas, III, fresh-water bivalves. Bull. Washburn College Lab. Nat. Hist., vol. 1, no. 3, pp. 93-97. 1885c. Contribution to a knowledge of the fresh-water Mollusca of Kansas, IV, Bull. Washburn College Lab. Nat. Hist., vol. 1, no. 4, pp. 115- 123. 1886. Fifth contribution to a knowledge of the fresh-water Mollusca of Kansas. Bull. Washburn College Lab. Nat. Hist., vol. 1, no. 6, pp, 177-183. 1887. SLxth contribution to a knowledge of fresh-water Mollusca of Kan- sas. Bull. Washburn College Lab. Nat. Hist., vol. 2, no. 8, pp. 11- 25. Chute, W. H. 1948. A list of common and scientific names of the better known fishes of the United States and Canada. Amer. Fish. Soc, special publ. no. 1, pp. 1-45. Clark, H. W., and Gillette, G. H. 1911. Some observations made on Little River, near Wichita, Kansas, with reference to the Unionidae. Proc. Biol. Soc. Washington, vol. 24, pp. 63-68. Clench, W. J., and Turner, R. D. 1956. Freshwater mollusks of Alabama, Georgia, and Florida from the Escambia to the Suwannee river. Bull. Florida State Mus., vol. 1, no. 3, pp. 99-239, 9 pis. COKER, R. E. 1919. Fresh-water mussels and mussel industries of the United States. Bull. U. S. Bur. Fish., vol. 36, no. 865, pp. 13-89, 46 pis. CoKER, R. E., Shira, a. F., Clark, H. W., and Howard, A. D. 1921. Natural history and propagation of fresh-water mussels. Bull. U. S. Bur. Fish., vol. 37, no. 893, pp. 77-181, 21 pis. Dundee, D. S. 1953. Formed elements of the blood of certain fresh-water mussels. Trans. Amer. Micros. Soc, vol. 72, no. 3, pp. 254-264. Ellis, M. M., Merrick, A. D., and Ellis, M. D. 1930. The blood of North American fresh-water mussels under normal and adverse conditions. Bull. U. S. Bur. Fish., vol. 46, no. 1097, pp. 509-542, 14 pis. Franzen, D. S., and Leonard, A. B. 1943. The Mollusca of the Wakarusa River Valley. Univ. Kansas Sci. Bull., vol. 29, pt. II, no. 9, pp. 363-439, pis. 28-32, figs. 1-5. Frye, J. C, and Leonard, A. B. 1952. Pleistocene geology of Kansas. Kansas Geol. Survey Bull., no. 99, 230 pp. 19.59 Flint Hills physiography. Field Conf., Kansas Geol. Soc, October, 1959, pp. 79-85. Goodrich, Calvin, and van der Schalie, H. 1944. A revision of the Mollusca of Indiana. Amer. Midi. Nat., vol. 32, no. 2, pp. 257-326. Grier, N. M., and Mueller, J. F. 1922. Notes on the naiad fauna of the upper Mississippi River. II, The naiads of the upper Mississippi drainage. Nautilus, vol. 35, pp. 46-49, 96-103. Haas, F. 1941. Records of large fresh-water mussels. Zool. Series, Field Mus. Nat. Hist. Chicago, vol. 24, no. 24, pp. 259-270. Unionid Mussels in Kansas 173 Howard, A. D. 1914. Experiments in propagation of fresh-water mussels of the Quadrula group. Bull. U. S. Bur. Fish., no. 801, 52 pp., 6 pis. ISLEY, F. B. 1914. Experimental study of the growth and migration of fresh-water mussels. Appendix III, Rep. U. S. Com. Fish., 1913, pp. 1-24, 3 pis., 4 figs. La Rocque, a. 1953. Catalogue of the recent Mollusca of Canada. Bull. Nat. Mus. Canada, no. 129, 377 pp. Lefevre, C, and Curtis, W. C. 1912. Studies on the reproduction and artificial isropagation of fresh- water mussels. Bull. U. S. Bur. Fish., vol. 30, no. 756, pp. 109-201, 17 pis. Lemche, H. 1957. A new living deep-sea moUusk of the Cambro-Devonian class Monoplacophora. Nature, vol. 179, pp. 413-416. Leonard, A. B. 1959. Handbook of gastropods in Kansas. Misc. Publ. Univ. Kansas Mus. Nat. Hist., 224 pp., 11 pis. Leonard, A. E. 1943. The Mollusca of Meade and Clark Counties, Kansas. Trans. Kansas Acad. Sci., vol. 46, pp. 226-240 Leonard, A. B., and Leonard, A. E. 1946. Mollusca from Greenwood County, Kansas. Univ. Kansas Sci. Bull., vol. 31, no. 6, pp. 115-122. Ortmann, a. E. 1912. Notes upon the families and genera of the najades. Mem. Carnegie Mus., vol. 8, pp. 222-365, 20 pis. 1919. A monograph of the naiades of Pennsylvania. Part HL Systematic account of the genera and species. Mem. Carnegie Mus., vol. 8, no. 1, pp. 1-384, 21 pis. Ortmann, A. E., and Walker, B. 1922. On the nomenclature of certain North American naiades. Occas. Papers Mus. Zool., Univ. Michigan, no. 112, 75 pp. Over, W. H. 1942. Mollusca of South Dakota. Nat. Hist. Studies, Uni\-. South Dakota, no. 5, 11 pp. Rafinesque, C. S. 1819. Prodrome de 70 nouveaux genes d'animaux decouverts dans I'in- terieur des Etats-Unis d'Amerique durant I'annee 1818. Jour, de Phys., de Chimie et d'Histoire Nat., vol. 89, pp. 426-427. 1820. Monographic des coquilles bivalves et fluviatiles de la ri\iere Ohio. Ann. Sci. Phys., Bruxelles, vol. 5, pp. 314-315. Scammon, R. E. 1906. The Unionidae of Kansas, part L Univ. Kansas Sci. Bull., vol. 3, pp. 279-373, pis. 52-86. Schoewe, W. H. 1949. The geography of Kansas, part H, physical geography. Trans. Kansas Acad. Sci., vol. 52, pp. 261-333. Simpson, C. T. 1900. Svnopsis of the naiades, or pearly fresh-water mussels. Proc. U. S. Nat. Mus., vol. 22, no. 1205, pp. 501-1044. 1914. A descriptive catalogue of the naiades or pearly fresh-water mussels. Detroit, 1540 pp. 174 University of Kansas Publs., Mus. Nat. Hist. Simpson, G. B. 1884. Anatomy and physiology of Anodonta fluviatilis. New York State Mus. Nat. Hist., pp. 169-191, 11 pis. Stains, H. J. 1956. The raccoon in Kansas. Misc. Publ. Univ. Kansas Mus. Nat. Hist., 76 pp., 4 pis. Storer, T. I., and Usinger, R. L. 1957. General Zoology. 3rd ed., McGraw-Hill Book Co., Inc., New York, pp. 522, 309 figs. Strecker, J. K. 1931. The distribution of the naiades or pearly fresh-water mussels of Texas. Baylor Univ. Mus., Special Bull. no. 2, 69 pp. SURBER, T. 1913. Notes on the natural hosts of fresh-water mussels. Bull. U. S. Bur. Fish., vol. 32, pp. 101-116, pi. 29-31, 1 fig. 1915. Identification of the glochidia of fresh-water mussels. Bull. U. S. Bur. Fish., 10 pp., 1 pi. Utterback, W. I. 1916a. The naiades of Missouri. Amer. Midi. Nat., vol. 4, nos. 1-10, 200 pp., 29 pis. 1916b. Parasitism among Missouri naiades. Amer. Midi. Nat., vol. 4, no. 12, pp. 518-521. VAN DER SCHALIE, HeNRY, and VAN DER SCHALIE, AnNETTE 1950. The mussels of the Mississippi River. Amer. Midi. Nat., vol. 44, no. 2, pp. 448-466. Walker, B. 1917. The method of evolution in the Unionidae. Occas. Papers Mus. Zool., Univ. Michigan, no. 45, 10 pp. 1918. A synopsis of the classification of the fresh-water Mollusca of North America, north of Mexico, and a catalogue of the more recently described species, with notes. Misc. Publ. Univ. Michigan, no. 6, 213 pp. GLOSSARY accessory denticle. — Small nacreous swelling on the hinge line of the bivalve shell, anterior or posterior to the main pseudocardinal teeth. accessory lateral tooth. — Small nacreous, lamellar swelling on the hinge line of the bivalve shell, ventral to the main lateral teeth. adductor muscle. — Muscle that closes the two valves of the shell; the muscle fibers extend from one valve to the opposing valve. ALATE. — Said of shells that have an anterior or posterior winglike projection of the two valves of the shell, extending dorsally above the hinge line. ANGULATE. — Said of shells having either the anterior or posterior margins form- ing a relatively acute angle. ANTERIOR adductor MUSCLE. — The largest of the anterior muscles. See ad- ductor muscle. anterior adductor muscle scar. — The largest of the nacreous impressions at the anterior end of the shell, forming the attachment of the anterior adductor muscle (Plate 2, fig. 5). anterior protractor muscle. — Muscle situated at the anterior end of the animal and aiding in extending the foot; the fibers extend from the visceral mass to the anterior part of the shell ( Plate 3, fig. 3 ) . Unionid Mussels in Kansas 175 ANTERIOR PROTRACTOR MUSCLE SCAR. — A relatively small, nacreous impression in the valve forming the attachment of the anterior protractor muscle (Plate 2, fig. 5). ANTERIOR RETRACTOR MUSCLE. — Musclc situated near the anterior end of the animal, and aiding in retraction of the foot. The fibers extend from the visceral mass to the shell (Plate 3, fig. 3). ANTERIOR RETRACTOR MUSCLE SCAR. — A relatively small, nacreous impression in the valve forming the attachment of the anterior retractor muscle (Plate 2, fig. 5). ANTERIOR SLOPE. — The area across the dorsal portion of the valve extending from the umbo to the anterior margin ( Plate 2, figs. 2 and 3 ) . ANTERIOR WING. — A dorsal projection of the shell anterior to the umbo (Plate 33). ARAGONiTE. — A Crystalline form of calcium carbonate that is one of two main components of the unionid shell ( see calcite ) . ARCUATE. — Said of shells having the anterior or posterior margins curved in the form of a bow. BEAK. See UMBO. BEAK SCULPTURE. — A pattern of ridges or pustules on the dorsalmost portion of the umbo. BRANCHIAL OPENING. — The modified posterior portion of the mantle that forms the incurrent and excurrent siphons through which water and suspended materials enter or leave the animal. CALCITE. — A crystalline form of calcium carbonate that is one of two main components of the unionid shell. It differs in molecular structure from aragonite. CARDINAL TOOTH. — An clevation on the hinge plate of one valve that interlocks with a corresponding depression on the opposing valve: situated directly ventral to the umbo. Not found in unionids ( see pseudocardinal TOOTH ) . CELT-SHAPED. — Said of some unionid glochidia in which the shape resembles an ax or chisel. CHEVRONS. — V-shaped marks or structures on the exterior of unionid shells. CICATRICE. — Scar or impression in the nacre of the unionid valve formed by the attachment of a muscle to the valve. CRENULATE. — Said of shells having a roughened or scalloped border. CRETACEOUS. — A geologic era starting approximately 12,5 million years ago and lasting approximately 55 million years. CRYSTALLINE STYLE. — A secrctcd, rodlike structure situated in a diverticulum of the stomach of some pelecypods; mediates the digestion of carbohy- drates. CTENDiUM. — A comblike respiratory organ or gill. CUSP. — See pseudocardinal tooth. DIOECIOUS. — Said of animals in which the male reproductive organs are in one individual and the female reproductive organs are in another. DORSAL hinge LIGAMENT. — An clastic, elongate, corneous structure that unites the two valves of the shell above the hinge plate ( Plate 2, fig. 5 ) . DORSAL MUSCLES. — Musclcs extending from the mantle into the umbonal cavity; these aid in retraction of the mantle. 176 University of Kansas Publs., Mus. Nat. Hist. DORSAL MUSCLE SCARS. — Impressions in the nacreous portion of the valve, formed by the attachment of the dorsal muscles; situated in the umbonal cavity (Plate 2, fig. 5). DORSOPOSTERIOR SLOPE. — The area across the dorsal portion of the valve ex- tending from the umbo to the posterior margin. EDENTULOUS. — Said of shells lacking both lateral and pseudocardinal teeth. EPIDERMIS. — The most external (and corneous) layer of the shell. See PERIOSTRACUM. EQUiPARTiTE. — Said of shells that have the umbo situated an equal distance from the anterior and from the posterior margins. EXCURRENT SIPHON. — The modified tubelike posterior portion of the mantle that allows the exit of water and wastes from the animal; situated dorsal to the incurrent siphon. FLUTiNGS. — A series of undulations on the dorsoposterior portion of certain unionid shells; example, Lasmigona costata (Plate 20). FOOT. — The ventralmost portion of a pelecypod; in unionid mussels it is essen- tially a locomotor organ ( Plate 3, fig. 1 ) . GAPE. — Said of certain unionid shells that have a noticeable space between the valves, when the two valves are appressed as closely as possible. GLOBOSE. — Said of shells that are nearly spherical or globular in shape. GROWTH LINES. — Indiscrete lines of temporarily arrested growth appearing on the epidennis of the shell. HINGE LINE. — The dorsal area of the shell that forms the pivot upon which the two valves rotate as they open. HINGE PLATE. — The dorsal tooth-bearing surfaces of the two valves. HYPOSTRACUM. — A layer of nacreous material deposited in the muscle scars of most unionids. IMPERFECT. — Said of unionids that do not have well developed lateral or pseudocardinal teeth; example, Strophitus rugosus. INCURRENT SIPHON. — The modified tubular posterior portion of the mantle that allows the entrance of water and food into the animal; situated ventral to the excurrent siphon. INEQUIPARTITE. — Said of shclls that have the umbones situated nearer to the anterior margin than to the posterior margin. INFLATED. — Said of unionid shells that are greatly swollen. INNER LAYER. — That portion of the shell deposited adjacent to the mantle. See NACRE. iNTERDENTUM. — A flattened area of the hinge plate between the pseudocardinal and lateral teeth of some unionid shells. KANSAN. — Applied to the second of a series of glacial stages in the Pleistocene epoch ( Ice Age ) . It may have begun as long ago as 500,000 years. KNOB. — Protuberance or boss (usually exceeding 3 mm. in height) on the surface of a unionid shell ( Plate 2, fig. 1 ) . Knobs usually are large in size and few in number, whereas pustules are usually small and numerous. LABIAL PALPI. — Fingerlike, cihated appendages on either side of the mouth of the unionid; they aid in transferring food to the mouth. LACHRYMOSE. — Said of pustules that are tear-drop in shape, as in Quadrula quadrula (Plate 2, fig. 3). LAMELLA. — A thin plate or layer. Unionid Mussels in Kansas 177 LANCEOLATE. — Said of structures that are narrow and that taper to a point at the apex. LATERAL RIDGE. — An elevated part of the unionid shell near the center of the valve, extending from the umbo to near the ventral margin. LIGAMENT. See DORSAL HINGE LIGAMENT. LATERAL TEETH. — Elongate, raised, interlocking structures on the hinge line of the valve, posterior to the umbo. LUNULE. — Depressed area of some unionid shells immediately anterior to the umbo. MALACOLOGiST. — One who studies mollusks. MANTLE. — Two lobes of tissue extending from the body of a mussel; they envelope the body and are applied to the inner surfaces of the two valves. MARSUPIAL SWELLING. — An enlarged or inflated ventroposterior part of the shell of the female unionid, that provides space for the expansion of the gills while they are carrying glochidia. MARSUPiUM. — In unionids, a brood-pouch for eggs and larvae, formed by one or more gills. MIDDLE LAYER. See PRISMATIC LAYER. MONOTYPic GENUS. — A gcuus that coutaius but one species. NACRE. — The iridescent, inner layer of a unionid shell, composed chiefly of calcium carbonate, and deposited in thin, overlapping lamellae (mother- of-pearl ) . NEPiONic SHELL. — The remains of the embryonic shell at the apex of the umbo. OBSOLETE. — Said of structures that are lacking or indistinctly developed. OPiSTHODETic. — Said of unionids in which the dorsal hinge ligament is posterior to the umbones. OSTIUM. — A small opening; here refers to the entrance of a water tubule in the gills of the unionid. OTOCYST. — An organ used by mussels for balance and orientation in the field of gravity. PALLiAL LINE. — A linear impression approximately parallel to the ventral margin of the valve produced by the attachment of muscles extending from the mantle to the shell. PAPILLA. — Small fingerlike projection on the posterior margin of the unionid mantle. PALLIUM. See MANTLE. PERIOSTRACUM. See EPIDERMIS. PLACENTA. — The gelatinous substance forming an envelope around one or more developing glochidia in a water tubule within the marsupial gill. PLANKTON. — Minute, aquatic plants and animals, having weak powers of loco- motion, that form the principal food of mussels. PLEISTOCENE. — A gcologic pcriod of time estimated to have begun approximately one million years ago and extending into recent times. PLICATIONS. — Parallel ridges on the surface of a unionid shell. PLIOCENE. — The geologic period of several million years duration immediately before the Pleistocene. POST-BASAL AREA. — That area of the shell near the posteroventral margin of the shell. 7—692 178 University of Kansas Publs., Mus. Nat. Hist. POSTERIOR ADDUCTOR MUSCLE. — The largest of the muscles at the posterior part of the unionid body; responsible, in part, for the closure of the valves (Plate 3, fig. 3). POSTERIOR ADDUCTOR MUSCLE SCAR. — A large impressiou in the nacreous layer near the posterior end of the valve, forming the attachment of the pos- terior adductor muscle (Plate 2, fig. 5). POSTERIOR RETRACTOR MUSCLE. — A Small muscle near the posterior end of the unionid animal that aids in retracting the foot of the mussel ( Plate 3, fig. 3). POSTERIOR RETRACTOR MUSCLE SCAR. — The smaller of the two impressions in the nacre near the posterior end of the valve, forming the attachment of the posterior retractor muscle ( Plate 2, fig. 5 ) . POSTERIOR RIDGE. — A low ridge on the shell, extending from the umbo to the posterior margin. POSTERIOR SLOPE. See DORSOPOSTERIOR SLOPE. POSTERIOR WING. — A dorsal extension of the shell above the hinge line, posterior to the umbo. PRISMATIC LAYER. — Tlic calcarcous, prismatically organized part of a unionid shell, situated between the outer corneous periostracum and the inner nacreous layers. PROTOTYPE. — An ( usually hypothetical ) ancestral form from which later groups of animals are derived. PSEUDOCARDINAL TOOTH. — Any One of the triangular hinge teeth near the dorsal margin of the shell; situated ventral to the umbo. PUSTULES. — Small, usually numerous, raised structures on the external part of the shell. RADULA. — A tonguelike organ in the mouth of gastropods having numerous transverse rows of minute teeth; it is lacking in pelecypods. REST PERIODS. — Thin, dark lines in the epidermis of the shell denoting ap- proximately one season's growth ( Plate 2, fig. 1 ) . RETUSE. — Said of shells that have a rounded anterior or posterior margin. RIB. — A raised portion of a unionid shell extending from the umbo to the ventral or ventroposterior margin. See lateral ridge (Plate 2, fig. 3). SCAR. — See cicatrice. SECCHi DISC. — A standard painted disc used for estimating the depth of the penetration of light into water. SECONDARY SEXUAL VARIATION. — Said of uuiouids that have the post-basal region of the shell of females inflated in contrast to the shells of males that are not inflated in the post-basal region. SINUS. — A cavity or hole. A feature of certain unionid shells that have a de- pression above or below the posterior ridge. SOLID. — Said of shells that are thick and heavy. SOLID HINGE LINE. — Said of shells having massive lateral and pseudocardinal teeth. SOLID TEETH. — Lateral and pseudocardinal teeth that are large and massive. SPECIES. — Groups of interbreeding natural populations that are reproductively isolated from other such groups. SUBSPECIES. — A geographically defined aggregate of local populations within a species that differs morphologically and or physiologically from other aggregations of local populations within the species. Unionid Mussels in Kansas 179 suLCATE. — Said of shells that have a deep depression between the two pseudo- cardinal teeth of a valve. suPRABRANCHiAL CHAMBER. — The dorsal portion of the gills of a mussel from which water and wastes are passed to the excurrent siphon. SURFACE SCULPTURE. — The pattern of ridges or pustules on the outer surface of the unionid shell. TERTIARY. — A geologic period of time starting approximately 70 million years ago and terminated by the beginning of the Pleistocene epoch. THERMOCLiNE. — Sudden drop in the temperature at lower depths of water in lakes and ponds. TRANs-MississiPPiAN. — That region of the United States west of the Mississippi River and east of the Rocky Mountains. TRiASSic. — A geologic period of time starting approximately 200 million years ago and lasting approximately 35 million years. TRUNCATE. — Said of shells having the end of the shell more or less at right angles to the dorsal and ventral margins. TUBERCLE. — Small, rounded, raised structure on the outer surface of the unionid shell. TYPE LOCALITY. — The prccisc locality from which a zoological object ( type ) that serves as the reference for the name of a species or subspecies is obtained. UMBO. — The dorsally raised, inflated area of the unionid shell ( Plate 2, fig. 1 ) . UMBONAL CAVITY. — The pit or cavity resulting from the dorsal extension of the umbones. It is situated lateral to the hinge line (Plate 2, fig. 4). vAL\E. — The right or the left half of a unionid shell. VENTRAL FOOD GROOVE. — The Specialized ventral portion of the unionid gill modified to transport food anteriorly to the mouth. WATER TUBES. — Channels in the gills of the unionid for transporting water dorsally through the gill. Transmitted August 25, 1961. INDEX Principal references are in bold-faced characters abbreviated synonomy, 37 aberti, Cyprogenia, 37, 38, 105, 106, 171 aberti, Unio, 105 accessory denticles, 18 Actinonaias, 108 carinata, 19, 28, 37, 108, 109 gibba, 111 adductor muscles, 19 muscle scar, 19 adult stage, 26 aesopus, Pleurobema, 68 Alasmodonta complanata, 87 confragosa, 96, 97 costata, 84, 86 alata, Lampsilis, 126 alata, Proptera, 18, 30, 35, 90, 126, 127 alatus, Unio, 126 Amblema, 45, 46 costata, 45, 49 olivaria, 161 ovalis, 45 Amphineura, 11 Anodon rugosus, 99 Anodonta, 90 arkansensis, 99 danielsii, 92, 165 edentula, 99 grandis, 18, 28, 30, 31, 35, 83, 91, 92, 93, 165, 170 imbecilis, 25, 26, 35, 93, 94, 95, 102, 104, 170 suborbiculata, 93, 158, 159, 160 undulata, 99 Anodontinae, 14 anodontoides, Lampsilis, 30, 31, 143, 144, 170 anodontoides, Unio, 143 anterior slope of shell, 15 anus, 24 Aplodinotus grunniens, 26 aragonite, 15 Arcidens, 96 confragosus, 34, 97, 98 arkansensis, Anodonta, 99 arteries, 25 asperrimus, Unio, 52 auricles, 25 bank conditions, 32 beak cavity, 56 beaks, 14 biological turbidity, 28 black crappie, 48 black sand mussel, 137 blood, 25 bluegill, 62, 111, 151 blue-point mussel, 47 bottom conditions, 29 temperature, 32 bowfin, 45 buckhorn mussel, 67 bullhead mussel, 68 butterfly mussel, 117 buttons, from shells, 9 byssus thread, 26 calcite, 15 calcium carbonate, 29 capax, Lampsilis, 132 capax, Proptera, 18, 35, 132, 133 capax, Unio, 132 carinata, Actinonaias, 19, 28, 37, 108, 109 carinatus, Unio, 108 Carunculina, 134 parva, 29, 36, 135, 137, 143, 170 catillus, Pleurobema cordatum, 37, 74,75 catillus, Unio, 74 Central Lowlands, 167 Cephalopoda, 11 cerebral ganglia, 25 channel catfish, 56 chemotactic sense receptors, 25 Chiton, 11 cicatrice, 18 circulatory system, mussels, 25 clams, 11 clintonense, Ptychobranchus, 163 coccinea, Quadrula, 71 coccineum, Pleurobema cordatum, 37, 42, 71, 72 coccineus, Unio, 71 common pond mussel, 140 complanata, Lasmigona, 30, 31, 33, 35, 87, 88, 90, 128, 170 Margaritana, 87 Symphynota, 87 condition of water, 28 confragosa, Alasmidonta, 96, 97 confragosus, Arcidens, 34, 97, 98 cordatum, Pleurobema, 37, 71, 72, 74, 75, 76, 77 cornutus, Unio, 103 costata, Alasmidonta, 84, 86 Amblema, 45, 49 Crenodonta peruviana, 28, 30, 34, 44, 46, 48, 49, 50 Lasmigona, 33, 85, 86 crassidens, Unio, 78 creek chub, 102 Crenodonta, 26, 29, 45 costata, 28, 30, 46, 48, 49, 50 peruviana, 34, 46, 47, 50 crowfoot hook, 31 (180) Index LSI Cryptochiton, 11 crystalline style, 24 ctenidia, 10 current, effect of, 30 cygneus, Mytiliis, 90 cylindrica, Quadrula, 35, 63, 64 cylindricus, Unio, 64 cyplia, Obliquaria, 68 cyphyus, Plethobasus, 35, 37, 68, 69 Cvprogenia, 105 ■ aberti, 37, 38, 105, 106, 171 danielsii, Anodonta, 92, 165 deer toe mussel. 111 Dentalium, 11 depth of water, effect of, 29 development, mussels, 26 digestive gland, 24 system, 24 dilatata, Unio, 80 dilatatus. Elliptic, 37, 79, 80 distribution maps, 38 donaciformis^ Plagiola, 114 Truncilla, 36, 114, 115, 116 Unio, 114 dorfeuilliana, Unio, 55 dorsal muscle scars, 19 dredge, for collecting, 32 Dysnomia, 155 triquetra, 36, 38, 116, 155, 156, 158 ebenus, Fusconaia, 41, 42 ecology of mussels, 27 economic value of shell, 38 edentula, Anodonta, 99 edentulus, Strophitus, 29, 99 elegans, Plagiola, 111 Unio, 111 ellipsis, Obovaria, 161 Unio, 161 Elliptio, 26, 78 dilatatus, 37, 79, 80 epidermis, 14 esophagus, 24 Eulamellibranchia, 11, 39 excurrent siphon, 25 fallaciosa, Lampsilis anodontoides, 36, 146, 147 fasciolare, Ptvchobranchus, 158, 159, 162, 163 ' fasciolaris, Obliquaria, 162 fat mucket mussel, 149 fawn's foot mussel, 114 Filobranchia, 11 flat-headed catfish, 45 flava, Fusconaia, 37 39, 42 flava, Obliquaria, 39 Flint Hills, 166 floater mussel, 92, 95 fluted mussel, 86 foot, of mussel, 24 fragile paper mussel, 120 fragilis, Leptodea, 29, 31, 35, 120, 121, 122 fragilis, Unio, 120 fragosa, Quadrula, 53 freshwater drum, 116, 125, 131 mussel, 11 Fusconaia, 26, 39 ebenus, 41, 42 flava, 37, 39, 42 undata, 41, 42 Gastropoda, 11 giant washboard mussel, 42 gibba, Actinonaias carinata. 111 gibbosus, Unio, 80 gigantea, Megalonaias, 29, 33, 34, 42,44 giganteus, Unio, 42 gills, of mussels, 24 glochidia, 24 glochidial larva, 10 glochidium, 26 gracilis, Lampsilis, 126 gracilis, Unio, 119, 120 grandis, Anodonta, 18, 28, 30, 31, 35, 83, 91, 92, 93, 165, 170 Great Plains, 167 green sunfish. 111 growth, of mussels, 27 growth lines, on mussel shells, 27 grunniens, Aplodinotus, 26 heel-splitter mussel, 158 heros, Quadrula, 42 lieros, Unio, 42 hickory-nut mussel, 161 hinge, of shell, 14 teeth, 18 houstonensis, Quadrula, 165 hypostracum, 15 Ictalurus punctatus, 26 imbecilis, Anodonta, 25, 26, 35, 93, 94, 95, 102, 104, 170 incurrent siphon, 24 industrial pollution, 28 insert map, 38 interdentum, 18 intestine, of mussel, 24 irroratus, Unio, 105 Juvenal mussel, 26 stage, 26 Kansas drainage systems, 166 kev, identification of mussels, 33 kidney, of mussel, 25 kidney-shell mussel, 162 labial palpi, 24 lachrymosa, Quadrula, 52 lachrymosus, Unio, 50 lady-finger mussel, 80 laevissima, Lampsilis, 123 Leptodea, 15, 35, 122, 123, 124, 170 Symphynota, 123 Unio, 123 182 Index Lampsilinae, 14 Lampsilis, 143 alata, 126 anodontoides, 30, 31, 36, 143, 144, 170 capax, 132 fallaciosa, 36, 146, 147 fallaciosus, 146 gracilis, 120 laevissima, 123 ligamentinus, 108 luteola, 149 ovata, 36, 152, 153 parvus, 135 purpurata, 129 radiata, 36, 149, 150, 152 recta, 137 satura, 154 siliquoidea, 149 subrostrata, 29, 140 ventricosa, 152 largemouth bass, 102 Lasniigona, 18, 26, 84 complanata, 30, 31, 33, 35, 87, 88, 89, 90, 128, 170 costata, 33, 85, 86 lateral tooth, of shell, 18 latissima, Ligumia recta, 28, 36, 81, 137, 138 Unio, 137 Leponiis niachrochirus, 78 Leptodea, 119 fragilis, 29, 31, 35, 120, 121, 122 laevissima, 15, 35, 122, 123, 124, 170 ligament, of shell, 15 ligamentinus, Lampsilis, 108 Unio, 108 light, effect on mussels, 30 Ligumia, 137 latissima, 28, 36, 81, 137, 138 recta, 28, 36, 81, 137, 138 subrostrata, 18, 28, 30, 36, 137, 140, 141 lilliput mussel, 135 Obliquaria, 117 lineolata, Plagiola, 28, 37, 117, 118 luteolus, Lampsilis, 149 Unio, 149 Lymnea, 11 Macrochirus, Lepomis, 78 mantle, of mussel, 19 cavity, 24 maple-leaf mussel, 50 Margaritana complanata, 87 rugosa, 86 marsupium, of mussel, 24 Megalonaias, 42 gigantea, 29, 33, 34, 42, 44 metanevra, Obliquaria, 61 Quadrula, 34, 60, 61 metanevrus, Unio, 61 Mollusca, 11,39 monkey-faced mussel, 61 Monoplacophora, 11 mouth, of mussel, 24 mucket mussel, 108 mucous string, 24 mussel, as animal food, 9 Mytilus cygneus, 90 Nautilus, 11 Neopilina, 11 nervous system, of mussels, 25 nigromaculatus, Pomoxis, 48 nitrogenous waste, elimination of, 25 nodulata, Obovaria, 58 Quadrula, 18, 34, 57, 58, 59, 105 nuclear shell, 14 Obliquaria, 102 cypha, 68 fasciolaris, 162 flava, 39 lineolata, 117 metanevra, 61 quadrula, 50 reflexa, 27, 34, 59, 103, 104 Obovaria, 161 ellipsis, 161 nodulata, 58 olivaria, 158, 159, 161, 162 occidens, Unio, 152 octupus, 11 olivaria, Amblema, 161 Obovaria, 161 organs, respiratory, 24 origin, Kansas unionid fauna ovalis, Amblema, 161 ovata, Lampsilis, 36, 152, 153 ovatus, Unio, 143 pallial line, of shell, 14 muscles, 19 pallium, 19 paper-shell mussel, 123 parasitic stage, of mussel, 26 parasitism, 30 parva, Carunculina, 29, 36, 135, 137, 143, 170 parvus, Lampsilis, 135 Unio, 134, 135 pealei, Unio, 161 pedal ganglion, 25 Pelecypoda, 11,39 pericardial fluid, 25 periostracum, of shell, 14, 27 peruviana, Crenodonta, 18, 34, 46, 47, 50 Unio, 47 phaseolus, Ptychobranchus, 162 Unio, 162 Physa, 11 pimple-backed mussel, 55 pink heel-splitter mussel, 126 pistol-grip mussel, 68 Plagiola, 116 donaciformis, 114 Index 183 elegans. 111 lineolata, 28, 37, 117, 118 securis, 117 plain pocketbook mussel, 152 plenus, Unio, 72 Plethobasus, 68 cyphyus, 35, 37, 68, 69 Pleurobema, 71 aesopus, 68 catillus, 37, 74, 75 coccineum, 37, 42, 71, 72 cordatum, 37, 42, 71, 72, 74, 76,77 pyramidatum, 37, 76, 77 plicatus, Quadnila, 47 Unio, 45, 47 pocketbook mussel, 132 Pomoxis nigromaculatus, 48 pond-horn mussel, 83 popenoi, Unio, 105 posterior slope, of shell, 15 predation, of mussels, 30 prismatic layers, of shell, 15 Proptera, 126 alata, 18, 30, 35, 90, 126, 127 capax, 18, 35, 132, 133 purpurata, 35, 128, 129, 130, 171 Protobranchia, 11 protractor muscles, 19 pseudocardinal denticles, 14, 18 Ptychobranchus, 162 clintonense, 163 fasciolare, 158, 159, 162, 163 phaseolus, 162 punctatus, Ictalurus, 26 purple shell mussel, 129 purpurata, Lampsilis, 129 Proptera, 35, 128, 129, 130, 171 Unio, 129 purpuratus, Unio, 129 pustulatus, Unio, 58 pustulosa, Quadrula, 30, 34, 37, 54, 55, 58, 59, 165 Unio, 55 pyramid pig-toed mussel, 76 pyramidata, Quadrula, 76 pyramidatum, Pleurobema cordatum, 37, 76, 77 pyramidatus, Unio, 76 Quadrula, 50 coccinea, 71 cylindrica, 35, 63, 64 fragosa, 53 heros, 42 houstonensis, 165 lachrymosa, 52 nodulata, 18, 34, 57, 58, 59, 105 plicata, 47 pustulosa, 30, 34, 37, 54, 55, 58, 59, 165 pvramidata, 76 quadrula, 18, 30, 34, 50, 51, 52 rubiginosa, 39 solida, 74 undulata, 49 quadrula, Obliquaria, 50 Quadrula, 18, 30, 34, 50, 51, 52 radiata, Lampsilis, 36, 149, 150, ]52 recta, Lampsilis, 137 Ligumia, 28, 36, 81, 137, 138 Unio, 137 rectum, of mussel, 24 rectus, Unio, 137 reflexa, Obliquaria, 27, 34, 59, 102, 103, 104 reproductive system, of mussel, 25 rest periods, of shell, 27 retractor muscle, 19 rock pocketbook mussel, 97 round pig-toed mussel, 71 rubiginosa, Quadrula, 39 rubiginosus, Unio, 39 rugosa, Margaritana, 86 rugosus, Anodon, 99 Strophitus, 18, 35, 99, 100, 101, 104 sapotalensis, Unio, 108 satura, Lampsilis ventricosa, 154 sauger, 62, 116, 154 Scaphopoda, 11 scars, muscle, 18 securis, Plagiola, 117 Septibranchia, 11 shell tongs, 32 short-nosed gar, 148 siliquoidea, LampsiHs, 149 siliquoides, Unio, 149 siphoning rate, of mussels, 10 slough sand mussel, 146 slugs, 11 small-mouthed bass. 111 snails, 11 snuffbox mussel, 155 solid pig-toed mussel, 74 solida, Quadrula, 74 solidus, Unio, 74 spectacle-case mussel, 64 Sphaeriidae, 14 Sphaerium, 11 squaw-foot mussel, 99 squids, 11 State Biological Sur\ey, 10 stream velocity, 32 Strophitus, 99 edentulus, 26, 99 rugosus, 18, 35, 99, 100, 101, 104 sturgeon, 148, 162 suborbiculata, Anodonta, 93, 158, 160 subrostrata, Lampsilis, 29, 140 Ligumia, 18, 28, 30, 36, 137, 140, 141 subrostratus, Unio, 140 184 Index suprabranchial chamber, 24 temperature, significance of, 30 tetralasmus, Unio, 81, 83 three horned wart-backed mussel, 103 three-ridged mussel, 49 topekaensis, Unio, 140 trigonus, Unio, 39 triqueter, Truncilla, 155 triquetra, Dysnomia, 36, 38, 116, 155, 156, 158 triquetra, TrunciHa, 155 Tritogonia, 29, 65 tuberculata, 67 verrucosa, 30, 34, 66, 67, 68, 170 trochophore larva, 10 truncata, Truncilla, 18, 36, 111, 112, 113, 114, 158 Truncilla, 111 donaciforniis, 36, 114, 115, 116 triqueter, 155 triquetra, 155 truncata, 18, 36, 111, 112, 113, 114, 158 Tryblium, 11 tuberculata, Tritogonia, 67 tuberculatus, Unio, 65, 67 type locality, 38 umbo, 15 umbonal cavity, 18 undata, Fusconaia, 41, 42 undulata, Anodonta, 99 Quadrula, 49 undulatus, Unio, 49 Unio aberti, 105 alatus, 126 anodontoidcs, 143 asperrimus, 52 camptodon, 83 capax, 132 carinatus, 108 catillus, 74 coccineus, 71 cornutus, 103 crassidens, 78 cylindricus, 64 dilatata, 80 donaciforniis, 114 dorfeuilliana, 55 elegans. 111 ellipsis, 161 fragilis, 120 gibbosus, 80 giganteus, 42 gracilis, 119, 120 heros, 42 irroratus, 105 lachrymosus, 50 laevissima, 123 latissima, 137 ligamentinus, 108 luteolus, 149 metanevrus, 61 occidens, 152 ovatus, 143 parvus, 134, 135 pealei, 161 peruviana, 47 phaseolus, 162 plenus, 76 plicatus, 45, 47 popenoi, 105 purpurata, 129 purpuratus, 129 pustulatus, 58 pustulosus, 55 pyramidatus, 76 recta, 137 rectus, 137 rubiginosus, 39 sapotalensis, 108 securis, 117 siHquoides, 149 solidus, 74 subrostratus, 140 tetralasmus, 81, 83 topekaensis, 140 trigonus, 39 tuberculatus, 65, 67 undulatus, 44, 49 ventricosus, 152 verrucosus, 67 zigzag, 114 Uniomerus, 81 tetralasmus, 28, 31, 36, 82, 83, 143, 170 Unionidae, 14, 38, 39 Unioninae, 14 valve, of shell, 14 vegetation, effect of, 30 veins, 25 veliger larva, 10 ventricle, 25 ventricosa, Lampsilis ovata, 36, 152, 153 ventricosa, Lampsilis satura, 154 ventricosus, Unio, 152 verrucosa, Tritogonia, 30, 34, 66, 67, 68, 170 verrucosus, Unio, 67 visceral ganglia, 25 mass, 24 Wabash pig-toed mussel, 39 walleye, 151 warty-backed mussel, 58 water temperature, 32 white bass, 45 crappie, 45, 48, 59, 125, 148, 154 heel-splitter mussel, 87 wing, 15 yellow, perch, 151 sand-shell mussel, 105 young fan-tailed mussel, 105 zigzag, Unio, 114 D -r — 1 2 <« « o X « 3 •4- "5 o z >, of -J o o 1> V) O- o- — 1: 11 is_ I University of Kansas Museum of Natural History, Miscellaneous Publications Institutional libraries interested in publications exchange may obtain this series from tlie Exchange Librarian, The University of Kansas, Lawrence, Kansas. Requests of individuals are handled instead by the Museum of Natural History, The University of Kansas, Lawrence, Kansas. There is no provision for sale of this series either by the Library or the Museum. However, when individuals request copies from the Museum, the amount indicated below should be included for the purpose of defraying some of the costs of producing. Nos. 6, 12, 17, and 27 obtainable only from the Arctic Institute. *1. The Museum of Natural Histoiy, the University of Kansas. By E. R. Hall and Ann Murray. Pp. 1-16, illustrated. January 5, 1946. *2. Handbook of Amphibians and Reptiles of Kansas. By Hobart M. Smith. Pp. 1-336, 233 figures in text. September 12, 1950. *3. In Memoriam, Charles Dean Bunker, 1870-1948. By E. Raymond Hall. Pp. 1-11, 1 figure in text. December 15, 1951. *4. The University of Kansas, Natural History Reservation. By Henry S. Fitch. Pp. 1-38, 4 plates, 3 figures in text. February 20, 1952. *5. Prairie Chickens of Kansas. By Maurice F. Baker. Pp. 1-68, 4 plates, 15 figures in te.\-t. March 10, 1953. 6. The Barren Ground Caribou of Keewatin. By Francis Harper. Pp. 1-163, 28 figures. October 21, 1955. Copies, paper bound, $1.50 postpaid from the Arctic Institute of North America, 1530 P Street NW, Washington 5, D. C. 7. Handbook of Mammals of Kansas. Bv E. Ravmond Hall. Pp. 1-303, illustrated. December 13. 1955. Paper bound, $1.50 postpaid (cloth $4.00). 8. Mammals of Northern Alaska, on the Arctic Slope. By James W. Bee and E. Ray- mond Hall. Pp. 1-309, Frontispiece colored, 4 plates, 127 figures in text. March 10, 1956. Pajjer bound $1.00 postpaid. 9. Handbook of Amphibians and Reptiles of Kansas. 2nd [revised] edition. By Hobart M. Smith. Pp. 1-356, 253 figures in text. April 20, 1956. Paper bound $1.50 posti^aid (cloth $4.00). *10. The Raccoon in Kansas. By Howard J. Stains. Pp. 1-76, 4 plates, 14 figures in text. Julv 6, 1956. 11. The Tree Squirrels of Kansas. By Robert L. Packard. Pp. 1-67, 2 plates, 10 fig- ures in text. August 20, 1956. 12. The Mammals of Keewatin. By Francis Harper. Pp. 1-94, 6 plates, 8 figures in text, 1 map. October 26, 1956. Copies, xjaper bound, 75 cents postpaid from the Arctic Institute of North America, 1530 P Street NW, Washington 5, D. C. *13. Museum of Natural Histoi-v . . . University' of Kansas. By Roy R. Moore and E. R. Hall. Pp. 1-8, illustrated. June 1, 1957. 14. Vernacular Names for North American Mammals North of Mexico. By E. Raymond Hall, Sydnev Anderson, J. Knox Jones, Jr., and Robert L. Packard. Pp. 1-16. June 19. 1957. *15. The Ecology of Bobwhites in South-central Kansas. By Thane S. Robinson. Pp. 1-84, 2 plates, 11 figures in text. September 6, 1957. 16. Natural History of the Prairie Dog in Kansas. By Ronald E. Smith. Pp. 1-36, 4 plates, 9 figures in text. June 17, 1958. 17. Birds of the Ungava Peninsula. By Francis Harper. Pp. 1-171, 6 plates, 26 maps. October 15, 1958. Paper bound, $2.00 postpaid from the Arctic Institute of North America, 1530 P Street NW, Washington 5. D. C. 18. Furbearers in Kansas: A Guide to Trapping. By Howard J. Stains and Rollin H. Baker. Pp. 1-100, illustrated. November 19, 1958. Paper bound, 50 cents post- paid. *19. Musevun of Natural Historv . . . University of Kansas. By Roy R. Moore and E. R. Hall. Pp. 1-S, illustrated. May 29, 1959. 20. Handbook of Gastropods in Kansas. By A. Bvron Leonard. Pp. 1-224, 11 plates, 87 figures in text. November 2, 1959. Paper bound, $1.00 (cloth $2.00) postpaid. 21. Management of Channel Catfish in Kansas. By Jackson Davis. Pp. 1-56, 8 figures in tex-t. November 2, 1959. 22. Hand-list of the Birds of Kansas. By Richard F. Johnston. Pp. 1-6. May 7, 1960. 23. Directoi-v to the Bird-life of Kansas. Bv Richard F. Johnston. Pp. 1-69, 1 figure. August 31, 1960. 24. Museum of Natural Histors' . . . Universits' of Kansas. Bv Roy R. Moore and E. R. Hall. Pp. 1-8, illustrated. October 19, 1960. 25. Guide to the Panorama of North American Mammals. By E. Raj-mond Hall et al. Pp. 1-32, illustrated, December 15, 1960. Paper bound, 50 cents postpaid. 26. Beaver in Kansas. Bv F. Robert Henderson. Pp. 1-85, illustrated. December 16, 1960. 27. Land and Fresh-water Mammals of the Ungava Peninsula. Bv Francis Harper. Pp. 1-17S, 8 plates. 3 figures. 45 maps. August 11, 1961. Paper bound, $2.00 postpaid from the Arctic Institute of North America, 1530 P Street NW, Washing- ton 5, D. C. , . „ 28. Handbook of Unionid Mussels in Kansas. By Harold D. Murray and A. Byron Leonard. Pp. 1-184, 45 plates, 42 figures in tex-t. May 10, 1962. Paper bound, $1.00 (cloth $2.00) postpaid. * Out of print. 574