The Neotropical Land Snail Genera: Labyrinthus and Isomeria
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LIBRARY OF THE UNIVERSITY OF ILLINOIS AT URBANA-CHAMPAIGN 530.5 FI v.50 CC2D.3 ..niUKAL HISTORY SURVEY t 50 THE NEOTROPICAL LAND SNAIL GENERA 3 LABYRINTHUS and ISOMERIA (PULMONATA, CAMAENIDAE) ALAN SOLEM FIELDIANA: ZOOLOGY VOLUME 50 Published by FIELD MUSEUM OF NATURAL HISTORY MAY 31, 1966 liB2A°Y FIELDIANA: ZOOLOGY A Continuation of the ZOOLOGICAL SERIES of FIELD MUSEUM OF NATURAL HISTORY VOLUME 50 FIELD MUSEUM OF NATURAL HISTORY CHICAGO, U.S.A. 1966 THE NEOTROPICAL LAND SNAIL GENERA LABYRINTHUS and ISOMERIA (PULMONATA, CAMAENIDAE) THE NEOTROPICAL LAND SNAIL GENERA LABYRINTHUS and ISOMERIA (PULMONATA, CAMAENIDAE) ALAN SOLEM Curator, Division of Lower Invertebrates FIELDIANA: ZOOLOGY VOLUME 50 Published by FIELD MUSEUM OF NATURAL HISTORY MAY 31, 1966 Library of Congress Catalog Card Number: 66-1 91 !& PRINTED IN THE UNITED STATES OF AMERICA BY FIELD MUSEUM PRESS / is F fX /LJL'O „ SO CONTENTS PAGE List of Text Figures 7 List of Tables 9 Introduction 11 General and Acknowledgments 11 Material 13 Life History and Ecology 19 Gross Anatomy 20 Classification 34 Fossil Record 36 Systematic Review Labyrinthus 37 Classification 42 Review of the species 44 Group of Labyrinthus isodon 44 Key to the Group 46 Labyrinthus triplicatus (von Martens) 47 Labyrinthus quadridentatus (Broderip) 50 Labyrinthus q. quadridentatus (Broderip) 51 L. q. biolleyi new subspecies 52 Labyrinthus tamsianus (Dunker) 53 Labyrinthus leucodon (Pfeiffer) 55 Labyrinthus I. leucodon (Pfeiffer) 55 Labyrinthus I. umbrus Thompson 58 Labyrinhus magdalenensis new species 59 Labyrinthus dunkeri (Pfeiffer) 60 Labyrinthus isodon (Pfeiffer) 62 Labyrinthus vexans (Dohrn) 65 Labyrinthus manueli Higgins 66 Group of Labyrinthus raimondii 67 Key to the Group 70 Labyrinthus sieversi (von Martens) 71 Labyrinthus sp 71 Labyrinthus ellipsostomus (Pfeiffer) 73 Labyrinthus diminutus Gude 75 Labyrinthus pronus Pilsbry 78 Labyrinthus leprieurii (Petit) 79 Labyrinthus tarapotoensis (Moricand) 80 Labyrinthus t. tarapotoensis (Moricand) 80 Labyrinthus t. baeri Dautzenberg 81 3 CONTENTS PAGE Labyrinthus raimondii (Philippi) 81 Labyrinthus furcillalus (Hup6) 86 Labyrinthus bifurcatus (Deshayes) 87 Group of Labyrinthus aenigmus 87 Key to the Group 89 Labyrinthus aenigmus (Dohrn) 89 Labyrinthus clappi Pilsbry 90 Labyrinthus otostomus (Pfeiffer) 91 Labyrinthus o. otostomus (Pfeiffer) 91 Labyrinthus o. bogotensis (Pfeiffer) 92 Labyrinthus o. assimilans E. A. Smith 93 Group of Labyrinthus unciger 94 Labyrinthus unciger (Petit) 95 Labyrinthus creveauxianus (Ancey) 100 Group of Labyrinthus otis (Lightfoot) 102 Key to the Group 103 Labyrinthus otis (Lightfoot) 105 Labyrinthus o. otis 105 Labyrinthus o. orthorhinus Pilsbry 108 Labyrinthus subplanatus (Petit) 114 Labyrinthus s. subplanatus (Petit) 114 Labyrinthus s. erectus (Mousson) 116 Labyrinthus s. sipunculatus (Forbes) 119 Labyrinthus plicatus (Born) 122 Labyrinthus marmatensis Pilsbry 124 Species of uncertain affinity 125 Labyrinthus euclausus Beddome 125 Labyrinthus sharmani (Gude) 126 Isomeria 126 Classification 130 Review of the species 131 Key to the Species Group 131 Group of Isomeria inexpectata 132 Isomeria inexpectata new species 132 Group of Isomeria bituberculata 134 Key to the Group 135 Isomeria medemi new species 136 Isomeria basidens (Mousson) 138 Isomeria b. basidens (Mousson) 138 Isomeria b. gudeana (Ancey) 139 Isomeria neogranadensis (Pfeiffer) 140 Isomeria bituberculata (Pfeiffer) 142 Isomeria juno (Pfeiffer) 145 Isomeria bourcieri (Pfeiffer) 148 Isomeria hartwegi (Pfeiffer) 150 Group of Isomeria subelliptica 151 Key to the Group 152 Isomeria morula (Hidalgo) 152 CONTENTS 5 PAGE Isomeria anodonta Pilsbry 153 Isomeria aloagana Jousseaume 155 Isomeria continua (Pfeiffer) 155 Isomeria minuta new species 157 Isomeria subelliptica (Mousson) 159 Group of Isomeria meobambensis 160 Key to the Group 161 Isomeria meobambensis (Pfeiffer) 161 Isomeria gealei (Smith) 162 Isomeria aequatoria (Pfeiffer) 164 Isomeria meyeri (Strubell) 164 Group of Isomeria oreas 165 Key to the Group 167 Isomeria oreas (Koch) 167 Isomeria kolbergi Miller 172 Isomeria triodonta (d'Orbigny) 174 Isomeria stoltzmanni (Lubomirski) 176 Isomeria jacksoni new name 178 Isomeria cymatodes (Pfeiffer) 180 Isomeria fordiana (Pilsbry) 183 Isomeria scalena (von Martens) 183 Isomeria aequatoriana (Hidalgo) 184 Isomeria globosa (Broderip) 190 Nomina dubia Isomeria calomorpha (Jonas) 193 Isomeria fauna (Philippi) 194 Isomeria equestrata (Moricand) 194 Zoogeography 196 Distribution of Labyrinthus 196 Altitudinal zonation 204 Distribution of Isomeria 206 Altitudinal zonation 209 Comparative remarks 211 Summary 213 References 215 Index 217 List of Illustrations PAGE 1. Jaws of Labyrinthus unciger and L. otis orthorhinus 22 2. Pallial regions of Labyrinthus otis orthorhinus and L. diminutus .... 24 3. Pallial regions of Isomeria globosa and Labyrinthus unciger 25 4. Genitalia of Isomeria globosa 26-27 5. Genitalia of Labyrinthus unciger 28 6. Genitalia of Labyrinthus otis orthorhinus 29 7. Genitalia of Labyrinthus diminutus 30 8. Interior of penial region in Labyrinthus diminutus 31 9. Apertural zones in Labyrinthus 39 10. Apertures of Labyrinthus triplicatus, L. quadridentatus and L. magdale- nensis 48 11. Shells of Labyrinthus triplicatus, L. quadridentatus and L. magdalenensis . 49 12. Apertures of Labyrinthus tamsianus, L. leucodon 56 13. Shells of Labyrinthus tamsianus, L. leucodon and L. dunkeri 57 14. Apertures of Labyrinthus dunkeri, L. isodon, L. vexans and L. manueli . 62 15. Shells of Labyrinthus isodon, L. vexans, L. manueli and L. sieversi ... 63 16. Apertures of Labyrinthus sieversi, L. leprieurii 72 17. Shells of Labyrinthus leprieurii and L. ellipsostomus 73 18. Aperture of Labyrinthus ellipsostomus 74 19. Shells of Labyrinthus sp. and L. diminutus 76 20. Labyrinthus pronus 78 21. Apertures of Labyrinthus diminutus, L. tarapotoensis, L. raimondii, L. fur- cillatus and L. bifurcatus 82 22. Shells of Labyrinthus tarapotoensis, L. raimondii, L. furcillatus and L. bi- furcatus 83 23. Apertures of Labyrinthus aenigmus, L. clappi, and L. otostomus .... 88 24. Shells of Labyrinthus aenigmus, L. clappi, and L. otostomus 89 25. Parietal lips in Labyrinthus otostomus 92 26. Labyrinthus otostomus assimilans and L. sharmani 94 27. Shells of Labyrinthus unciger and L. creveauxianus 96 28. Aperture of Labyrinthus creveauxianus 97 29. Palatal tooth variation in Labyrinthus unciger 98 30. Shells of Labyrinthus otis 110 31. Supraperipheral grooves in Labyrinthus otis Ill 32. Apertures in Labyrinthus subplanatus and Labyrinthus plicatus . .114 33. Shells of Labyrinthus subplanatus 115 34. Apertures of Labyrinthus sipunculatus and L. plicatus 120 35. Variation of peripheral notch in Labyrinthus sipunculatus 121 36. Shells of Labyrinthus plicatus and L. marmatensis 124 37. Isomeria inexpectata 133 38. Isomeria medemi 137 39. Isomeria basidens 140 7 8 LIST OF ILLUSTRATIONS PAGE 40. Apertures of Isomeria basidens, I. neogranadensis and /. bituberculata . 141 4 1 . Shells of Isomeria neogranadensis, I. bituberculata, I. juno and /. hartwegi . 146 42. Shells of Isomeria juno, I. hartwegi and /. bourcieri 147 43. Isomeria aloagana 154 44. Shells of Isomeria anodonta and /. continua 156 45. Isomeria minuta 158 46. Isomeria subelliptica 159 47. Shells of Isomeria meobambensis, I. gealei and J. aequatoria 163 48. Shells of Isomeria oreas, I. kolbergi and /. triodonta 170 49. Apertures of Isomeria oreas, I. kolbergi and /. triodonta 171 50. Isomeria stoltzmanni 177 51. Shells of Isomeria jacksoni, I. cymatodes and J. fordiana 180 52. Apertures of Isomeria jacksoni, I. cymatodes and /. fordiana 181 53. Shells of Isomeria aequatoriana, I. globosa, I. scalena and /. fauna . 186 54. Apertures of Isomeria aequatoriana, I. scalena and /. fauna 187 55. Isomeria aequatoriana 189 56. Apertures of Isomeria globosa 192 57. Distribution of L. isodon complex 197 58. Distribution of Labyrinthus raimondii complex 199 59. Distribution of Labyrinthus otis complex 201 60. Altitudinal range in Labyrinthus 203 61. Altitudinal range in Isomeria 210 List of Tables PAGB I. Specimens examined of Labyrinthus 13 II. Specimens examined of Isomeria 14 III. Representation in collections 15 IV. Size and shape variation in Labyrinthus triplicatus, L. q. quadridentatus, L. q. biolleyi, L. tamsianus, L. I. leucodon and L. magdalenensis ... 54 V. Size and shape variation in Labyrinthus dunkeri and L. manueli ... 64 VI. Size variation in Labyrinthus raimondii complex 69 VII. Size and shape variation in Labyrinthus ellipsostomus, L. diminutus, L. tarapotoensis baeri and L. furcillatus 77 VIII. Size and shape variation in Labyrinthus raimondii, L. clappi, L. oto- stomus bogotensis and L. creveauxianus 85 IX. Size and shape variation in Labyrinthus unciger 101 X. Size and shape variation in Labyrinthus o. otis and L. o. orthorhinus . 107 XI. Size and shape variation in Labyrinthus o. orthorhinus 109 XII. Size and shape variation in Labyrinthus s. subplanatus, L. s. sipuncu- . latus and L. plicatus 118 XIII. Size and shape variation in Isomeria inexpectata, I. juno, I. bituberculata and J. meobambensis 143 XIV. Size and shape variation in Isomeria oreas, I. kolbergi and /. jacksoni . 169 XV. Size and shape variation in Isomeria cymatodes, I. aequatoriana and /. globosa 185 XVI. Transandean distributions in Central Ecuador 207 INTRODUCTION 1 This review of the mainland Neotropical Camaenidae had its ori- gins in an attempt to evaluate infra-specific variation in shells of Panamanian Labyrinthus. It soon developed into a synoptic