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ENDODONTOE) LAND SNAILS FROM PACIFIC 
ISLANDS 

(Mollusca: Pulmonata: Sigmurethra) 

PartH 

Families Punctidae and Charopidae, Zoogeography 

Alan Solem 



Field Museum of Natural History 
Chicago, Illinois 



. 



UNlVLhSiTY OF 

ILLINOIS LIBRARY 

AT. URBANA -CHAMPAIGN 



ENDODONTOID LAND SNAILS FROM PACIFIC 
ISLANDS 

(Mollusca: Pulmonata: Sigmurethra) 



Part II 

Families Punctidae and Charopidae, Zoogeography 

Alan Solern 

Curator of Invertebrates 

Field Museum of Natural History 






Field Museum of Natural History 
Chicago, Illinois 



Submitted December 29, 1978 

Published by: 

Field Museum of Natural History 

December 30, 1982 

Mailed January 7, 1983 



Library of Congress Catalog Card No.: 76-9516 
ISSN 0015-0754 



PRINTED IN THE UNITED STATES OF AMKRICA 



II 









CONTENTS 



LIST OF FIGURES V 

LIST OF TABLES VIII 

INTRODUCTION 1 

Acknowledgements 2 

PREVIOUS STUDIES 3 

MATERIAL STUDIED 4 

METHODS OF ANALYSIS 5 

PATTERNS OF MORPHOLOGICAL VARIATION 8 

Size and shape variations 8 

Shell sculpture 9 

Apertural barriers 15 

Parietal barriers 19 

Columellar barriers 21 

Palatal barriers 21 

Patterns of barrier variation 23 

Summary of barrier variations 23 

Summary of shell variations 23 

Gross anatomy 23 

Genital system 24 

Ovotestis 24 

Hermaphroditic duct 24 

Talon and carrefour 25 

Albumen gland 25 

Prostate and uterus 25 

Terminal male genitalia 25 

Terminal female genitalia 30 

Pallial complex 30 

Digestive system 32 

Free muscle system 34 

Nervous system 34 

External body features 34 

Patterns of visceral hump reduction 36 

Summary of anatomical variation 37 

CHAROPID-ENDODONTID CONCHOLOGICAL COMPARISONS 38 

HABITAT RANGE AND EXTINCTION 45 

PHYLOGENY AND CLASSIFICATION 46 

FAMILY CLASSIFICATION OF THE ENDODONTOIDS 47 

PHYLOGENETIC PROCEDURES 48 

GENERIC CLASSIFICATION 49 

SYSTEMATIC REVIEW 50 

List of the taxa 50 

Geographic keys to the genera 51 

Family Punctidae 53 

Genus Punctum Morse, 1864 57 

Family Charopidae 59 

Subfamily Rotadiscinae 70 

Genus Microcharopa, new genus 70 

Subfamily Charopinae 72 

Genus Discocharopa Iredale, 1913 74 

Genus Sinployea, new genus 81 

III 



IV 

Society Island Sinployea 85 

Cook Island Sinployea 94 

Samoan and Swains Island Sinployea 117 

Western Polynesian and Micronesian Sinployea 133 

Fijian Sinployea 145 

Melanesian Sinployea 166 

Genus Ba, new genus 172 

Genus Maafu, new genus 174 

Genus Lauopa, new genus 177 

Genus Tuimalila, new genus 178 

Genus Lagivala, new genus 183 

Genus Vatusila, new genus 191 

Genus Graeffedon, new genus 200 

Subfamily Trukcharopinae, new subfamily 205 

Genus Trukcharopa, new genus 207 

Genus Kubaryiellus, new genus 210 

Genus Russatus, new genus 214 

Genus Roimontis, new genus 217 

Genus Palikirus, new genus 219 

Genus Jokajdon, new genus 222 

Genus Palline, new genus 228 

Subfamily Semperdoninae, new subfamily 235 

Genus Semperdon, new genus 236 

Genus Ladronellum, new genus 255 

Genus Himeroconcha, new genus 259 

Incertae sedis 266 

ZOOGEOGRAPHY 268 

Patterns of family and generic distributions 268 

Patterns of diversity 275 

Hawaiian Islands 275 

Marquesas Islands 277 

Tuamotu Archipelago 279 

Mangareva, Gambier Islands 280 

Rapa Island 282 

Austral Islands 287 

Society Islands 288 

Cook Islands 289 

Samoa 290 

Tonga 292 

Lau Archipelago 292 

Fiji, Main Islands 295 

Ellice Islands 295 

Marshall Islands 295 

Caroline Islands 297 

Mariana Islands 297 

Palau Islands 298 

Summary of specific and generic diversity 301 

Shell size and geography 305 

Hypothesized origin and radiation of Pacific Island endodontoid snail fauna 307 

Patterns of species diversity 310 

Summary of zoogeography 315 

SUMMARY 316 

REFERENCES 317 

APPENDIX: Anatomical terms 324 

INDICES . 325 



LIST OF FIGURES 

1. Shell microsculpture inSinployea modicella 10 

2. Shell microsculpture inSinployea peasei and Tuimalila pilsbryi 11 

3. Shell microsculpture inPalline biakensis 12 

4. Shell microsculpture in Microcharopa mimula 13 

5. Details of shell sculpture inDiscocharopa aperta 16 

6. Microstructure of largest palatal barrier inJokajdon callizonus 18 

7. Surface of largest palatal barrier in Graeffedon graeffei 19 

8. Structure of apertural barriers in Semperdon xyleborus 20 

9. Anatomy ofCharopa coma 26 

10. Interior of lower vas deferens and epiphallus in Charopa coma 27 

11. Anatomy of Phenacohelix pilula 28 

12. Radular teeth ofPunctum minutissimum 33 

13. Radular teeth of Tuimalila pilsbryi 35 

14. Radular teeth of Tuimalila pilsbryi 36 

15. Mean shell height distribution in the Endodontidae and Charopidae 38 

16. Mean shell diameter distribution in the Endodontidae and Charopidae 39 

17. Mean height/diameter ratio distribution in the Endodontidae and Charopidae 39 

18. Mean whorl count distribution in the Endodontidae and Charopidae 39 

19. Mean diameter/umbilical width ratio distribution in the Endodontidae and Charopidae 40 

20. Mean spire height/body whorl width ratio distribution in the Endodontidae and Charopidae 40 

21. Mean ribs on body whorl distribution in the Endodontidae and Charopidae 40 

22. Mean ribs/mm, on the body whorl distribution in the Endodontidae and Charopidae 41 

23. Patterns of whorl width increment in selected Endodontidae and Charopidae 42 

24. Partial anatomy ofPunctum minutissimum 54 

25. Anatomy ofLaoma leimonias 55 

26. Partial anatomy ofParalaoma aprica 57 

27. Anatomy of "Paralaoma" coesa 58 

28. Punctum polynesicum 60 

29. Anatomy of Flammulina zebra 64 

30. Anatomy ofAmphidoxa marmorella 65 

31. Anatomy ofStephanoda binneyana 67 

32. Anatomy of Pseudocharopa lidgbirdi 69 

33. Microcharopa mimula 71 

34. Anatomy of Discocharopa aperta 75 

35. Parietal barrier variation inDiscocharopa aperta 76 

36. Discocharopa aperta from the New Hebrides 80 

37. Australian Discocharopa aperta 81 

38. Sinployea modicella and S. tahitiensis 87 

39. Anatomy of Tahitian and Moorean Sinployea 90 

40. Sinployea lamellicosta 91 

41. Sinployea montana andS. neglecta 93 

42. Anatomy of Society and Cook Island Sinployea 96 

43. Anatomy of Sinployea avanaensis 97 

44. Sinployea andrewi andS. atiensis 100 

45. Sinployea peasei and S. avanaensis 103 

46. Sinployea proximo and S. planospira 107 

47. Sinployea decorticata andS. rudis 109 

48. Sinployea harveyensis and S. youngi 113 

49. Sinployea canalis 115 

50. Sinployea otareae and S. tenuicostata 116 

V 



VI 

51. Sinployea clista andS. clausa 120 

52. Sinployea aunuuana andS. intermedia 123 

53. Anatomy of Sinployea aunuuana, S. clista, and S. intermedia 124 

54. Sinployea allecta allecta and S. a. tauensis 126 

55. Anatomy of Sinployea allecta allecta 127 

56. Sinployea complementaria 130 

57. Anatomy of the Samoan Sinployea complementaria 132 

58. Sinployea vicaria vicaria and S. v. paucicosta 136 

59. Shell sculpture details of Sinployea vicaria vicaria 137 

60. Anatomy of the Tongan Sinployea vicaria vicaria 138 

61. Sinployea pseudovicaria andS. rotumana 139 

62. Sinployea ellicensis ellicensis and S. e. nukulaelaeana 141 

63. Sinployea kusaieana and Trukcharopa trukana 144 

64. Sinployea recursa andS. angularis 148 

65. Sinployea princei andS. irregularis 151 

66. Sinployea inermis inermis, S. i. meridionalis, and S. i. lakembana 156 

67. Anatomy of Fijian, Caroline Island, and New Hebridean Sinployea 158 

68. Sinployea lauensis andS. navutuensis 160 

69. Sinployea godeffroyana andS. adposita 162 

70. Sinployea euryomphala 168 

71. Sinployea solomonensis 169 

72. Sinployea kuntzi 170 

73. Sinployea novopommerana and S. descendens 171 

74. Ba humbugi 173 

75. Anatomy ofBa humbugi 175 

76. Maafu thaumasius 176 

77. Lauopa mbalavuana 178 

78. Tuimalila pilsbryi and T. infundibulus 180 

79. Anatomy of the Tongan Tuimalila pilsbryi 182 

80. Lagivala vivus andL. minusculus 186 

81. Lagivala microglyphis andL. macroglyphis 188 

82. Vatusila kondoi and V. nayauana 193 

83. Vatusila tongensis and V. niueana 197 

84. Partial anatomy of Vatusila tongensis 198 

85. Vatusila uaitupuensis and V. eniwetokensis 199 

86. Graeffedon graeffei 201 

87. Anatomy of the Samoan Graeffedon graeffei 203 

88. Graeffedon savaiiensis and G. pricei 204 

89. Kubaryiellus kubaryi andRussatus nigrescens 213 

90. Anatomy ofRussatus, Kubaryiellus, and Trukcharopa 216 

91. Roimontis tolotomensis andPalikirus cosmetus 218 

92. Jokajdon callizonus and J. tumidulus 224 

93. Anatomy of Jokajdon andPalline 227 

94. Palline notera notera and P. n. palauana 230 

95. Palline notera gianda andP. micramyla 233 

96. Palline biakensis 235 

97. Semperdon uncatus 239 

98. Semperdon kororensis andS. xyleborus 242 

99. Semperdon heptaptychius andS. rotanus 246 

100 Anatomy of Semperdon 249 

101. Ladronellum mariannarum 256 

102. Anatomy of Ladronellum mariannarum 258 

103. Anatomy ofHimeroconcha 261 

104. Himeroconcha rotula and//, lamlanensis 262 

105. Himeroconcha quadrasi and//, fusca 264 

106. Distribution of Pacific Island endodontoid families 269 

107. Distribution of most generalized and most specialized endodontid genera 270 

108. Ranges of Lagivala and Vatusila 271 

109. Range of Mautodontha and areas where umbilical brood chambers evolved 272 

110. Range of Sinployea . 273 



VII 

111. Ranges ofGraeffedon, Palline, and Semperdon 274 

112. Collecting localities on smaller islets of Mangareva, Gambier Islands 280 

113. Collecting localities on Mangareva Islet, Mangareva, Gambier Islands 281 

114. Collecting localities on Rapa Island 283 

115. Major geographic features on Rurutu, Austral Islands 287 

116. Collecting localities on Tubuai, Austral Islands 288 

117. Collecting localities on Raivavae, Austral Islands 289 

118. Collecting localities on Rarotonga, Cook Islands 290 

119. Collecting localities on Upolu, Samoa 291 

120. Collecting localities on Savai'i, Samoa 291 

121. Main islands of Tonga 292 

122. Collecting localities on Vava'u, Tonga 293 

123. Collecting localities on Tongatapu, Tonga 294 

124. Collecting localities on Eua, Tonga 295 

125. Islands of the Lau Archipelago 296 

126. Collecting localities on Kusaie, Caroline Islands 298 

127. Collecting localities on Ponape, Caroline Islands 298 

128. Islets of the Truk Group, Caroline Islands 298 

129. Collecting localities on Moen Islet, Truk, Caroline Islands 298 

130. Collecting localities on Tol Islet, Truk, Caroline Islands 299 

131. Collecting localities on Dublon Islet, Truk, Caroline Islands 299 

132. Collecting localities on Guam, Mariana Islands 299 

133. Islands of the Palau Group 300 

134. Collecting localities on Peleliu, Palau Group 300 

135. Collecting localities on Koror and Auluptagel, Palau Group 300 

136. Collection station on Angaur, Palau Group 301 

137. Species diversity in island groups for the Endodontidae and Charopidae 303 

138. Generic diversity in island groups for the Endodontidae and Charopidae 304 

139. Median mean and mean ranges of adult shell diameter in island groups for the Endodontidae 
and Charopidae 306 

140. Species-area relationships for total land snail faunas of selected Pacific Islands 312 

141. Species-area relationships for selected land snail families on Polynesian and Micronesian islands 313 

142. Species diversity in islands grouped by size 313 

143. Species diversity in islands grouped by elevation 313 



LIST OF TABLES 



I. Range of variation in the Pacific Island Charopidae 8 

II. Body whorl contour in Endodontidae and Charopidae 8 

III. Shell diameter and rib spacing in the Charopidae 14 

IV. Degree of apertural narrowing by barriers 18 

V. Percentage distribution of columellar barrier numbers 21 

VI. Percentage distribution of palatal barrier numbers 22 

VII. Palatal barrier numbers in constricted aperture Charopidae 22 

VIII. Phyletic representation of dissected taxa 24 

IX. Relative length of kidney lobes in the Pacific Island Charopidae 31 

X. Conchological comparisons of Endodontidae and Charopidae from Pacific Islands 38 

XI. Species numbers of Endodontoid land snails from selected geographic areas 53 

XII. Range of variation in Punctum, Discocharopa, and Microcharopa 59 

XIII. Geographic variation in Discocharopa aperta 79 

XIV. Ratio of penis length to shell diameter in Sinployea 83 

XV. Mean species measurements in Sinployea 83 

XVI. Range of variation in Society Island Sinployea 85 

XVII. Local variation in Society Island Sinployea, 1 89 

XVIII. Local variation in Society Island Sinployea, II 95 

XIX. Range of variation in Cook Island Sinployea 98 

XX. Local variation in Cook Island Sinployea, 1 101 

XXI. Local variation in Sinployea peasei 104 

XXII. Local variation in Cook Island Sinployea, II Ill 

XXIII. Local variation in Cook Island Sinployea, III 112 

XXIV. Range of variation in Samoan and Swains Island Sinployea 118 

XXV. Local variation in Samoan Sinployea, 1 121 

XXVI. Local variation in Samoan Sinployea, II 128 

XXVII. Range of variation in Western Polynesian and Micronesian Sinployea 133 

XXVIII. Local variation in Sinployea vicaria 135 

XXIX. Local variation in Western Polynesian Sinployea 142 

XXX. Range of variation in Fijian Sinployea 146 

XXXI. Local variation in Fijian, Melanesian, and Micronesian Sinployea 152 

XXXII. D/U ratio frequency in Sinployea inermis subspecies 153 

XXXIII. Ribs/mm, frequency in Sinployea inermis subspecies 153 

XXXIV. Rib frequency in Sinployea inermis subspecies 153 

XXXV. Local variation in Sinployea inermis, 1 154 

XXXVI. Local variation in Sinployea inermis, II 155 

XXXVII. Local variation in Sinployea adposita andS. irregularis 164 

XXXVIII. Range of variation in Melanesian Sinployea 167 

XXXIX. Range of variation in Fijian and Tongan endemic genera 181 

XL. Range of variation inLagivala 184 

XLI. Local variation inLagivala demani 191 

XLII. Range of variation in Vatusila 194 

XLIII. Range of variation in Graeffedon 202 

XLIV. Sympatry of Charopidae on Ponape, Caroline Islands 208 

XLV. Range of variation in Micronesian Charopinae, 1 209 

XLVI. Local variation in Trukcharopa 211 

XLVII. Age distribution in Trukcharopa trukana 211 

XLVIII. Local variation in Kubaryiellus andRussatus 215 

XLIX. Local variation inPalikirus andJokajdon 220 

L. Range of variation in Micronesian Charopinae, II 223 

VIII 



IX 



LI. Sculptural variation inJokajdon tumidulus 226 

LII. Local variation inPalline 232 

LIII. Range of variation in Semperdoninae 237 

LIV. Local variation in Palau Island Semperdon 240 

LV. Summary of barrier variation in Mariana Islands Semperdon 241 

LVI. Local variation in Semperdon rotanus 248 

LVII. Barrier variation in Semperdon rotanus 248 

LVIII. Local variation in types and large form of Semperdon heptaptychius 252 

LIX. Local variation in Semperdon heptaptychius 253 

LX. Sculpture variation in Semperdon heptaptychius 254 

LXI. Frequency distribution of diameter in populations of Semperdon heptaptychius 254 

LXII. Frequency distribution of ribs in populations of Semperdon heptaptychius 254 

LXIII. Frequency distribution of ribs/mm, in populations of Semperdon heptaptychius 254 

LXIV. Barrier variation in Semperdon heptaptychius 255 

LXV. Local variation in Ladronellum and Himeroconcha 257 

LXVI. Generic distribution of Hawaiian Endodontidae and Punctidae 276 

LXVII. Geographic distribution of species level taxa in Hawaiian Endodontidae 276 

LXVIII. Diversity of Hawaiian land snails 276 

LXIX. Land snail fauna of Rapa Island 284 

LXX. Ages of Pacific Islands 307 

LXXI. Pacific Island land snail families with comprehensive distributional data available 311 

LXXII. Land snail faunas of Lord Howe, Upolu, and Viti Levu 312 

LXXIII. Calculated and observed maximum density of land snail species 312 

LXXIV. Correlation between island size, elevation, and number of land snail species 314 

LXXV. Land snail diversity on islands of same size but different distances from Indonesian-Melane- 

sian core 314 

LXXVI. Diversity of land snails on small, isolated islands 314 






INTRODUCTION 



This is the second of two monographs revising the 
endodontoid land snails of Polynesia, Micronesia, and 
Fiji. Part I, covering the older and larger family En- 
dodontidae, reviewed 185 species-level taxa (Solem, 
1976b). Subsequently (Solem, 1977a) I described Pro- 
toendodonta laddi and Cookeconcha antiquus from 
Late Pleistocene strata in core drillings on Midway 
Atoll and have three new subfossil Pseudolibera from 
Makatea (Solem, in preparation B). Together with the 
98 species-level taxa belonging to the Families Punc- 
tidae and Charopidae diagnosed below, a total of 290 
species-level taxa have been reviewed during this pro- 
ject. Describing the extensive Hawaiian Island 
monophyletic radiation of up to 290 unnamed, now 
mostly extinct, endodontid taxa preserved in the Ber- 
nice P. Bishop Museum is a task left to others. 

The basic descriptive work essentially was com- 
pleted in 1969, based on research supported by Na- 
tional Science Foundation grants G-16419, GB-3384, 
and GB-6779. This has been supplemented by modest 
materials received subsequently. As a by-product of 
the basic monographs, papers on a number of prob- 
lematic groups (Solem, 1964a-b, 1968b, 1969b, 1972b, 
1973c-d), extralimital endodontoids (Solem, 1958b, 
1961, 1968c, 1970a, c, 1975, 1976a, 1977c, in prepara- 
tion A), biogeography (Solem, 1958a, 1959a, 1968a, 
1969c, 1973c, 1979a-b, 1981), endodontoid biology 
(Solem, 1969a, d, 1974), and fossil land snails have 
been prepared (Solem, 1977a, in preparation B; Solem 
& Yochelson, 1979). 

In the early 1970s, the new availability of scan- 
ning electron microscopy (hereafter SEM) added a 
major new dimension to this project. A series of reports 
(Solem, 1970b, 1972a, c, 1973a-b, f, 1976a, 1977a-b) 
explored its potential, and extensive additional data 
has been incorporated into the major reviews. The first 
monograph was submitted for publication in De- 
cember, 1973, and was issued in due course (Solem, 
1976b). 

Finishing Part II has involved making very dif- 
ficult decisions as to where and when to terminate 
additional information seeking. The Endodontidae are 
restricted to the Pacific Islands and thus this mono- 
graph was easily circumscribed. Both the Punctidae 
and Charopidae have their main centers of diversity 
elsewhere, have been liberally supplied with generic 
and family-level names based on shell features and 
hunches, frequently have been rather poorly described 
and often never illustrated (particularly Iredale, 1913, 
1933, 1937a-c, 1939, 1941a-c, 1942, 1944, 1945), and 



essentially are unknown anatomically except for the 
data supplied by Hutton (1884a), Suter (see Refer- 
ences), and Climo (1969a-b, 1970, 1971a-b, 1978) on 
New Zealand taxa and contributions on South African 
taxa summarized by Connolly (1939). I have dissected 
and partially illustrated more than 75 extralimital 
species in addition to those included below. This 
attempt to work out extralimital relationships and to 
assign subfamily and generic names is only a start 
toward producing an overall phylogeny and classifica- 
tion. 

The limits finally adopted in Part II were: (1) to 
include all extralimital species known to belong in 
genera endemic to Polynesia, Micronesia, and Fiji; and 
(2) to include anatomical data on extralimital taxa 
only when needed to establish the subfamily units used 
in the systematic review and biogeography. Thus, 10 of 
the 98 taxa discussed here are found only in such areas 
as the New Hebrides, Solomon Islands, Bismarck Ar- 
chipelago, West Irian, and Indonesia (six species of 
Sinployea, three species of Lagivala, and Palline 
biakensis), whereas another species, Discocharopa 
aperta (Mollendorff, 1888), occurs sporadically in the 
area covered (Fiji, Samoa, Austral and Society Islands) 
but has its main distribution from the Philippines to 
Australia. Data are presented on type genera or species 
of several nominate subfamilies from diverse areas 
(New Zealand, Juan Fernandez), but no attempt is 
made to provide an overall classification of the 
Charopidae in this monograph. There still are too 
many gaps in our knowledge of anatomy and shell 
structure. 

By the early 1970s there still remained a few prob- 
lem taxa. Microcharopa and Discocharopa showed 
many differences from the other Pacific Island taxa, 
but I had not been able to determine their affinities 
because only empty shells were available. In addition, 
there was the problem of Stenopylis, a second Philip- 
pine Islands to Australia taxon. Both Stenopylis and 
Discocharopa were known to occur in Central Aus- 
tralia. Planned fieldwork in Western and Central Aus- 
tralia in early 1974 hopefully would yield live mate- 
rial. Early dry season collecting near Alice Springs 
produced only dead shells, but museum records sug- 
gested that both genera had a wide distribution in 
North Australia. The probability of subsequently ob- 
taining live materials seemed excellent. In addition, 
the possibility of field survey work in Micronesia for 
the Office of Endangered Species arose, which could 
have greatly expanded the anatomical coverage of the 



SOLEM: ENDODONTOID LAND SNAILS 



Trukcharopinae and Semperdoninae. This prospect did 
not materialize, but in 1976 and 1977 additional field 
and dissection work in Australia permitted allocating 
Discocharopa, Stenopylis, and Microcharopa to family 
units. The writing of introductory sections and the 
biogeographic review, assigning table and plate num- 
bers, and final editing were completed in 1978. 

It is with a mixture of delight and regret that I 
conclude this systematic project delight at the new 
information and ideas produced, regret that the hand 
of man has so altered the Pacific Island environment 
that future work on most of these taxa and testing of 
the new ideas presented has been rendered impossible 
by extinctions. The charopids apparently have been 
less affected than the Endodontidae, but even with 
these it is probable that many already are a vanished 
group of organisms. 

Concluding this monograph is a detailed biogeo- 
graphic review of the endodontoid land snails. This 
project spanned the time from the dominance of fixed 
continents as a biogeographic axiom to the current 
dogma of wandering continents. Land snails have not 
been versed in biogeographic controversies, and their 
patterns of distribution continue to fit neither model 
exclusively. 

The endodontoid land snails were the most spe- 
ciose land snail group on the Pacific Islands (Solem, 
1976b, pp. 1-2). They still are the most speciose group 
in Australia and New Zealand (table XI), areas in 
which they seem to be flourishing despite human ac- 
tivities. The data and ideas presented in these volumes 
hopefully will inspire more investigations of these 
taxa. 



ACKNOWLEDGEMENTS 

In addition to the many individuals listed by 
Solem (1976b, pp. 5-6), the following people have con- 
tributed significantly to the success of this project. For 
access to specimens in collections, suggestions as to 
collecting localities, loan of materials, and many help- 
ful ideas, I am indebted to Dr. Frank Climo, National 
Museum of Natural History, Wellington, New Zealand; 
Drs. Edmund Gittenberger and A. C. van Bruggen, 
Rijksmuseum van Natuurlijke Historie, Leiden; Dr. 
Jackie van Goethem, Institut Royal des Sciences 
Naturelles de Belgique, Brussels; Dr. Simon Tillier, 
Museum National d'histoire Naturelle, Paris; Dr. 
Peter Mordan, British Museum (Natural History), 
London; Dr. Barry Wilson, Dr. Fred Wells, Ms. Shirley 
Slack-Smith, Mr. George Kendrick, Western Austra- 
lian Museum, Perth; Dr. Winston Ponder, Australian 
Museum, Sydney; and Mr. and Mrs. Fred Aslin, Mt. 
Gambier, South Australia. 

For assistance in the field, or supplementary 
fieldwork by, I continue to be deeply in debt to Mr. 
Laurie Price, Kaitaia, New Zealand, Field Associate of 
Field Museum of Natural History. Mr. Carl Christen- 



sen, University of Arizona; Dr. Barry Wilson, Shirley 
Slack-Smith, George Kendrick, Clayton Bryce, and 
many other staff members of the Western Australian 
Museum; Fred and Jan Aslin, Mt. Gambier, South 
Australia; Dr. Brian J. Smith, National Museum of 
Victoria, Melbourne; Dr. Winston Ponder, Australian 
Museum, Sydney; and Mr. Ron Kershaw, Launceston, 
Tasmania, have assisted in Australian fieldwork or 
have provided critical help with collecting localities. 

Charts and figures in the introductory sections 
have been prepared new or reworked by Ms. Elizabeth 
Liebman, Illustrator, Division of Invertebrates, Field 
Museum of Natural History. Invaluable help with 
mounting and labeling was provided by Mrs. Dorothy 
Karall, Associate, Division of Invertebrates. Illus- 
trations for the systematic review were done primarily 
by the several artists cited by Solem (1976b, p. 6). The 
credit for the individual figures is indicated by the fol- 
lowing abbreviations: Ms. Nelva Bonucchi (NB); Mrs. 
Jane Calvin (JC); Mrs. Carole W. Christman (CW); Mr. 
Sam Grove (SG); Mrs. Lynda Hanke (LH); Mr. Sander 
Heilig (SH); Dr. Harold J. Walter (HJW); Mr. E. John 
Pfiffner (EJP); Mrs. Patricia Rill Smiley (PS); Ms. 
Marcia Oddi (MO); and the late Mrs. Margaret Anne 
Moran McKibben (MM). In particular, the work by the 
late Margaret Anne Moran McKibben was critical to 
the success of this project. 

Tabular material in the systematic review was 
typed by Mrs. Sandra Rendleman and Ms. Jayne 
Freshour. Tabular material in the introductory sec- 
tions was typed by Mrs. Valerie Connor-Jackson, Sec- 
retary, Division of Invertebrates. 

Grateful acknowledgement is made to the Na- 
tional Science Foundation, whose generous support 
through grants G- 16419, GB-3384, GB-6779, and DEB 
75-20113 in support of fieldwork, illustration prepara- 
tion, and museum study of types has made this study 
possible. The establishment of a scanning electron 
microscope laboratory at Field Museum of Natural 
History was funded by National Science Foundation 
grant BMS 72-02149. Publication of Part I (Solem, 
1976b) was in large part made possible by National 
Science Foundation grant DEB 75-14048. 

Much help with final editing and complete typing 
of the manuscript was given by Ms. Sharon Bacoyanis 
and Mrs. Valerie Connor-Jackson. Their efficient in- 
volvement in every phase of final preparation was crit- 
ical. Proofreading and indexing was greatly aided by 
Valerie Connor-Jackson. 

As with Part I, the years of labor by the late C. 
Montague Cooke, and by Yoshio Kondo, plus the 
facilities provided by the Bernice P. Bishop Museum 
during 1961-1962 and their long loan of needed mate- 
rials were instrumental to the initiation and comple- 
tion of this project. Dr. Cooke had not reviewed any of 
the taxa covered in this report, so all errors are my 
responsibility. 



PREVIOUS STUDIES 



A more extensive historical review regarding the 
Pacific Island taxa was given in Solem (1976b, pp. 7-8) 
and should be consulted for background comments. 
Previously described charopid taxa date from the 
studies of Ferussac (1840), Hombron & Jacquinot 
(1841), Mousson (1865, 1869, 1870, 1871), Cox (1870), 
Pease (1870), Garrett (1872, 1881, 1884, 1887a-c), 
Liardet (1876), Tapparone-Canefri (1883), Mollendorff 
(1888, 1900), Beddome (1889), Quadras & Mollendorff 
(1894), Smith (1897), Clapp (1923), Rensch (1937), Dell 
(1955), Ladd (1958, 1968), and Solem (1959a, 1960). Of 
the 45 previously described species-level taxa, 11 were 
named by Garrett, six by Mollendorff, five by Mousson, 
four by Quadras & Mollendorff, and four extralimital 
species by Rensch. The remaining 15 species are scat- 
tered among the other authors. The dates of species 
descriptions were summarized in Solem (1976b, p. 3, 
table III). The cluster of 1890 and 1900s taxa in the 
Charopidae reflects the activities of Mollendorff and 
his field associate, J. F. Quadras. The 10 taxa described 
since 1920 include the highly significant fossils from 
Eniwetok, Vatusila eniwetokensis (Ladd, 1958), and 
Funafuti, Lagivala davidi (Ladd, 1968), plus basically 
extralimital taxa (Solomon Islands, New Hebrides, 
Bismarck Archipelago) that happen to belong to gen- 
era also represented on the Pacific Islands. As with the 
Endodontidae, the greatest period of descriptive activ- 
ity was in the 1870s and 1880s, reflecting the publica- 
tions of Mousson and Garrett. 

No data had been recorded in the literature con- 
cerning the anatomy of the Pacific Island taxa except 
for a few notes on the tail, jaw, and radula ofRussatus 
nigrescens (Mollendorff, 1900, p. 108, figs, 1-3) and the 
radula and penis complex exterior of Sinployea 
euryomphala (Solem, 1959a, pi. 6, figs. 7-8). 

Because the main center of the charopid radiation 
is extralimital to the Pacific Islands, extensive refer- 
ence has had to be made to taxa from other parts of the 
world. New Zealand, Australian, and New Caledonian 



taxa in particular have yielded important comparative 
data. Rather than detail the many reports here, cita- 
tions are given under the organs or species referred to 
at that place in the text. Here it is necessary only to 
point out that these are diverse faunas with a long 
record of study. The literature is highly fragmentary 
and widely scattered, but fortunately there are sum- 
mary references that enable relatively quick access to 
the literature. For New Zealand, the classic mono- 
graph of Suter (1913) and the checklist of Powell (1976) 
summarize the work done prior to the highly signifi- 
cant studies of Climo (1969a-b, 1970, 1971a-b, 1978). 
For Australia, the checklist of Iredale (1937a-c) and 
his faunistic reviews of nearby areas as well as Aus- 
tralia (Iredale, 1939, 1941a-c, 1942, 1944, 1945) pro- 
vide essential but very difficult to use reports. Franc 
(1957) and Solem (1961) supplement the excellent re- 
view of New Caledonian land mollusks given by Crosse 
(1894). 

Early observations on the anatomy and radular 
structure of some Australian (Hedley, 1891, 1893) and 
New Zealand (Hutton, 1884a, Suter, 1890, 1891a-b, 
1892a-c, 1893a-d, 1894a-d, 1901, 1903, 1913) 
charopids were accurate within the limits of optical 
observation and the state of knowledge concerning 
molluscan anatomy extant at that time, but have little 
value today in deciding generic and family limits. 

Even more than for the Endodontidae, the classifi- 
cation of the charopids used previously was based on 
form "pigeon holes," and the assignments made to gen- 
era were based upon superficial conchological simi- 
larities. Of the 21 genera reviewed below, only two 
were named previously, and both of them, Punctum 
Morse, 1864, and Discocharopa Iredale, 1913, are gen- 
era whose main distributions lie outside the Pacific 
Island area. All the remaining species belong to unde- 
scribed, endemic taxa, mostly not yet known from 
other than the fringing islands and Pacific Island area 
proper. 



MATERIAL STUDIED 



The 98 taxa formally described or alluded to as 
probably representing new taxa were represented by 
about 7,500 specimens. Since the summary presented 
by Solem (1976b, p. 9), a number of charopids from the 
Lau Archipelago of Fiji collected by Price in 1970 and 
Cernohorsky in 1977, Society Islands by Solem in 1977, 
Western Australia by Solem, L. Price, and Carl Chris- 
tensen in 1976 and 1977, and near Alice Springs in the 
Northern Territory in 1974 and 1977 have become 
available. Data from them has been incorporated into 
the systematic review. No attempt at recalculating the 
data in Solem (1976b, pp. 9-10, tables IV- V) has been 
made because the numbers added are small in com- 
parison to the previously listed materials. 

Throughout the text, the following abbreviations 
are used to indicate the repository of the cited speci- 
mens: 

AIM Auckland Institute and Museum, Auckland 

AMS Australian Museum, Sydney 

ANSP Academy of Natural Sciences, Philadelphia 

BMNH British Museum (Natural History), London 

BPBM Bernice P. Bishop Museum, Honolulu 

DMW National Museum of Natural History, Wellington 

FMNH Field Museum of Natural History, Chicago 

IRB Institut Royal des Sciences Naturelles de Belgique, 

Brussels 

JDCP Collection of Journal de Conchyliologie, Paris 
MCZ Museum of Comparative Zoology, Harvard Univer- 
sity, Cambridge 
MSNG Museo Civico di Storia Naturale "Giacomo Doria," 

Genova 

NHM Naturhistorisches Museum, Basel 
NMWC National Museum of Wales, Cardiff 
RNHL Rijksmuseum van Natuurlijke Historic, Leiden 
RSM Royal Scottish Museum, Edinburgh 
SAM South Australian Museum, Adelaide 
SMF Natur-Museum Senckenberg, Frankfurt-a.-M. 
UMMZ University of Michigan, Museum of Zoology, Ann 

Arbor 

USNM National Museum of Natural History, Smithsonian 
Institution, Washington, D.C. 



WAM Western Australian Museum, Perth 

ZMA Zoologisch Museum, Amsterdam 

ZMB Zoologisches Museum der Humboldt-Universitat, 

Berlin 
Zurich Zoologisches Institut der Universitat, Zurich 

Most of the material in the BPBM resulted from 
the historical activities of Andrew Garrett plus speci- 
mens obtained during three major expeditions in the 
1930s. The Mangarevan Expedition from April 15 
through October 28, 1934 (Cooke, 1935), Micronesian 
Expedition from December 8, 1935, to June 10, 1936 
(Gregory, 1936, p. 40), and the Henry G. Lapham Ex- 
pedition to Fiji from June 27 through September 28, 
1938 (Buck, 1939, pp. 29-30), contributed major mate- 
rials for this study. Additional collections made in Fiji 
by Harry G. Ladd, plus miscellaneous samples taken 
by a variety of BPBM staff members, combined with 
the above to provide the bulk of material analyzed. 

Fieldwork sponsored by FMNH and National Sci- 
ence Foundation in Rarotonga (1964 and 1965), West- 
ern Samoa (1965), Fiji (1965 and 1970), Tonga (1965- 
1966), New Caledonia (1967), and New Hebrides (1972) 
by Laurie Price and/or Alan Solem was designed to 
sample areas that were underrepresented in the BPBM 
collections. Fieldwork in New Zealand (1962, 1965) 
and various parts of Australia (1962, 1965, 1973, 1974, 
1976-77) provided critical comparative materials that 
enabled assigning subfamily names and indicating 
possible phylogenetic trends within the Arionacea. 
Only a few of these dissections are illustrated or cited 
here because this report is deliberately restricted to the 
Pacific Islands, except where a genus extends extralim- 
itally. 

Special features regarding the methods of collect- 
ing and specimen storage used by the BPBM staff were 
reviewed by Solem (1976b, p. 10) and are not repeated 
here, although applying in full to the studied materials 
used in this report. 



METHODS OF ANALYSIS 



The basic data on distribution of size and shape 
variation within a population, how to recognize an 
adult shell, the limits of measurement reliability, and 
definitions of the standard measurements used were 
fully explained in Solem (1976b, pp. 11-15). 

It is necessary to emphasize some data concerning 
sample bias because rather significant mean size 
differences among museum lots of previously described 
species are sometimes ignored. Prime examples of this 
are in such Rarotongan taxa as Sinployea proxima 
(Garrett, 1872) (table XX), S. rudis (Garrett, 1872) 
(table XXIII), S. harveyensis (Garrett, 1872) (table 
XXII), and the Funafuti Atoll S. ellicensis ellicensis 
(table XXIX). The pattern of trading samples from 
larger series to other museums or collectors still is 
common practice among conchologists. Larger speci- 
mens will tend to be retained by the original source, 
with smaller examples being dispersed widely. The 
possible extent of such dispersal is perhaps best illus- 
trated by the endodontid land snail species Libera 
tumuloides (Garrett, 1872). Although "over 300 speci- 
mens" were collected originally, only 72 could be lo- 
cated for this study, indicating dispersion and loss or at 
least absence from major museum collections of more 
than 228 examples in less than 100 years (Solem, 
1976b, p. 430). In contrast, the endodontid Mauto- 
dontha (M.)zebrina (Garrett, 1874) was known from "a 
half dozen specimens," all of which were located 
(Solem, 1976b, p. 162). Hence the reliability of mean 
measurements made on rare species is apt to be much 
greater than the means for common species where 
trading bias probably has distorted the data base. The 
exact extent of this distortion is unknown because the 
original measurements were generally of only a single 
specimen and were cited to the nearest millimeter. It 
must be recognized that sizes cited here for pre-1900 
species may be larger than the means of the popula- 
tions from which they came, but how much larger is 
unknown. 

Data from the BPBM and FMNH collections in- 
volved measuring field samples that had not been di- 
luted by trading activities. Thus, these results are di- 
rectly comparable with each other. 

For each adult specimen, shell height, diameter, 
whorl count, umbilical width, number of radial ribs on 
the body whorl, number of apical spiral cords, and any 
information concerning number and position of aper- 
tural barriers were recorded. The only difference in 
data recording from that used in Solem (1976b) is the 
consistent counting of body whorl ribs and apical cords. 



The lack of apertural barriers in many charopids 
suggested greater use of sculptural features, and thus 
body whorl ribs were counted wherever possible. Solem 
(1976b, p. 14, fig. 5; p. 43, fig. 32; p. 52, fig. 36) defined 
each of these parameters except for the apical spiral 
cord count because the Endodontidae normally lacks 
this type of major sculpture. This count was taken on a 
suture-to-suture transect just before the end of the 
apex, provided that portion of the shell was unworn. At 
the start of the apex the exposed portion of the whorl 
profile is narrower than at the end, producing a sig- 
nificantly lower spiral ridge count. In Microcharopa 
mimula (fig. 4a), for example, the apex tip shows 15 
ribs, and the apex end, 20 ribs. Usually, if a count could 
not be made near or at the usual position, the apex was 
sufficiently worn so that no count could be obtained. 
Apical cord count could not be tallied on many speci- 
mens. It was, in general, subject only to minor error in 
tallying where countable. 

Height/Diameter (hereafter H/D) ratios, Diameter/ 
Umbilical Width (hereafter D/U) ratios, and Ribs/mm, 
of body whorl circumference were calculated, the latter 
by the formula: 

rib count on body whorl 
Ribs/mm. = 

TT x shell diameter in mm. 

The validity of this index was discussed by Solem 
(1976b, pp. 42-43). 

Means, ranges, and standard errors of the means 
were computed for each parameter except whorl count 
for each population. Some of these results are pre- 
sented in tables headed "Local Variation in ... ." 
The total variation within each species or subspecies is 
summarized in tables headed "Range of Variation in 
. . . ." Means are given, but no standard errors of the 
mean were calculated because of the wide temporal 
span (1820 to 1977) of the materials represented. 

At a later stage in this review, from eight to 12 
examples of each species, where sufficient materials 
were available, were measured as to spire elevation 
and body whorl width, with a suitable ratio (SP/BWW) 
calculated and included in the "Range of Varia- 
tion. . . ." tables. This gives an index of actual spire 
elevation undistorted by differences in the degree of 
body whorl descension behind the lip (see Solem, 
1976b, pp. 24-26). At the same time these additional 
measurements were made, the pattern of the micro- 
sculpture on the early portion of the body whorl (see 
Solem, 1976b, p. 42, fig. 32b) and the pattern of rib 
spacing were determined and added to the draft diag- 
noses and descriptions. 



SOLEM: ENDODONTOID LAND SNAILS 



Dissections of as many taxa as possible were car- 
ried out between 1961 and 1977, with several rechecks 
of structure as my understanding of the variation pat- 
terns increased. Because only fragmentary preserved 
animals were available for many species (see Solem, 
1976b, p. 19), often only data on the terminal portions 
of the genitalia could be presented. Where possible, 
usually both the entire genitalia and the penis complex 
interior plus the pallial region were illustrated. Data 
on body color, tail structure, muscle system, and radular 
structures were obtained, but have not been illustrated 
because of lesser information content for classification 
and phylogeny. Unfortunately, the bulk of the dissec- 
tion, illustration, and descriptive work was done prior 
to the availability of scanning electron microscopy as a 
research tool. Hence only a modest number of SEM 
observations are included in this study, although ex- 
tensive use has been made of the SEM in working with 
materials from extralimital areas. 

The data base consists of many shell observations 
and measurements, data from the pallial region, 
genitalia, radula, and external body features. In the 
previous report, some use was made of computer- 
generated phylogenies (Solem, 1976b, pp. 114117, 
figs. 59-61). No such attempts were made in studying 
the Charopidae, because major features such as the 
apertural barriers obviously arose independently in 
several lineages (Solem, 1973b, p. 305), there were 
sharp anatomical differences between major groups 
with highly similar shell features, and the resulting 
convergences would utterly confuse any phenetic pro- 
gram unless de post facto elimination of convergences 
was undertaken. The benefits of the computer model- 
ing outlined by Solem (1976b, pp. 114-117) were 
applied to this analysis without the work of program- 
ming and running various combinations. Mean mea- 
surement data was key-punched and numerous print- 
outs prepared, but this was essentially an aid to the 
phylogenetic analyses. Cladistic analysis was not at- 
tempted, because there are still numerous gaps in 
knowledge of the extralimital taxa. Outgroup compari- 
sons would have been sketchy or incomplete at best, 
since the large charopid and punctid faunas of New 
Zealand, Australia, New Caledonia, and associated is- 
lands occur under conditions of complex sympatry, 
with many interspecific adjustments in penial chamber 
features that make interpreting relationships exceed- 
ingly difficult. 

The result is that the proposed phylogeny has been 
arrived at by phylogenetic methodology after looking 
at a large number of characters in several organ sys- 
tems and interpreting these according to the basic 
three-tiered pattern of evolutionary events outlined in 
Solem (1978b). I have used a pragmatic rather than an 
ideologic approach. 

The problem of how to determine species limits 
when allopatric island populations are involved is 
quite difficult (Solem, 1976b, pp. 15-17). Sympatric 
congeneric species pairs or trios are invaluable in pre- 



dicting the probability that two allopatric populations 
are or are not reproductively compatible. Since the 
Charopidae, as do the Endodontidae, use the surface 
features of the penis chamber and lower female tract 
for species recognition when two closely related species 
are sympatric, there usually are demonstrable dif- 
ferences in this region of the reproductive tract. Corre- 
lated with this will be minor to prominent shifts in 
conchological structures. By determining the "mini- 
mum conchological difference" between sympatric con- 
generic species pairs, a pragmatic standard is available 
against which to measure the degree of conchological 
difference between allopatric populations. If they equal 
or exceed the "minimum conchological difference," 
they are presumed to have diverged genetically to the 
level of species. If they show slight differences that are 
below the "minimum conchological difference," they 
are judged to be conspecific or at most subspecifically 
separable. Under conditions of allopatry there are few 
to no selective pressures for changes in the species rec- 
ognition features of the genitalia. Indeed, there proba- 
bly is rather strong selective pressure for maintaining 
a conservative pattern of species recognition features 
because departure from the pattern might be a mating 
disadvantage. 

This methodology does not presume genetic link- 
age between conchological differences and species rec- 
ognition features. Quite the contrary pattern is 
hypothesized. The conchological differences are linked 
to physiological or behavioral traits that have been 
selected for by environmental pressures, producing 
eventual basic genetic incompatibility. Upon becoming 
microsympatric with a congener, strong selective pres- 
sure for species recognition changes would exist, if, as 
seems probable, nonmultiple mating is a standard pat- 
tern for the charopids. Although we have no data on 
the frequency of mating, the absence of any evidence of 
more than one set of transferred sperm during dissec- 
tion of several hundred individuals suggests that lim- 
ited mating, rather than multiple mating, is more 
probable. 

There are far fewer clear examples of congeneric 
sympatry with adequate materials for dissection in the 
Charopidae than in the Endodontidae (see Solem, 
1976b, pp. 80-83, tables LII-LIV). Partly this reflects 
the difference in the distribution and probable age 
within the area. Partly this reflects a greater vertical 
range in habitat for the Charopidae, with tree trunk to 
semiarboreal habitat available even at dryer eleva- 
tions for the charopids as contrasted with the terres- 
trially limited endodontids. Partly it is a matter of the 
Charopidae, except for Sinployea, showing rather low 
speciation but high generic diversity within the Pacific 
Island area. The one island on which an apparent mul- 
titude of species in one genus existed, Rarotonga, ap- 
parently has nine of 11 charopid species extinct. 

The example that comes closest to being sympatric 
and showing minimal conchological differences in- 
volves Sinployea peasei and S. avanaensis on Raro- 



METHODS OF ANALYSIS 



tonga. The shell of S. avanaensis (fig. 45d-f) has more 
than twice the number of radial ribs on the body whorl, 
a distinctly narrower umbilicus, and averages about 
one-quarter whorl less, although it is nearly identical 
in size and shape. Anatomically, the penis of S. 
avanaensis is 1.41.6 mm. long, compared with 1.1-1.3 
mm. in S. peasei; the verge (PV, fig. 43e) and muscle 
collar of S. avanaensis are larger than the equivalent 
structures in S. peasei (fig. 42d-e); and the pallial re- 
gion of S. avanaensis (fig. 43b) is slightly shorter, with 
a noticeably shorter and broader rectal kidney arm 
than in S. peasei (fig. 42c). The two species apparently 
have ecological separation, with S. peasei found almost 
without exception under rotting wood and stones, 
whereas, except at a very dry station, S. avanaensis 
lived in moss and lichens on tree trunks. More detailed 
collecting in Avana Valley on Rarotonga is needed to 
determine if they are actually sympatric at any point 
or if they are microallopatric in their distribution. 

In the Lau Archipelago of Fiji, for example, both S. 
inermis (Mousson, 1870) and S. adposita (Mousson, 
1870) have been collected on Lakemba, Aiwa, and 
Yangasa Levu, and fragmentary material adequate for 
dissection of both species from a single island was 
available. The penis length of S. inermis is 0.8-1.0 
mm. compared with 1.65 mm. for S. adposita, and the 
shells are very distinctive. The differences are larger 
than in the Rarotongan example. Sinployea adposita 
has a normal vergic papilla and a very small stimula- 



tory pad, whereas S. inermis has a large stimulatory 
pad and a small vergic papilla. 

On Mt. Lamlan, Guam (Station 137), both Himer- 
oconcha lamlanensis and H. fusca (Quadras & Mollen- 
dorff, 1894) were taken alive. The former has a short- 
ened penis (2.0 mm.) with major pilasters forming 
halfway from epiphallic pore to atrium, and the latter 
has a longer (2.7 mm.) penis with the major pilasters 
starting just below the epiphallic pore (compare fig. 
103a-b, d, f). These differences would serve quite effec- 
tively as species recognition devices. 

Fortunately, differences between species in the 
Pacific Island Charopidae appear to be rather striking 
and uniform, although often difficult to characterize 
verbally because of extensive local variation. 

Basically the same procedures and philosophical 
approach were used to study both the Charopidae and 
Endodontidae, so that the results are directly compar- 
able in terms of species discrimination, classification, 
and phylogenetic interpretations. The greater generic 
diversity and usually lower specific diversity in the 
Pacific Island Charopidae are interpreted as resulting 
from multiple invasions from other areas at a more 
recent time than for the Endodontidae, plus the proba- 
ble fact that the Helicarionidae and Euconulidae may 
have been invading only slightly later in time than the 
Charopidae. Conceivably, this often arboreal group of 
Limacacea could have denied the opportunity for ex- 
tensive arboreal radiation to the Charopidae. 



PATTERNS OF MORPHOLOGICAL VARIATION 



Initial studies from 1961 through 1969 on both 
shell and anatomy were carried out at 100 x magnifica- 
tion or less using a Leitz dissecting microscope. Radu- 
lar mounts in Euparol were examined unstained using 
a Leitz Ortholux compound microscope under bright- 
field, dark-field, and phase-contrast illuminations. 
Subsequently, SEM observations on both shell and 
radular features resulted in a series of reports (Solem, 
1970b, 1972a-c, 1973a-b, d, 1974, 1975, 1976a, 
1977a-b) whose results are partially incorporated here. 

None of the available anatomical material was 
suitable for histological investigation. In many cases 
the apical genitalia of the BPBM material could not be 
studied because of the preservation technique utilized 
(Solem, 1976b, p. 19). 

The following survey of shell and anatomical vari- 
ation is less detailed than that presented for the En- 
dodontidae (Solem, 1976b, pp. 19-99). Instead of one 
phylogenetic lineage being represented, the Pacific Is- 
land Charopidae include several independent lineages. 
Instead of a monophyletic origin for the apertural bar- 
riers, several independent origins are involved. Thus, 
certain of the questions asked in the endodontid review 
are not relevant and were omitted. In terms of shell 
structure, the Charopidae on the Pacific Islands simply 
are more conservative. 

SIZE AND SHAPE VARIATIONS 

At least one specimen was seen of all the species 
treated in this monograph, so that the discussion that 
follows includes all the named taxa. Specimens that 
were indicated as probably representing new taxa, but 
which were not formally named, have been omitted 
from the comments and statistical analyses. No at- 
tempt at use of factor or multivariate analysis has been 
attempted because only the relatively crude standard 
shell measurements were available. The overall pat- 
tern of mean measurements of each species follows that 
of the Endodontidae (Solem, 1976b, pp. 20-21, figs. 
6-10) except that they are more centrally clustered. 
Table I summarizes the basic distribution for nine var- 
iables or ratios. Although the extremes are well re- 
moved from the median, the distance between the first 
and third quartiles is relatively restricted. For shell 
height, diameter, and H/D ratio the first to third quar- 
tile range is 18%- 19% of the total range, for the whorl 
count it is only 13%, for apical cords only 16%, and for 
the number of ribs on the body whorl (assuming the 
maximum as 200) and body width it is 22.5%. Consid- 



TABLE I. - RANGE OF VARIATION IN THE PACIFIC ISLAND CHAROPIDAE. 



1st 
Minimum Quartile 



3rd 
Quartile 



Means of: 






















Height 


0. 


48 


1. 


23 


1. 


51 


1 


.81 


3, 


69 


Diameter 


1. 


07 


2. 


27 


2. 


76 


3 


,49 


7, 


52 


H/D Ratio 


0. 


365 


0. 


486 


0. 


523 





.566 


0. 


801 


Whorls 


3 


1/8+ 


3 


7/8 


4 


1/8- 


4 


3/8- 


6 


5/8 


D/U Ratio 


2. 


03 


3. 


44 


3. 


94 


4. 


73 


CLOSED 


Ribs on 
body whorl 


18 


.9 


74 


.0 


91 


.1 


115.0 


REDUCED 



Ribs /mm. 

Body Whorl 
Width 



1.70 
0.32 



Apical Cords 6.5 



7.66 

0.76 

9.5 



11.25 

0.90 

10.5 



14.61 

1.15 

12.0 



REDUCED 
2.06 

22.0 



ering that four phyletic lines are clumped, this is a 
quite narrow basic range of variation. It is also much 
less than in the Endodontidae (Solem, unpublished 
data). The extent to which this results from distortions 
of the brood chamber formation as opposed to the much 
larger number of taxa and greater age of the Endodont- 
idae is unknown. 

Body whorl contour in the Charopidae shows the 
same type of variation found in the Endodontidae but 
differs in frequencies. Table II contrasts the two 
families, using the same definitions as for the char- 
acter states in the Endodontidae except for omitting 
the brood chamber condition that has no Charopidae 
equivalent. Lateral compression, angulation, or cari- 
nation of the periphery is far more prevalent in the 

TABLE II. - BODY WHORL CONTOUR IN ENDODONTIDAE AND CHAROPIDAE. 



Total Number of Taxa: 



Endodontidae 
38 
43 
21 



Body Whorl Contour 

Laterally compressed 
Evenly rounded 

Flattened above and/or 
below rounded periphery 

Angled periphery 14 

Carinated periphery 28 

State not recorded for several taxa. 



Charopidae 

6 

41 
39 

1 
3 



PATTERNS OF MORPHOLOGICAL VARIATION 



Endodontidae. The Charopidae almost all have either 
an evenly rounded periphery or a mild to prominent, 
Sinployea canalis (Garrett, 1872) (fig. 49b), lateral flat- 
tening above a rounded periphery. A very few species, 
such as Sinployea planospira (Garrett, 1881) (fig. 46e), 
have a clearly laterally compressed periphery. Only 
Sinployea angularis (fig. 64e) has an angled periphery, 
whereas Himeroconcha quadrasi (Mollendorff, 1894) 
(fig. 105b), H. fusca (Quadras & Mollendorff, 1894) (fig. 
105e), and especially Maafu thaumasius (fig. 76b) have 
carinated peripheries. These differences in peripheral 
contour obviously are not exclusive and have no value 
in trying to characterize the families, except to point 
out another area in which their average pattern of 
growth diverges. In the Endodontidae there was a 
noticeable effect on shell diameter and H/D ratio with 
changes in body whorl contour (Solem, 1976b, p. 23, fig. 
14). The same pattern applies to the few Charopidae 
with drastically altered contours, but the numbers are 
too few to warrant statistical treatment. 

Because there are significant differences between 
the families in terms of spire protrusion, a comparison 
is given on pp. 38-40. The only really high-spired 
charopid is Ba humbugi (fig. 74b). The only truly 
sunken spire in the Charopidae is found in Roimontis 
tolotomensis (fig. 91a), three additional species have 
the spire depressed below the body whorl (Maafu 
thaumasius, fig. 76a-b; Himeroconcha lamlanensis, 
fig. 104d-e; Kubaryiellus kubaryi, fig. 89a-b), and 11 
taxa have the spire barely emergent to slightly de- 
pressed or actually variable in the case of Lagivala 
demani (Tapparone-Canefri, 1883). The differences in 
spire elevation are so minor that no analyses of possi- 
ble effects on basic measurements are presented. 

Umbilical shape in the Charopidae is monoton- 
ously V-shaped, becoming saucer-shaped when more 
widely open as in Discocharopa (fig. 36c) and nearly 
U-shaped in the few species where it is narrowed (figs. 
41c, 51f). There is none of the rich variation seen in the 
Endodontidae (Solem, 1976b, p. 27, fig. 16), hence only 
a few comments were presented in the family compari- 
sons (p. 40). 

Many of the Charopidae tend to develop a columel- 
lar and basal callus upon cessation of growth. This has 
the effect of covering over threadlike apertural bar- 
riers on the columellar wall and basal lip. Much of the 
variation in presence or absence of a threadlike col- 
umellar barrier recorded in Table V probably is di- 
rectly caused by the degree of basal callus formation. It 
shows particularly clearly in the illustrations of Pal- 
line biakensis (fig. 96b), P. micramyla (fig. 95d), P. no- 
terapalauana (fig. 94e), and Vatusila kondoi (fig. 82b). 
In complete contrast, taxa such as Kubaryiellus 
kubaryi (Mollendorff, 1900) and Russatus nigrescens 
(Mollendorff, 1900) (fig. 89b, e) totally lack any trace of 
such a callus. 

A final comment is required concerning the de- 
velopment of sulci on the whorls and the very unusual 
canalization of the sutural area. In the Endodontidae 



(Solem, 1976b, pp. 51-52) development of a subsutural 
and/or subperipheral sulcus is primarily associated 
with keel formation. In the two sharply keeled Char- 
opidae, Maafu thaumasius (fig. 76a-c) and Himero- 
concha quadrasi (Mollendorff, 1894) (fig. 105b), quite 
prominent sulci are present both above and below the 
periphery. In H. fusca (Quadras & Mollendorff, 1894) 
(fig. 105e), which is much less sharply keeled, only a 
weak supraperipheral sulcus is present. In the one 
species with angulated periphery, Sinployea angularis 
(fig. 64e), there is a clearly defined subsutural sulcus, 
and in both Vatusila kondoi (fig. 82b) and Sinployea 
rudis (fig. 47e), which have rounded peripheries, there 
are distinct supraperipheral sulci. They are the only 
equivalent to the unusual sulcus formation reported in 
the endodontid genus Anceyodonta (Solem, 1976b, p. 
51, figs. 81d-e, 83, 86a, c, 88a, c, 89a). The functional 
significance, if any, of this last development is un- 
known. 

A most unusual modification involves canalization 
of the suture. This is more clearly seen in Sinployea 
proxima (Garrett, 1872) (fig. 46a-b) where it is accom- 
plished by slight detachment of at least part of the body 
whorl and creation of a narrow channel at the suture. 
This also occurs in Lagivala macroglyphis (Rensch, 
1937) (fig. 81d-e), where the channel is formed by de- 
tachment to a greater angle and thus is shallower and 
wider, and to a barely detectable extent in L. micro- 
glyphis (Rensch, 1937). The same phenomenon is seen 
in the New Caledonian species Andre francia alveolus 
(Gassies, 1881) (Solem, 1961, pp. 454-456, fig. 12) and 
to a lesser extent in some Australian and New Zealand 
taxa. There is no equivalent change in the Endodont- 
idae. 

SHELL SCULPTURE 

Data on the basic method of sculpture formation, 
apical sculpture type, and sculpture spacing is pre- 
sented on pp. 40-41 as part of the family comparison 
discussion. In general the postapical sculpture in the 
Charopidae is relatively uniform in composition. It 
consists of major ribs, a few microradials in between, 
and fine microspiral segments that act as lateral sup- 
ports to the microradials (figs. Id-e, 2b, d), much in the 
same way as the buttresses to the microradials found 
in Ptychodon microundulata (Suter, 1890) (Solem, 
1970b, pi. 59, figs. 9-11). The latter are quite different 
from the microspirals found in the Endodontidae, but 
this difference can be investigated only with the SEM. 
In some of the charopid species with reduced radial 
sculpture, such as Himeroconcha, these microspirals 
become enlarged into essentially continuous spiral 
cords. Again, this is far less specialized than in some of 
the situations found in the Endodontidae. 

As an example of the specializations in microspiral 
elements, compare Figures Id-e, 2d, and 3b, e. In 
Sinployea modicella (Ferussac, 1840) (fig. Id-e) the 
microspirals are on both sides of the radial riblets and 
tend to be joined medially and form minor cordlets. In 




FIG. 1. Shell microsculpture in Sinployea modicella (Ferus- 
sac). Station 893, Faatoai Valley, Moorea, Society Islands. BPBM 
150377: a-b, apical and early postnuclear sculpture, note radial 
element on apex in a (295x) and secondary spiral "cording" in b 
(310x); c, detail of apical sculpture, note minute radial undula- 
tions and even nature of spiral cords (970 x); d, postnuclear 
sculpture between two major ribs on body whorl (960 x); e, detail 
of microsculpture, note uneven nature of microspiral buttresses 
(2,950x). 



PATTERNS OF MORPHOLOGICAL VARIATION 



11 



contrast, Tuimalila pilsbryi (fig. 2d) shows some of the 
same kind, but in the same inter-rib area it can have 
the microspirals as buttresses on only one side of the 
microradial. Palline biakensis (fig. 3b, e) and Micro- 
charopa mimula (fig. 4a) have very minor actual 
surface microspiral elements (fig. 3e), simply a very 
narrow support ridge leading up to the top of the mi- 
croradial whose upper edge then expands into an elon- 



gated bead (fig. 3e) upward along the riblet. These 
"beads" appear, when viewed at lower magnification 
(fig. 3b), to form microspiral "cords" that at optical 
magnification are equivalent in aspect to the serrated 
edges on the microriblets in the endodontid Austral- 
donta raivavaeana Solem (1976b, p. 33, fig. 23a-d). In 
both situations the spiral sculpture is barely visible at 
100 x magnification, and determination of the compo- 







FIG. 2. Shell microsculpture in Sinployea peasei and Tuimalila pilsbryi: a-b, Sinployea peasei. Station 14, Avana Valley, Rarotonga, Cook 
Islands. FMNH 144613. a, apex and early postnuclear sculpture (320 x); b, suture between apex and first postnuclear whorl, note beaded 
secondary spiral "cording" and waved effect of apical cords caused by conforming to minute radial undulations on surface ( 1,065 x); c-d, 
Tuimalila pilsbryi. Station T-22, east side of Eua, Tonga. FMNH 152378. c, Apical and early postnuclear sculpture, note definite, widely spaced 
large radial swellings on apex, reduced size of spiral cords (320x); d, postnuclear sculpture on body whorl, note varying height of microspiral 
buttresses (l,065x). 





FIG. 3. Shell microsculpture in Palline biakensis. Hospitaal- 
grot, Biak, West Irian. Holotype. Rijksmuseum van Natuurlijke 
Historie, Leiden: a, apical and early postnuclear sculpture (275 
x); b, 1st and 2nd postnuclear whorls (280x); c, detail of apical- 
1st postnuclear whorl suture (930x); d, end of apical shell 
sculpture (920x); e, detail of sutural area between 1st and 2nd 
postnuclear whorls (910 x). 



12 







FIG. 4. Shell microsculpture in Microcharopa mimula. Cen- 
tral peak, Mothe, Lau Archipelago, Fiji. BPBM 78585: a, apex 
and part of 1st postnuclear whorl (295x); b, suture between apex 
and 1st postnuclear whorl (970x); c, center of apex (980x); d, 
center of apex (2,935x); e, detail of apical ridge (9,850x). 




13 



14 



SOLEM: ENDODONTOID LAND SNAILS 



nents creating the visual effect requires l,000x SEM 
viewing. 

The transition between apical and postapical 
sculpture is quite abrupt (fig. 3d). There is a very short 
distance of stuttering apical growth, then a start of the 
typical postapical sculpture immediately. No dif- 
ferences in this pattern were seen in other species 
examined with the SEM. 

Rib counts were made on the body whorl of all 
unworn adult specimens. The pattern of mean rib 
counts and rib spacing is presented in Table I and also 
discussed in the family comparison section. The ques- 
tion of the utility of rib counts is covered by Solem 
(1976b, pp. 39-44), and justification of this is not re- 
peated here. Data on shell diameter and rib spacing 
correlations are presented in Table III. The correlation 
between shell diameter and rib spacing is less tight 
than in the Endodontidae (Solem, 1976b, p. 44, table 
XV), probably reflecting in part the smaller number of 
species involved. Rib reduction is discussed elsewhere 
(p. 9) and also is not as tightly correlated with size as in 
the Endodontidae (Solem, 1976b, pp. 47-50). 

In the Endodontidae and the Charopidae there are 
a few situations in which the major ribs have become 
grossly enlarged and widely spaced. Although none of 
the charopids equals the spectacular "wings" of Zyz- 
zyxdonta alata Solem (1976b, p. 466, fig. 198a-c), one 
does come close to the huge ribs of Cookeconcha stell- 
ulus (Gould, 1844) (Solem, 1976b, p. 218, fig. 93). 
Palikirus cosmetus (fig. 91d-f) from Ponape is the one 
example of spectacular rib enlargement in the 
Charopidae. Even when a keel is formed and the shell 
enlarged as in Maafu thaumasius (fig. 76a-c), the ribs 
themselves remain relatively small. When there is 
great size enlargement of the shell in such endodontid 
genera as Gambiodonta (Solem, 1976b, pp. 435-442, 
figs. 186-189), sometimes (G. mirabilis Solem, 1976; G. 
grandis Solem, 1976) there is clear rib enlargement. 

Variation in the number of apical cords between 
charopid species is relatively small and generally is 



TABLE III. - SHELL DIAMETER AND RIB SPACING IN THE CHAROPIDAE. 



Ribs/nun. 

LESS THAN 2 

2.0-2.99 

3.0-3.99 

4.0-4.99 

5.0-5.99 

6.0-6.99 

7.0-7.99 

8.0-9.99 

10.0-12.99 

13.0-19.99 

MORE THAN 20 



Number of 
Taxa 



2 
1 
2 
5 
4 
6 
5 

14 

26 

14 

6 



Mean and Range 
of Shell Diameter 
in mm. 



6.06(4 

2.40 

4.00(3 

4.88(3 

3.29(2 

3.45(2 

2.69(1 

3.06(1 

2.61(1 

2.29(1 

1.91(1 



59-7.52) 

29-4.70) 
49-5.68) 
11-4.30) 
17-5.23) 
75-3.49) 
65-4.63) 
73-3.74) 
40-2.95) 
07-3.21) 



greatly exceeded by the variation within species. A 
range of four to six within a population or species is not 
unusual where the mean is 11 or 12, dropping to a 
range of three to four where the mean is eight or nine 
cords. Such variation in part reflects accidents of post- 
nuclear whorl attachments that can change the count 
by one or two cords, which is partly what happens in 
terms of cord fragmentation near the upper suture and 
is partly actual variation in cord width and spacing. 
Because the intrapopulational variation is so large, lit- 
tle importance is attached to minor mean differences. 
Data on 83 of the 95 taxa were available. Of the re- 
maining 12, Discocharopa has only radial apical 
sculpture, three had an apical sculpture too fine to 
count optically, five had the apex too worn, and three 
had the apical whorls missing or were worn fossils. 
Table I presents the dispersal of the species means, 
which shows a rather tight central cluster. Of the five 
species with a mean of less than 8.5 spiral cords, three 
(Roimontis tolotomensis, Himeroconcha lamlanensis, 
and Kubaryiellus kubaryi) have a moderately de- 
pressed to sunken spire, Jokajdon callizonus (Mollen- 
dorff, 1900) (fig. 92a-c) may be involved in secondary 
size reduction, and only Vatusila vaitupuensis (fig. 
85a-c) shows no special features explaining the reduc- 
tion in number of apical cords. At the other extreme, 
Lauopa mbalavuana (fig. 77a-c) with 22 cords, Mi- 
crocharopa mimula (fig. 4a) with a mean of 17.8, 
Sinployea irregularis (Garrett, 1872) (fig. 65d-f) with 
18.5, S. recursa (fig. 64a-c) with 15.9, S. angularis (fig. 
64d-f) with 14.6, and Palline biakensis (fig. 96a-c) 
with 14.0 cords show no clear indication of why the 
numbers have increased. Even the very elevated spire 
of Ba humbugi (fig. 74b) did not make a noticeable 
difference in apical cords (mean 12.0). The number is 
undoubtedly much higher in the species of Tuimalila 
(fig. 2c), but these cords are nearly suboptical in size 
and are partly obscured by the radial swellings. 

Variation in the height of the apical cords can be 
studied easily with the SEM. A few examples are 
shown in Figures 1-3. The situation in Sinployea 
modicella (Ferussac, 1840) is typical (fig. la-c), with 
the height of each cord (mean 9.2 cords) being slightly 
less than its width. There is an obscure radial rugosity 
as a secondary feature. Within the same genus, 
Sinployea peasei (fig. 2a-b) (mean 11.4 cords) shows a 
reduction in both height and width of each cord, al- 
though the underlying rugosity is just as clearly out- 
lined. In Tuimalila pilsbryi (fig. 2c) the spiral cords are 
about the same size as in Sinployea peasei, but the 
secondary radial swellings confuse the viewer using 
optical equipment. Finally, in Palline biakensis (fig. 
3a-d) the spiral cords are much higher and narrower, 
and there is no trace of the radial rugosities seen in the 
Sinployea. 

Only two Pacific Island charopids show a different 
apical sculpture. Microcharopa mimula (fig. 4a-e) has 
spiral apical ribbing, but this is broken up into a series 
of short segments with peculiar open ends (fig. 4d-e). 



PATTERNS OF MORPHOLOGICAL VARIATION 



15 



At optical magnifications the sculpture appears to be 
simple spiral cords, but obviously it is much more com- 
plex. The same sculpture is found in the North Ameri- 
can Radiodiscus millecostatus (Pilsbry & Ferriss, 1906) 
(Solem, 1977b, p. 152, figs. 7-8) and an unnamed 
Western Australian genus (Solem, unpublished data). 
These genera are grouped in the charopid subfamily 
Rotadiscinae H. B. Baker (1927, p. 228). Discocharopa 
aperta (Mollendorff, 1888) (fig. 5a-c) has an apical 
sculpture of fine radial ribs that become very crowded 
(fig. 5a) near the end of the apex. Higher on the apex 
(fig. 5b-c) there are irregularly spaced periostracal 
spiral folds that add a partial spiral element. The post- 
apical whorls (fig. 5d-f) show no trace of any spiral 
sculpture. 

In summary, the Pacific Island Charopidae mostly 
have a very characteristic and conservative pattern of 
sculpture. The apex has a variable number of spiral 
cords to which radial swellings are occasionally added. 
The postapical whorls have major radials whose spac- 
ing and number are somewhat correlated with shell 
diameter, finer microradials in between, with very fine 
microspiral units that buttress or connect the microra- 
dials. Most of this sculpture is formed by the perio- 
stracum (p. 137). Reduction in prominence or loss of 
sculptural features is an apparently rare event. 

The situation is much more complex in relation to 
the New Zealand and Australian charopid taxa. 

APERTURAL BARRIERS 

Traditionally, those endodontoid taxa with aper- 
tural barriers were placed in the form genus En- 
dodonta, and those without barriers, in the form genus 
Champa (Solem, 1976b, pp. 118-119). The discovery 
that taxa with apertural barriers showed family-level 
differences in anatomy is one of the more significant 
results of this study. Solem ( 1973b) reviewed the bar- 
rier frequency and structure in the two families and 
concluded that although the endodontoid barriers are 
of common origin, those found in the Pacific Island 
Charopidae have evolved at least four times. Thus, in- 
stead of considering monophyletic variation, as in the 
equivalent discussion of the endodontid barriers 
(Solem, 1976b, pp. 52-72), an obviously polyphyletic 
situation exists. 

Such barriers are a common phenomenon in many 
land snail families (Solem, 1972c), but the timing of 
their appearance in ontogeny, methods of formation, 
state in adulthood of the individual, positions in the 
aperture, length, and numbers differ radically. They 
are analogous rather than homologous developments. 
Retention of specialized terms for the different 
structures found in separate families is fully justified. I 
have chosen to use a uniform terminology for the bar- 
riers in the Endodontidae and Charopidae, despite 
their independent origins, for ease in comparisons and 
because the barriers have the same patterns of change 
and growth. In both families the barriers appear at or 



very shortly after hatching, are added to anteriorly and 
resorbed posteriorly during subsequent growth, and 
often effectively narrow the aperture in adulthood. 

Recognition of parietal, columellar, and palatal 
zones in the aperture, the differences between major 
barriers and small traces, and the numbering sequence 
[parietals (upper to lower), columellars (upper to 
lower), palatals (lower to upper), all in numerical se- 
quence regardless of possible homology] have been 
given by Solem (1976b, pp. 52-54, figs. 36-38). 

The occurrence of barriers in the Charopidae is 
neither taxonomically nor geographically random: 32 
(33.7%) of the 95 taxa have parietal and 27 (28.4%) 
have palatal barriers. 1 These are clustered in 11 of the 
20 genera. Lauopa, Discocharopa, Ladronellum, and 
Roimontis are monotypic, Jokajdon and Palikirus have 
two species, Graeffedon and Palline have three, Sem- 
perdon has five, and Lagivala and Vatusila have six 
species. No species in these genera are without bar- 
riers, except for Palikirus ponapicus (Mollendorff, 
1900), which is known from a single, fungus-covered 
specimen and was associated with Palikirus as a con- 
venience, and great secondary reduction of the barriers 
in Vatusila niueana (fig. 83e). Five of the 32 species are 
basically extralimital: Lagivala macroglyphis (Rensch, 
1937) and L. microglyphis (Rensch, 1937) from New 
Britain; L. demani (Tapparone-Canefri, 1883) from 
Ambon, Timor, and parts of West Irian; Palline biaken- 
sis from West Irian; and Discocharopa aperta (Moll- 
endorff, 1888), which ranges from the Philippines to 
Society Islands. The three Graeffedon are limited to 
Western Samoa; the Lau Archipelago has two species 
(Vatusila kondoi and V. nayauana) on Nayau and 
Lauopa mbalavuana on Vanua Mbalavu and two 
species on Viti Levu (Lagivala vivus and L. minus- 
culus). Vatusila tongensis on Eua, Tonga; Lagivala 
davidi (Ladd, 1968) fossil on Funafuti and Vatusila 
vaitupuensis live on Vaitupu, Ellice Islands; V. 
niueana on isolated Niue Island; and V. eniwetokensis 
(Ladd, 1958) fossil on Eniwetok in the Marshall Islands 
complete the western distribution. A second cluster of 
taxa occurs on some of the Caroline, Palau, and 
Mariana Islands. On Ponape there are Palline mi- 
cramyla, Jokajdon callizonus (Mollendorff, 1900), J. 
tumidulus (Mollendorff, 1900), Palikirus cosmetus 
(Mollendorff, 1900), and Roimontis tolotomensis. On 
the islands of Palau, the three subspecies of Palline 
notera, Semperdon uncatus, S. xyleborus, and S. 
kororensis (Beddome, 1889), are found. In the Mariana 
Islands S. heptaptychius (Quadras & Mollendorff, 
1894) and Ladronellum mariannarum (Quadras & 
Mollendorff, 1894) live on Guam, and Semperdon 
rotanus occurs on Rota and the northern tip of Guam. 



'This differs from the figure cited by Solem (1973b, p. 301) be- 
cause Discocharopa aperta (Mollendorff, 1888) has some populations 
with and many without a parietal barrier, so this species was omitted 
from that calculation. 




FIG. 5. Details of shell sculpture in Discocharopa aperta (Mollendorff). Busuanga, Philippine Islands. FMNH 18725: a, shell sculpture at 
apical-postnuclear boundary (l,000x); b, apical and postapical sculpture (l,035x); c, details of apical sculpture (3,000x); d, postnuclear 
sculpture (l,015x); e, postnuclear sculpture showing serrated edging (3,000x); f, detail of postnuclear sculpture showing absence of spiral 
elements (9,950x). 

16 



PATTERNS OF MORPHOLOGICAL VARIATION 



17 



Except for the scattered occurrence of Dis- 
cocharopa aperta (Mollendorff, 1888) in the Austral Is- 
lands and on Borabora, Society Islands, the entire Cook 
and Society Islands area of great charopid diversity is 
free from barrier-bearing genera. Only the one taxon is 
known from Tonga, although several Sinployea and 
Tuimalila live in this group. Both the Lau Archipelago 
and Viti Levu have more taxa without than with bar- 
riers, as does Samoa. In Micronesia, Guam has the en- 
demic barrierless Himeroconcha, and Ponape has two 
genera, Kubaryiellus andRussatus, that lack barriers. 
The Palau group is unique in having only barrier- 
equipped taxa in both the Endodontidae and Charo- 
pidae. On some other Caroline Islands, there are 
barrier-free taxa, and on both Funafuti and Vaitupu 
there are species of Sinployea. The extension of 
barrier-equipped taxa to the Marshall Islands repre- 
sents their only penetration beyond the barrier-free 
taxa. 

A summary of the evidence that the barriers in 
these genera represent independent derivations was 
given in Solem (1973b). The uniform pattern of micro- 
armature and growth in the Endodontidae was re- 
viewed both in Solem (1973b) and in more detail sub- 
sequently (Solem, 1976b, pp. 54-66, figs. 39-42). In 
brief, the tops of the endodontid barriers normally are 
modestly expanded into a cordlike shape, with very 
fine triangular points (Solem, 1976b, figs. 39e, 40, 
41a-c, 42a-f) that are added to the surface. They are 
not parts of basic crystal layers that simply extend 
outward, but are an extra feature. Checking the re- 
sorption edge of a barrier (Solem, 1976b, p. 64, fig. 40) 
shows that they, at times, will be covered up by sub- 
sequent growth as the height of the barrier is in- 
creased. 

The patterns in the Charopidae are only partly 
studied. At the time that most of the material used was 
on loan for study (1962-1967), the SEM was not yet 
available. In some of the genera, Lagiuala, Vatusila, 
and Graeffedon, specimens are few in number and the 
barriers too deeply recessed for adequate SEM study 
without breaking the shell. Thus the following com- 
ments must be viewed as only introductory. 

In Jokajdon callizonus (Mollendorff, 1900) (fig. 
6a-d) from Ponape, views of the third palatal barrier 
demonstrate the basic structures in one species com- 
plex. In vertical view of the posterior descending face, 
crest, and upper anterior descending face (fig. 6a), the 
etched resorption plane is on the left, whereas the right 
shows the larger platelets on the upper anterior slope 
of the barrier. At higher magnification (fig. 6b), the 
mixture of large and small platelets on the upper an- 
terior face is seen in vertical view. In Figure 6d, the 
same area is shown in lateral view to demonstrate that 
the platelets are simply prolonged crystals from the 
growth face of the barrier. Figure 6c, at the point 
where the sharp anterior descension of the barrier 
ends, shows that there is an abrupt shift from large to 
small platelets. At this point, the combined major func- 
tion of gripping surface for extending the body from the 



shell aperture and providing setal catching projection 
against arthropod entry becomes much less significant, 
and hence the probable shift from large to small 
platelets. 

Looking at the first palatal barrier in the Western 
SamoanGraeffedongraeffei (Mousson, 1869) (fig. 7a-c) 
shows a second pattern of structure. The specimen was 
taken alive, and the shell was partly fractured during 
dissection. The barrier surface was not worn, so that 
the structure of long, straight, projecting ridges across 
the anterior descending face of the barrier is normal. 
There is distinct variation in the length of the ridges 
(fig 7b-c). The most intriguing aspect is seen on the 
side of the barrier (fig. 7a). The individual pockets hold 
platelets at a variety of angles, sometimes protruding, 
sometimes almost vertical in orientation. I consider it 
possible that they represent fragments from the ad- 
sorption surface that have been transported and re- 
fixed in a near anterior position. Unfortunately, 
specimens of Graeffedon are quite rare, and it is im- 
probable that material sufficient to test this idea will 
become available for study. 

The Palau Island Semperdon xyleborus (fig. 8a-e) 
has long parietal barriers, with the upper parietal 
posteriorly bifurcate (fig. 8a). The upper edge of the 
third parietal (fig. 8b-c) shows a few broad, projecting 
plates that point anteriorly. Some of them stretch 
across almost the entire edge. In lateral view (fig. 8d), 
the edge of the fourth parietal shows that the plates 
become much smaller on the side of the barrier and 
that the large plates are growing from the barrier sur- 
face as it slants slightly downward. They are partly 
elevated from the surface. 

The basic difference between the three types can 
be summarized as: ( 1 ) crystal layers simply continued 
directly forward and broken into microplatelets (Jo- 
kajdon); (2) anterior added ridges that may be in part 
transported as large crystals (Graeffedon); and (3) 
broad ridges angled up from the surface layers (Sem- 
perdon). Despite similar appearance and function, 
these are actually three different ways of reaching the 
same end result, i.e., a series of microprojections that 
provide gripping surfaces for the advancing mantle col- 
lar during emergence of the body from a fully with- 
drawn position. They are just as effective as the fourth 
solution, the triangular points in the Endodontidae, 
but are not as elegantly constructed. Coupled with the 
very obvious size and position differences of the bar- 
riers in the three charopid examples cited, which are 
typical of their anatomical groupings, I conclude that 
they are of independent origin. When compared with 
the apertural barrier microstructure in other families 
(Solem, 1972c), the charopid experiments show greater 
individual variation and are much less "finished" in 
appearance, even though the use of angled plates is a 
more common pattern than the addition of points or 
barbs on the surfaces. 

The presence of barriers in the Charopidae is a 
polyphyletic and secondary character, rather than the 
norm as in the Endodontidae. Thus, the comparative 



SOLEM: ENDODONTOID LAND SNAILS 




FIG. 6. Microstructure of largest palatal barrier mJokajdon callizonus (Mollendorff). Station 118, Ponape, Caroline Islands. BPBM 154161: 
a, vertical view onto top of largest palatal at l,030x, note weak etch marks on upper left and the clear cross-strata pattern on entire posterior 
(left margin); b, area of sharp anterior slope in vertical view at 3,200x; c, area at bottom of sharp anterior descension showing transition from 
large crystal plates to small crystals at 3,125x; d, lateral view at 3,050x of large crystal area showing how these plates are continuations of the 
vertical growth layers. 



treatment will be different than in Solem (1976b). 
Correlations among barrier condition and other shell 
features are not as simple and direct as in the En- 
dodontidae. For example, the relative degree of aper- 
ture narrowing in the Endodontidae (Solem, 1976b, p. 
71, table XLVIII) and Charopidae (table IV) is quite 
different. The percentages of the barrier-possessing 
taxa that are intermediate in closure differ greatly. In 
the Endodontidae, 57.5% are intermediate, but only 
25.0% in the Charopidae. The latter have the barriers 



TABLE IV. - DEGREE OF APERTURAL NARROWING BY BARRIERS 



Strong 

Moderate 

Weak 

Not 

Total Taxa 



Endodontidae 

19.9% 
40.9% 
16.6% 
22.6% 
181 



Charopidae 

43.7% 

9.4% 

15.6% 

31.3% 

32 



PATTERNS OF MORPHOLOGICAL VARIATION 



19 







FIG. 7. Surface of largest palatal barrier in Graeffedon graeffei 
(Mousson). Station 39, Mt. Solaua, Upolu, Western Samoa. FMNH 
153420: a, angled view of front edge and inner side showing different 
lateral and anterior surface texture (305 x); b, anterior edge at 1,100 
x showing projecting platelets on surface of descending edge; c, ver- 
tical view onto descending edge surface at 3,150x . 



either strongly constricting the aperture (figs. 98b, e, 
99b, d, e), or they are essentially ridgelike remnants 
(figs. 85b, lOlb). 

In both families, more species have parietal bar- 
riers than either columellar and palatal, and no species 
has either of the latter without also having the former. 

Parietal barriers 

The number of parietal barriers ranges from the 
single barrier that may be present or absent in Dis- 
cocharopa, to the four large barriers plus three traces 
found in Semperdon xyleborus (fig. 98e). The only vari- 
ation in major barrier numbers occurs in Semperdon 
heptaptychius (Quadras & Mollendorff, 1894), which 
normally has three parietals and one accessory trace. 
Occasionally (5.1%), the trace and third parietal are 
missing, rarely (0.3%), a fourth parietal is developed, 
and rarely (3.3%), the trace is missing. In Semperdon 
xyleborus, 84.6% have three traces, 15.4% have two 
traces. Occasionally, a parietal trace will be present in 
Palline notera notera and P. n. gianda, and there is one 
superior trace present in S. rotanus (fig. 99d). Other- 
wise, the number of parietal barriers is constant. 

Barrier length partly correlates with height. The 
method of determining length is explained by Solem 
(1976b, p. 70, fig. 43). Where the parietal is a thread- 
like remnant, it is Vie to Vs whorl long (figs. 83e, 85b, 
95d, lOlb), except in Palline biakensis (fig. 96b) and 
Roimontis tolotomensis (fig. 91b) where the length 
reaches 3 /ie of a whorl. In Vatusila nayauana (fig. 82e), 
Semperdon uncatus (fig. 97b), and Palikirus cosmetus 
(fig. 91e), the parietal is a raised ridge, but still short. 
The only taxa with high, short barriers are the races of 
Palline notera (figs. 94b, e, 95a) and Semperdon 
xyleborus (figs. 8a, 98e). The remaining barriers are V* 
to 3 /ie of a whorl long, except for several species of 
Lagivala where they extend to or beyond the line of 
vision from the aperture (L. microglyphis, fig. 81b; L. 
macroglyphis, fig. 81e; L. minusculus, fig. 80e). These 
species are very small in size, with mean diameters of 
1.55-1. 73 mm. 

Structure of the barriers is highly varied. The 
threadlike and raised ridge taxa present no unusual 
features and have been mentioned above. Where there 
is a single high barrier, such as Lauopa mbalauuana 
(fig. 77b), Discocharopa aperta (Mollendorff, 1888) (fig. 
35a-b), and Lagivala minusculus (fig. 80e-f), it is a 
high thin blade, usually with abrupt anterior descen- 
sion. When there are two or three high lamellae, the 
typical shape found in most of the Endodontidae is 
usual: an abrupt posterior resorption face, elevated 
middle with slightly to strongly expanded upper edge, 
and then gradual anterior descension. The second and 
fourth parietals in Semperdon xyleborus (fig. 8a) are 
typical. Several taxa have a modification of the first 
parietal that also was found in some Endodontidae. It 
can become deflected downward on the posterior por- 
tion in Jokajdon callizonus (Mollendorff, 1900) (fig. 
92b), Semperdon heptaptychius (Quadras & Mollen- 




FIG. 8. Structure of apertural barriers in Semperdon 
xyleborus. Station 184, Peleliu. Palau Islands: a, parietal barriers 
at 120x, note posterior bifidity of upper (1st) parietal: b, upper 
edge of 3rd parietal at 950x; c, upper edge of 3rd parietal at 
2,850x; d, upper edge of 4th parietal at 2,910x; e, lower palatal 
barriers at 326x . Anterior side at left margin in each photograph. 



20 



PATTERNS OF MORPHOLOGICAL VARIATION 



21 



dorff, 1894) (fig. 99b), and S. rota nut* (fig. 99d); it is 
bifid in Vatusila tongensis (fig. 83b) or posteriorly 
bifurcated in V. kondoi (fig. 82b), Jokajdon tumidulus 
(Mollendorff, 1900) (fig. 92e), Semperdon kororensis 
(Beddome, 1889) (fig. 98b), and S. xy/ebon/s (fig. 8a, 
upper barrier). Similar bifurcation was found also in 
three lineages of the Endodontidae several Cooke- 
concha sand Endodonta (Solem, 1976b, p. 379, fig. 167a, 
g), some Min idon ta (Solem, 1976b, p. 138, fig. 65b,d-e, 
g; p. 149, fig. 71c), and some Anceyodonta (see Solem, 
1976b, p. 57). The reason for these repetitive bifurca- 
tions, usually of the upper parietal barrier, is un- 
known. 

The basic similarity in shape between the larger 
parietals of the Charopidae and Endodontidae proba- 
bly indicates that this is an efficient shape, not only in 
terms of continual anterior addition and posterior re- 
sorption, but also in functioning during body extension 
of the animal. 

None of the Charopidae show the splitting into 
threadlike traces of the barriers that was common in 
several lines of the Endodontidae (Solem, 1976b, p. 57; 
p. 332, fig. 144a-c). 

The Charopidae show instead a quite conservative 
shape and number pattern of the parietal barriers. 

Columellar barriers 

Of the 32 taxa with parietal barriers, 13 lack any 
columellar barrier. This part of the shell is missing in 
Vatusila eniwetokensis (Ladd, 1958), so that its situa- 
tion is unknown. Semperdon xyleborus and S. kororen- 
sis (Beddome, 1889) (fig. 98b-c, e) have two prominent 
columellar barriers, and the former species normally 
has two accessory traces above the upper columellar. 
Six taxa, Lagivala macroglyphis (Rensch, 1937) (fig. 
81e-f), L. microglyphis (Rensch, 1937) (fig. 81b), 
Jokajdon callizonus (Mollendorff, 1900) (fig. 92b), J. 
tumidulus (Mollendorff, 1900) (fig. 92e-f), Graeffedon 
savaiiensis (fig. 88b), and Pa/line notera notera (fig. 
94b) have only one columellar. This varies from a 
threadlike ridge in the last two, to high and sometimes 
sharply ascending barriers in the other four species. 
The remaining 10 taxa (table V) show intraspecific 
variation in the number or absence of columellar bar- 
riers. When the variation is present or absent, in all 
but Lagivala demani (Tapparone-Canefri, 1883), the 
barrier is minute and threadlike. Its absence would 
mostly result from being covered by the basal callus. In 
Lagivala demani (Tapparone-Canefri, 1883) the varia- 
tion is geographic, with a large columellar present in 
specimens from Timor, medium-sized in Ambon and 
Aru Islands shells, and very small to absent in Biak 
and West Irian examples. In Semperdon rotanus and S. 
heptaptychius (Quadras & Mollendorff, 1894) (fig. 99e) 
the second columellar, when present, is a small thread- 
like trace above the larger columellar that is a high 
ridge parallel to the plane of coiling. A large form of 
the latter species (fig. 99b) occurs in which the columel- 
lar barrier is absent. 



TABLK V. - PERCKNTACK DISTRIBUTION OF COLUMKI.I.AR BARRIER NUMBERS. 



Vatusila kondoi 
V. nayauana 
Lagivala demani 
Graeffedon graeffei 



Palline notera gianda 



P. notera palauana 



Number of Columellar Barriers 






1 


5.6 


94.4 


40.0 


60.0 


* 


* 


30.0 


70.0 


84.0 


16.0 


95.0 


5.0 


83.0 


17.0 




46.5 


0.3 


42.1 


3.5 


96.5 



53.5 

56.4 



Semperdon uncatus 

S . rotanus 

^. heptaptychius 

Ladronellum roar iannarum 

* Geographically variable. 



In most taxa the columellar barrier lies parallel to 
the plane of coiling. Semperdon xylcborus and S. 
kororensis (Beddome, 1889) have one barrier slanting 
downward (fig. 98b, e); Jokajdon tumidulus (Mollen- 
dorff, 1900) (fig. 92e), Lagivala microglyphis ( Rensch, 
1937), and L. macroglyphis (Rensch, 1937) (fig. 81b, 
e-f) have it twisted up toward or almost onto the 
parietal wall; and in others, such as Jokajdon cal- 
lizonus (Mollendorff, 1900) (fig. 92b), it is a deeply re- 
cessed, triangular knob. 

Thus, the 18 species of Charopidae known to have 
a columellar barrier show almost the same total range 
of variation reported for the Endodontidae (Solem, 
1976b, pp. 57-59). 

Palatal barriers 

Five of the 32 taxa with parietal barriers lack 
palatals Lauopa mba/auuana (fig. 77b),Discocharopa 
aperta (Mollendorff, 1888) (fig. 36b), Palline biakensis 
(fig. 96b), Roimontis tolotomensis (fig. 91b), and 
Palikirus cosmetus (fig. 91e). Except for the Roimontis. 
which has two threadlike parietals, all of them have 
only a single, short lamellar parietal. Because of the 
missing palatal wall in the holotype and only known 
example, the palatal barrier situation in Vatusila 
eniwetokensis (Ladd, 1958) is unknown. In Ladronel- 
lum mariannarum (Quadras & Mollendorff, 1894) (fig. 
lOlb), many Vatusila nayauana (fig. 82e), and V. 
niueana (fig. 83e) there is a modest ridge to barely 
detectable trace on the middle of the palatal wall. 
Semperdon uncatus (fig. 97b) normally has one low 
barrier on the basal lip, but in 17% of the individuals 
this is missing, and Palline micramyla (fig. 95d) has 
two low lamellar barriers on the basal lip. Lagivala 
minusculus (fig. 80e) has a rather unique ridge with a 
lateral buttress on the basal lip, and Vatusila tongensis 
(fig. 83b) has a very large basal lip barrier that is 
bifurcated or split into two parts plus a very unusual 



22 



SOLEM: ENDODONTOID LAND SNAILS 



upper raised callus that is 0.25-0.36 mm. wide. Vat- 
usila vaitupuensis (fig. 85b) has a midpalatal thread- 
like trace and a broader ridge on the basal lip very near 
the columellar margin. 

The remaining species have a more typical array 
of normally three to seven palatal barriers, sometimes 
with additional accessory traces present. The position- 
ing of these barriers varies from at the lip edge in Sem- 
perdon (figs. 98b-c, e, 99b, d, e),Palline notera and its 
subspecies (figs. 94b, e, 95a), Graeffedon (figs. 88b, e, 
86b), and Lagivala microglyphis (Rensch, 1937) (fig. 
81b), to partly recessed in Jokajdon callizonus (Moll- 
endorff, 1900) (fig. 92b), progressively recessed in J. 
tumidulus (Mollendorff, 1900) (fig. 92e), moderately 
recessed in Vatusila kondoi (fig. 82b) and Lagivala 
macroglyphis (Rensch, 1937) (fig. 81e), and deeply re- 
cessed in both L. vivus (fig. 80b) and L. davidi (Ladd, 
1968) (Ladd, 1958, pi. 30, figs. 13-15). 

The shape of the barriers is rather variable among 
these species, although constant within a particular 
species. A simple crescent-shape with modestly to 
moderately expanded upper edge is found in Lagivala 
vivus (fig. 80b), L. macroglyphis (Rensch, 1937) (fig. 
81e), andL. demani (Tapparone-Canefri, 1883); a more 
expanded upper edge, in L. microglyphis (Rensch, 
1937) (fig. 81b) and Graeffedon pricei (fig. 88e); typical 
structure, in Vatusila kondoi (fig. 82b); very high, in 
Graeffedon graeffei (Mousson, 1869) (fig. 86b) and G. 
savaiiensis (fig. 88b); typical, in the races of Palline 
notera (figs. 94b, e, 95a); and bulbous-edged, again in 
Semperdon rotanus and S. heptaptychius (Quadras & 
Mollendorff, 1894) (fig. 99b, d-e). In Semperdon xyle- 
borus (figs. 8e, 98e), the crescent is anteriorly elon- 
gated, a process continued in the two species of Jokaj- 
don (fig. 92b, e,) and reaching the typical parietal bar- 
rier shape in Semperdon kororensis (Beddome, 1889) 
(fig. 98b). 

The predominance of the crescent-shaped palatal 
barriers outlined above is very different from the situa- 
tion in the Endodontidae. In that family (Solem, 1976b, 
p. 60, table XXXII) only five species had crescentic 
palatals, whereas 130 had bladelike barriers. In the 
Charopidae, only the two Jokajdon, Semperdon koro- 
rensis (Beddome, 1889), and possibly S. xyleborus could 
be considered to have a bladelike palatal barrier form. 

Intraspecific variation in the number of palatal 
barriers is summarized in Table VI. The great varia- 
tion found in the two Semperdon exceeds that found in 
any of the Endodontidae (Solem, 1976b, p. 67, table 
XL). None of the ratios in Table VI convincingly 
suggests genetic dominance. Assigning the most fre- 
quent percentage to represent that species, the pattern 
of palatal barrier numbers is summarized in Table VII. 
It is evident that there is considerable variation within 
genera, particularly Lagivala, Graeffedon, Palline, and 
Semperdon. Even within genera, there is no clear cor- 
relation between number of barriers and shell size. 
Only Palline notera shows a partial correlation, in that 
the two larger subspecies have fewer and smaller-sized 



TABLE VI. - PERCENTAGE DISTRIBUTION OF PALATAL BARRIER NUMBERS 



Taxon Number of Palatal Barriers 

012345 

Vatusila kondoi 11.1 88.9 

V. nayauana 40.0 60.0 

Lagivala demani * 



20.0 80.0 

10.0 75.0 10.0 5.0 
16.0 68.0 16.0 

1.2 3.0 19.8 55.0 

1.2 80.0 13.1 3.9 1.5 

92.0 



21.0 
0.3 
8.0 



Palline notera 
palauana 

V_. ri. gianda 

Semperdon uncatus 17.0 83.0 

S. rotanus 

S. heptaptychius 

. xyleborus 

* Geographically variable. 



palatals than the nominate race, which averages 
0.24-0.41 mm. less in diameter than the others. There 
is also no clear correlation between number of palatals 
and degree of apertural narrowing. 

The presence of palatal traces is sporadic. Several 
taxa will have one or two traces between lower pairs of 
palatal barriers: Lagivala microglyphis (Rensch, 1937), 
Graeffedon graeffei (Mousson, 1869), G. pricei, Palline 
notera gianda, Semperdon rotanus, and S. hepta- 
ptychius (Quadras & Mollendorff, 1894). Other taxa will 
have three traces above the upper palatal: Jokajdon 
callizonus (Mollendorff, 1900), Semperdon kororensis 
(Beddome, 1889), and most examples of S. xyleborus. A 
few of the latter will have four (7.5%) or five (7.5%) 
traces above. Jokajdon tumidulus (Mollendorff, 1900) 
normally has three traces above the upper palatal and 
three (rarely two) between lower pairs. The same pat- 

TABLE VII. - PALATAL BARRIER NUMBERS IN CONSTRICTED APERTURE CHAROPIDAE. 



3 Palatals 
Vatusila kondoi 
Lagivala demani 
Jokajdon callizonus 
J_. tumidulus 

4 Palatals 
Lagivala davidi 



Graeffedon pricei 
Palline notera gianda 
P_. _n. palauana 
Semperdon heptaptychius 



5 Palatals 

Lagivala microglyphis 
U macroglyphis 
Graeffedon graeffei 
Palline notera notera 

6 Palatals 
Lagivala vivus 



7 Palatals 

Graeffedon savaiiensis 
Semperdon rotanus 
S. xyleborus 
S. kororensis 



PATTERNS OF MORPHOLOGICAL VARIATION 



23 



tern of traces above the upper palatal is seen in the 
Endodontidae (Solem, 1976b, p. 59) and was inter- 
preted as a space-preserving function, because the 
great size of the first parietal and the need to keep this 
space open for withdrawal and extension of the buccal 
mass and foot combine to make reduction in size of 
palatal projections in this area advantageous. 

As in the Endodontidae, the normal pattern of size 
change is for the palatal barriers to decrease in height 
as they ascend the palatal wall. An obvious exception 
will be reduced size of the first palatal in situations 
where it is right at the columellar margin (fig. 99d) or 
where there is large size to the columellar and/or lower 
parietal barriers (figs. 86b, 92b, e). So few taxa show 
either recession of the palatals or major upper edge 
expansion of the barriers that discussion of shell corre- 
lations separately is not attempted. Most of the com- 
ments made about general features of the palatal bar- 
riers in the Endodontidae (Solem, 1976b, pp. 59-63) 
seem to apply to the few Charopidae also. 

Despite only 27 charopids having palatal barriers, 
they show a wide range of variations that mostly paral- 
lel the situations found in the Endodontidae. The 
biggest difference lies in the basic shape of the bar- 
riers: bladelike in 86.1%, ridged in 10.6%, and 
crescent-shaped in 3.3% of those Endodontidae with 
palatal barriers compared with bladelike in 15.4% 
(four), a lamellar or unusual ridge in 23.1% (six), and 
crescent-shaped in 61.5% (16) of the 26 charopid taxa 
for which data on the palatal barrier structure were 
available. Judging from the size of the parietal barriers 
(fig. 85e), Vatusila eniwetokensis (Ladd, 1958) probably 
had large palatal barriers, but the entire outer wall of 
the only known specimen is missing, and thus the state 
of the barriers is unknown. 

Patterns of barrier variation 

Despite the lack of size and shape correlations 
with variation in the barriers found in the Charopidae, 
a few comments can be made concerning general pat- 
terns within a particular group. In Lagivala (figs. 
80-81) the variation is geographic, with the Bismarck 
Archipelago species having the strongest apertural 
barriers, progressive reduction occurring in both L. 
demani (Tapparone-Canefri, 1883) to the west and in 
the Fijian and Funafuti taxa to the east (fig. 80b, e). 
Vatusila shows no size correlation, with the largest (V. 
tongensis, fig. 83b) and one of the smaller (V. 
eniwetokensis, fig. 85e) having quite large barriers, 
and both large (V. vaitupuensis, V. niueana, figs. 83e, 
85b) and small (V. nayauana, fig. 82e) having reduced 
barriers. There does appear to be some geographic con- 
cordance, with Tonga and Lau species having larger 
barriers than taxa from the more isolated Vaitupu and 
Niue localities. The fossil record of a large-barriered 
form on Eniwetok in the Upper Miocene gives an his- 
torical perspective to the group that otherwise is lack- 
ing in the Charopidae. 

The reasons for the reduction in barrier size in the 



races of Palline notera on Koror (palauana, fig. 94e) 
and Babelthuap (gianda, fig. 95a) compared with the 
nominate race on Peleliu (fig. 94b) is unknown, al- 
though slight size differences are involved. Barrier size 
variation in Semperdon shows no correlations that I 
could detect. The unconstricted aperture of the mean 
diameter 2.83 mm. S. uncatus (fig. 97b) contrasts with 
the highly constricted aperture in the mean diameter 
2.88 mm. S. xyleborus (fig. 98e) and mean diameter 
4.59 mm. S. kororensis (Beddome, 1889) (fig. 98b). All 
of these species have been dissected and all occur on 
Koror. The differences in barrier prominence cannot be 
explained at present. 

Summary of barrier variations 

Despite the different microstructure on the barrier 
edges and the evident polyphyletic origin of the aper- 
tural barriers in the Charopidae, in general they paral- 
lel those of the Endodontidae. The retention of parietal 
barriers extending to the lip edge, general shape, and 
multiple bifurcations of the parietals all suggest paral- 
lel function. Given the similar habitat and overlapping 
shell size, plus the same basic growth structure, this is 
not surprising. The difference in shape frequency of the 
palatal barriers, a greater tendency for loss or reduc- 
tion in the barriers, and the general shorter length of 
the charopid barriers are real average differences but 
cannot be used as criteria for family-level separation 
because there is so much overlap between families. 

SUMMARY OF SHELL VARIATIONS 

With few exceptions the range of variation in the 
shells of the Charopidae is less than that found in the 
Endodontidae. Major differences between the two fam- 
ilies are reviewed on pp. 40-44. Here it is sufficient 
to point out that the formation of sculpture by the 
periostracum rather than the calcareous layers, lack of 
umbilical and growth modifications to form a brood 
chamber, and secondary derivation of apertural bar- 
riers in the Charopidae have had profound effects on 
the range of shell variation. 

GROSS ANATOMY 

Including the Punctum sp. from Tahiti, Sinployea 
sp. from Borabora, and Sinployea sp. from Saipan, 
there are 98 species-level taxa considered in this 
monograph. Of these, 51 (52.0%>) belong to one genus, 
Sinployea. It was possible to see at least fragmentary 
soft parts for 43 (43.9%) of the total taxa and 21 (41.1%) 
of the Sinployea. The taxonomic distribution of the dis- 
sected material is given in Table VIII. Of the 21 gen- 
era, six genera, five of them monotypic, could not be 
dissected. Only Lagivala, with six species, represents 
an undissected speciose group. The taxonomic coverage 
of dissected material is broader than that given the 
Endodontidae (Solem, 1976b, p. 73, table L), where 
only 58 (32.4%) of the 179 taxa seen were dissected. 
This reflects both a lesser degree of extinction for the 



24 



SOLEM: ENDODONTOID LAND SNAILS 



TABLE VIII. - PHYLETIC REPRESENTATION OF DISSECTED TAXA. 



Total 
Taxa 



Number 
Dissected 



Family Punctidae 

Punctum 
Family Charopidae 



1 
1 

51 
1 
1 
1 
2 
6 
6 
3 
1 
1 
1 
1 
2 
2 
5 
5 
1 
A 

98 



21 
1 



1 
1 
1 
1 
1 

1 
2 
2 
5 
1 
3 



Charopidae and a difference in areas of species abun- 
dance. 

In part, the apparent greater amount of anatomi- 
cal data for the Charopidae is illusory because many 
taxa were represented only by pulled fragments. This 
reflects the method of processing and specimen storage 
used at the BPBM (see Solem, 1976b, p. 19). Most fre- 
quently, a break occurs so that the digestive gland and 
ovotestis do not pull out of the shell, but sometimes 
only the head and terminal genitalia will be extracted. 
Thus, in only 21 of the 43 taxa do I have data on the 
ovotestis structure. For 14 additional taxa I could re- 
cord data on the postapical genitalia and the entire 
pallial region. For four taxa most of the terminal 
genitalia was extracted, but only part or none of the 
pallial region, and for four taxa it was possible to re- 
cord data on penial structure only. Only partial review 
of the anatomical patterns is possible. 

The sequence of discussion follows that used for 
the Endodontidae (Solem, 1976b, pp. 72-99). Expla- 
nations and data presented there are not repeated, al- 
though fully cross-referenced. In certain cases, refer- 
ence is made to extralimital taxa that have bearing on 
subfamily assignments. Where appropriate, their 
anatomy is illustrated here to provide standards 
against which to measure the variations found in the 
Pacific Island taxa. Illustration of the anatomy for new 
Australian and Lord Howe Island taxa will be pre- 
sented elsewhere, although data from them are used to 
establish subfamily limits. 



GENITAL SYSTEM 

The organs are discussed in approximate order 
from apex of the coiled visceral hump to the atrium. 
The terminology is modified from that originated by H. 
B. Baker (1938b, pp. 6-10, 92) and follows exactly the 
usage in Solem (1976b) with a few additions for new 
structures. 

OVOTESTIS (G) The hermaphroditic gland or 
ovotestis in the Charopidae normally consists of one or 
two clumps of palmately clavate alveoli that usually lie 
buried in the digestve gland above the stomach- 
intestine junction. The only Pacific Island species that 
departs from this pattern is Discocharopa aperta (M611- 
endorff, 1888) (fig. 34a), where the bilobed ovotestis 
(G) lies alongside the stomach rather than apically in 
the digestive gland. In Sinployea (figs. 39b, g, 43c, 53g, 
57b, 60b) there may be one or two lobes, basically de- 
pending on exactly where the last branching of the 
hermaphroditic duct occurs. Generally the branching 
of the ducts is simple, but in Graeffedon graeffei (Mous- 
son, 1869) (fig. 87d) there are lateral short branches off 
the individual alveoli after the initial bifurcation. In 
Ba humbugi (fig. 75c) the ovotestis is shortened consid- 
erably. Jokajdon, Kubaryiellus, andLadronellum have 
one clump, Palline and Trukcharopa, two clumps 
typical differences are seen in Figure 90f, i. Species of 
Semperdon (fig. lOOc, g) have one or two clumps. In 
every case these clumps lie essentially parallel to the 
plane of coiling and extend apically with only an initial 
angling outward from the parietal wall where the start 
of the hermaphroditic duct lies. 

This is a very different pattern from the perpen- 
dicular to slanted orientation and division into a larger 
number of lobes seen in the Endodontidae (Solem, 
1976b, p. 74, fig. 44). Similar orientation and multipli- 
cation of ovotestis lobes is found in some Australian 
and New Zealand Charopidae, but among the Pacific 
Island Charopidae and Endodontidae genera the ovo- 
testis differences are diagnostic. 

HERMAPHRODITIC DUCT (GD) As in nearly 
all pulmonates, this duct serves to transport sex prod- 
ucts from the mass of the ovotestis to the fertilization 
area below the expanded stomach. Its length is directly 
correlated with the length of the stomach, which, in 
turn, is dependent upon the whorl count and cross- 
sectional whorl area in this part of the visceral hump. 
In Ba humbugi, which averages only 3% whorls and 
has a comparatively wide aperture (fig. 74b), the her- 
maphroditic duct (fig. 75c) is very short compared with 
most other taxa. Whether the ovotestis starts after 
union of the ovotestis collecting tubules with a sudden 
or gradual expansion appears to be possibly seasonal. 
In no species did I have samples from the same popula- 
tion taken in different months of the year available in 
order to check whether swelling is a sign of reproduc- 
tive activity. None of the Charopidae show the "kink- 
ing" of the hermaphroditic duct seen in the Endodont- 



PATTERNS OF MORPHOLOGICAL VARIATION 



25 



idae (Solem, 1976b, p. 75, fig. 45). The charopids do 
show a characteristic purple-red, iridescent sheen to 
the hermaphroditic duct that is not seen in the en- 
dodontids. 

TALON (GT) and CARREFOUR (X) The talon 
(GT) varies from a simple swelling just above the en- 
trance of the hermaphroditic duct (GD) (fig. 39e) to a 
circular head on a stalk (fig. 103e). The carrefour (X) is 
a slight swelling in the tract after reflexion of the duct. 
The same pattern holds in all the species examined 
from the Pacific Islands. As usual in both the 
Charopidae and Endodontidae, there is a partial reflex- 
ion of the hermaphroditic duct before it enters the car- 
refour. This permits the compaction of apical organs 
when the animal has retracted into the shell. The gen- 
eral pattern of a circular head on a short stalk in the 
Pacific Island charopids contrasts with the more slen- 
der head pattern in the endodontids (Solem, 1976b, p. 
85, fig. 49a; p. 373, fig. 164b; p. 471, fig. 199c), but the 
differences are small in many cases, and the variation 
in extralimital charopids is extensive. In both families 
the collecting ducts of the albumen gland empty into 
the carrefour below the level of the hermaphroditic 
duct entrance (fig. 39e; Solem, 1976b, p. 471, fig. 199c). 
No histological studies have been made of structures in 
this area. 

ALBUMEN GLAND (GG) The distance between 
the apex of the pallial cavity and the base of the 
stomach is less in the Charopidae than in the Endodont- 
idae, since the median whorl counts are, respectively, 
4Va and 5V2 + . In the Endodontidae, there is more of a 
tendency for the albumen gland to be elongated and to 
occupy only part of this area. In the Charopidae, the 
albumen gland is almost shapeless, roughly globular, 
with the surface deeply indented by intestinal loops, 
head of the spermatheca (S), esophagus, arteries, and 
stomach base. Rarely does the albumen gland dissect 
out intact, and in order to check the shape of the talon 
and carrefour plus the point of entrance for the her- 
maphroditic duct, some of the albumen gland tissue 
has to be teased away before illustration. Thus, only 
rarely can the actual shape be shown. The size of the 
alveoli appears larger in the Charopidae than in the 
Endodontidae, but no quantification of this was at- 
tempted. 

PROSTATE (DG) and UTERUS (UT) In the En- 
dodontidae (Solem, 1976b, pp. 77-78; p. 373, fig. 164d; 
p. 471, fig. 199c) the prostate and uterus are completely 
separate tubes that are only lightly bound together by 
connective tissue. In the Charopidae they are partly 
(some Rotadiscinae) to completely (most Charopinae) 
fused, sharing a common lumen, with the prostate 
channel a lateral outpocket (fig. 102e) into which enter 
the prostatic alveoli. These are much fewer in number 
than in the Endodontidae, longer, and not arranged in 
distinct rows. Generally the prostate is as long as the 
expanded uterine upper section and extends slightly 
down along the even more expanded uterine lower 



chamber. The latter tapers into the free oviduct (UV) 
without clear demarcation, so the exact relationship 
between prostate and uterine length remains to be de- 
termined by histological studies. 

The uterus has a clear separation into an upper 
(UT 1 ) thin-walled narrower chamber and a lower (UT 2 ) 
thick-walled, more expanded chamber (fig. 43c-d). 
Presumably, actual egg encapsulation takes place in 
the lower chamber. This is simpler than the situation 
in the Endodontidae (Solem, 1976b, p. 449, fig. 191b) 
where four sections could be distinguished in the 
uterus. Comparative studies on the function and his- 
tology of this area in both families would be quite 
worthwhile. Solem (1972b, pp. 108-112) reviewed the 
pattern of fused versus separated pallial gonoducts in 
the Pulmonata and concluded that the separated condi- 
tion found in the Endodontidae was more primitive 
than the fused condition found in the Charopidae and 
many European-North American families. 

TERMINAL MALE GENITALIA The organs 
that may be present in this complex are the vas defer- 
ens (VD), epiphallus (E), penial retractor muscle (PR), 
penis (P), penis sheath (PS), atrium (Y), and internal 
structures of the epiphallus and penis proper. The lat- 
ter regions are used in species recognition by the 
snails, so that highly significant variations can be 
found within genera, particularly under conditions of 
sympatry. The discussion of variations in the Endodont- 
idae (Solem, 1976b, pp. 78-83) should be read as 
background information on the general patterns before 
attempting to deal with the complexities in the 
Charopidae, where few of these organs are constant in 
shape. 

The vas deferens (VD) starts from the end of the 
prostate alveoli attachment and tapers for a variable 
distance before becoming a narrow tube that reflexes 
at the penioviducal angle to ascend the penis. The area 
of tapering can be very short as in Russatus (fig. 90b), 
long in Ladronellum (fig. 102a), extend nearly to the 
penioviducal angle as in Tuimalila (fig. 79b), or highly 
variable as in Sinployea (fig. 67a, e-f, h). The ascend- 
ing arm of the vas deferens is without unusual features 
until it reaches the next structure, which differs from 
subfamily to subfamily of the Charopidae. 

In the Charopinae the vas deferens (VD) passes 
into an epiphallic enlargement that may be apicad or 
anterior of the penial retractor muscle insertion. In 
Charopa coma (Gray, 1843) (figs. 9b, d, 10) there is an 
enlargement of the vas deferens about one-third of the 
way from the penial retractor muscle insertion. Inter- 
nally (fig. 10) it can be seen that the actual differentia- 
tion occurs some distance after the swelling has begun. 
Internal pilasters of the vas deferens (VD) enlarge, 
terminating in finger-like projections that surround a 
central cavity. Below this point, which is the start of 
the epiphallus, two high pilasters and a lower ridge 
line the inside of the epiphallic chamber down to the 
point of entry into the penis chamber (fig. 9d). The 



IE 




PC 



FIG. 9. Anatomy of Charopa coma (Gray). Waiwera-Puhoi Road, north of Auckland, North Island, New Zealand. A. Solem! 11-10-1962. 
FMNH 135420: a, left side of foot showing size relation to shell; b, genitalia; c, details of pallial genitalia; d, internal structures of penis and 
lower female tract; e, pallial region; f, details of pneumostomal area; g, bottom and top views of anterior digestive system. Scale lines as marked. 

(PS). 



26 



PATTERNS OF MORPHOLOGICAL VARIATION 



27 







FIG. 10. Interior of lower vas deferens and epiphallus in Charopa 
coma (Gray). Waiwera-Puhoi Road, north of Auckland, North Island, 
New Zealand. A. Solem! 11-10-1962. FMNH 135420. Greatly en- 
larged. (NB). 



penis retractor muscle (PR) inserts as a U-shaped fan 
around the lower part of the epiphallus (E). Various 
modifications occur extralimitally in such taxa as 
Phenacohelix pilula (Reeve, 1852) (fig. 11). Here the 
penial retractor muscle inserts on the head of the 
epiphallus, and the vas deferens enters almost later- 
ally. The shift in muscle attachment is relatively 
insignificant, but does make external recognition of 
the epiphallus presence difficult if not actually impos- 
sible. 

The Pacific Island Charopinae mostly show, in- 
sofar as it could be checked, a specialized vas defer- 
ens-epiphallus junction (figs, 57e, 79c). Exceptions are 
found in Graeffedon and the provisionally assigned 
Discocharopa (see below). A circular sphincter sur- 
rounds the epiphallic pore (DE), and a Y-shaped plug 
extends through the pore to attach onto the wall of the 
epiphallus as a circular ridge. In general, the walls of 
the epiphallus are without sculpture or have the typi- 
cal ridges (fig. 87c) seen in Charopa coma (Gray, 1843) 
(fig. 10). Because of size and preservation problems, 



only a few of the species could be checked for this 
structure, and illustrations are given only for 
Sinployea complementaria (Mousson, 1865) (fig. 57e) 
and Tuimalila pilsbryi (fig. 79c). It was observed in 
several other Sinployea. Normally the head of the 
epiphallus expands abruptly, indeed it can be bulbous 
(fig. 67a), so that external recognition is easy. 

In the Semperdoninae (figs. lOOb, e-g, 102a, 103a, 
c-d), the vas deferens remains a thin tube during its 
ascent and enters laterally into the epiphallus, with 
the penial retractor muscle inserting directly onto the 
head of the epiphallus. The exact point of entry of the 
vas deferens into the epiphallus differs. In Himero- 
concha and Ladronellum the entry occurs slightly 
below the rounded head (fig. 102d), whereas in Sem- 
perdon the entry is right next to one edge of the penial 
retractor muscle insertion (fig. lOOi). The epiphallus is 
a double-walled tube, sometimes coiled within the 
muscle sheath. The outer wall is a thin muscle sheath, 
and the inner a thick glandular tube usually rolled 
inward on one side. 

In the Trukcharopinae (figs. 90b-d, f-g, i-j, 93b-c, 
f, h) the vas deferens enters directly into the penis 
through a simple pore. In all examined genera except 
Jokajdon (fig. 93b-c) the vas deferens passes through 
the muscle fan before entering the penis. In Jokajdon 
the muscle insertion is directly on the head of the 
penis, and the vas deferens enters lateral to the inser- 
tion. This is interpreted as a secondary modification 
probably resulting from reduction in size. The Truk- 
charopinae thus have no trace of an epiphallus. 

In the Rotadiscinae (H. B. Baker, 1927, pi. 16, fig. 
14; pi. 17, figs. 22, 27) the epiphallus is less clearly 
differentiated externally, and it is either before 
(Radioconus) or after (Rotadiscus, Radiodiscus) inser- 
tion of the penial retractor muscle. The enlarged pilas- 
ters of the epiphallus are the main differentiating fea- 
tures. 

In all of the Pacific Island Charopidae examined to 
date the penial retractor muscle originates from the 
diaphragm near the apex of the pallial cavity and in- 
serts onto the penis or epiphallus. There is no shift to 
the columellar muscle equivalent to that seen in the 
Endodontidae (Solem, 1976b, pp. 81-83), which per- 
mitted elongation of the penis in such taxa as En- 
dodonta and Australdonta. In addition, generally the 
penial retractor muscle in the Charopidae is very 
short, at times scarcely a tuft connecting the dia- 
phragm and penis or epiphallus head. This is another 
correlative of the reduced whorl count in the 
Charopidae as compared with the Endodontidae. 

With a few exceptions, the external appearance of 
the penis in the Pacific Island Charopidae is for a bul- 
bous apical section that then tapers either abruptly or 
quickly to a slender stalk of variable length that joins 
the vagina to form the short atrium. The degree of the 
apical bulge depends directly upon the size and com- 
plexity of the internal pilasters and vergic structures. 
Figure 67 gives a fair sample of shape variations, and 



KD 




GD 



FIG. 11. Anatomy of Phenacohelix pilula (Reeve). Church Road, Kaitaia, Northland, North Island, New Zealand. L. Price! X-1962. FMNH 
135421: a, pallial region; b, extended foot showing caudal horn (CH) and pedal grooves (FS); c, genitalia; d, junction of hermaphroditic duct (GD) 
and talon (GT); e, internal structure of penis. Scale lines as marked. (PS). 



28 



PATTERNS OF MORPHOLOGICAL VARIATION 



29 



this is within a single genus. The major exception is 
the Semperdoninae (figs. 100, 102, 103) where a penis 
sheath, narrowed collar between epiphallus and penis, 
plus a quite different pattern of internal structure have 
combined to produce a more tubular appearance. It is 
not possible to assign a species to a genus on the basis 
of the external penis appearance. The internal struc- 
tures must be examined. They differ radically from 
subfamily to subfamily, and within a genus are subject 
to extensive modifications for species recognition pur- 
poses. A review of structure on a subfamily by subfam- 
ily basis is required. 

In the Charopinae from Australia, New Zealand, 
New Caledonia, Lord Howe Island, South Africa, St. 
Helena, and South America there is an amazing 
amount of variation that will have to be dealt with 
elsewhere. Comments here will be restricted to the 
type genus, Charopa Albers, 1860, Phenacohelix Suter, 
1892, and the Pacific Island taxa. Charopa coma (Gray, 
1843) has the penis with only a modestly swollen ex- 
terior (fig. 9b) and a very short basal stalk portion. 
Internally (fig. 9d) it shows an apical verge (PV) with 
near terminal slitlike pore, below this is a pocket 
stimulator (PC), followed by a series of variable-sized 
circular pilasters that occupy a quite low position. In 
addition, the upper and middle walls of the penis 
chamber have raised, soft pustules. Phenacohelix 
pilula (Reeve, 1852) has a much elongated and more 
slender penis (fig. lie), with shifted position of the 
epiphallus (E) to below the insertion of penial retractor 
muscle (PR). Internally (fig. lie) a small verge (PV) 
with terminal pore lies above a series of corrugated 
longitudinal pilasters and a vague swelling that I 
interpret as a remnant of the pocket stimulator. The 
verge and ridge correspond to the external bulge on the 
penis section. The sculpture of the wall, position and 
number of circular ridges, size of pocket stimulator, 
and verge size differ between Phenacohelix and 
Charopa, but the fact of a basic set of structures is the 
important data. 

In most of the Pacific Island Charopinae there is 
simple variation in the relative sizes of the verge, 
pocket or modified pocket stimulator, and the number 
and prominence of the circular bands. The wall 
sculpture found in the New Zealand taxa seems to be 
absent in the Pacific Island taxa. The combination of 
very small size (see table XV), limited material, and 
preparation of illustrations at an early stage in the 
study means that treatment of the variations in this 
portion of the study is inadequate. Specific comments 
on differences between geographically close species are 
given in appropriate places in the systematic review. 
Here I will mention only a few more obvious changes. 
The verge varies from a small spade-shaped papilla 
with open side (fig. 42e, S. peasei), to a conical projec- 
tion with terminal pore (fig. 43e, S. auanaensis), or it is 
reduced to a swelling attached to one wall (fig. 55e, S. 
allecta), or a globular swelling (fig. 57c, S. com- 
plementaria, fig. 75h, Ba humbugi), a protruding lobe 
with laterally apical pore (fig. 79c, Tuimalila pilsbryi), 



or the bulbous structure seen in Vatusila tongensis (fig. 
84b). 

The circular ridge can be split into several and low 
(fig. 42d-e, Sinployea peasei), doubled and thick (fig. 
43e, S. auanaensis), single and thin (figs. 51h, S. inter- 
media; 57c, S. complementaria; 67j, S. euryomphala), 
or very thick and partly altered into pads (figs. 55e, S. 
allecta, 79c, Tuimalila pilsbryi). 

The pocket stimulator is particularly variable and 
with few exceptions has been inadequately illustrated 
and interpreted. Comparison of the doughnut shape in 
Figure 39h, split "U" in Figure 42d, raised globular 
mass in Figure 53c, low "U" in Figure 53h, lobular 
structure in Figure 55e, "U" ridge in Figure 67d, 
widely open pocket in Figure 67j, greatly reduced and 
altered structure in Figure 75h, and nearly closed 
pocket in Figure 79d gives an idea of the great extent of 
variation in this structure, but this is only an introduc- 
tion to its complexity. 

This study demonstrates that extensive variation 
occurs in verge, circular ridges, and pocket stimulator 
in the Pacific Island Charopinae, but analysis of the 
patterns of variation is beyond the scope of this report. 
Because Sinployea is evidently in a highly active stage 
of speciation, study of this area might yield important 
data, but is left for others. 

A major modification of this pattern occurs in 
Graeffedon (fig. 87c). The epiphallus (E) opens through 
a simple pore that is surrounded by a huge circular 
pilaster (PP). Immediately below this are three large 
pilasters: an upper, free-tipped vergic stimulator, a 
long and low semicircular ridge oriented longitudi- 
nally in the penis chamber, and a broad, low ridge with 
slightly free upper tip that occupies the lower half of 
the penis. Coupled with the absence of the valve at the 
vas deferens-epiphallus junction, these structures 
mark Graeffedon as remote from the other Charopinae, 
but it is better accommodated in this subfamily than 
assigned to one of its own. 

The Trukcharopinae (fig. 90c-d, g, i) have a simple 
pore opening from the epiphallus (Russatus), a low 
vergic papilla (Kubaryiellus), or even a tubular verge 
(Palline). Various pilasters and a possibly highly mod- 
ified pocket stimulator are found in some of the taxa. 
These variations are discussed more fully on pp. 205- 
207. Many of these represent additive structures com- 
pared with the Charopinae, but basically they could be 
derived from some of the Charopinae conditions in ex- 
tralimital taxa (Solem, unpublished data). 

The Semperdoninae (figs. lOOd, f, j, 102d, 103b, f) 
have the epiphallus entering the penis through a nar- 
row muscular collar, with high lamellar pilasters ex- 
tending down and then usually coalescing into three, 
high glandular pilasters. The whole penis is sur- 
rounded by a muscular sheath. Exceptions to the basic 
pilaster pattern are Himeroconcha rotula (Quadras & 
Mollendorff, 1894), which lacks the initial radiating 
pilasters although retaining the three basal ones, and 
Ladronellum mariannarum (Quadras & Mollendorff, 



30 



SOLEM: ENDODONTOID LAND SNAILS 



1894) in which the initial radiating pilasters coalesce 
into a huge horseshoe-shaped pilaster (fig. 102d) that is 
inflatable with fluids. The enclosure of both epiphallus 
and penis by a muscle sheath is as great a change as is 
the pilaster pattern. These make the Semperdoninae 
the most isolated of the charopid subfamilies in terms 
of penis structure. 

In the few dissected Rotadiscinae (H. B. Baker, 
1927; Solem, unpublished data) the epiphallus enters 
through a pore, papilla, or short verge into a very 
thick-walled chamber with longitudinal pilasters. 
There may or may not be accessory organs associated 
with the penis and/or atrium. This represents yet 
another series of experiments in the Charopidae. 

Finally, the situation in Discocharopa (fig. 34c) re- 
quires comment. A short, tubular penis with lateral 
entrance of the vas deferens has the penial retractor 
muscle inserting apically. Apparently there are at 
least two longitudinal pilasters inside the penis, but 
the available material was not adequate to work out 
the details (see p. 75). Although this departs from the 
reported pattern for the Charopinae (see above), the 
genus is temporarily placed in the Charopinae pending 
revision of the extralimital groups. It probably will be 
split into another subfamily unit, but available data 
are inadequate to propose a more appropriate classifi- 
cation. 

The atrium, in all dissected material, is a simple, 
short tube opening externally behind and above the 
right rhinophore. It varies somewhat in length but 
shows no really significant changes. 

TERMINAL FEMALE GENITALIA The post- 
uterine or free oviduct (UV), spermatheca (S), and va- 
gina (V) in the Endodontidae are very slender tubes 
without unusual structures. They show variation pri- 
marily where the spermathecal shaft inserts penis, 
atrium, or free oviduct (Solem, 1976b, pp. 83-84). In 
the Charopidae the situation is quite different. Nor- 
mally, the spermathecal base and at least parts of the 
vagina are greatly enlarged and with complex internal 
pilasters. The free oviduct, most of the vagina, and the 
spermathecal shaft above the swelling will be quite 
slender, with an ovate, expanded head of the sperm- 
atheca buried in the albumen gland-prostate margin. 
The shape and proportions shown by Sinployea avan- 
aensis (fig. 43c) are typical. The shifted expanded 
area in S. aunuuana (fig. 53b) or expanded free oviduct 
(UV) in S. allecta (Cox, 1870) (fig. 55c), shortened va- 
gina in S. inermis lakembana (fig. 67f), and generally 
greater elongation in the Semperdoninae (figs. lOOb, e, 
g, 102a, 103a, c-d) are the typical minor variations. 
The nearest thing to a major variation is found in 
Jokajdon (fig. 93bc) where the expanded area is re- 
stricted to the base of the vagina and atrium, and the 
normally expanded areas are slender in comparison. In 
addition there is an accessory muscle attaching to the 
penioviducal angle. These changes may correlate with 
the very restricted shell aperture and large barriers of 
Jokajdon. 



Internally there appear to be two basic patterns of 
sculpture of the upper vagina and lower spermathrca. 
In the typical Charopinae, such as S. complementaria 
(Mousson, 1865) (fig. 57f), S. allecta (Cox, 1870), and 
Ba humbugi, there are weak longitudinal pilasters in 
the vagina, the free oviduct (UV) has a constricting 
pilaster with central pore (UVO), and the opening of 
the spermatheca (S) has a central pore (SO) through a 
circular pilaster with one edge a free flap. Because of 
size problems and the early stage in the study at which 
this area was examined, only the one has been illus- 
trated. The second pattern is seen quite clearly in 
Graeffedon (fig. 87c) where the opening to the free 
oviduct is a simple pore and there are huge longitudi- 
nal pilasters lining the walls of the spermatheca and 
vagina. Essentially the same pattern is seen in 
Palikirus, Palline, Semperdon uncatus, S. hepta- 
ptychius (Quadras & Mollendorff, 1894), Ladronellum 
mariannarum (Quadras & Mollendorff, 1894), 
Himeroconcha lamlanensis, and H. fusca (Quadras & 
Mollendorff, 1894), thus indicating that this general 
type occurs not only in some of the Charopinae, but 
also in the Trukcharopinae and Semperdoninae. A de- 
tailed comparative study of this region would be --veil 
worthwhile but is beyond the scope of this monograph. 

PALLIAL COMPLEX 

A discussion of the functional significance that the 
closed and complete ureter found in most Charopidae 
has in relation to water conservation as contrasted to 
the incomplete posteriorly opening ureter of the En- 
dodontidae was given previously (Solem, 1976b, pp. 
84-87). Some of the New World Rotadiscinae show 
only a partial closed ureter (H. B. Baker, 1927, pi. 16, 
fig. 17, pi. 17, figs. 24, 30), and there are Australian 
equivalents (Solem, unpublished data). All of the 
Pacific Island Charopidae that have been dissected 
show a complete ureter with the ureteric pore opening 
next to the anus just inside the pneumostome. This 
contrasts immediately with the Endodontidae where 
the ureteric pore opens at the posterior part of the pal- 
lial cavity near where the anterior margin of the rectal 
kidney lobe touches the hindgut (Solem, 1976b, p. 85, 
fig. 49c). Thus, even a glance at the anterior part of the 
pallial cavity is sufficient to tell whether a Pacific Is- 
land species belongs to the Endodontidae or 
Charopidae. 

A typical charopid configuration is seen in 
Sinployea allecta allecta (Cox, 1870). Viewed from the 
outside (fig. 55a), the bilobed nature of the kidney (K) 
is obvious, with the longer, cigar-shaped rectal lobe 
(lower in figure) definitely overlapping the hindgut 
(HG). The shorter, irregularly shaped pericardial lobe 
(upper in figure) partly overlies and is cupped partly 
around the heart (H). The kidney base (left in figure) 
stops short of the downward twist of the hindgut as 
intestine and is abutted by the lobules of the digestive 
gland. The primary ureter (KD) originates from near 
the anterior margin of the pericardial lobe of the kid- 



PATTERNS OF MORPHOLOGICAL VARIATION 



31 



ney, follows the upper margin of this lobe posteriorly, 
lying partly on the pallial cavity roof and partly on the 
kidney itself, reflexes abruptly and as the secondary 
ureter follows the lower margin of the rectal kidney 
lobe anteriorly to its termination, then lies next to the 
hindgut until both disappear under the mantle collar 
(MC). The principal pulmonary vein (HV) extends an- 
teriorly from the heart along the pallial roof, but in the 
smaller species shows no sign of branching, fading out 
from visual observation well short of the mantle collar. 

The bilobed kidney, or in a torn and extracted 
specimen, the tip of the rectal lobe of the kidney, plus 
the presence of the secondary ureter as a distinct tube 
next to the hindgut, are sufficient to immediately iden- 
tify a species as a member of the Charopidae or 
Punctidae rather than the Endodontidae. In the latter 
family (Solem, 1976b, p. 459, fig. 195a) there is at most 
a short rectal lobe with the ureter ending in a ureteric 
pore (KX) as it reaches the hindgut. In a few 
Charopidae, such as Phenacohelix pilula (Reeve, 1852) 
(fig. lla), the rectal arm is as short as in the Endodon- 
tidae, but the clear presence of the secondary ureter 
along the hindgut is sufficient for family separation. 
Exceptions to this in some Australian taxa will be con- 
sidered elsewhere. 

When dissected out (fig. 55b) and viewed from an 
inside view of the pallial cavity, the typical charopid 
pallial region shows only minor additional features. 
Remnants of a retractor muscle to a rather strongly 
developed muscle come off the parietal-palatal margin 
near the kidney base. A weaker version of the same 
muscle can be seen in many species of Endodontidae 
(Solem, 1976b, p. 459, fig. 195a). The heart in this view 
clearly lies on top of the kidney, the rectal kidney lobe 
extends partly under the hindgut, thus overlapping on 
both sides of this tube, and both the hindgut and ureter 
terminate just inside the mantle collar. Quite possibly 
this area is involved in water resorption and is a pre- 
cursor of the more elaborate structures seen in higher 
aulacopods, such as an apparent bladder in Deroceras 
reticulatum (Miiller, 1774) (Runham & Hunter, 1970, 
pp. 77-79, figs. 32-33). 

Modifications of this general pattern involve addi- 
tion of new structures, changes in the relative size of 
the kidney lobes, differences in the amount of space 
between the two lobes, and compactional alterations 
correlated with reduction in whorl counts. Data on at 
least part of the pallial complex was recorded for 36 of 
the 43 taxa for which at least some anatomical mate- 
rial was available. Illustrations are presented for 20 of 
these. The only additive structure seen in a Pacific Is- 
land taxon is the extensive intrusion of mantle gland 
tissue onto the pallial roof in Graeffedon graeffei 
(Mousson, 1869) (fig. 87a, MG) and a slight extension 
in Semperdon xyleborus. This phenomenon is far more 
common in extralimital taxa, with the intrusion rang- 
ing from a massive and sharply defined area in 
Champa coma (Gray, 1843) (fig. 9a) to the short area in 
Phenacohelix pilula (Reeve, 1852) (fig. lla) and the 



elongated finger in Laoma leimonias (Gray, 1850) (fig. 
25a). The function of this extension is unknown. 

Although there is a modest amount of intraspecific 
variation in the relative length of the two kidney lobes, 
much of this seems to be caused by differential contrac- 
tion and compaction when the animal withdraws into 
the shell. Dissections made from deeply retracted 
specimens tended to have the rectal kidney lobe un- 
changed in shape and position, but pulled back further 
relative to the heart and pericardial kidney lobe. The 
latter would tend to be somewhat twisted and distorted 
(for example see fig. 34d). I made no exact measure- 
ments as to the relative lengths because the results 
would not be strictly comparable. Table IX summarizes 
the relative lengths in the Pacific Island species. Only 
in Discocharopa aperta (Mollendorff, 1888) (fig. 34d) is 
the rectal lobe somewhat reduced to as great an extent 
as in the New Zealand Phenacohelix pilula (Reeve, 
1852) (fig. lla). The general pattern is for the rectal 
lobe of the kidney to be much longer than the pericar- 
dial. This reaches its greatest extent in such taxa as 
Graeffedon graeffei (Mousson, 1869) (fig. 87a) and 
Semperdon heptaptychius (Quadras & Mollendorff, 
1894) (fig. lOOa). In both of these taxa the pericardial 
lobe is reduced to a small fraction of the length and 
volume of the rectal. 

The length relationship varies within a genus, 
since in Sinployea there are nine species in which the 

TABLE IX. - RELATIVE LENGTH OF KIDNEY LOBES 
IN THE PACIFIC ISLAND CHAROPIDAE. 



Subequal or 
Equal 



Sinployea tahitiensis 

^. lamellicosta 

j>. avanaensis 

^. intermedia 

S^. allecta allecta 

ji. vicaria vicaria 

^. kusaieana 

j>. inermis inermis 

J[. i_. lakemba 

^. adposita 

Ba humbugi 

Tulmalila pilsbryi 

Russatus nigrescens 

Himeroconcha fusca 

H. lamlanensis 



Rectal Much 
Longer 

Sinployea modicella 
^>. montana 
S, neglecta 
S. peasei 
S^. aunuuana 
^. clista 
S_. complement aria 
S^. ir_regulaj^is_ 
S^. eu r y ompha 1 a. 
Graeffedon graeffei 
Kubaryiellus kubaryi 
Trukcharopa trukana 
Palikirus cosmetus 
Jokajdon calllzonus 
Palline np_t era not era 
P_. micramyla 
Semperdon uncatus 



j^. xyleborus 

^. heptaptychius 

S_. rotanus 

Ladronellum raariannarum 



Pericardial 
Much Longer 

Discocharopa aperta 



32 



SOLEM: ENDODONTOID LAND SNAILS 



kidney lobes are equal or nearly equal in length and 
nine in which the rectal is significantly longer. I could 
detect no conchological correlations in size, shape, or 
whorl count with this variation in kidney lobe length 
concerning Sinployea. The other Charopinae have a 
secondarily shortened and very fat kidney correlated 
with whorl count reduction (Ba humbugi, fig. 75a), 
subequal lobes in the gigantic Tuimalila pilsbryi (fig. 
79a), or the very elongated rectal lobe in Graeffedon 
graeffei (Mousson, 1869) (fig. 87a). In the Truk- 
charopinae only Russatus nigrescens (Mollendorff, 
1900) (fig. 90a), whose whorl count is reduced to a 
mean of 3Vs + , has a shortened kidney, whereas in the 
Semperdoninae only the large-sized Himeroconcha 
have the kidney lobes equal or subequal in length. 

In both the Trukcharopinae and the Semper- 
doninae the arms of the ureter are tightly pressed to- 
gether without any pallial roof tissue visible between 
them. The pattern is somewhat unusual in Russatus 
nigrescens (Mollendorff, 1900) (fig. 90a), with the 
abrupt angling of both ureter arms at the anterior 
margins of the kidney lobes. I interpret this as the 
result of shortening and secondary thickening of the 
kidney in relation to whorl and pallial cavity reduc- 
tion. In the Charopinae, the same compaction is seen in 
Ba humbugi (fig. 75a), with only slightly less abrupt 
and extensive angling of the ureter. Of the examined 
Sinployea, all taxa except S. allecta allecta (Cox, 1870) 
(fig. 55a-b) and S. irregularis (Garrett, 1887) have at 
least a narrow strip of pallial roof tissue visible be- 
tween the ureter arms, as does Tuimalila pilsbryi (fig. 
79a). Extralimital Charopinae are variable in this fea- 
ture (see figs. 9e, lla). 

Shortening of the pallial cavity is more common 
than elongation. The latter seems to have occurred 
only in Jokajdon callizonus (Mollendorff, 1900) (fig. 
93a) where it extends % of a whorl apically. This prob- 
ably correlates with the narrowed cross-section of the 
body whorl in this species (fig. 92b). Major shortening 
has occurred in Ba humbugi (fig. 75a) and Russatus 
nigrescens (Mollendorff, 1900) (fig. 90a). Their respec- 
tive mean whorl counts of 3% and 3Vs+ are the lowest 
in the Pacific Island Charopidae, except for the rotadis- 
cine Microcharopa mimula (mean whorl count 3V4-). 
In both Ba and Russatus the width of the kidney is 
almost equal to its length, and the arms of the ureter 
are tightly compacted and overlap each other between 
the kidney lobes. The process has been carried further 
in Russatus, with V4 whorl length to the pallial cavity, 
than inBa, where the length is still Vz whorl. The same 
type of compaction has occurred in both genera, and 
they form a contrast to the situation in the New 
Guinea charopids Pilsbrycharopa and Paryphantopsis 
(Solem, 1970a, p. 250, fig. 2a, f) in which kidney com- 
paction resulted in progressive increase of the angle 
between the arms of the ureter, and the kidney is ro- 
tated away from the hindgut. Yet another pattern is 
seen in the New Zealand flammulinid and Maori- 
concha groups. 

The pallial cavities in the Pacific Island Endodon- 



tidae and Charopidae have radically different ureter 
and kidney structures. Within the Pacific Island 
Charopidae there is a relatively simple pattern of vari- 
ation in relative lengths of the kidney lobes and the 
extent to which lung roof tissue is visible between the 
arms of the ureter. 

DIGESTIVE SYSTEM 

Allowing for the generally reduced whorl counts 
(table X) in the Charopidae, the gross features of the 
endodontid (Solem 1976b, pp. 372-373, figs. 163-164) 
and charopid (figs. 9e, g, 75a) digestive tracts are the 
same. The digestive glands (OG) are in contact above 
the esophagus (BE), which continues past the pallial 
cavity as a slender tube. In some taxa, such as the 
larger Sinployea, Ladronellum mariannarum (Mollen- 
dorff, 1900), and Kubaryiellus kubaryi (Mollendorff, 
1900), the salivary glands are fused above the 
esophagus, whereas in Ba humbugi, they do not even 
touch posteriorly. I am not certain if there is any sys- 
tematic significance to these changes. Intestinal loops 
above the pallial cavity apex occupy about Vie of a 
whorl, instead of the Vs whorl in the Endodontidae. The 
charopid stomach expansion averages about % of a 
whorl, with the digestive gland shorter and more com- 
pact than in the Endodontidae. Except for these 
changes associated with the reduced whorl count, the 
gross features of the digestive tract are the same in the 
two families. 

Radular features are one of the best guides to 
separating the Endodontidae, Charopidae, and Punc- 
tidae on the Pacific Islands. The endodontid radula 
(Solem, 1976b, pp. 88-94, figs. 51-54) has a tricuspid 
central, several bicuspid laterals, and marginals that 
have the ectocone fragmented and the endocone in- 
creasing almost to the size of the mesocone. There are 
very noticeable differences in the angling of the indi- 
vidual teeth, but their small size and difficulties in 
mounting and preparation have prevented full study of 
their variation. 

The punctid radula (fig. 12) presents a number of 
unusual features. The central tooth (fig. 12a, upper) 
has a long slender mesocone and two very slender, 
much shorter ectocones. The laterals (fig. 12b-e) have 
two slender cusps and three much shorter accessory 
cusps, whereas the outermost lateromarginals (fig. 12f) 
are broader and with the outer large cusp reduced in 
size. The accessory cusps are at to below the limit of 
optical microscope examination, depending upon the 
quality of the equipment and illumination. As far as I 
am aware, the only optical microscopist to detect and 
illustrate these accessory cusps was H. B. Baker (1927, 
pi. 16, fig. 11), also reprinted in Pilsbry (1948, p. 642, 
fig. 349, d). 

In addition to the obvious cusp edges and numbers 
there are fundamental differences in the nature of the 
cusps and basal plates in the two families. The en- 
dodontid basal plate (Solem, 1976b, p. 89, fig. 52d) has 
a typical interlocking relationship with the next pos- 




FIG. 12. Radular teeth ofPunctum minutissimum (Lea). Cedar bog, Woodburn Road, 4 miles southwest of Urbana, Champaign County, Ohio. 
E. Keferl! X-20-1969. FMNH 151102: a, central (upper) and 1st lateral (23,000x); b, early lateral teeth (10,800x); c, single midlateral tooth 
(21,500x); d, late lateromarginal teeth (15,900x); e, low angle view of lateromarginal teeth (10,700x); f, outer lateromarginal tooth showing 
cusp reduction (28,000x). 

33 



34 



SOLEM: ENDODONTOID LAND SNAILS 



terior tooth, is square to rectangular in shape (Solem, 
1976b, p. 91, fig. 54b, e), and, at least for the early 
lateral teeth, the cusps are sharply pointed and ele- 
vated at a high angle (Solem, 1976b, p. 88, fig. 51a-b; 
p. 89, fig., 52a-b, d). In the Punctidae (fig. 12c, e) the 
basal plate is long and tapering, has no apparent inter- 
locking relationships, and the cusps are bluntly 
rounded and point almost directly forward. It is unfor- 
tunate that these differences can be seen only with the 
scanning electron microscope, a fact that reduces their 
routine utility in identification and classification. 

The Pacific Island Charopidae have a very stan- 
dard pattern of structure (figs. 13-14). As pointed out 
earlier (Solem, 1976b, p. 93), extralimital charopids 
show a variety of structure, but review of these is out- 
side the scope of this monograph. The tricuspid central 
tooth (figs. 13b, 14a) is slightly narrower and shorter 
than the adjacent laterals, which are tricuspid. When 
viewed at a low angle, it is evident that the side cusps 
of both central and laterals are raised above the eleva- 
tion plane of the mesocone (fig. 14a), whereas in the 
Endodontidae (Solem, 1976b, p. 88, fig. 51a) they are in 
the same elevation plane. The outer laterals and early 
marginals in the Charopidae (fig. 14b) do not have the 
elevated side cusps. The marginals may retain the 
tricuspid pattern (fig. 13c-f), or rarely the outer mar- 
ginals may become multicuspid. Climo (1969a, figs. 
31-34; 1970, figs. 11-14) gives a number of radular 
transects for New Zealand charopids. 

Because only a few species could be examined with 
the SEM and their pattern of structure was quite 
uniform, results from optical viewing mostly have been 
omitted from the text in this volume. 

The bicuspid laterals of the Endodontidae, tricus- 
pid laterals of the Charopidae, and multicuspid 
lateromarginals of the Punctidae found on Pacific Is- 
lands thus present clearcut differences among the 
families. The Austro-Zelandic charopids, however, 
show a great variety of radular structures, so that the 
endodontid-charopid distinction does not hold for that 
area. 

Jaw structure in the Charopidae was not studied 
in detail because the pattern of partial plate fusion in 
larger taxa paralleled the situation found in the En- 
dodontidae (Solem, 1976b, p. 94). 

FREE MUSCLE SYSTEM 

As in the Endodontidae (Solem, 1976b, p. 94), all 
dissected charopids had the right ommatophoral re- 
tractor passing through the penioviducal angle and 
joining the right rhinophoral retractor that passes out- 
side the penioviducal angle posteriorly. Unlike the En- 
dodontidae, in all examined Charopidae the penial re- 
tractor muscle arises from the diaphragm and inserts 
onto the penis or epiphallus. Only a few alterations in 
the common pattern of unions were observed. Russatus 
nigrescens (Mollendorff, 1900) has the tentacular re- 
tractors fusing with the tail fan at the columellar 
muscle rather than earlier as in most taxa. InJokajdon 



callizonus (Mollendorff, 1900) a new muscle attaches at 
the penioviducal angle, joining the tail fan much later. 
With the elongated (% whorl) pallial cavity and nar- 
rowed aperture with large barriers (fig. 92b) in this 
species, the added muscle may play a major role in 
successful body retraction. 

In Sinployea complementaria (Mousson, 1865) 
there is a muscle from the right tentacular retractor 
that inserts on the apex of the free oviduct, whereas in 
Tuimalila pilsbryi a muscle runs from the columellar 
retractor to the apex of the free oviduct. It is possible 
that a weaker version of this same muscle is present in 
smaller species of Pacific Island Charopidae and was 
overlooked in earlier phases of this study. It was not 
possible to recheck all taxa for this feature. Extralimi- 
tal taxa, such asStephanoda binneyana (Pfeiffer, 1847) 
(fig. 31c), show a vaginal retractor muscle (VRM). It is 
probable that a number of such experiments in added 
muscles exist, but have not been observed. 

NERVOUS SYSTEM 

Wherever possible, the enervation of the penis 
from the right cerebral ganglion was confirmed, but 
because of difficulties in handling the small-sized 
material, preservation in alcohol, and heavy covering 
of connective tissue over the ganglia, no attempt at 
working out the details of the nervous system was 
made. Climo (1970, fig. 21B) illustrated the central 
nervous system ofPhenacharopa novoseelandica (Pfeif- 
fer, 1853). 

EXTERNAL BODY FEATURES 

In the Trukcharopinae and Semperdoninae, except 
for Semperdon heptaptychius (Quadras & Mollendorff, 
1894), Ladronellum mariannarum (Quadras & Mol- 
lendorff, 1894), and Himeroconcha fusca (Quadras & 
Mollendorff, 1894), the body color is yellow-white, 
without darker markings. In the latter three species 
there are gray to reddish gray markings on the neck, 
ommatophores, and mantle collar. In the Charopinae, 
all live-collected Cook, Society, and Samoan Island 
species of Sinployea, except the Swains Island S. in- 
termedia, have light to dark gray markings. All of the 
Sinployea with gray markings have been taken in 
semiarboreal situations. The purely terrestrial S. 
kusaieana, S. euryomphala, S. inermis, S. adposita, 
and S. irregularis (Garrett, 1887) have yellow-white 
body color. Tuimalila pilsbryi also shows the darker 
mantle and neck coloration. The dark body color ap- 
pears to be a correlative of semiarboreal habitat. 

Many of the arboreal New Zealand and Australian 
charopid taxa have a strongly developed mucus appa- 
ratus at the hind end of the foot. A typical "mucus 
pore" or "caudal foss" (CF) is present in Phenacohelix 
pilula (Reeve, 1852) (fig. lib), with the pore overhung 
by a caudal horn (CH). This is effectively an intensifi- 
cation of the point where the foot grooves unite above 
the tail plus a mucus-secreting gland concentrating at 





FIG. 13. Radular teeth of Tuimalila pilsbryi. Station T-22, 1,000ft. elevation, Eua, Tonga. FMNH 152378: a, near middle of radula at 280 x; 
b, central tooth and 1st lateral on right side of radula at 2,820 x ; c, early laterals on right side of radula at 2,900 x ; d, transition from laterals to 
marginals on left side of radula at 2,950x; e, middle marginals from left side at 2,875x; f, outermost marginals at 2,875x. All views nearly 
vertical. 



35 



36 



SOLEM: ENDODONTOID LAND SNAILS 




FIG. 14. Radular teeth ofTuimalilapilsbryi. Station T-22, 1,000 ft. elevation, Eua, Tonga. FMNH 152378: a, 45 angle view of central (c) and 
early laterals at 3,200 x showing pattern of cusp elevation; b, same view of transitional zone between laterals and marginals at 3,125x. 



this point. This apparatus is absent in Charopa coma 
(Gray, 1843) (fig. 9a) and all of the dissected Pacific 
Island Charopidae (fig. 43a). The presence of this pore 
is associated with arboreal snails (Solem, 1976b, pp. 
105-106) and is not of major phyletic importance, as 
was also confirmed by Pilsbry (1892a-b, 1893a-b) and 
Climo (1969a, pp. 148-150). 

As in the Endodontidae (Solem, 1976b, p. 94), the 
mantle collar normally is without protrusions, the 
gonopore is located below and slightly behind the right 
ommatophore, and the slime network is rather weakly 
defined. Neither family shows substantial external 
modifications in the Pacific Island taxa. 

PATTERNS OF VISCERAL HUMP REDUCTION 

In the Endodontidae (Solem, 1976b, pp. 94-98) one 
of the major repetitive changes is an increase in the 
number of whorls and thus elongation of the visceral 
hump. Not only do the Pacific Island Charopidae have 
a much lower average whorl count than the Endodont- 
idae (median means 4Vfe and 5V2 + , respectively), but 
of the Charopidae only Semperdon kororensis (Bed- 
dome, 1889) with 5Vs whorls and the Rarotongan 
Sinployea planospira (Garrett, 1881) with 6% whorls 
exceed a mean count of five whorls. The latter has not 
been taken in this century, and of the former only 
fragmentary extracted pallial collars and terminal 
genitalia were available. Thus I can offer no data of 
any changes associated with elongation of the visceral 
hump. 

Reduction in whorl count and thus in the total 
length of the visceral hump is a common pattern in 
charopids from many areas of the world. Such taxa as 



the Juan Fernandez A mphidoxa (fig. 30a, c), New Zea- 
land Flammulina, Maoriconcha, and Otoconcha 
(Climo, 1971a), a few of the South African Trachycystis 
(Connolly, 1939), and some New Caledonian taxa 
(Solem, 1961) show varying degrees of visceral hump 
reduction. This is carried furthest in the New Zealand 
Otoconchinae and the semislug Ranfurlya. Discussion 
of their changes is beyond the scope of this review, but 
the general pattern is for zonal compaction of organs in 
several systems (see Solem, 1966, for a discussion of 
this in the Thailand Helicarionidae). 

In the Pacific Island Charopidae whorl reduction 
takes two forms simple decrease in mean whorl count 
without any change in whorl profile (Microcharopa 
mimula, Z l A-;Lagivala minusculus, 3%; Discocharopa 
aperta, 3%; and Palikirus ponapicus, 3%) or reduction 
in whorl count accompanied by a drastic increase in 
cross-sectional areas of the body whorl (Russatus ni- 
grescens, 3Vs + ',Ba humbugi, 3%). 

Either the former taxa have not been dissected, or, 
in the case of Discocharopa, there are no close relatives 
known with which anatomical comparisons can be 
made. Thus, comments here must be restricted to the 
latter situation, whorl count reduction combined with 
whorl cross-section increment. The degree of whorl 
changes can be judged by comparing whorl increment 
rates and whorl profiles in Kubaryiellus and Russatus 
(fig. 89a-b, d-e) and then in Sinployea irregularis 
(Garrett, 1887) (fig. 65d-e) and Ba humbugi (fig. 
74a-b). Anatomically, the typically trukcharopinine 
half whorl pallial region of Kubaryiellus (fig. 90e) can 
easily be altered to the nearly square kidney (K) and 
one quarter whorl pallial cavity of Russatus (fig. 90a) by 
shortening and widening of the kidney, ventral flexing 



PATTERNS OF MORPHOLOGICAL VARIATION 



37 



of the intestinal loop, and a slight ventral movement of 
the anterial part of the pericardial kidney lobe over the 
heart (H) and principal pulmonary vein (HV). Compar- 
ing the gross genitalia (fig. 90b, f), the proportionately 
shortened free oviduct (UV), spermatheca (S), prostate 
(DG), and uterus (UT) ofRussatus are obvious. There is 
no detectable change in the penis (P) and vagina (V). 
The pallial region of Sinployea irregularis (Garrett, 
1887) was not illustrated, but it has the typical generic 
pattern of extending % of a whorl apically, there is no 
lung roof space visible between the arms of the ureter, 
and the rectal lobe is distinctly longer than the 
pericardial. In Ba humbugi (fig. 75a) the anterior half 
of the half whorl pallial cavity has undergone shorten- 
ing and change, with the anterior margins of the kid- 
ney flared laterally, but the posterior portion of the 
kidney and the intestinal loops are essentially unmod- 
ified when compared with the degree of change seen in 
Russatus (fig. 90a). Contrasting the genitalia of S. ir- 
regularis (fig. 67a, drawn from a deeply retracted 
specimen) and So (fig. 75b-e, h) shows a rather drastic 
folding of the prostate and uterus plus altered insertion 
of the penial retractor muscle in the latter taxon. Ba 
also shows shortening of the vagina and spermatheca. 
Thus, changes in the visceral hump length of 
Pacific Island Charopidae involve selective shorten- 
ings in portions of the organ systems lying in the pal- 
lial region. Unless there is clear shortening of the neck 
region (area between ommatophores and pallial collar 
edge when animal is crawling), the prostate-uterine 
area and spermathecal-free oviduct sections are more 
apt to be involved than the penis-vagina, and the kid- 
ney area, than the gas exchange surfaces of the pallial 
roof. More extended comments on these changes are 
postponed pending completion of studies on Australian 
and New Zealand taxa. 



SUMMARY OF ANATOMICAL VARIATION 

The Pacific Island Charopidae show a number of 
minor variations in the terminal genitalia and kidney 
configurations. The former involve species recognition 
factors, the latter are of uncertain significance except 



in cases of obvious elongation or shortening of the pal- 
lial cavity. These terminal genitalia patterns are not 
continuously variable, but fall into rather discrete 
general patterns that indicate multiple colonizations of 
the island areas and are used in part to recognize sub- 
family units. 

The major and minor anatomical differences be- 
tween the Pacific Island Charopidae and Endodontidae 
were summarized by Solem (1976b, pp. 97-98, tables 
LVIII-LIX). Consideration of the complexities in the 
Australian, New Zealand, and New Caledonian taxa 
must be deferred. There are no clear unitary anatomi- 
cal trends within the Pacific Island Charopidae com- 
pared with those outlined for the Endodontidae (Solem, 
1976b, pp. 98-99), although the addition of structures 
among the various charopid subfamilies is striking. 

A major correlative of the "looser" whorl coiling 
pattern (pp. 41-43) in the Charopidae compared with 
the Endodontidae is that the cross-sectional area of the 
body whorl in particular is greater in the former fam- 
ily. This has important implications on the size of the 
anterior body and its contained organs. The wider area 
in the Charopidae permits widening of the terminal 
genitalia. The much fatter penis and vagina-free 
oviduct-spermathecal union area in the Charopidae is 
possible directly because of this extra space provided 
by the looser coiling. Whereas in the Endodontidae 
these organs all are thin tubes, in most Charopidae 
they are thick and with complex internal structures. 

The only Pacific Island charopid known to have 
clearly narrowed structures in this area is Jokajdon 
callizonus (Mollendorff, 1900) (fig. 93a-b), whose dras- 
tically narrowed shell aperture and large apertural 
barriers (fig. 92a) closely approach the typical en- 
dodontid condition. Even though species of Palline 
(figs. 94b, e, 95d) have large apertural barriers and 
somewhat narrowed apertures, the thick terminal 
genitalia (fig. 93f, h) contrasts with that of Jokajdon 
and is in the typical charopid pattern. 

It is quite probable that the variety of genital 
structures seen in the Charopidae as opposed to the 
Endodontidae are in large part the result of simple 
space availability for experimentation. 



CHAROPID-ENDODONTIDCONCHOLOGICAL 

COMPARISONS 



Despite almost complete overlap with regard to 
most conchological characteristics and their close simi- 
larity when viewed with the naked eye, there are a few 
clear differences between the Endodontidae and 
Charopidae in meristic and structural features. Data 
in Tables I-III and X and Figures 15-23 summarize 
both similarities and differences in some basic para- 
meters. Figures 15-22 were prepared several years ago 
and omit data on the Lau Archipelago endodontids, 
Priceconcha tuuuthaensis Solem (1973d) and Thauma- 
todon spirrhymatum Solem (1973d). Their inclusion 
would not have changed the results significantly. In 
addition, four taxa of endodontids were not seen or 
measured. These two exceptions account for the 
difference between the 185 taxa listed by Solem 
(1976b, p. 9, table IV) and the 179 listed as measured in 
this tabular comparison. 

For ease in visual comparison, in Figures 15-22 
the actual numbers of species for each graph unit have 
been converted into percentages of the total measured 
within that family so that the graphs will be directly 
comparable. Otherwise the difference between 179 en- 
dodontids and 95 charopids would make visual com- 
parisons difficult. 

Table X shows the median mean value and total 
range of mean values for the species measured within 
each family. The distinctly larger size and higher 
whorl count of the Endodontidae is evident, whereas 
their near identity in H/D ratios, D/U ratios, and mean 
rib counts is surprising. The greater ribs/mm, in the 
Charopidae directly correlate with their smaller size. 

TABLE X. - CONCHOLOGICAL COMPARISONS OF ENDODONTIDAE AND CHAROPIDAE 



Number of species 
level taxa measured 

Median mean height (range) 
Median mean diameter (range) 

Median mean whorl count 
(range) 

Median mean H/D ratio 
(range) 

Median mean D/U ratio 
(range) 

Median mean ribs on body 
whorl (range) 

Median mean ribs/mm. 
Taxa with ribs reduced 



PACIFIC ISLANDS. 
Endodontidae 

179 

1.48(0.92-7.26) 
3.77(1.7-12.3) 

5 1/2+ 

(3 5/8-8 1/8) 

0.531 

(0.344-0.789) 

3.94 

(1.68-closed) 

91.0(19-202) 

9.2(1.4-40) 
39(21.8%) 



Charopidae 

95 

1.51(0.48-3.69) 
2.76(1.07-7.52) 

4 1/8- 

(3 1/8-6 5/8) 

0.523 

(0.365-0.801) 

3.94 

(2.03-closed) 

91.2(19-225) 

11.3(1.7-37) 
10(10.5%) 



The greater degree of rib reduction in the Endodon- 
tidae and the reduction in percentage of taxa with 
apertural barriers in the Charopidae also indicate 
major differences. 

More detailed comments can be made from the 
data in Figures 15-23. Mean height distribution (fig. 
15) shows a slight Endodontidae offset for most species, 
then an extended high-spired portion that far sur- 



Charopidae 
Endodontidae 




175 275 375 

Mean Height 



4.75 575 over 65 



38 



FIG. 15. Mean shell height distribution in the Endodontidae and 
Charopidae. 

passes the maximum height recorded for the Char- 
opidae. The three highest spired Charopidae are 
Lauopa mbalauuana, Tuimalila infundibulus (Hom- 
bron & Jacquinot, 1841), and T. pilsbryi. All three have 
average to only slightly increased whorl counts, and 
only the latter has a high SP/BWW ratio (mean 2.23). 
Their large height is the result of size increase alone, 
rather than change in shell form. In contrast, the 
high-shelled Endodontidae mostly are species with 
umbilical brood chambers and increased whorl counts, 
plus the few Nesodiscus and pre-brood chamber En- 
dodonta (Solem, 1976b, pp. 27-30). 

Mean diameter distribution (fig. 16) shows a 
greater similarity between the two families, both hav- 
ing a significant right extension of the frequency 
curve. The sharper peak for the Charopidae relates to 



CHAROPID-ENDODONTID CONCHOLOGICAL COMPARISONS 



39 



30 



25 - 



20 - 



Charopidae 
Endodontidae 




1.75 



2.75 3.75 



475 
Mean Diameter 



5-75 6.75 7.75 over 

8.0 



FIG. 16. Mean shell diameter distribution in the Endodontidae 
and Charopidae. 

the speciose genus Sinployea, which accounts for more 
than half of the total taxa reviewed. The secondary 
peak around 4.75 mm. results from some of the large 
Rarotongan and Samoan Sinployea, plus Himero- 
concha, Russatus, and large Semperdon from Micro- 
nesia. Both families have a few taxa that exhibit 
gigantism compared with the average species. 

Mean H/D ratio distribution (fig. 17) differs only 
because of the brood chamber taxa in the Endodontidae 
(Solem, 1976b, p. 29, fig. 19), producing a slight bulge 



30 
25 

20 

~ 
I 
o 15 



- 
I 



-Charopidae 
Endodontidae 




0.325 



0.425 



0.525 0-625 

Mean H/D Ratio 



0.725 



0.825 



FIG. 17. Mean height/diameter ratio distribution in the En- 
dodontidae and Charopidae. 



in the higher ratios for that family. The only very 
high-spired charopids are the nearly scalariform Ba 
humbugi (fig. 74b), Ladronellum mariannarum (Qua- 
dras & Mollendorff, 1894) (fig. lOlb), and Semperdon 
kororensis (Beddome, 1889) (fig. 98b). Only Ladronel- 
lum has a normally open umbilicus (fig. lOlc), whereas 
the others have a closed umbilicus that normally in- 
creases the H/D ratio dramatically (Solem, 1976b, p. 
25, fig. 15). 

The most dramatic difference is shown by the 
mean whorl count distribution (fig. 18). The only 
charopids to exceed a mean whorl count of five whorls 



Charopidae 
Endodontidae 



( I 



10 




3 1 '. 



4'i 



5'', 6'* 

Mean Whorls 



FIG. 18. Mean whorl count distribution in the Endodontidae and 
Charopidae. 



are the Micronesian Semperdon kororensis (Beddome, 
1889) (fig. 98a) with 5V& average, and the Rarotongan 
Sinployea planospira (Garrett, 1881) (fig. 46d) with 6% 
whorl average. Although there is considerable overlap 
in mean whorl counts of five whorls or less, the drama- 
tic increase in higher mean whorl counts in the En- 
dodontidae is quite clear. A fair portion of this increase 
to the right correlates with brood chamber formation 
(Solem, 1976b, p. 29, fig. 20). 

Means of D/U ratio for the Endodontidae in Figure 
19 exclude the brood chamber taxa entirely, hence the 
two families show very similar patterns. Because egg 
laying in the umbilicus is a family characteristic in the 



40 



SOLEM: ENDODONTOID LAND SNAILS 



n 



25- 



20- 



Charopidae 
Endodonlidae 




225 325 4.25 525 6.25 7- 11- 21- crack cloud 

10 20 50 

X D/U Ratio 

FIG. 19. Mean diameter/umbilical width ratio distribution in the 
Endodontidae and Charopidae. 



Endodontidae (Solem, 1976b, pp. 100-101), extreme 
narrowing or closure of the umbilicus should be a rarer 
event in the Endodontidae than in the Charopidae. 
Surprisingly enough, it is not. The only Charopidae 
with a closed or nearly closed umbilicus are Semperdon 
kororensis (Beddome, 1889) (fig. 98c) from Palau, 
Sinployea clista (fig. 51c) from Samoa, and Be humbugi 
(fig. 74c) from Viti Levu, Fiji. Sinployea clausa (fig. 
5 If) from Samoa has an extremely narrowed um- 
bilicus. Most other species have the umbilicus much 
more widely opened, and only in the Lau Archipelago 
Sinployea adposita (Mousson, 1870) (fig. 69f) andRuss- 
atus nigrescens (Mollendorff, 1900) (fig. 89f) from 
Ponape is the umbilicus very narrow. In the Endodon- 
tidae (Solem, 1976b, pp. 26, 491-492) there are 10 taxa 
with closed and 10 with barely perforate umbilici. Most 
of these are found on Rapa and Mangareva, areas far 
from the main areas of distribution and possibly from 
the natural occurrence of potential egg predators that 
occupy the leaf litter. Hence, the unexpected high 
occurrence of closed or barely perforate umbilici in the 
Endodontidae is a geographic phenomenon. 

The degree of spire protrusion, as measured by the 
SP/BWW ratio, is quite different for the two families 
(fig. 20). The Endodontidae are in general much higher 
spired. This relates primarily to the taxa reaching the 
Nesodiscus and brood chamber levels of organization. 
The most elevated Charopidae are Ba humbugi from 
Fiji (fig. 74b), Ladronellum mariannarum (Quadras & 
Mollendorff, 1894) from Guam (fig. lOlb), Sinployea 
angularis from Fiji (fig. 64e), and Vatusila nayauana 
from Fiji (fig. 82e). Because so few Pacific Island 
Charopidae have elevated spires, the type of analysis 
done for the Endodontidae (Solem, 1976b, p. 25, fig. 15) 
in which spire protrusion was correlated with varia- 



Cfiaropidae -.- 
Endodontidae . 




depressed Hal 0.05 0.15 0.25 035 04 055 065 075 085 0.95 overlO 
Spire height/body whorl width ratio 

FIG. 20. Mean spire height/body whorl width ratio distribution 
in the Endodontidae and Charopidae. 



tions in shell height, diameter, H/D ratio, and D/U 
ratio is not presented since the differences were so 
slight. Comparatively few Charopidae have a flat or 
clearly depressed spire. Only Roimontis tolotomensis 
has a clearly sunken spire. 

The striking similarity in median mean rib counts 
on the body whorl (table X) and the very similar distri- 
bution of mean rib counts in the two families (fig. 21) 
was not anticipated. It does suggest that the hypoth- 
esized function of the ribbing to reduce adherence of 
particles to the shell surface (Solem, 1976b, p. 50) may 
be correct. Rib reduction in the Endodontidae is 
primarily size correlated. Species with a mean diame- 
ter of 4.75 mm. or more frequently show a marked de- 
gree of rib reduction (Solem, 1976b, pp. 46-50, tables 



Cliaiofxd* 
Endodontidae 



10 - 




190 over reduce) 
200 



70 110 ISO 

Ribs on Body Whorl 

FIG. 21. Mean ribs on body whorl distribution in the Endodon 
tidae and Charopidae. 



CHAROPID-ENDODONTID CONCHOLOGICAL COMPARISONS 



41 



XVII- XIX). There are comparatively few taxa in the 
Charopidae that show major rib reduction. The 10 are 
not phyletically correlated and show reduction in 
different degrees and ways. In Himeroconcha rotula 
(Quadras & Mollendorff, 1894) and H. lamlanensis 
from Guam, plus Sinployea rudis (Garrett, 1872) and 
S. harveyensis (Garrett, 1872) from Rarotonga, the 
sculpture is reduced to irregularity on the body whorl, 
with the major ribs becoming too crowded and too ir- 
regular to count well before the lip. In Sinployea ir- 
regularis (Garrett, 1887) and Ba humbugi from Fiji 
plus Russatus nigrescens (Mollendorff, 1900) from 
Ponape, the postapical sculpture is highly irregular, 
and occasionally a rib with a high lamellar extension 
appears. In Sinployea recursa from Fiji and 
Himeroconcha quadrasi (Mollendorff, 1894) and H. 
fusca (Quadras & Mollendorff, 1894) from Guam, the 
sculpture is reduced in prominence on both the spire 
and body whorl and rapidly becomes indistinguishable 
from growth lines after an initial portion where the 
ribs are large enough to be counted. The four 
Himeroconcha do show a pattern of size-associated re- 
duction in rib prominence, since the two smaller 
species have more prominent spire sculpture (fig. 
104a-f) than do the two larger taxa (fig. 105a-f). There 
is no data available concerning the ecology of 
Himeroconcha, so that the reasons for this pattern of 
sculpture reduction are unknown. Russatus, Ba, and 
Sinployea irregularis (Garrett, 1887) are known to be 
terrestrial in habitat, and Garrett (1872, pp. 227-228) 
reported that both S. rudis (Garrett, 1872) and S. har- 
veyensis (Garrett, 1872) were collected under rotting 
wood. Only dead examples of the Lau Archipelago 
Sinployea recursa are known, although I have specu- 
lated that it could be an arboreal species. 

The overall pattern of rib reduction in the 
Charopidae is less clearly size linked than in the En- 
dodontidae. Three of the 10 species, S. recursa, S. ir- 
regularis (Garrett, 1887), and Ba humbugi are within 
0.11 mm. of the median mean diameter, the other 
seven (38.9%) are among the 18 over 3.75 mm. in 
diameter, and three (37.5%) are among the eight 
species that average more than 4.75 mm. in diameter. 
They are Russatus nigrescens (Mollendorff, 1900), 
Himeroconcha fusca (Quadras & Mollendorff, 1894), 
and H. quadrasi (Mollendorff, 1894). In the Endodon- 
tidae, 50% of the taxa averaging over 4.76 mm. in 
diameter have reduced sculpture (Solem, 1976b, p. 47, 
table XVIII). 

The variation in mean ribs/mm, on the body whorl 
(fig. 22) is equivalent in the two families, once allow- 
ance is made for the smaller mean size of the Charo- 
pidae. As in the Endodontidae (Solem, 1976b, pp. 
44-45, tables XV- XVI), the smaller the mean diame- 
ter, in general the more numerous are the radial ribs 
(see table X). 

In summary, basic shell size, shape, and radial 
sculpture spacing and frequency in both families are 
more similar than dissimilar, once allowance is made 



Charopidae 
[ndodonlidae 




J I 



H 15 

Ribs/mm 



I I 



over reduced 
22 



FIG. 22. Mean ribs/mm, on the body whorl distribution in the 
Endodontidae and Charopidae. 



for the changes in the Endodontidae caused by the for- 
mation of a brood chamber. The Endodontidae are 
larger and with a higher whorl count than the 
Charopidae, but in basic form and character of the ra- 
dial sculpture they are very similar. This is not sur- 
prising because both are basically found in litter, 
under stones, and in or under rotting wood. A much 
higher number of Charopidae than Endodontidae has 
been taken in semiarboreal to arboreal situations. 

In addition to gross size, shape, and sculpture 
comparisons, an important question is whether there is 
a different pattern of growth between the two families. 
With the complications introduced by variations in 
spire protrusion, body whorl descension, umbilical 
width, whorl counts, and sculptural protrusions, it is 
difficult to find a simple index of whorl increment pat- 
tern that will at least indicate the pattern of whorl 
width increase. Figure 23 attempts this through use of 
a crude measure of size increment plotted against 
whorl count of the figured individual. To try and 
minimize difficulties in interpretation, brood chamber 
taxa and the very high-spired members ofAaadonta in 
the Endodontidae plus the high-spired Ba humbugi in 
the Charopidae have been omitted. Inclusion of these 
would have added data points to the lower right of the 
figure. Because the increase in shell diameter is less 
when the spire is elevated, the results would have been 
artificially skewed toward the right. 

The basic data for Figure 23 were taken from the 
top views of shells as published in Solem (1976b) and 
this report. Because top views were not published for 
species of Anceyodonta, nearly all Cookeconcha, Opan- 
ara, Rhysoconcha, Ruatara, Orangia, Taipidon, 
Planudonta, Rikitea, and Nesophila, this is only a par- 
tial sampling of the Endodontidae. In contrast, nearly 
all of the Charopidae have been figured in top view. On 
each illustration, two measurements were made: the 
width of the first whorl from suture to suture and then 
the diameter of the entire shell. In addition, the 



42 



SOLEM: ENDODONTOID LAND SNAILS 



27- 

26 

25- 

24 

23 

22 

21 

20 

19 

18 



I 16 



Endodontidae 
Charopidae 



D 

D D 

O D 
D DD 

DO D D 

DD Q 

OOD 6 
D 

5 



2 



D 



D D D 

3 



n 



D DO OOD dS 



13 



12 





11 

D 



DQ D rm 
4 

D I I OT 

D DO DQ 

D DD 

D D 



10 

D 



an 



Whorl count 

FIG. 23. Patterns of whorl width increment in selected Endodontidae and Charopidae. Brood chamber, very high spired, and unillustrated 
top views of species in the Endodontidae are omitted, as is the high-spired Ba humbugi in the Charopidae. Unusual taxa are: (1) Minidonta 
manuaensis Solem, 1976; (2) Mautodontha aoraiensis Solem, 1976; (3) Australdonta pseudplanulata Solem, 1976; (4) A. pharcata Solem, 1976; (5) 
Cookeconcha stellulus (Gould, 1844); (6) Thaumatodon laddi Solem, 1976; (7) Zyzzyxdonta alata Solem, 1976; (8) Aaadonta kinlochi Solem, 1976; 
(9)Sinployea canalis (Garrett, 1872); (10) S . planospira (Garrett, 1881); (11) Russatus nigrescens (Mollendorff, 1900); (12) Semperdon kororensis 
(Beddome, 1889); and (13) Minidonta micraconica Solem, 1976. 



number of whorls in the illustrated specimen was re- 
corded. The width of the first whorl was divided by the 
total diameter to give a percentage of the total diame- 
ter taken up by the initial nuclear whorl. This is, of 
course, partly inaccurate, because the suture-to-suture 
distance will be less than the periphery-to-periphery 
distance. The error will be approximately the same for 
each species, so that the degree of error will be in the 
same direction. Use of a percentage index of total 
diameter plotted against the actual whorl count gives 
an approximate indication of basic whorl width incre- 
ments. 



It is evident that members of the Endodontidae 
have a tighter pattern of coiling at the same whorl 
count than do the Charopidae. The higher the percent- 
age of the first nuclear whorl width, the slower is the 
rate of whorl width increment for succeeding whorls. 
There is, in general, a rather clear separation between 
the two family units. The exceptions from the general 
pattern in the Charopidae are a few species with un- 
usual coiling patterns: (number 9 on fig. 23) Sinployea 
canalis (Garrett, 1872) (fig. 49a-c), which has a later- 
ally compressed whorl profile and increased whorl 
count; (number 10) S. planospira (Garrett, 1881) (fig. 



CHAROPID-ENDODONTID CONCHOLOGICAL COMPARISONS 



43 



46d-h), which is laterally compressed, has increased 
whorl count, and very tight coiling; and (number 12) 
Semperdon kororensis (Beddome, 1889) (fig. 98a-c), 
with its raised spire, lateral compression, and closed 
umbilicus. At the other extreme, (number 11) Russatus 
nigrescens (Mollendorff, 1900) (fig. 89d-f), which has a 
flat spire, reduced whorl count, and reduced sculpture, 
is offset to the left considerably. The unusual Endodon- 
tidae include (number 1) Minidonta manuaensis Solem 
(1976b, p. 131, fig. 62a) and (number 13) Minidonta 
micraconica Solem (1976b, p. 138, fig. 65a-c) with re- 
duced whorl counts and abnormally flat-spired taxa in 
normally more elevated genera such as (number 2) 
Mautodontha aoraiensis Solem (1976b, p. 160, fig. 
74d-f), (number 3) Australdonta pseudplanulata Solem 
(1976b, p. 295, fig. 127d-f), and (number 4) A.pharcata 
Solem (1976b, p. 313, fig. 137a-c). Taxa whose diame- 
ter was increased by greatly enlarged radial ribs in- 
clude (number 5) Cookeconcha stellulus (Gould, 1844) 
(Solem, 1976b, p. 218, fig. 93a-c) and (number 7) Zyz- 
zyxdonta alata Solem (1976b, p. 466, fig. 198a-c). 
Aaadonta kinlochi Solem (1976b, p. 486, fig. 208a-c, 
number 8) is a flat-spired species in a genus that is 
normally highly elevated, whereas (number 6) 
Thaumatodon laddi Solem (1976b, p. 452, fig. 193d-f) 
is a flat-spired species with slightly protruded 
periphery. Thus, all exceptions are readily explained 
as representing special situations. 

Correlated with the more rapid whorl width in- 
crement in the Charopidae is a greater increase in 
cross-sectional area of the whorl profile. Without sec- 
tioning shells and measuring the areas of the whorls, 
quantification of this difference is not feasible. The fact 
of the greater cross-sectional area and concomitant in- 
crease in linear wall distance at any given point un- 
doubtedly has had major effects on the placement of 
pallial organs in relation to each other and their 
length. It also has permitted thickening of genital or- 
gans over the endodontid condition. Discussion of these 
changes was dealt with under the patterns of anatomi- 
cal variation. 

On the submicroscopic level, there are three major 
differences between the Endodontidae and Charopidae 
found on the Pacific Islands. The apical sculpture of the 
shell, the method of forming the postapical shell 
sculpture, and the way in which the apertural barriers 
are formed and armed with microdenticles are 
different. These differences have been reviewed in part 
elsewhere (Solem, 1969d). Only a brief summary is in- 
cluded here. 

In the Endodontidae, the apical sculpture consists 
of prominent radial ribs, with or without microriblets 
in between, plus very fine spiral cords that are best 
termed "squiggly" (Solem, 1976b, pp. 35-41, figs. 
25-31). In one genus, Aaadonta Solem (1976b, pp. 
38-39, figs. 28-29), the major radial sculpture has 
been lost, only the microradials are left on the postapi- 
cal whorls, and only the squiggly spiral cords are left 
on the apex. In the Pacific Island Charopidae, the basic 



apical sculpture is of strong spiral cords, typically as 
seen in Sinployea modicella (Ferussac, 1840) (fig. 
la-c). They can be reduced in prominence and in- 
creased in number, as in Sinployea peasei (fig. 2a-b), or 
combined with a secondary sculpture of low radial 
swellings, as in Tuimalila pilsbryi (fig. 2c). The only 
exceptions on the Pacific Islands concern the rotadis- 
cine genus Microcharopa, in which the spiral apical 
cords are broken up into short, twisted segments (fig. 
4a-e), and the widespread Discocharopa (fig. 5), in 
which both apical and postapical sculpture consists of 
fine radial ribs and there is no trace of spiral sculpture. 
The situation becomes much more complicated in the 
charopid taxa of Australia and New Zealand where 
there have been many experiments in shell sculpture 
(Solem, unpublished data). However, in relation to the 
Pacific Island taxa, the sharp division into taxa with 
spiral cords (Charopidae) and taxa with radial ribs 
(Endodontidae) holds with a few secondary exceptions. 
Under optical magnification, Aaadonta would be con- 
fused with the charopid condition and Tuimalila with 
the endodontid condition, but SEM studies show that 
these apparent exceptions are secondary modifications. 

Except for obvious periostracal setae and exten- 
sions in such taxa as Cookeconcha decussatulus (Pease, 
1866) (Solem, 1976b, p. 36, fig. 26a-c; p. 40, fig. 30b), 
the postapical sculpture in the Endodontidae appar- 
ently is formed by a thin template of periostracum, 
with most of the sculpture thickness consisting of un- 
derlying calcium layers. This holds even for the fine 
apical features (Solem, 1976b, p. 35, fig. 25d). In con- 
trast, the microsculpture and much of the major rib 
projection in the Charopidae consists entirely of perio- 
stracal materials. Frequently the only calcareous 
sculptural element will be a swelling underneath each 
major radial rib. An example of this is seen in 
Sinployea uicaria vicaria (Mousson, 1871) (fig. 59). The 
same pattern of structure seems to hold true for at least 
many of the New Zealand and Australian Charopidae. 
This makes the similarity in gross sculpture effect be- 
tween the two families even more remarkable. 

An initial review of the apertural barrier 
differences between the Endodontidae and Charopidae 
was given in Solem (1973b). The following summary is 
taken from that paper, Solem (1976b, pp. 52-72), and 
pp. 15-23. In the Endodontidae, the barriers show a 
uniform pattern of structure, growth, and microden- 
ticulation. The parietal and columellar walls will have 
weak pustulations, as is typical of many taxa in a vari- 
ety of families (Solem, 1972c), but the characteristic 
feature on the barrier tops is a series of additive tri- 
angular microdenticulations (Solem, 1973b, figs. 7-13, 
23-24; Solem, 1976b, p. 55, fig. 39d-e; p. 64, figs. 40, 
41a-c). It apparently does not matter whether the ex- 
panded upper edge of the barrier is continuous or 
broken up into a series of expanded beads as in 
Thaumatodon, Aaadonta, and Zyzzyxdonta (Solem, 
1976b, p. 457, fig. 194b-e; p. 466, fig. 198b; p. 475, fig. 
203b, e). The pattern of minute triangular denticles 



44 



SOLEM: ENDODONTOID LAND SNAILS 



that face toward the aperture and are additions to the 
surface of the barrier, rather than an outgrowth from 
it, holds. The only known exceptions are in certain of 
the very large species of Cookeconcha and Endodonta, 
where the denticulations on the upper edge of the bar- 
riers are blunt-tipped (Solem, 1976b, p. 66, fig. 42d-f) 
although retaining their triangular shape on the sides 
of the barriers, and in other genera such asNesodiscus, 
where the barriers are greatly reduced in both size and 
number. These barriers may be secondarily without 
denticulations. I have thus concluded that the barriers 
seen in members of the Endodontidae had a common 
origin and that reduction or loss of barriers is a secon- 
dary phenomenon in that family. 

In the Charopidae (pp. 15-17), in contrast, three of 
the four major groups with barrier-equipped genera 
found on the Pacific Islands show barriers that differ in 
size, position, growth, and microdenticulations (Solem, 
1973b, pp. 303-304, figs. 4-6, 14-22). The latter basi- 
cally are crystals growing out of the barrier surface 
and are blunt tipped. It is thus concluded that aper- 
tural barriers have evolved independently in each of 
these lineages, despite the fact that SEM observations 
could not be made on the Smp/ojea-derivative genera. 
Their barriers show such obvious differences from the 
other three groups (figs. 80-83, 85) that I have no 
doubt of their independent origin. More detailed dis- 
cussion of the barrier structure and variation has been 



given in the Patterns of Morphological Variation (pp. 
15-23). 

In the Endodontidae the great majority of the 
species have barriers on the palatal wall, with a 
bladelike shape and gradual anterior descension. In 
the Charopidae only four species have bladelike 
palatal barriers, and 13 have purely crescent-shaped 
barriers. 

The frequency of barrier occurrence is lower in the 
Charopidae than in the Endodontidae. In the latter, 
including taxa not seen but illustrated in older litera- 
ture, 184 out of 185 (99.5%) have parietal barriers and 
163 (88.1%) have palatal barriers. In the Charopidae 
32 (33.7%) of 95 taxa have parietals, and 27 (28.4%) 
have palatal barriers. Thus the families differ signifi- 
cantly not only in whether or not barriers are present 
in the aperture but also in how they are formed, the 
normal shape of the palatals, and the nature of the 
microdenticulations on the barrier edges. 

Despite great optical similarity in size, color, 
shape, presence of apertural barriers in many species, 
and presence of fine radial ribbing in at least the 
smaller taxa, differences in whorl count (fig. 18), 
growth pattern of whorl width (fig. 23), apical 
sculpture, method of formation for the postapical 
sculpture, and details of the apertural barrier 
structure and microdenticulations serve to distinguish 
the two families on conchological grounds. 



HABITAT RANGE AND EXTINCTION 



In contrast with the Endodontidae (Solem, 1976b, 
pp. 100-101), the Charopidae have been able to exploit 
the semiarboreal and arboreal habitats as well as re- 
maining well represented in the ground stratum. Their 
closed secondary ureter has been hypothesized (Solem, 
1976b, p. 100) as the reason the charopids have been 
able to become successfully semiarboreal. A second fac- 
tor in their success is the apparent lack of umbilical 
egg deposition by the charopids. Of all the Pacific Is- 
land charopid specimens studied, only one example of 
Tuimalila infundibulus (Hombron & Jacquinot, 1841) 
had a snail egg capsule inside the umbilicus, whereas 
this is the routine pattern in the Endodontidae. Of ex- 
tralimital taxa, the New Zealand charopids, Fectola 
marsupialis Powell, 1941, and Aeschrodomus worleyi 
Powell, 1928, have the umbilical egg-laying pattern 
(see Climo, 1969a, pp. 218-219; 1970, p. 306). 

On such drastically ecologically altered islands as 
Viti Levu, Tahiti, and Moorea, specimens ofSinployea 
and Ba have been collected from lowland vegetational 
remnants in the 1960s and 1970s. They have not, how- 
ever, been found where the ground stratum has been 
severely and regularly disturbed by chickens and/or 
pigs. In parts of Samoa and on the Lau Archipelago of 
Fiji the charopids still are common and relatively 
abundant. Only on Rarotonga is there clear evidence of 
major charopid extinctions within a century. Two sepa- 
rate collecting efforts on Rarotonga in the mid-1960s 
failed to find nine out of 10 Sinployea species collected 
in the 1860s and 1870s and five out of six endodontids 
reported from the same island. All of the missing taxa 
were described as ground stratum inland species. The 



one collected endodontid occupies the storm line coral 
boulder habitat, and the one previously described 
Sinployea is at least occasionally semiarboreal. 

The actual egg-laying habits of the Pacific Island 
Charopidae are unknown, but their nondependence 
upon the umbilicus for egg-laying probably has effec- 
tively lowered ant predation, which apparently was a 
major factor in the mass extinction of the Pacific Island 
Endodontidae. The semiarboreal to arboreal habitat 
range extension of the Charopidae also has lessened 
human impact through introductions, because arboreal 
snails of the Pacific Islands, in general, have been able 
to adapt to plantations of bananas and even coffee, 
whereas strictly ground-dwelling taxa are absent from 
these human-introduced environments. 

The charopids are abundant in Australia, South 
Africa, New Zealand, New Caledonia, and parts of 
South America all areas in which ants are common. I 
thus expect that the ability of the Pacific Island 
charopids to survive despite the introduction of preda- 
cious ants is a carryover from conditions in their areas 
of greater abundance rather than a new adaptation. 
Since ants first appeared in the Cretaceous fossil rec- 
ord (Wilson et al., 1967), and have an extensive fossil 
record of living tribes and genera from the Oligocene 
and Miocene, their coexistence with the basic charopid 
stock seems highly probable. 

The charopids thus occupy a wider range of 
habitats than do the Endodontidae and persist in fair 
variety despite the great ecological alterations of the 
past century on the Pacific Islands. 



45 



PHYLOGENY AND CLASSIFICATION 



Reviews of the hypothesized early evolution and 
progressive trends among land snails have been given 
previously (Solem, 1974; 1976b, pp. 102-104; 1978b). 
The position of the Endodontidae as the most gen- 
eralized extant group of the Sigmurethra, the absence 
of any family unit from which the endodontid anatomi- 
cal structures can be derived, and the positioning of 
both the Charopidae and Endodontidae within the 
Arionacea complex have been discussed earlier and 
will not be rejustified here. 

The Charopidae are more advanced than the En- 
dodontidae in having a closed secondary ureter, fused 
prostate-uterine tubes with a common lumen, a re- 
duced number of lobes in the ovotestis, and in transfer- 
ring sperm in a packet (see Ba humbugl, fig. 75f-g), 
which correlates with the general presence of an 
epiphallus. Charopids are more complex than the En- 



dodontidae in having great elaboration of terminal 
genital structures and in the variety of their radular 
structures. Recent dissections of some undescribed 
Western Australian taxa have revealed genera that 
are partly transitional between the Endodontidae and 
Charopidae. These will be described and discussed 
elsewhere. 

It is premature to discuss in detail the probable 
phylogenetic relationships among the endodontoid 
families. After synoptic coverage of the Austro- 
Zelandic taxa, such a discussion will be profitable. At 
this time I choose to end the discussion of interfamily 
phylogeny by emphasizing that the Charopidae are 
more advanced than and derivable from the Endodont- 
idae. The phylogenetic position of the Punctidae in re- 
lation to the other families is still uncertain. 



46 



FAMILY CLASSIFICATION OF THE ENDODONTOIDS 



A list of subfamily and family names available for 
members of this complex was given in Solem (1976b, p. 
105). I recognize five families and group the assignable 
names as below. The name Patulinae Tryon (1866, p. 
243) is discarded (see Solem, 1976b, p. 105). The af- 
finities of the Indian Thysanotinae Godwin-Austen, 
1907, are still unknown. Available literature informa- 
tion on their anatomy is not adequate, and I have not 
been able to obtain preserved materials for dissection. I 
have not dissected one of Iredale's family units, the 
Hedleyoconchidae, but on the basis of shell structures 
have no hesitation in lumping it with the Charopidae 
pending further study. Definitions and discussion of 
the Helicodiscidae (see Solem, 1975) and Discidae 
(Solem, in preparation A) are not included here, since 
both families are extralimital to this study. Definitions 
of both the Punctidae and Charopidae are included 
below in the systematic review; the Endodontidae were 
defined by Solem (1976b, p. 121). 

The proposed synonymization of family level units, 
in chronological order, is: 

Family Punctidae (Morse, 1864, p. 27) 

+ Laominae Suter (1913, p. 732) 

+ Paralaomidae Iredale (1941a, p. 263) 
Family Charopidae Mutton (1884b, p. 199) 

+ Phenacohelicidae Suter (1892a, p. 270) 

+ Otoconchinae Cockerel! (1893, pp. 188, 205) 

+ Flammulinidae Crosse (1894, p. 210) 

+ Rotadiscinae H. B. Baker (1927, pp. 226, 230) 

+ Amphidoxinae Thiele (1931, p. 575) 



+ Dipnelicidae Iredale (1937b, pp. 22-23) 

+ Hedleyoconchidae Iredale (1942, pp. 34-35) 

+ Pseudocharopidae Iredale (1944, p. 312) 
Family Endodontidae Pilsbry, 1895 (Pilsbry, 1893-1895, p. 

xxviii) 

Family Helicodiscidae Pilsbry, 1927 (in Baker, 1927, pp. 226, 
230) 

+ Stenopylinae Thiele (1931, p. 569) 
Family Discidae Thiele ( 1931, p. 578) 

+ Goniodiscinae Wagner (1927, p. 305) 

Although I do not accept strict nomenclatural 
priority for names of families and higher level taxa, the 
above usage comes close to following this precept. I 
reject both Patulinae Tryon (1866) and Goniodiscinae 
(Wagner, 1927); the former name because Patula 
mostly has been associated with members of the land 
snail family Oreohelicidae, a totally unrelated group, 
until finally assigned as a synonym of Discus. 
Goniodiscus Fitzinger, 1833, is now ranked as a sub- 
genus or synonym of Discus (see discussion by Forcart, 
1957), and I consider it inappropriate to use as a 
family-level name. 

The morphological gaps between these families 
are of the same order of magnitude or greater than the 
gaps separating recognized families of the Limacacea 
(see Solem, 1976b, p. 107), although the gaps between 
the family units certainly are not equal in size. 

Discussion of subfamily and generic units is de- 
ferred to the family discussions of the Punctidae and 
Charopidae. 



47 



PHYLOGENETIC PROCEDURES 



I am a pragmatic phylogeneticist in the sense of 
Mayr and have used neither Hennigian cladistics nor 
phenetic manipulations in this study. The criteria used 
are those cited by Solem (1976b, p. 108) and the three- 
tiered approach to evolutionary change outlined in 
Solem (1978b). These are suited to the peculiarities of 
molluscan shell growth, with the ontogenetic develop- 
ment from embryo on visible in old age, the water- 
dependent ecology of land snails, and their use of ter- 
minal genitalia for species recognition. 



The absence of many fixed reference points within 
the snail's body and the difficulties of making precise 
and meaningful measurements on the shell combine to 
make quantification of many features impossible and 
thus use of the more sophisticated mathematical ma- 
nipulations of data matrices impractical. Similarly, in 
the absence of equivalently detailed analyses of most 
other land snail families and with the uncertainty as to 
the overall phylogeny of either characters or taxa, use 
of cladistic methodology is premature. 



48 



GENERIC CLASSIFICATION 



The same type of "pigeon-hole" generic units cited 
for the Endodontidae (Solem, 1976b, pp. 118-119) 
existed for the Pacific Island Charopidae. If there were 
apertural barriers present, the species was referred to 
the form genus Endodonta or to one of its sections, and 
if there were no such barriers present, it was placed in 
the New Zealand genus Charopa. Although Pilsbry 
(1893-1895, p. 21) had pointed out the artificiality of 
this division, no significant changes were introduced 
subsequently. Reference of a new species to Flam- 
mulina (Russatus nigrescens) by Mollendorff (1900) or 
attempts to use other described genera than Charopa 
(Solem, 1959a, 1960) did not alter the essential absence 
of any meaningful classification on the generic level. 



Of the 21 genera in the Punctidae and Charopidae 
reviewed below, only two, Punctum and Discocharopa, 
have available names. The other 19 are named here. 
The same criteria for generic separation are used here 
in the Charopidae as were used in classifying the En- 
dodontidae into genera (see Solem, 1976b, pp. 119- 
120). Thus the generic concepts are directly compar- 
able and the minimum generic gap equally applied. 
The difference in frequency of monotypic genera in the 
two families and presence of one particularly speciose 
genus in the Charopidae was summarized by Solem 
(1976b, p. 120). This is a real difference in evolutionary 
patterns and not an artifact of classification. 



49 



SYSTEMATIC REVIEW 



Because the two families are so disparate in the 
number of species reviewed and the complexities of 
charopid classification require considerable reference 
to extralimital taxa, the discussion of charopid 
phylogeny and classification is subtended to the family 
review. 

Following are a formal list of the taxa reviewed 
and geographic keys to the genera. These sections fol- 
low the format used in Solem (1976b) as do the species 
and generic accounts. 

LIST OF THE TAXA 

Family Punctidae Morse ( 1864, p. 27) 

Genus Punctum Morse, 1864 
Punctum sp. Society Islands: Tahiti, Mt. Aorai 
Punctum polynesicum, new species Austral Islands: Tubuai, 
Raivavae 

Family Charopidae Hutton (1884b, p. 199) 
Subfamily Rotadiscinae H. B. Baker (1927, pp. 226, 228) 

Genus Microcharopa, new genus 

Microcharopa rnimula, new species Fiji Islands: Viti Levu; Lau Ar- 
chipelago (Munia, Mothe, Wangava, Nayau, Namuka, Yangasa 
Levu) 

Subfamily Charopinae Hutton ( 1884b, p. 199) 

Genus Discocharopa Iredale, 1913 

Discocharopa aperta (Mollendorff, 1888) Philippines, Indonesia, 
New Guinea, Western Australia, Northern Territory, Queensland, 
Bismarck Archipelago, New Hebrides, Fiji Islands, Samoan Is- 
lands, Austral Islands (Rurutu), Society Islands (Borabora) 

Genus Sinployea, new genus 

Sinployea modicella (Ferussac, 1840) Society Islands: Moorea 
Sinployea tahitiensis, new species Society Islands: Tahiti 
Sinployea lame/licosta (Garrett, 1884) Society Islands: Tahiti 
Sinployea montana, new species Society Islands: Tahiti 
Sinployea neglecta, new species Society Islands: Huahine 
Sinployea sp. Society Islands: Borabora 

Sinployea atiensis (Pease, 1870) Cook Islands: Atiu, Aitutaki 
Sinployea andrewi, new species Cook Islands: Mangaia 
Sinployea peasei, new species Cook Islands: Rarotonga 
Sinployea avanaensis, new species Cook Islands: Rarotonga 
Sinployea proximo (Garrett, 1872) Cook Islands: Rarotonga 
Sinployea planospira (Garrett, 1881) Cook Islands: Rarotonga 
Sinployea rudis (Garrett, 1872) Cook Islands: Rarotonga 
Sinployea decorticata (Garrett, 1872) Cook Islands: Rarotonga 
Sinployea harveyensis (Garrett, 1872) Cook Islands: Rarotonga 
Sinployea youngi (Garrett, 1872) Cook Islands: Rarotonga 
Sinployea canalis (Garrett, 1872) Cook Islands: Rarotonga 
Sinployea otareae (Garrett, 1872) Cook Islands: Rarotonga 
Sinployea tenuicostata (Garrett, 1872) Cook Islands: Rarotonga 
Sinployea clausa, new species Samoan Islands: Manu'a Group 
(Tau) 



Sinployea clista, new species Samoan Islands: Tutuila, Upolu 

Sinployea aunuuana, new species Samoan Islands: -Aunuu off 
Tutuila, possibly Manu'a Group (Tau) 

Sinployea allecta allecta (Cox, 1870) Samoan Islands: Upolu, 
Savai'i 

Sinployea allecta tauensis, new subspecies Samoan Islands: Manu'a 
Group (Tau) 

Sinployea complementaria (Mousson, 1865) Samoan Islands: Upolu 

Sinployea intermedia, new species Swains Island 

Sinployea vicaria vicaria (Mousson, 1871) Hoorn Islands: Futuna; 
Tonga Islands: Vava'u, Tongatapu, Eua, Ha'apai Group 

Sinployea vicaria paucicosta, new subspecies Tonga Islands: 
Vava'u 

Sinployea rotumana (Smith, 1897) Rotuma 

Sinployea ellicensis ellicensis, new species and subspecies Ellice Is- 
lands: Funafuti 

Sinployea ellicensis nukulaelaeana, new subspecies Ellice Islands: 
Nukulaelae 

Sinployea pseudovicaria, new species Ellice Islands: Vaitupu 

Sinployea kiisaieana, new species Caroline Islands: Kusaie 

Sinployea sp. Mariana Islands: Saipan (possibly introduced) 

Sinployea angularis, new species Fiji Islands: Lau Archipelago 
(Namuka) 

Sinployea recursa, new species Fiji Islands: Lau Archipelago 
(Wangava) 

Sinployea princei (Liardet, 1876) Fiji Islands: Taveuni, Kandavu 

Sinployea inermis inermis (Mousson, 1870) Fiji Islands: Lau Ar- 
chipelago (Vanua Mbalavu, Mango, Kimbombo) 

Sinployea inermis meridionalis, new subspecies Fiji Islands: Lau 
Archipelago (Yangasa Levu, Navutu-i-Loma, Aiwa) 

Sinployea inermis lakembana, new subspecies Fiji Islands: Lau Ar- 
chipelago (Lakemba) 

Sinployea lauensis, new species Fiji Islands: Lau Archipelago 
(Nayau, Yangasa Levu, Navutu-i-Loma, Wangava) 

Sinployea navutuensis, new species Fiji Islands: Lau Archipelago 
(Navutu-i-Loma, ? Oneata) 

Sinployea adposita (Mousson, 1870) Fiji Islands: Lau Archipelago 
(Oneata, Mothe, Munia, Lakemba, Aiwa, Karoni, Nayau, Yangasa 
Levu) 

Sinployea irregularis (Garrett, 1887) Fiji Islands: Viti Levu 

Sinployea godeffroyana, new species Fiji Islands: Viti Levu 

Sinployea euryomphala (Solem, 1959) New Hebrides: Espiritu 
Santo, Maewo, Gaua, Vanua Lava 

Sinployea solornonensis (Clapp, 1923) Solomon Islands: Ugi Island 
off San Cristobal 

Sinployea kuntzi (Solem, 1960) Solomon Islands: Florida Island off 
Tulagi 

Sinployea nissani (Dell, 1955) Solomon Islands: Nissan (Green Is- 
land), north of Bougainville 

Sinployea novopommerana (Rensch, 1937) Bismarck Archipelago: 
New Britain 

Sinployea descendens (Rensch, 1937) Bismarck Archipelago: New 
Britain 

Genus Ba, new genus 
Ba humbugi, new species Fiji Islands: Viti Levu 

Genus Maafu, new genus 

Maafu thaumasius, new species Fiji Islands: Lau Archipelago 
(Nayau) 



50 



SYSTEMATIC REVIEW 



51 



Genus Lauopa, new genus 

Lauopo mbalavuana, new species Fiji Islands: Lau Archipelago 
(Vanua Mbalavu) 

Genus Tuimalila, new genus 

Tuimalila infundibulus (Hombron & Jacquinot, 1841) Tonga Is- 
lands: Vava'u 
Tuimalila pilsbryi, new species Tonga Islands: Eua 

Genus Lagivala, new genus 

Lagivala davidi (Ladd, 1968) Ellice Islands: Funafuti (fossil) 
Lagivala vivus, new species Fiji Islands: Viti Levu 
Lagivala minusculus, new species Fiji Islands: Viti Levu 
Lagivala macroglyphis (Rensch, 1937) Bismarck Archipelago: New 

Britain 
Lagivala microglyphis (Rensch, 1937) Bismarck Archipelago: New 

Britain 
Lagivala demani (Tapparone-Canefri, 1883) Indonesia (Timor, 

Ambon, Aru Islands); West Irian (Misool) 

Genus Vatusila, new genus 

Vatusila kondoi, new species Fiji Islands: Lau Archipelago (Nayau) 
Vatusila nayauana, new species Fiji Islands: Lau Archipelago 

(Nayau) 
Vatusila eniwetokensis (Ladd, 1958) Marshall Islands: Eniwetok 

(fossil) 

Vatusila tongensis, new species Tonga Islands: Eua 
Vatusila vaitupuensis, new species Ellice Islands: Vaitupu 
Vatusila niueana, new species Niue Island 

Genus Graeffedon, new genus 

Graeffedon graeffei (Mousson, 1869) Samoan Islands: Upolu 
Graeffedon savaiiensis, new species Samoan Islands: Savai'i 
Graeffedon pricei, new species Tonga Islands: Tongatapu 

Subfamily Trukcharopinae, new subfamily 

Genus Trukcharopa, new genus 

Trukcharopa trukana, new species Caroline Islands: Truk, 
Lukunor 

Genus Kubaryiellus, new genus 
Kubaryiellus kubaryi (Mollendorff, 1900) Caroline Islands: Ponape 

Genus Russatus, new genus 
Russatus nigrescens (Mollendorff, 1900) Caroline Islands: Ponape 

Genus Roimontis, new genus 
Roimontis tolotomensis, new species Caroline Islands: Ponape 

Genus Palikirus, new genus 

Palikirus cosmetus, new species Caroline Islands: Ponape 
Palikirus ponapicus (Mollendorff, 1900) Caroline Islands: Ponape 

Genus Jokajdon, new genus 

Jokajdon tumidulus (Mollendorff, 1900) Caroline Islands: Ponape 
Jokajdon callizonus (Mollendorff, 1900) Caroline Islands: Ponape 

Genus Palline, new genus 
Palline notera notera, new species and subspecies Palau Islands: 

Peleliu 

Palline notera palauana, new subspecies Palau Islands: Koror 
Palline notera gianda, new subspecies Palau Islands: Babelthuap 
Palline biakensis, new species West Irian: Biak 
Palline micramyla, new species Caroline Islands: Ponape 

Subfamily Semperdoninae, new subfamily 

Genus Semperdon, new genus 
Semperdon uncatus, new species Palau Islands: Angaur, Aulup- 

tagel, Koror 
Semperdon xyleborus, new species Palau Islands: Angaur, Koror, 

Ngemelis, Peleliu 
Semperdon kororensis (Beddome, 1889) Palau Islands: Koror 



Semperdon rotanus, new species Mariana Islands: Rota, northern 

tip of Guam 
Semperdon heptaptychius (Quadras & Mollendorff, 1894) Mariana 

Islands: Guam 

Genus Ladronellum, new genus 

Ladronellum mariannarum (Quadras & Mollendorff, 1894) 
Mariana Islands: Guam 

Genus Himeroconcha, new genus 

Himeroconcha lamlanentiis, new species Mariana Islands: Guam 
Himeroconcha rotula (Quadras & Mollendorff, 1894) Mariana Is- 
lands: Guam 
Himeroconcha quadrasi (Mollendorff, 1894) Mariana Islands: 

Guam 

Himeroconcha fusca (Quadras & Mollendorff, 1894) Mariana Is- 
lands: Guam 

Incertae sedis 

Helix filiola Ferussac, 1840 Tonga Islands 
Helix oceanica Le Guillou, 1842 Society Islands: Tahiti 
Helix minutialis Deshayes, 1851 Society Islands: Tahiti 
Helix multispirata Hombron & Jacquinot, 1852 Tonga Islands: 

Vava'u 
Helix rotula Hombron & Jacquinot, 1852 (not Lowe, 1831) Gambier 

Islands: Mangareva 
Pithys verecunda Pease, 1870 Marquesas 



GEOGRAPHIC KEYS TO THE GENERA 

The following artificial keys supplement those 
presented for the Endodontidae in Solem (1976b, pp. 
124-126). They are designed to enable identification of 
adult shells. Endodontid taxa are cross-referenced 
below but usually are not keyed down to the generic 
level in order to save space and printing costs. Because 
there are no charopids or punctids known from the 
Mangareva, Marquesas, Rapa, or Tuamotu Islands, 
these areas are not included in this set of keys; nor are 
keys presented for extralimital areas. Additions to this 
set of keys are the Caroline, Mariana, and main Fijian 
Islands. For islands not included in the main groups, 
reference to the list of "odd island" taxa at the end of 
the keys may help. 

AUSTRAL ISLANDS 

1. Apical whorls with radial ribs 2 

Apical whorls with spiral cords 

Punctum polynesicum (p. 58) 

2. Shell minute, diameter less than 1.8 mm.; at most 1 deeply 

recessed parietal barrier 

Discocharopa aperta (Mollendorff, 1888) (p. 76) 
Shell normally much more than 1.8 mm. in diameter; usually 

several apertural barriers 

Endodontidae (Solem, 1976b, p. 125). 



CAROLINE ISLANDS 

1. Aperture without barriers 2 

Aperture with 1 or more barriers 6 

2. Whorls increasing rapidly in width (fig. 89d); mean D/U ratio 

about 10; mean whorl count about 3 l /e 

Russatus nigrescens (Mollendorff, 1900) (p. 215) 

Whorls increasing less rapidly in width (fig. 89a); mean D/U 

ratio 3.40-4.00; mean whorl count usually more than 3'/2. ...3 



52 



SOLEM: ENDODONTOID LAND SNAILS 



3. TrukorKusaie 4 

Ponape 5 

4. Kusaie; spire elevated (fig. 63a); mean rib count on body whorl 

about 91 Sinployea kusaieana (p. 143) 

Truk; spire barely elevated (fig. 63d); mean rib count on body 
whorl about 150 Trukcharopa trukana (p. 208) 

5. Spire fiat or sunken; ribs on body whorl about 132; mean diame- 

ter about 3.75 mm 

Kubaryiellus kubaryl (Mbllendorff, 1900) (p. 212) 
Spire slightly elevated; ribs on body whorl about 46; mean 

diameter about 2.2 mm 

Palikirus ponapicus (Mollendorff, 1900) (p. 221) 

6. No columellar or palatal barriers 7 

Palatal and/or columellar barriers present 8 

7. Only 1 parietal barrier; spire elevated; 25 large ribs on body 

whorl Palikirus cosmetus (p. 219) 

8. Parietal barriers 2; spire sunken; about 60 low ribs on body 

whorl Jtoimontis tolotomensis (p. 217) 

Parietal and palatal barriers many and large 9 

Barriers consist of 1 low parietal, 2 low palatals 

Palling micramyla (p. 234) 

9. Spire more elevated; ribs larger; body whorl rounded (fig. 

92e) Jokajdon tumidulus (Mollendorff, 1900) (p. 223) 

Spire barely elevated; ribs smaller; body whorl laterally com- 
pressed behind aperture (fig. 92b) 

Jokajdon callizonus (Mollendorff, 1900) (p. 226) 

COOK ISLANDS 

1. Apical whorls with radial ribs; at least 1 apertural barrier 

Endodontidae (Solem, 1976b, p. 125) 

Apical whorls with spiral cords; no apertural barriers 

Sinployea, key on p. 98 



ELLICE ISLANDS 

1. Apical whorls with spiral cords 2 

Apical whorls with radial ribs 

Endodontidae, Thaumatodon decemplicata (Mousson, 1873) 

(Solem, 1976b, p. 451) 

2. Aperture with barriers 3 

Aperture without barriers Sinployea, see p. 134 

3. Vaitupu; living; 3 small barriers 

Vatusila vaitupuensis (p. 196) 

Funafuti; fossil; 6 large barriers 

Lagivala davidi (Ladd, 1968) (p. 184) 



FIJI, MAIN ISLANDS 

1. Apical whorls with spiral cords 2 

Apical whorls with radial ribs 

Discocharopa aperta (Mollendorff, 1888) (p. 76) 

2. Aperture without barriers 3 

Aperture with barriers 5 

3. Diameter less than 1.3 mm Microcharopa mimula (p. 71) 

Diameter more than 2.0 mm 4 

4. Spire greatly elevated (fig. 74b); umbilicus closed 

Ba humbugi (p. 172) 

Spire nearly flat or slightly elevated; umbilicus open 

Sinployea, see key on p. 147 

5. Parietal barriers 3; D/U ratio about 2.80; ribs on body whorl 

about 62 Lagivala uiuus (p. 185) 

Parietal barrier 1; D/U ratio about 5.25; ribs on body whorl 
about 120 Lagivala minusculus (p. 185) 



HAWAIIAN ISLANDS 

Apical whorls with radial ribs, or, in larger species, smooth 

Endodontidae (Solem, 1976b, p. 125) 



Apical whorls with spiral cords; shell minute 

Punctum horneri (Ancey, 1904) (p. 57) 

LAU ARCHIPELAGO, FIJI 

1. Apical whorls with radial ribs 2 

Apical whorls with spiral cords 3 

2. Several apertural barriers; diameter more than 2.5 mm 

Endodontidae (Solem, 1976b, p. 125) 

One or no apertural barriers; diameter less than 2 mm 

Discocharopa aperta (Mollendorff, 1888) (p. 76) 

3. Aperture without barriers 4 

Aperture with 1 or more barriers 6 

4. Diameter more than 2.0 mm 5 

Diameter less than 1.3 mm Microcharopa mimula (p. 71) 

5. Periphery protruded into a threadlike keel (fig. 76b); ribs on 

body whorl about 44 and very large 

Maafu thaumasius (p. 176) 

Periphery at most angulated (fig. 68e); ribs reduced in size and 
usually more numerous Sinployea, see key on p. 147 

6. Diameter less than 2.0 mm 7 

Diameter more than 7.0 mm.; 1 parietal barrier 

Lauopa mbalavuana (p. 177) 

7. Palatal barriers 3; parietal barrier bifid 

Vatusila kondoi (p. 192) 

Palatal barrier 1; parietal barrier simple 

Vatusila nayauana (p. 194) 

MARIANA ISLANDS 

1. Aperture without barriers 2 

Aperture with barriers 3 

2. Shell small, diameter less than 2.5 mm.; Saipan 

Sinployea sp. (p. 145) 
Shell large, 3.8-7.2 mm.; Guam Jfimeroconcha (p. 259) 

3. Aperture with 3 very small barriers (fig. lOlb) 

Ladronellum mariannarum (Quadras & Mbllendorff, 1894) 

(p. 255) 

Aperture with 6 to many barriers (fig. 99b, d-e) 

Semperdon rotanus and S. heptaptychius (Quadras & Mollen- 
dorff, 1894) (pp. 245, 247) 

MARSHALL ISLANDS 

1. Apical whorls with spiral cords 

Vatusila eniwetokensis (Ladd, 1958) (p. 195) 

2. Apical whorls with radial ribs 

Endodontidae (Solem, 1976b, p. 125) 



PALAU ISLANDS 

1. Postapical whorls with microradial riblets only, no major radial 

ribs Endodontidae, Aaadonta (Solem, 1976b, p. 473) 

Postapical whorls with major ribs and microsculpture 2 

2. Shell diameter less than 2.6 mm.; mean whorl count less than 

4V* 3 

Shell diameter more than 2.7 mm.; mean whorl count more 
than4V2 5 

3. Palatal barriers 4, low ridges (figs. 94e, 95a); Koror and Babel- 

thuap 4 

Palatal barriers 5, high blades (fig. 94b); Peleliu 

Palline notera notera (p. 229) 

4. Palatal barriers 2; Koror Palline notera palauana (p. 231) 

Palatal barriers 3; Babelthuap. ..Palline notera gianda (p. 231) 

5. Palatal barriers many (fig. 98b, e) 6 

Palatal barriers 0-1 (fig. 97b) Semperdon uncatus (p. 238) 

6. Mean diameter about 4.6 mm.; umbilicus a very narrow 

slit Semperdon kororensis (Beddome, 1889) (p. 244) 

Mean diameter 2.9 mm.; umbilicus widely open 

Semperdon xyleborus (p. 24P 



SYSTEMATIC REVIEW 



53 



TABLE XI. - SPECIES NUMBERS OF 
ENDODONTOID LAND SNAILS FROM SELECTED GEOGRAPHIC AREAS 



PUNCTIDAE 



CHAROPIDAE 



Polynesia, Fiji 
Hawaii 1 



Known 

1 

1 



Melanesia 
New Guinea 
Indonesia 
New Caledonia'* 
Lord Howe Id . 
Norfolk Id. 
Kermadec Is. 8 



Australia 
Tasmania 
Victoria 
N. S. Wales 
Queensland 
S. Australia 
N. Territory 



7 

2 

80 

14 
9 
3 

4 
1 
6 



St. Helena 

Neotropica and 
Juan Fernandez 



New Known 



29 
16 
11 
9 

49 

19 

8 

7 

147 

38 
18 

28 
14 
2 
3 

10 

114 

11 

55(?) 

2 






TOTALS 



646 



Solem (1976b, this monograph, in preparation) 
Solem (1959a, 1960, 1962, 1963, this monograph) 

3 Solem (1958b, I970a) 

4 Solem (1961) 

Collecting by L. Price and P. Bouchet 
Iredale (1944) and subsequent collecting 
Iredale (1945) and subsequent collecting 

8 Iredale (1913) 

9 Powell (1976) 

Iredale (1937a, 1937c, 1939) 

U Burch (1976) 

Collecting by A. Solem 



New 



ENDODONTIDAE 
Known New 



143 
31 
11 



50(7, 5 ' 12 



?? 



14 



3 

290 



293 



Connolly (1939); Solem (1970c) 
Solem (1978a) 
Various papers and Odhner (1922) 
Pilsbry (1948) and various papers 
F. Clirao, personal communication 



SAMOAN ISLANDS 

1. Apical whorls with radial ribs 2 

Apical whorls with spiral cords 3 

2. Aperture without palatal barriers; umbilicus very wide 

Discocharopa aperta (Mollendorff, 1888) (p. 76) 



Aperture with palatal barriers; umbilicus rather narrow 

Endodontidae (Solem, 1976b, p. 126) 

3. Aperture with prominent barriers 4 

Aperture without barriers Sinployea, see key on p. 119 

4. Shell diameter about 3.3 mm.; whorls 4Vi; Savai'i 

Graeffedon savaiiensis (p. 202) 

Shell diameter over 5.0 mm.; whorls about 5; Upolu 

Graeffedon graeffei (Mousson, 1869) (p. 200) 

SOCIETY ISLANDS 

1. Apical whorls with radial ribs 2 

Apical whorls with spiral cords 3 

2. Shell diameter less than 2.0 mm.; at most 1 deeply recessed 

parietal barrier 

Discocharopa aperta (Mollendorff, 1888) (p. 76) 
Shell diameter over 2.25 mm.; usually several parietal barriers 

or shell diameter over 3.0 mm 

Endodontidae (Solem, 1976b, p. 126) 

3. Shell minute, diameter less than 1.5 mm....Punctum sp. (p. 57) 
Shell larger, diameter over 2.4 mm Sinployea, see p. 86 

TONGA 

1. Apical whorls with spiral cords predominating, at most low ra- 

dial swellings 2 

Apical whorls with narrow radial ribs 

Endodontidae (Solem, 1976b, p. 126) 

2. Aperture without barriers 3 

Aperture with prominent barriers 5 

3. Diameter less than 3.5 mm Sinployea, see key on p. 134 

Diameter more than 5.0 mm 4 

4. Eua; mean diameter about 6.7 mm. ..Tuimalila pilsbryi (p. 179) 
Vavau; mean diameter about 5.7 mm 

Tuimalila infundibulus (Hombron & Jacquinot, 1841) (p. 181) 

5. Palatal wall bearing a broad ridge (fig. 83b); Eua 

Vatusila tongensis (p. 196) 

Palatal wall with 3 barriers (fig. 88e); Tongatapu 

Graeffedon pricei (p. 205) 

A few species are known from islands that do not 
fall into the major groups. These species are: 

Futuna Sinployea vicaria vicaria (Mousson, 1871) (p. 134) 
Niue Vatusila niueana (p. 198) 
Rotuma Sinployea rotumana (Smith, 1897) (p. 138) 
Swains Sinployea intermedia (p. 131) 

It must be emphasized that the above keys are 
adequate for identification of adult specimens in which 
the sculpture and apertural barriers can be observed. 
They are less adequate for juveniles and quite in- 
adequate for very worn examples. It still will be possi- 
ble to collect new species in areas such as the Caroline, 
Palau, Mariana, Lau, and main Fijian Islands that 
may or may not key out to a correct genus. 



FAMILY PUNCTIDAE 

Generally minute to small endodontoids, primitively with spiral 
apical sculpture and a combination of macro- and microradial post- 
apical sculpture that is periostracal in origin. Sculpture altered in 
larger taxa. Whorl counts and shape highly variable, particularly in 
New Zealand taxa. Umbilicus open, regularly decoiling to nearly 
closed. Color monochrome to highly tessellated. A few taxa develop 
apertural barriers, but most lack barriers. Foot of animal undivided. 
Prominant pedal and suprapedal grooves unite above tail without 
forming a caudal foss. Radula with small tricuspid central. 



54 



SOLEM: ENDODONTOID LAND SNAILS 



lateromarginals bicuspid with three minute accessory cusps, major 
cusps becoming reduced in prominence on outer teeth, but not tend- 
ing to split into additional cusps. Basal plates long and slender, cusps 
tending to point directly forward. Jaw of separated, square plates, 
partly fused in larger species. Pallial complex with a bilobed kidney, 
lobes generally equal in length, primary ureter originating at apex of 
pericardia! kidney lobe, reflexing posteriorly as secondary ureter, 
opening at ureteric pore alongside to moderately behind the anus 
and just inside pneumostome. Mantle collar often without lobes or 
lap>. but a glandular extension onto pallial roof present in some taxa. 
Genital system variable, but apical sections typically as follows: 
ovotestis of two closely appressed, variously subdivided, teardrop- 
shaped lobes lying just above stomach apex; hermaphroditic duct a 
simple, uncoiled tube entering laterally into carrefour swelling; 
talon with globular head on a short stalk entering apex of carrefour; 
albumen gland elongate-ovate, deeply indented by loops of intestine 
and head of spermatheca. Prostate-uterus partly fused to fused, free 
oviduct short to long, uterus with different glandular zones. Sper- 
matheca with lower shaft expanded, upper shaft more slender after 
tapering, head ovate, expanded, lying imbedded in base of albumen 
gland above pallial cavity apex. Vas deferens a slender tube entering 
penis complex near insertion of penial retractor muscle to well below 
muscle insertion. Penis complex with a penis-derived epiphallic sec- 
tion in many taxa, interior of penis with long and irregular pilasters, 
often with a short vergic papilla. Epiphallic section, when present, 
with slightly different textured pilasters. Entire complex slender and 
tuhelike. Penial retractor muscle arising from diaphragm, inserting 
on head of penis or epiphallic section, without complexities of inser- 
tion. Interior of free oviduct and base of spermathecal shaft with 
irregular folds in the few examined. Digestive and free muscle sys- 
tems agreeing with endodontid and charopid patterns. No data are 
recorded on the nervous system. 

The family unit Punctidae is used here in a re- 
stricted sense, not as the umbrella for the endodontoids 
as suggested by Climo (1969a, 1971a, 1978). Pending 
anatomical confirmation, I include the Punctidae or 
Punctinae of Northern Hemisphere workers, Laominae 
in the sense of Suter (1913) and Gabriel (1930), and 
Paralaomidae of Iredale (1941a, 1944). As a rough es- 
timate, about 159 named taxa from the southern parts 
of Australia, New Zealand, subantarctic islands, 
Holarctic, and scattered African localities would be in- 
cluded. Only a very few taxa have had other than jaw 
and radula data recorded in the literature, and it is 
quite probable that some of the included species and 
genera will prove to be charopids and some taxa as- 
signed to the charopids will prove to be punctids. Par- 
ticularly when shell sculpture becomes reduced, a 
common pattern in the Austro-Zelandic taxa, assign- 
ment to a family group on shell features alone becomes 
extremely hazardous. 

The anatomy o{Punctum Morse (1864) was illus- 
trated and discussed by Baker (1927, pp. 227-228, pi. 
16, figs. 8-12) then reprinted in Pilsbry (1948, pp. 
641-643, fig. 349a-f). Data on the shell sculpture for- 
mation was given by Solem (1977b, pp. 150-152, figs. 
4-6, 11-12). Anatomical data are set forth here on 
Punctum (fig. 24a-b), the New Zealand Laoma 
leimonias (Gray, 1850) (fig. 25a-h), Western Austra- 
lian Paralaoma aprica Iredale, 1939 (fig. 26a-c), and 
Tasmanian rr Paralaorna" coesa (Legrand, 1871) (fig. 
27a-d). These extralimital taxa are illustrated to show 
the essential unity of structure within the Punctidae 
and to provide data for comparing charopid patterns. 




FIG. 24. Partial anatomy ofPunctum minutissimum (Lea). Cedar 
bog, Woodburn Road, 4 miles southwest of Urbana, Champaign 
County, Ohio. E. Keferl! X-20-1969. FMNH 151102: a, pallial region; 
b, apical genitalia with albumen gland removed. No scale lines pre- 
pared. (CW). 



To my knowledge, no other taxa have been illustrated 
in equivalent detail. An anatomical description of 
Laoma leimonias follows to provide detailed compara- 
tive data. 

Laoma leimonias (Gray, 1850). 

Description of soft part*. Foot and tail slender, rounded pos- 
teriorly, not tapering noticeably. Sole undivided, without corruga- 
tions i fig. 25d). Pedal grooves high on foot, suprapedal very much 
smaller than pedal, uniting above tail. Pedal groove passing around 
tail with a distinct constriction on top of tail. No protruding caudal 
horn, but top of tail indented, with subpedal region extending pos- 
teriorly. No middorsal groove. Slime network very inconspicuous. 
Body roundly truncated anteriorly. 

Body color in preservative yellow white, ommatophores black. 
Surface of pallial roof and visceral hump with frequent black speck- 
les, less frequent white marks. 

Mantle collar (MC) elongated (fig. 25a), with bluntly rounded 
anterior edge. Basal margin with long intrusion of mantle gland onto 
pallial roof, corresponding with position of large left anterior mantle 
lobe (MA I at parietal-palatal angle. Anus (A) opening at inner edge 
of mantle collar (fig. 25b), slightly anterior to external urinary pore 
(KX). Urinary chamber (LK) a sharply defined groove that is im- 
mediately united with the V-shaped anal channel passing forward in 
the mantle collar. 

Pallial region (fig. 25a-b) very elongated, extending P/i-2 
whorls apically from aperture. Basal Vi whorl with glandular exten- 
sion of mantle collar (MG). longest where major palatal lamella is 
formed and reabsorbed. Kidney (K) U-shaped, pericardia! and rectal 
branches equal in length. Rectal arm of kidney lapping completely 
under hindgut (HG) and extending onto parietal margin of pallial 
cavity (fig. 25c). Thus, rectal arm lies on both parietal and upper 
palatal margins of pallial cavity. Pericardia! arm of kidney appears 
more slender in Figure 25a, but solely because pericardium lies un- 
derneath kidney. Hindgut (HG) paralleling parietal-palatal margin 
well past apex of pallial region. Loop of intestine (I) with apical base 




KD ^-^\ Q 

""" "p^~" 




HV 



2 mm 





1mm 






x 




^ 



^r 2 



GD 



FIG. 25. Anatomy of Laoma Icimonian iGrayi. Herekino Gorge, Kaitaia, Northland, North Island, New Zealand. L. Price! XI-1962. FMNH 
135401: a, pallia! region; b, detail of pneumostomal openings; c, detail showing extent to which kidney <K) overlaps hindgut (HG); d, extended 
foot showing pedal grooves and lack of a caudal horn; e, genitalia; f, junction of hermaphroditic duct (GO) and talon <GT); g, detail of uterine 
structure; h, internal structure of penis and epiphallus. Scale lines as marked. (PS). 



55 



56 



SOLEM: ENDODONTOID LAND SNAILS 



of kidney extended above it for a considerable distance. Ureter (KD) 
sigmurethrous, arms without any lung roof visible between them, 
not expanded on any portion. When first removed, apical rectal arm 
of ureter hidden by parietal extension of kidney, but in Figure 25a 
this is stretched outward. Heart (H) comparatively large, about half 
as long as kidney, lying partly under pericardia! arm of kidney, 
reaching pallial roof only on basal margin of whorl. Principal pulmo- 
nary vein (HV) narrow, reaching mantle gland, without noticeable 
secondary venation. 

Ovotestis (G) composed of 1-2 clumps of finger-like lobes (fig. 
25c), buried in apical section of digestive gland above stomach- 
intestine reflexion, lighter in color than digestive gland. Hermaph- 
roditic duct (GD) slender at first, gradually becoming a thick, lumpy, 
muscular tube, slightly iridescent in tone. Duct narrows abruptly 
just at base of albumen gland then reflexes up along albumen gland 
(GG) before entering duct of talon (GT). Latter a slender tube with 
expanded head (fig. 25f), entering head of prostate-uterus without 
differentiated carrefour. Prostate (DG) composed of a few relatively 
large acini fastened to wall of upper uterine chamber. Lower part of 
prostate squeezed between uterine sections. Uterus (UT) three- 
chambered: upper section (UT,) a very thin-walled tube, opening into 
a second chamber (fig. 25g) with glandular walls and two or more 
low, rounded pilasters (UT 2 ). Lower chamber (UT,) barely distin- 
guished externally from second chamber, a thin-walled sac with 
lamellar plates on the wall. 

Vas deferens (VD) arising from narrowed base of prostate, a 
slender tube becoming moderately expanded, then narrowed again 
before inserting laterally on epiphallus (E) between arms of a narrow 
U-shaped stimulator (fig. 25h). Vas not tightly bound to penioviducal 
angle. Epiphallus weakly differentiated from penis, a short swollen 
tube with a U-shaped stimulator and a narrow neck before joining 
the penis proper (P). At apex of penis lies a high muscular stimulator 
(PP), ovate in outline and attached at both ends, its apical tip lying 
between the arms of the epiphallic stimulator (fig. 25h). 

Below penial stimulator, there are two weak "pilasters" caused 
by folding of the penis wall and very faint spiral pustulations. Bas- 
ally, penis tapering to a very slender tube before entering short at- 
rium ( Y). Gonopore a short slit behind and below right ommatophore. 

Free oviduct (UV) long with thick muscular wall, a glandular 
collar separating its base from spermathecal pore. Spermatheca (S) 
long, expanded head lying partly in albumen gland and partly in 
digestive gland, lying above pallial cavity. Lower part of sper- 
mathecal stalk slightly larger than oviduct, thin-walled with distinct 
longitudinal pilasters, narrowing at base of prostate-uterus to a very 
slender tube, partly buried in acini of prostate. Vagina (V) very 
short, not differentiated internally from atrium and base of penis. 

Buccal mass of average length, high, strongly tapered an- 
teriorly, generative sac small. Buccal retractor massive, not split, 
inserting at about posterior third of buccal mass in U-shaped fan. 
Esophagus initially large in diameter, then slender for length of 
pallial cavity, inserting slightly behind midpoint of buccal mass. 
Interior of esophagus with weak longitudinal pilasters, becoming 
fainter apically. Esophagus extends about 1 A whorl past head of pal- 
lial cavity, entering stomach along lower parietal margin of shell. 
Stomach very large, reaching surface of parietal and upper palatal 
margins, extending apically for slightly more than one whorl. Intes- 
tine reflexing along columellar margin to lower margin of kidney 
base, looping up along kidney base to just below hindgut, reflexing 
posteriorly and diagonally downward to base of stomach, then loop- 
ing up to parietal-palatal margin and running forward as hindgut. 
Hindgut follows parietal-palatal margin along edge of pallial cavity 
to anus. Loop of intestine abutting kidney projects into kidney, which 
extends outside of intestine apically, thus separating body wall from 
pallial roof. 

Salivary glands white, lying alongside esophagus, touching me- 
dially, near posterior end. Salivary ducts straight, entering laterally 
on base of esophagus. 

Digestive gland extending from apex of soft parts to apex of 
pallial cavity, reduced in region of stomach, distinctly darker in color 
than ovotestis. 



Free muscle system simple: buccal retractor massive, not split 
anteriorly, uniting with tail fan to form columellar retractor about 
two whorls above aperture. Right ommatophoral retractor passing 
between penioviducal angle. Rhinophoral retractors very short, unit- 
ing with ommatophorals just before union with tail fan. Entire free 
muscle system short and massive, extending apically less than 2M> 
whorls from aperture. 

(Based on 4 adult examples from south side of Herekino Gorge, 
10 mi. southwest of Kaitaia, Northland, New Zealand (35 17' S, 173 
12' E), FMNH 135401. Collected by L. Price, November 1962.) 

Of the dissected species, Punctum minutissimum 
(Lea, 1841) andParalaoma aprica (Iredale, 1939) are in 
the 1.5-2.0 mm. size range; Laoma leimonias (Gray, 
1850) is a very high-spired shell, about 2.3 mm. in 
diameter and 2.7 mm. high with 7 whorls; and "Para- 
laoma" coesa (Legrand, 1871) is the largest species, 
about 3.5-4.0 mm. in diameter, with 4Vz whorls. As a 
systematic comment, Punctum and Paralaoma may 
well be synonyms, but "Paralaoma" coesa probably 
does not belong to the same genus. There are 
nomenclatural problems in dealing with this species. 
The very long, free oviduct and penis of Laoma is re- 
lated to the greatly increased whorl count of that 
species and has no special systematic significance. 

The basic anatomical features of the Punctidae 
are: (1) the bicuspid lateromarginal teeth with minute 
accessory cusps (fig. 12b-e), long basal plates, and for- 
ward pointing of the rounded cusps; (2) the ovotestis of 
two teardrop-shaped lobes that may be appressed (figs. 
24b, 25e, 26c) or modestly separated (fig. 27b) and are 
subdivided to various extents (compare figs. 24b and 
27b with Baker, 1927, p. 16, fig. 8); (3) the bilobed 
kidney of approximately equal lobe length, with (fig. 
27a) or without (figs. 24a, 25a, 26a) lung roof visible 
between the ureter arms; (4) the slender penis complex 
with simple insertion of the penial retractor muscle 
that internally has a penis-derived epiphallic section 
followed by vergic papilla and smooth regular to ir- 
regular pilasters; and (5) normally rather slender ter- 
minal female genitalia with tapering lower sper- 
mathecal stalk, internally with free oviduct, vagina, 
and spermatheca sharing smooth pilaster sculpture. 

The Charopidae contrast in having: (1) tri cuspid 
laterals and marginals with rectangular basal plates, a 
simplified interrow tooth support system during feed- 
ing, and upward-slanting pointed cusps (figs. 13-14); 
(2) the ovotestis consisting of one to two clumps of sev- 
eral long follicles that lie parallel to the shell axis (figs. 
90i, lOOb-c, g); (3) a generally bilobed kidney in which 
the lobes may be equal in length (figs. 39c, 90a), the 
rectal much longer (figs. 53a, e, 90e), or infrequently 
the pericardial lobe is longer (fig. lla), and there may 
(fig. 39c) or may not (fig. 90e) be lung roof visible be- 
tween the arms of the ureter; (4) the penis complex 
normally is short and fat (fig. 90b, f, i), with simple to 
complex insertion of the penial retractor muscle, the 
penis frequently with a verge, complex stimulators and 
stimulator pockets, a vas deferens-derived epiphallus 
in many taxa, and the lower penis area without 
sculpture; and (5) normally thick terminal female 



SYSTEMATIC REVIEW 



57 












FIG. 26. Partial anatomy of Paralaoma aprica Iredale. Station 
WA-98, north end of townsite reserve, Nangeenan, west of Merredin, 
Western Australia. A. Solem and L. Price! 11-22-1974. FMNH 
182222: a, pallial region; b, postapical genitalia; c, apical genitalia. 
Scale lines as marked. (EL). 



genitalia with very thick spermathecal base that 
rather abruptly narrows to the mid stalk, internally 
often with very prominent pilasters that may be corru- 
gated (fig. 9d) or smooth but very sharply defined (fig. 
87c). 

The Endodontidae contrast to both in having: (1) 
bicuspid laterals and marginals with the marginal ec- 
tocones split (Solem, 1976b, p. 91, fig. 54a-f), rectangu- 
lar basal plates with interrow support system (Solem, 
1976b, p. 89, fig. 52d), and rather sharp elevation of the 
pointed cusps (Solem, 1976b, p. 89, fig. 52b); (2) the 
ovotestis of multiple clumps angled to the shell axis 
(Solem, 1976b, p. 372, fig. 163c); (3) the kidney with at 
most a weak rectal lobe developed and only a primary 
ureter that opens at the front edge of the rectal kidney 
lobe near the posterior of the pallial cavity (Solem, 
1976b, p. 459, fig. 195a); (4) a slender penis that nor- 
mally has only two simple longitudinal pilasters, never 
a verge or vergic papilla, only rarely a penis chamber- 
derived epiphallic section (Solem, 1976b, p. 396, fig. 
171b-c, h); and (5) very thin terminal female genitalia 
without special internal sculpture and with highly var- 



iable insertion of the spermathecal stalk (Solem, 
1976b, p. 84, fig. 48a-c). In addition, the complete sep- 
aration of the prostate and uterus and apparent ab- 
sence of a sperm packet in the Endodontidae are major 
differences from either of the other families. 

The changes outlined above are noncorrelated, and 
their different states in the three families indicate that 
the families represent three separate patterns of diver- 
sification. These differences are greater than those 
seen between limacacean families and thus merit rec- 
ognition as family units. Whether the punctid radula is 
correlated with specialized feeding habits as opposed to 
those of the Charopidae and Endodontidae is unknown. 

The Punctidae are only marginally present in the 
Pacific Island area, with a scattering of records in 
Hawaii, a single record from the mountains of Tahiti, 
and a new species, Punctum polynesicum, described 
here from the Austral Islands. Thus, no attempt is 
made at revising the generic classification or discuss- 
ing detailed patterns of distribution. Reference of these 
Pacific specimens to Punctum is based strictly on con- 
chological similarity in the absence of preserved 
specimens. The distinction between Paralaoma and 
Punctum may not be maintained in the future, but the 
shells of the Pacific Island taxa are far more similar to 
the Holarctic Punctum than to the Austro-Zelandic 
Paralaoma. In view of the limited material, no diag- 
nosis is presented of the genus Punctum. 

Genus Punctum Morse, 1864 

Type species. Helix minutissima Lea, 1841, by 
original designation. 

There are about 50 sets of Punctum from various 
Hawaiian Islands in the Bernice P. Bishop Museum 
which may be lumped temporarily as Punctum horneri 
Ancey (1904, p. 66, pi. 5, figs. 11-12), but no material 
has been reported previously from Polynesia. A single 
specimen (BPBM 145298) from Station 865, Mt. Aorai 
Trail, Tahiti, collected between 5,600 and 6,300 ft. ele- 
vation, can be identified as a Punctum, but is too frag- 
mentary for any attempt at giving it a specific name. 
The high spire and narrow umbilicus of the fragment 
combine with the finer and less protracted sculpture to 
place it in Punctum rather than Striatura. 

Euconulus (Monoconulus) conoidea H. B. Baker 
(1941, pp. 214-215), the only non-Hawaiian Pacific Is- 
land Euconulus known to date, was taken at the same 
collecting station. Presence of these otherwise Ha- 
waiian and Holarctic taxa in Tahiti at high elevations 
could be explained by the deus ex machina of accidental 
transport on the feathers of migratory birds. 

More abundant material of quite different species 
from the Austral Islands is described below as 
Punctum polynesicum. Although somewhat atypical in 
whorl count and heavy sculpture, it is nearer to 
Punctum than any other genus known to me. Only 
dead specimens were available. Measurements are 
summarized in Table XII. 



58 



SOLEM: ENDODONTOID LAND SNAILS 



UT 




FIG. 27. Anatomy of "Paralaoma" coesa (Legrand). Station TA-38, Circular Head, near Stanley, North Tasmania. L. Price! XI-8-1966. 
FMNH 154974: a, pallial region; b, genitalia; c, interior of penis; d, interior of free oviduct and spermatheca. Scale lines as marked. (NB). 



Punctum polynesicum, new species. Figure 28a-c. 

Diagnosis. Shell minute, diameter 0.95-1.02 mm. (mean 0.99 
mm.), with 3V8-3%+ rather tightly coiled whorls. Apex and spire 
moderately and evenly elevated, body whorl descending more 
rapidly, spire protrusion about V4 body whorl width (0.30-0.34 mm.), 
H/D ratio 0.563-0.597 (mean 0.580). Apical whorls IMs, sculpture of 
about 15 fine spiral cords, whose interstices are 3-4 times their 



width. Postnuclear whorls with narrow, rounded, strongly protrac- 
tively sinuated, prominent radial ribs, 72-83 (mean 77.3) on the 
body whorl, whose interstices are about twice their width until end of 
body whorl where ribbing becomes crowded. Ribs/mm. 23.2-25.9 
(mean 24.9). Microsculpture of fine radial riblets, 2-3 between each 
pair of major ribs, crossed by equally fine spiral riblets. Umbilicus 
shallow, broadly V-shaped, regularly decoiling, contained 3.33-3.44 
times (mean 3.37) in the diameter, margins rounded. Sutures deep, 



SYSTEMATIC REVIEW 



59 



TABLE XII. - RANGE OF VARIATION IN PUNCTUM, DISCOCHAROPA, AND MICROCHAROPA. 



NUMBER OF 
SPECIMENS 



NAME 

Punctum 

polynesicum 6 

Discocharopa 

aperta(Mlldf.) 310 



Microcharopa 
mlmula 



RIBS 

77.3(72-83) 
110.6(77-148) 



RIBS/MM. 
24.9(23.2-25.9) 
26.9(18.0-40.4) 



126.6(117-136) 36.5(32.3-40.5) 



HEIGHT DIAMETER H/D RATIO 

0.58(0.56-0.61) 0.99(0.95-1.02) 0.580(0.563-0.597) 

0.59(0.48-0.74) 1.36(1.18-1.84) 0.438(0.341-0.615) 

0.48(0.43-0.53) 1.07(0.95-1.13) 0.449(0.400-0.469) 



WHORLS UMBILICUS D/U RATIO 

polyn. 3 1/4(3 1/8-3 3/8) 0.29(0.28-0.30) 3.37(3.33-3.44) 
aperta 3 l/2-(3 1/8-4 1/4) 0.54(0.43-0.92) 2.58(2.00-3.21) 



APICAL 
CORDS 



MAJOR 
RADIALS 



SPIRE 

ELEVATION 



BODY WHORL 
WIDTH 



SP/BWW 

0.08(0.07-0.10) 0.32(0.30-0.34) 0.255(0.216-0.316) 
0.07(0.03-0.15) 0.40(0.35-0.44) 0.170(0.086-0.286) 



mimula 3 l/4-(3 1/8-3 3/8) 0.40(0.35-0.46) 2.68(2.43-3.05) 17.8(16-20) 0.05(0.04-0.07) 0.32(0.28-0.34) 0.186(0.118-0.289) 



whorls strongly rounded, with slight lateral compression above 
periphery and on basal margin. Aperture circular, inclined about 30 
from shell axis. Color light yellow-horn, without darker markings. 

Punctum polynesicum is the smallest species yet 
described within the genus. Its very prominent radial 
ribbing and rather wide umbilicus separate it from 
most species. Punctum vitreum H. B. Baker, 1930, is 
perhaps nearest in strength of ribbing, but is a much 
larger shell with a lower H/D ratio. 

Description. Shell minute, with slightly more than 3V4 rather 
tightly coiled whorls. Apex and spire moderately and evenly ele- 
vated, body whorl descending much more rapidly, H/D ratio 0.589. 
Apical whorls I'/i, sculpture partly eroded. Postnuclear sculpture 
and microsculpture as in diagnosis, 83 ribs on the body whorl. Um- 
bilicus broadly V-shaped, regularly decoiling, contained 3.35 times 
in the diameter. Sutures and whorl contour typical, aperture shape 
and inclination as in diagnosis. Height of holotype 0.59 mm., diame- 
ter 1.02 mm. 

Holotype. Austral Islands: Raivavae, Station 
622, Ahuoivi Point. Collected dead under hau leaves by 
Y. Kondo, D. Anderson, and C. M. Cooke, Jr., on Au- 
gust 9, 1934. BPBM 147099. 

Range. Tubuai and Raivavae, Austral Islands. 

Paratypes. Raivavae: west side Vaiaunana Pe- 
ninsula (Station 587), 100 yd. inland at 200 ft. eleva- 
tion, under dead leaves (three specimens, BPBM 
146725). Tubuai: southwest of Murivai (Station 698), 
300-400 yd. inland under Barringtonia trees (two 
specimens, BPBM 147679). 

Remarks. Without preserved material for dissec- 
tion, classification of this species will remain uncer- 
tain. The presence of endodontoid reticulated micro- 
sculpture between the major ribs, rather than the 
transverse growth wrinkles seen in Striatura (Solem, 
1977b, p. 151, fig. 1) has dictated my placing this 
species in Punctum. In several respects P. polynesicum 
agrees more with Striatura. The latter averages fewer 
whorls (3 l /4-3%) and has a wider umbilicus than does 
Punctum with its 3%-4% whorls and generally nar- 
rower umbilicus. Apical sculpture in Striatura is gen- 
erally more prominent and narrower, whereas in 
Punctum it is wider and lower. The low whorl count 



and fine apical sculpture of P. polynesicum thus appear 
somewhat anomalous. 

Neither genus has been reported from South 
Pacific Islands, although both live in Hawaii. A typi- 
cally sculptured Punctum is reported from Tahiti 
above, but this species is very different in form, color, 
shell thickness, and ribbing from P. polynesicum. Mi- 
crocharopa could be confused on the basis of size, but 
that species has radically different apical sculpture 
(see fig. 4, p. 13), a much lower H/D ratio (mean 0.449), 
and many more, much finer radial ribs (117-136 on the 
body whorl). 

An alternative possibility is that this species rep- 
resents the outermost Polynesian representative of the 
Charopidae and is related to Sinployea. I consider it 
much more probable that it is a Punctum accidentally 
introduced by migratory birds. In the absence of dissec- 
tions, no subgeneric reference would be meaningful. 

FAMILY CHAROPIDAE 

Small to large endodontoids, apical sculpture of shell very di- 
verse, postapical whorls primitively with major radial ribs and a 
microsculpture of radial riblets and secondary spiral elements, most 
sculpture features composed of periostracal elements only. Many 
taxa with sculpture secondarily reduced to lost. Whorl counts and 
shell shape highly variable. Umbilicus widely open to closed. Color 
monochrome to brightly tessellated. Apertural barriers evolved in- 
dependently in several lineages, most species without trace of bar- 
riers. Visceral hump and shell greatly reduced to almost lost in a few 
taxa. Foot of animal undivided, prominent pedal and suprapedal 
grooves uniting above tail, in some taxa forming a caudal foss or 
mucus pore. Radula with tricuspid central slightly to distinctly 
smaller than normally tricuspid laterals, rarely bicuspid or with only 
one cusp. Marginals frequently tricuspid, sometimes with ectoconal 
splitting. Basal plates of central and marginal teeth rectangular, 
interrow support system well developed, latter absent on marginals. 
Cusps of major teeth sharp and tending to point upward with ec- 
tocones elevated much more than in Endodontidae. Jaw of separated, 
squarish plates in smaller species, becoming partly to completely 
fused in larger taxa. Pallial complex usually with a distinctly bilobed 
kidney, relative size of lobes quite variable among species, plus a 
closed secondary ureter that normally opens just inside pneumo- 
stome. Some taxa with only a partial secondary ureter that opens far 



60 



SOLEM: ENDODONTOID LAND SNAILS 






h H 

FIG. 28. a-c, Punctum polynesicum, new species. Station 622, 
Ahuoivi Point, Raivavae, Austral Islands. Holotype. BPBM 147099. 
Scale line equals 1 mm. (LH). 



posteriorly in the pallial cavity, and/or rectal kidney lobe greatly 
reduced. Mantle collar normally without prominent lobes or laps, a 
glandular extension onto pallial roof found in some taxa. Genital 
system highly variable. Ovotestis typically of one or two clumps of 
long follicles that lie parallel to the outer wall of the body cavity and 
occur sequentially. Some taxa with the ovotestis broken up into a 
series of lobes that are perpendicular to the outer wall of the body 
cavity and widely separated from each other, a few taxa with follicles 
shortened and resembling those of the Punctidae. Hermaphroditic 
duct a simple unkinked, usually iridescent tube entering usually 
laterally onto apex of carrefour. Talon a finger-like projection or 
swollen head on a short stalk. Albumen gland irregular in shape, 
deeply indented by loops of intestine and head of spermatheca. Pros- 
tate and uterus fused with partial common lumen, uterus with sepa- 
rate and sequential glandular sections. Free oviduct short to long, 
thin, base of spermatheca and upper vagina normally swollen. Sper- 
mathecal shaft tapering abruptly, head above apex of pallial cavity, 
an expanded oval, to elongate oval structure. Vas deferens slender, 
normally entering a clearly differentiated epiphallus, often through 
a complex valvular pore. Many taxa with epiphallus secondarily 
compressed into penis or absent. Penis usually fat, at least in upper 
section, not usually long and slender, lower part often a thin tube. 
Interior of penis frequently with a well-developed verge, circular 
ridges, stimulatory pilasters, and pocket stimulators. Penis retractor 
arising from diaphragm, usually very short, inserting in simple to 
complex fashion on the penis or epiphallus. Interior of lower female 
tract simple to with very large longitudinal simple to corrugated 
pilasters, openings to free oviduct and/or spermathecal shaft varying 
from simple pores to with complex valves. Penis, so far as observed, 
enervated from right cerebral ganglion. Digestive and free muscle 
system typically endodontoid. 

The Charopidae, as rather broadly defined here, 
includes the Phenacohelicidae, Otoconchinae, Flam- 
mulinidae, Rotadiscinae, Amphidoxinae, Dipnelicidae, 
Hedleyoconchidae, and Pseudocharopidae of authors. 
In terms of species, it is the dominant land snail group 
of New Zealand (Powell, 1976; Climo, 1969a-b, 1970, 
1971a-b, 1978), New Caledonia (Franc, 1957; Solem, 
1961), parts of Australia (Tasmania, Victoria, less 
abundant in New South Wales, South Australia, 
southwestern Western Australia, sparse in the North- 
ern Territory, well represented in the rain forest areas 
of Queensland; Iredale, 1937a-c, 1939, 1941a-b; 
Burch, 1976; Solem, in preparation C), Lord Howe Is- 
land (Iredale, 1944), Norfolk Island (Iredale, 1945), and 
Kermadec Islands (Iredale, 1913). The Charopidae is 
only sparsely represented in Indonesia (Solem, 1958), 
New Guinea (Solem, 1958, 1959b, 1970a), Solomon Is- 
lands (Solem, 1960), and New Hebrides (Solem, 1959a). 
Charopids occur with varying abundance on the is- 
lands surveyed in this report. 

Extralimital to the Pacific Basin, species of 
charopids are common to dominant in southern Africa 
(Connolly, 1939; Solem, 1970c), were common on St. 
Helena (Solem, 1977c), and Notodiscus (Solem, 1968c) 
is found on the Subantarctic Islands (Kerguelen, Pos- 
session, Heard, Marion, Amsterdam). There is a mod- 
est Juan Fernandez radiation (Ohdner, 1922) and 
many scattered descriptions and records from southern 
South America, the Andes, and Central America. Two 
species occur in western North America as far north as 
Idaho and Montana (Pilsbry, 1948). 

Including the taxa discussed and described below, 
approximately 646 named species are included in the 



SYSTEMATIC REVIEW 



61 



Charopidae. This is a very conservative estimate of ac- 
tual extant diversity because Climo (personal com- 
munication) has large numbers of undescribed New 
Zealand taxa, a rain forest patch survey of Queensland 
and Northern New South Wales revealed approxi- 
mately 30 undescribed taxa (Burch, 1976), and my own 
1974 and 1976-1977 field collections in southwestern 
Western Australia produced at least 15 new species 
(Solem, in preparation C). In contrast, several years of 
intensive fieldwork in Victoria by Brian Smith and 
associates from the National Museum of Victoria have 
confirmed the described diversity and greatly extended 
ranges but have not produced new taxa. It is thus im- 
possible to present an accurate estimate of diversity in 
the Charopidae at the present time, except to state that 
the currently recorded taxa are significantly less in 
number than the extant taxa. 

A breakdown on estimated endodontoid species 
numbers is given in Table XI. The grand total of 1,494 
described and collected but undescribed taxa contrasts 
with the species estimate given by Boss (1971, p. 108) 
of only 502 species of endodontoid land snails. A 
critique of Boss's assumptions and procedures was in- 
cluded in Solem (1978b). 

Because this monograph deals with taxa from only 
one area of species abundance, it is inappropriate to try 
for a worldwide revision of higher categories. More im- 
portant, data in the literature is not adequate to make 
higher level taxonomic decisions, and only part of the 
material needed to work out these problems is avail- 
able, dissected, and illustrated. Illustrations and de- 
scriptions needed to justify most synonymization of 
family names and to emphasize the broad distributions 
involved are published, but work on the Western Aus- 
tralian taxa (Solem, in preparation C), including the 
Dipnelicidae, is omitted at this time. 

Where possible, actual type species have been dis- 
sected and illustrated, but in taxa such as Paralaoma 
(figs. 26-27) this was not possible. Anatomical illus- 
trations and descriptions ofCharopa coma (Gray, 1843) 
(figs. 9-lO),Phenacohelixpilula (Reeve, 1852) (fig. 11), 
Flammulina zebra (Le Guillou, 1842) (fig. 29), Am- 
phidoxa marmorella (Pfeiffer, 1845) (fig. 30), 
Stephanoda binneyana (Pfeiffer, 1847) (fig. 31), and 
Pseudocharopa lidgbirdi (Etheridge, 1889) (fig. 32), 
based on new dissections, are presented here. Litera- 
ture data on the anatomy of Rotadiscus (Baker, 1927, 
pp. 228-230, pi. 16, figs. 13-20) and the Otoconchinae 
(Climo, 1969a, fig. 2C; 197 la) allow handling of all the 
included subfamily names except the Hedleyoconch- 
inae. This Australian group, fairly common in New 
South Wales and Queensland, with a possible relative 
on Lord Howe Island (Iredale, 1944, pp. 317-318), has 
not been dissected, but probably is another example of 
shell sculpture reduction and spire elevation in an ar- 
boreal taxon. External body features, jaw, and radula 
were figured by Hedley (1888, fig.; 1889, pi. 15) and 
copied by Pilsbry (1893-1895, pi. 9, figs. 27-29). These 
features are typically charopid in aspect. 



Australian and New Zealand workers have vari- 
ously used the names Flammulinidae, Charopidae, 
Phenacohelicidae, and Otoconchidae for subgroups of 
this complex (Gabriel, 1930; Iredale, 1937a et seq.; 
Climo, 1969a-b, 1970, 1971a-b, 1978; Powell, 1976). 
In large part this was based on early work of Henry 
Suter, which was summarized in his magnificent 
monograph (Suter, 1913). The ancient question as to 
the significance of the caudal pore or foss was discussed 
above and well summarized by Climo (1969a). It has no 
major taxonomic utility. Climo (1969a) focused on the 
presence or absence of an epiphallus as a significant 
subfamily key character. The externally easily recog- 
nizable epiphallus ofCharopa (fig. 9b, d) does contrast 
with the situation in Phenacohelix (fig. lie, e) where 
the insertion of the penial retractor has shifted to the 
epiphallus-vas deferens junction, rather than inserting 
on the penis-epiphallus junction as in Charopa. In 
Flammulina (fig. 29b, e-j) the situation is complicated 
by visceral hump reduction and drastic increase in 
whorl profile cross-sectional area. In Flammulina 
zebra, the genotype, the epiphallus has been compacted 
back into the penis (fig. 29e-h) and is thus completely 
undetectable from external view (fig. 29b). In other 
taxa referred to Flammulina (Climo, 1969a, fig. 2A) 
and the Otoconchinae (Climo, 1969a, fig. 2B-C; 1971a, 
fig. 1E-F) greater compaction has taken place in the 
region of the prostate and uterus, whereas the penis 
and epiphallus region are maintained as separate en- 
tities. 

Anatomical descriptions of the family units' type 
genera follow to provide comparative data with the 
later dissections and to record structural variations for 
consideration by future revisors of extralimital taxa. 
The descriptions follow the same sequence as the illus- 
trations Charopa coma (Gray) (figs. 9-10), Phe- 
nacohelix pilula (Reeve) (fig. 11), Flammulina zebra 
(Le Guillou) (fig. 29),Amphidoxa marmorella (Pfeiffer) 
(fig. 30), Stephanoda binneyana (Pfeiffer) (fig. 31), and 
illustrations only of Pseudocharopa lidgbirdi 
(Etheridge) (fig. 32). Discussion of anatomical varia- 
tion in that species is deferred to a review of the pat- 
terns in Lord Howe Island taxa (Solem, in prepara- 
tion). Nomenclatural details and a shell description of 
Amphidoxa marmorella are included because so little 
has been recorded about this species. 

Charopa coma (Gray, 1843). 

Description of soft parts. Foot and tail slender, rounded pos- 
teriorly, but not tapering. Sole undivided and without any corruga- 
tions (fig. 9a). Pedal grooves not particularly high on foot, suprapedal 
less distinct than pedal, both grooves uniting over tail. No caudal 
horn or foss. Middorsal groove weak. Slime network rectangular, 
fairly inconspicuous. Head extended well in front of foot, which is 
squarely truncated anteriorly. Ommatophores long. 

Body color in preservative light yellow-white, ommatophores 
black. 

Mantle collar (MC) rather narrow (fig. 9e f) with a large glandu- 
lar extension (MG) encroaching on lung surface. Pneumostome (LP) 
masked by quite small left anterior mantle lobe (MA) at parietal- 
palatal angle. Anus (A) opening slightly anterior to external ureteric 



62 



SOLEM: ENDODONTOID LAND SNAILS 



pore (KX) and sharing single opening with pneumostome (LP). Uri- 
nary chamber (LK) a narrow groove angling past anus. 

Pallial region (fig. 9e) varying in length from %-l whorl, elon- 
gated, anterior portion filled with mantle gland (MG). Lung roof 
clear, except for occasional white speckles near kidney tip. Kidney 
(K) U-shaped, rectal arm equal in length to pericardia!, but dis- 
tinctly more slender, lapping under hindgut. Hindgut (HG) parallel- 
ing parietal-palatal margin past top of pallial cavity, slender. Kidney 
abutting on loop of intestine (I). Ureter (KD) sigmurethrous, not 
expanded at any portion, both arms compreseed between lobes of 
kidney, then bordering hindgut to external ureteric opening (KX). 
Heart (H) proportionately large, much shorter than kidney. Prin- 
cipal pulmonary vein following edge of kidney, then running forward 
to edge of mantle gland. No conspicuous secondary venation. 

Ovotestis (G) embedded in apical lobe of liver (fig. 9b), variable 
in form and spacing, consisting of a few finger-like lobes, often split 
one or more times. Individual lobes with slender pink rods. Mass of 
ovotestis may be concentrated at one point or divided into two masses 
separated by part of digestive gland. Hermaphroditic duct (GD) 
straight, thick-walled, iridescent pink, sometimes with one or more 
knobs on surface, narrowing slightly to radically just before reaching 
albumen gland (GG) and entering midway on talon (fig. 9b, GT). 
Latter a blind pouch buried in albumen gland, head enlarged and 
pink, duct slender, passing straight into head of prostate-uterus. 
Prostate (DG) of large rectangular glandular lobes pressed against 
surface of uterine sector. Uterus (UT) an extremely thin-walled cir- 
cular tube partly filled with spongy tissue opening through a con- 
stricted collar into a thick-walled, white, expanded glandular pore 
with smooth inner walls except for a single large pilaster. Muscle 
fibers pass from uterine base to columellar retractor. 

Vas deferens (VD) arising from narrowing of prostate along 
glandular section of uterus, first part expanded, then continuing as 
narrow tube along free oviduct and vagina to penioviducal angle, 
then up penis to epiphallus (E). Epiphallus short, thick-walled, ex- 
panded, internally (fig. 10) with two longitudinal pilasters, refiexed 
and entering penis (P) through a small verge (fig. 9d, PV). Penial 
retractor muscle (PR) arising on diaphragm, attached to apex of 
penis in a U-shaped fan. Penis expanded, internally with thick pus- 
tulose glandular lining extending from apex to near base which ter- 
minates in a thickened collar. Verge small, with terminal pore, not 
developed in juveniles. Below level of verge, a blind pocket with 
smoothly muscled walls develops. This "penial caecum" (PC) is 
strictly internal and not visible on outside of penis. Glandular collar 
considerably thicker than rest of penis interior and gives false ap- 
pearance of being a verge when atrium (Y) opened. 

Length of free oviduct (UV) highly variable, apparently corre- 
lated with length of pallial region. In figured individual it is ex- 
tremely short; in others almost equal to vagina in length. Base of free 
oviduct constricted by a glandular papilla with central pore (UVO). 
Spermatheca (S) with very slender duct, bound to surface of prostate, 
passing into visceral mass above pallial cavity and expanding into a 
distinct head. Albumen gland also in mass of digestive gland and 
intestinal loop above pallial cavity. Vagina (V) long, swollen me- 
dially, with a prominent tubercle, or set of tubercles (VS), located 
between oviducal pore and the several longitudinal pilasters that 
extend to atrium (Y). Latter short, thin-walled, opening into gono- 
pore, a transverse slit located just behind and above right 
rhinophore. 

Free muscle system simple. Tentacular retractors (TP) uniting 
with tail fan (FR) well in front of union with buccal retractor (BR). 
Point of union between ommatophoral (TER) and rhinophoral (TVR) 
retractors variable, usually about midway along distance to tail fan. 
Right ommatophoral retractor passing between penioviducal angle. 

Buccal mass (B) short and high (fig. 9g), with very slender 
generative sac. Buccal retractors (BR) insert as U-shaped fan about 
Vt of way from posterior end. Esophagus (BE) very long and slender, 
originating high on buccal mass only slightly behind midpoint and 
well in front of buccal retractor insertion. Interior of esophagus with 
longitudinal ribs. Stomach originating just above pallial cavity, ex- 
tending apically for % of a whorl and with reflexion of stomach at 
point well below first lobe of ovotestis. First arm of intestine follow- 



ing lower inner margin of whorl to base of kidney, looping below 
cephalic aorta and across base of kidney, then reflexing for less than 
1 k length of stomach, rising to upper parietal-palatal margin and 
passing forward as hindgut. 

Digestive gland extending from apex of soft parts to just above 
intestinal loop, lower parts finger-like because of the huge stomach. 
Main artery, granular white in color, empties primarily into stomach 
and secondarily into recurved lower loop of intestine. 

Salivary glands (OG) elongate, situated on sides and top of 
esophagus, closely associated with prostate-uterus in body cavity. 
Salivary ducts (OD) straight, inserting on each side at base of 
esophagus. 

(Based on FMNH 135420, Waiwera-Puhoi Road, North of Auck- 
land, North Island, New Zealand, A. Solem! February 10, 1962. Sev- 
eral adults and juveniles.) 

Phenacohelix pilula (Reeve, 1852). 

Description of soft parts. Foot long and slender, tapering 
gradually to a sharply rounded posterior margin. Sole undivided, 
without corrugations. Pedal grooves very high on foot, suprapedal 
distinctly less impressed than pedal, both grooves uniting over tail. 
Caudal horn a very long, tapering projection (fig. lib, CH). Middor- 
sal groove absent. Slime network very weak, barely visible. Head 
extending only slightly in front of foot, which is bluntly rounded 
anteriorly. 

Body color in preservative yellow-white, with faint grayish tinge 
to upper part of tail and caudal horn, ommatophores reddish black. 
Dark color markings on pallial roof visible through shell. 

Mantle collar (MC) narrow (fig. lla), with moderate glandular 
intrusion (MG) onto pallial roof. Pneumostome without clearly de- 
fined lappets. Anus (A) opening above and slightly anterior to exter- 
nal ureteric pore (KX), urinary chamber (LK) a shallow groove lead- 
ing past anus and fading out toward mantle collar margin. 

Pallial region (fig. lla) rather short and compact. Lung roof with 
numerous scattered black speckles and much less frequent white 
patches. Kidney (K) bilobed, pericardia! arm much longer than rec- 
tal, 3.5 times length of pericardium (H). Base of kidney abutting on 
loop of intestine. Rectal arm of kidney lapping under hindgut, less 
than half length of pericardial arm. Ureter (KD) sigmurethrous, 
pericardia! arm slightly expanded over that along hindgut (HG). A 
clear section of lung roof visible between arms of ureter. Neither 
ureter nor hindgut expanded during anterior passage toward 
pneumostome. Hindgut (HG) paralleling parietal-palatal angle well 
past apex of pallial cavity. Heart (H) short, angled to plane of 
hindgut. Principal pulmonary vein (HV) simple, extending forward 
to edge of mantle gland intrusion, without conspicuous secondary 
venation. 

Ovotestis (fig. lie, G) buried in apical lobes of digestive gland 
above stomach-intestine reflexion, consisting of one or two clumps of 
pinkish iridescent finger-like alveoli joined by slender ducts in a "Y" 
leading to main hermaphroditic duct (GD). Latter a slender duct 
tapering gradually toward an enormously swollen portion, about 1 A 
of duct length, shortly before approach to albumen gland, then be- 
coming an extremely slender tube before joining talon (fig. lid, GT), 
which is buried in albumen gland (GG). Latter a blunt, finger-like 
projection lying buried in visceral hump above pallial cavity and 
composed of quite large acini with a yellow-orange tinge. Basal por- 
tion of hermaphroditic duct reaching base of albumen gland, refiexed 
apically along surface before entering laterally upon enlarged head 
of talon, which is partly buried within albumen gland. Narrowed 
shaft of talon running to head of prostate-uterus, no expanded car- 
refour visible. Prostate (GD) of much larger, lighter colored acini 
than those of albumen gland, fastening to thin-walled, upper uterine 
chamber. This is a flat, smooth tube gradually tapering from a nar- 
row apex to a broad base where the second uterine chamber (UT 2 ), a 
glandular walled structure, expands. Lower part of uterus a thin- 
walled chamber with internal lamellar plates, opening into a bul- 
bously expanded free oviduct. 

Vas deferens (VD) arising as groove from base of prostate, partly 
hidden by third uterine chamber, passing down as a thin-walled tube 



SYSTEMATIC REVIEW 



63 



to penioviducal angle (to which it is lightly bound by fibers), reflex- 
ing along penis and becoming progressively larger in diameter and 
more muscular before joining epiphallus laterally. Penial retractor 
(PR) inserting directly on head of epiphallus (E), a short, solid, 
globose tissue mass arising on diaphragm. Epiphallus an expanded 
muscular tube entering penis (P) through a smooth muscular verge 
(fig. lie, PV). Walls of penis with weak crenulated pilasters and 
pustulations basally, apical portion with glandular collar just below 
tip of verge. Externally, position of glandular collar and verge indi- 
cated by swelling in penis-epiphallus. Atrium (Y) shortened and 
leading to gonopore located below and a little behind right om- 
matophore. 

Free oviduct (UV) a short, globosely expanded structure narrow- 
ing slightly to point of union with spermathecal stalk. Spermatheca 
(S) with very slender stalk, partly buried between acini of prostate, 
passing apically to small expanded head lying above apex of pallial 
cavity in visceral mass together with albumen gland and talon. Va- 
gina (V) much longer than free oviduct with a very thin-walled ac- 
cessory pocket that was badly torn in all dissected specimens. The 
exact relationship of the vagina, oviduct, and accessory vaginal 
pocket was not determined. 

Buccal mass rather short, strongly curved on top, generative sac 
very small. Buccal retractors insert in U-shaped fan about '/-.< of way 
from posterior of buccal mass. Esophagus inserting nearly at mid- 
point of upper surface of buccal mass, very slender and thin-walled, 
extending through visceral hump past apex of pallial cavity. At this 
point, the esophagus expands into a funnel shape, entering stomach 
yet allowing room for positioning of intestinal loops. Esophagus en- 
tering stomach on lower inner whorl margin just above apex of pal- 
lial cavity. Stomach occupies nearly entire whorl, grossly expanded 
to cover outer and upper margins, extending apically to point below 
"Y" of hermaphroditic duct, intestine reflexing along base of whorl, 
about midway between inner and outer margin. Intestine running 
apically to apex of pallial cavity, looping across outer whorl margin 
and abutting on kidney, reflexing backward and downward along 
expanding margin of stomach, then looping upward on outer wall to 
parietal-palatal margin and passing forward as hindgut 

Digestive gland extending from tip of soft parts to apex of pallial 
cavity, greatly reduced in region occupied by stomach. 

Salivary glands slender, white, lying along and surrounding 
esophagus. Salivary ducts entering buccal mass-esophagus junction 
laterally. 

Free muscle system simple. Columellar retractor very large, 
buccal retractor uniting with tentaculars shortly before joining main 
tail fan. 

(Based on FMNH 135421, Church Road, Kaitaia, Northland, 
New Zealand, L. Price! October 1962, several adults.) 



Flammulina zebra (Le Guillou, 1842). 

Description of soft parts. Foot and tail about equal in length to 
shell diameter, truncated anteriorly, rapidly tapering from about 1 
mm. behind visceral stalk posterior. Sole undivided longitudinally, 
with weak transverse corrugations extending from sides of foot. 
Pedal grooves high on foot, suprapedal slightly weaker than pedal, 
united above tail, no caudal horn, or middorsal groove present. Slime 
network irregularly rectangular, relatively weak. Head and om- 
matophores without unusual features. Gonopore on right side of 
head, directly behind ommatophore. 

Body color dark gray on head and tail, lighter between pedal 
grooves, soles and sides of foot yellow- white. 

Mantle collar (fig. 29a, MC) without lobes or laps, no glandular 
extension onto pallial roof. Anus (A) opening slightly anterior to 
external ureteric pore (KX). Urinary chamber (LK) not dif- 
ferentiated. 

Pallial region (fig. 29a) extending about V:> whorl apically, quite 
broad, lung roof without color markings. Kidney (K) bilobed, short, 
quite wide. Ureter (KD) thick, arms sharply angled, with angle ap- 
proaching 80. Heart (H) half length of kidney. Principal pulmonary 
vein (HV) weakly branched, stopping well short of mantle collar. 



Hindgut (HG) slender, reaching parietal-palatal margin opposite 
point where ureter reflexes, then running forward to anus. 

Ovotestis two large clumps of palmately clavate alveoli buried in 
digestive gland above stomach apex. Clumps with central orienta- 
tion perpendicular to plane of coiling, laterally fanning from center. 
Hermaphroditic duct narrow, variously kinked and coiled, extending 
to side of prostate, reflexed upward and straightened to angled union 
with carrefour (fig. 29c-d). Albumen gland large, of slender acini, 
tightly packed around carrefour region. Talon (GT) very short, with 
circular head, buried in albumen gland. Carrefour (X) with only 
slight enlargement from talon shaft (fig. 29c-d). Prostate (DG) of 
comparatively few acini along sides of uterus, exact structure of pas- 
sage from carrefour to vas deferens not determined. Possibly a nar- 
row part of the uterine lumen serves to transfer sperm. Uterus (UT) 
grossly enlarged, twisted, with thick glandular walls. Vas deferens 
(VD) thicker for first V:i of length, then narrowed for last %, entering 
apex of epiphallus next to point where penial retractor muscle in- 
serts, tightly bound to penis surface (fig. 29b). Epiphallus compacted 
into upper penial chamber (fig. 29e-i), with complex valvular en- 
trance from vas deferens (fig. 29i-j). Penial retractor (PR) very short, 
scarcely a separate muscle zone. Penis (P) almost circular, about 1.3 
mm. long, apex (fig. 29e) with compacted epiphallus opening through 
a corrugated verge (fig. 29e-f, PV), lower portion with a complex set 
of pilasters, highly modified from standard charopid pattern. Atrium 
(Y) long and slender, thin-walled, without internal sculpturing (fig. 
29b). 

Free oviduct (UV) short, twisted, fusing indistinctly with lower 
organs. Spermatheca (S) expanded basally, shaft following side of 
uterus, curving up and around carrefour region, head partly next to 
exposed section of prostate, partly next to albumen gland, oval in 
shape. Vaginal and lower penis area fused into a common chamber. 

Free muscle system short, characterized by lack of fusion be- 
tween muscles. Right ommatophoral retractor (TER) not passing 
through penioviducal angle, arising laterally from columellar mus- 
cle. Right rhinophoral retractor (TVR) arising from columellar mus- 
cle just inside origin of ommatophoral retractor. 

Buccal mass shortly oval, proportionately very large, with small, 
generative sac. Buccal retractors split, each half inserting laterally 
near posterior margin of buccal mass, inserting at apex of columellar 
muscle. Esophagus short, not enlarged, arising from middle of buccal 
mass upper surface. Stomach starting expansion before apex of pal- 
lial cavity, occupying '/z whorl, then reflexing to intestine, which 
loops down and forward along lower whorl margin, looping inward 
and upward along side of pericardium and pericardial kidney, then 
making a curved U-loop backward and inward before angling for- 
ward as hindgut to parietal-palatal margin. Hindgut slanting down 
from parietal-palatal margin at point of ureter reflexion. 

Digestive glands typical. Salivary glands lying on top and on 
either side of esophagus, partly fused medially. 

(Based on FMNH 153217, Okuti Valley, Banks Peninsula, near 
Christchurch, New Zealand, F. Climo! March 7, 1965. Three dis- 
sected adults.) 

Amphidoxa marmorella (Pfeiffer, 1845). 

Helix marmorella Pfeiffer, 1845, Proc. Zool. Soc. London, 1845, p. 
125- Island of Juan Fernandez; Pfeiffer, 1847, Monog. helic. viv., 
1, p. 66; Albers, 1850, Die Heliceen, Isted., p. 110; Pfeiffer, 1852, 
Syst. Conchol. Cab., I, 19(2), pp. 129-130, pi. 88, figs. 17-19 
(plate issued in 1850); Reeve, 1852, Conchol. Icon., Helix, pi. 113, 
fig. 649. 

Helix (Amphidoxa) marmorella Pfeiffer, Tryon, 1887, Man. 
Conchol., (2), 3, p. 46, pi. 20, fig. 54. 

Amphidoxa (Amphidoxa) marmorella (Pfeiffer), Pilsbry, 1893, 
Man. Conchol., (2), 9, pp. 39-40, pi. 7, figs. 10-12. 

Amphidoxa marmorella (Pfeiffer), Odhner, 1922, Nat. Hist., Juan 
Fernandez and Easter Island, 3, pp. 229, 230-231, figs. 9-10 
(radula and jaw). 

Description of shell. Shell small, with slightly more than 3 
rapidly enlarging whorls (fig. 30a). Apex slightly emergent, penul- 
timate and body whorl flatly coiled, H/D ratio 0.550. Apical whorls 




FIG. 29. Anatomy of Flammulina zebra (Le Guillou). Okuti Valley, Banks Peninsula, near Christchurch, New Zealand. F. Climo! HI-7-1965. 
FMNH 153217: a, pallial region; b, postapical genitalia; c-d, views of the hermaphroditic duct-talon-carrefour junction; e, interior of penis 
chamber; f-h, detail of vas deferens-epiphallus-penis junction; i-j, detail of vas deferens-epiphallus junction. Scale lines equal 1 mm. foro-6, e; 
other figures greatly enlarged. (CW). 



64 







HG 



KO 






PR 




1 mm 



FIG. 30. Anatomy of Amphidoxa marmorella (PfeifTer). 500 m. elevation, northeast slope, Portazuelo, Masatierra, Juan Fernandez. B. 
Malkin! IV-7-1962. FMNH 135424: a, apical view of shell prior to dissection; b, basal view of shell; c, pallial region; d, genitalia; e, interior of 
penis. Scale lines as marked. (PS). 



65 



66 



SOLEM: ENDODONTOID LAND SNAILS 



1%, sculpture of 20 protractively angled large radial ribs over a mi- 
croreticulated lattice visible only under 96 x magnification. Remain- 
ing whorls with protractively sinuated, lamellar radial ribs, 32 on 
the body whorl, whose interstices are more than 8 times their width. 
Ribs elevated greatly by periostracal extensions. Microsculpture a 
lattice of microradial and microspiral riblets, the former slightly 
stronger. Sutures deep, whorls rather flatly rounded above evenly 
rounded periphery and base. Color light yellow-white with very 
vague reddish flammulations, waved, becoming strongly retractive 
on base of shell, apparently not extending into umbilicus. Umbilicus 
narrow, V-shaped, regularly decoiling, contained 8.3 times in the 
diameter. Aperture large, subcircular, slightly flattened laterally 
above periphery, inclined about 30 from shell axis. No apertural 
barriers. Height of shell 1.64 mm., diameter 2.99 mm. 

Description of soft parts. Body completely retracted within 
shell (fig. 30b), soft parts relatively hardened and difficult to dissect. 
Foot relatively short and broad, rounded behind, apparently not ta- 
pering, truncated anteriorly. Sole undivided longitudinally, with 
rather strong transverse corrugations near tail, corrugations coming 
from slime network. Pedal grooves deep, high on foot, suprapedal as 
marked as pedal, both grooves uniting over tail. Because of contrac- 
tion, presence or absence of middorsal groove could not be deter- 
mined. Slime network a sharply defined irregularly rectangular 
network. Ommatophores short and chunky. 

Body color yellow-white, mantle collar brownish. Eyespots small 
and black, upper part of stalk with black speckles. 

Mantle collar (fig. 30c, MC) thick, glandular, without any man- 
tle gland extension onto lung roof. Pneumostome masked by bulges 
in mantle collar, but no distinctive lobes present. Anus (A) opening 
at inner edge of mantle collar, a slight groove running from it to 
outer edge of collar. 

Pallial region (fig. 30c) extending about '/n whorl, shape and 
positioning of structures probably altered by extreme contraction of 
animal. Lung roof clear, without dark speckling except in region of 
ureter and pericardium and a very few along hindgut near anus. 
Kidney (K) bilobed, main lobe pericardial, with a narrow tongue 
extending along hindgut (HG). Main lobe twisted in front of heart 
(H) extending on both sides of ureter (KD) tip. Base of kidney extend- 
ing outside of intestinal loops along outer whorl edge (partly an 
artifact of contraction?). Entire first loop of intestine and part of loop 
to hindgut lying next to kidney. Ureter (KD) following anterior mar- 
gin of pericardial kidney arm upward and slightly backward into 
small pocket between pericardial arm of kidney and hindgut then 
ending in a pore pointing apically. No groove leads toward pneumo- 
stome nor is there any trace of a secondary ureter. Heart (H) rather 
large, constricted by retraction. Principal pulmonary vein (HV) fol- 
lowing edge of kidney and tip of ureter, then passing forward to 
mantle collar with no conspicuous lateral venation. 

Ovotestis (fig. 30d, G) two clusters of finger-like lobes, some 
bifurcated, imbedded in digestive gland above stomach, a Y-fork col- 
lecting tubule attaching them to hermaphroditic duct (GD). Latter 
extending along stomach at basal margin of whorl, a thick, nodular, 
opaque, iridescent tube becoming slender near albumen gland and 
running to stalk of talon. Albumen gland (GG) finely textured, oc- 
cupying visceral area between stomach base and apex of pallial cav- 
ity inside of intestinal loops. Head of spermatheca (S) lying in angle 
between prostate head and albumen gland base. Talon (GT) long 
with slender, slightly tapering head and broader base that is buried 
in albumen gland. Hermaphroditic duct entering talon below apex of 
slender stalk. Talon opening into apices of prostate and uterus. Pros- 
tate (DG) of large acini opening into a closed duct that is partly 
enfolded by uterus (UT1. Lower part of prostate squeezed by ex- 
panded uterus and opening into slender vas deferens (VD). Uterus 
(UT) a slender, thin-walled tube above, opening into a grossly ex- 
panded chamber with thicker glandular walls and weak traces of 
ridging. 

Vas deferens (VD) a very slender tube passing to penioviducal 
angle, around right ommatophoral retractor, then up to head of penis 
(P). Penial retractor (PR) arising on diaphragm, inserting on head of 
epiphallus (E). Vas deferens inserting laterally on epiphallus, which 
contains longitudinal pilasters surrounding the vas pore and leading 
into the verge (fig. 30e, PV). Penis proper bulbous above, tapering to 



a slender duct before joining vagina (V) to form atrium (Y). Inter- 
nally (fig. 30e) penis with a large apical verge (PV) whose tip is 
bifurcate; a large circular muscular collar (PP,) separated into two 
parts; a broad, thin pilaster (PP 2 ) tapering to the slender basal stalk; 
and an apparently doughnut-shaped pilaster (PP :1 ) that is partly 
folded over upon itself in the available specimen. Atrium (Y) short, 
rather large in diameter. 

Free oviduct (fig. 30d, UV) short, only slightly narrowed from 
uterus, internally with glandular walls and vague pilasters. Sper- 
matheca (S) with basal part of shaft slightly expanded, tapering 
along surface of uterus and prostate, expanded head lying in angle 
between prostate and albumen gland. Vagina (V) very large, ex- 
panded, internally with broad longitudinal glandular pilasters that 
continue into atrium (Y). No special pilasters or collars mark sper- 
mathecal or oviducal terminations. A small vaginal diverticulum 
(VC) is a blind pocket opening between two major pilasters that 
continue up into spermatheca. 

Buccal mass ovoid, strongly contracted, generative sac rather 
large. Buccal retractor inserting in U-shpaed fan about Vy of way 
from posterior end, not split. Esophagus entering top of buccal mass 
slightly behind midpoint, a rather thick tube with longitudinal pilas- 
ters following inner margin of whorl past apex of pallial cavity and 
entering stomach very shortly past that point. Stomach extending 
about '/4 whorl, expanded to fill parietal and upper palatal walls 
almost immediately, narrowing about Vh of way from apex, the nar- 
rowing resulting in both a descension from the parietal-palatal angle 
and basal narrowing. At apex, stomach reflexes downward and turns 
near basal margin before joining intestine. At stomach-intestine 
junction, ducts of digestive gland enter on each side. Intestine follow- 
ing basal-palatal margin of whorl forward to base of kidney (fig. 30c, 
K), looping across kidney base, nearly touching hindgut, looping di- 
rectly downward, then backward and upward as hindgut to par- 
ietal-palatal margin, finally following that margin to anus. Extreme 
sharpness of intestinal looping may be result of contraction. 

Digestive gland extending from apex of soft parts to apex of 
pallial cavity; a broad band along lower palatal wall between intes- 
tine and outer part of stomach, reduced in amount in area of intesti- 
nal loops and albumen gland. Openings into stomach-intestine junc- 
tion very prominent. 

Salivary glands white, finely textured, lying lateral to esopha- 
gus, touching above. Ducts of salivary glands entering buccal mass 
on each side of esophageal insertion. 

Free muscle system short and massive. Buccal retractor nearly 
as broad as long when contracted. Right ommatophoral retractor 
passing between penioviducal angle, uniting with rhinophoral re- 
tractor just before joining tail fan. Tentacular retractors joining tail 
fan subapically and laterally, buccal retractor joining medially and 
apically. 

Jaw fragmented during mounting, composed of about 20 delicate 
plates that are weakly striated and do not strongly overlap. 

Radula with 75 rows. Central tooth very small, with weak side 
cusps. Lateral teeth large, tricuspid, endoconal size gradually in- 
creasing during shift to marginals, ectocones on outer marginals 
split. 

(Based on FMNH 135424, northeast slope at 500 m. elevation, 
above Portazuelo, Masatierra, Juan Fernandez, B. Malkin! April 7, 
1962. One adult specimen.) 

Stephanoda binneyana (Pfeiffer, 1847). 

Description of soft parts. Foot narrow and rather elongated, not 
tapering posteriorly, bluntly rounded behind (fig. 31f). Pedal grooves 
deeply and equally impressed, high on sides of foot. Sole undivided 
longitudinally, but strongly corrugated perpendicular to body axis, 
such corrugations being continuations of lateral grooves below pedal 
grooves. Tail with middorsal groove, bell-shaped in cross section, 
sloping down to point where pedal grooves unite. No caudal foss or 
horn developed. Slime network fine, irregular, oblong to rectangular 
patches. 

Body color in preservative pale yellow-white, sides of head and 
mantle with gray markings, ommatophores black. 

Mantle collar (MC) thick with tapering edge (fig. 31a). Pneumo- 



SYSTEMATIC REVIEW 



67 




FIG. 31. Anatomy of Stephanoda binneyana (Pfeiffer). Rio Cisnes (44 45' S, 72 W), Prov. Aysen, Chile. L. E. Pena! 11-1961. FMNH 135428: 
a, pallial region; b, genitalia; c, free muscle system; d, interior of terminal genitalia; e, detail of epiphallus-penis junction; f, rear portion of tail; 
g, detail of anterior mantle cavity; h, detail of hermaphroditic duct-talon-carrefour junction; i, lobe of ovotestis. Scale lines as marked, detail 
sketches greatly enlarged. (PS). 



stome (LP) masked by small left anterior mantle lobe (MA). Edge of 
collar just anterior of pneumostome has a weak thickening forming 
an accessory mantle lobe (AMA). Anus, excretory pore, and 
pneumostome sharing common opening (fig. 31g). Anus (A) slightly 
anterior to external ureteric pore (KX). Urinary chamber (LK) a 
shallow groove running past anus and continuing (although less 
sharply defined) nearly to edge of mantle collar. 

Pallial region (fig. 31a) elongated, but in view of contracted state 
no accurate estimate of length possible. Lung roof with variable col- 
oring, speckled with widely scattered black and white flakes to 
nearly coal black. Kidney (K) short, elongately triangular with a 



narrow, hooked basal extension abutting hindgut (HG). Base of kid- 
ney against loop of intestine. Ureter (KD) slender, sigmurethrous, 
flaring slightly midway along hindgut, arms of ureter widely sepa- 
rated by lung roof. Heart (H) slightly more than '/2 length of kidney. 
Principal pulmonary vein (HV) angling toward pneumostome, 
nearly reaching mantle collar, without conspicuous side branches. 

Ovotestis (G) imbedded in digestive gland above stomach- 
intestine reflexion, consisting of 8 sets (4 shown in fig. 31b) of 
clumped palmately clavate alveoli (fig. 31i) strung linearly along 
very thin hermaphroditic duct (GD). Ovotestis distinctly lighter in 
tone than digestive gland. Collecting tubule of hermaphroditic duct 



68 



SOLEM: ENDODONTOID LAND SNAILS 



very thin, bilobed. Main section a round muscular tubule, slightly 
iridescent in tone, narrowing just before base of albumen gland (GG), 
reflexing abruptly and ascending to apex of bilobed talon (GT) (see 
fig. 31h). Albumen gland elongately ovate, blunt-tipped, lying com- 
pletely above pallia! cavity, very fine-textured. Talon (GT) buried in 
albumen gland, quite small, white. Duct from talon opening into 
head of prostate-uterus without differentiated carrefour. Prostate (D) 
equal in width to upper portion of uterus, individual acini much 
larger than those of albumen gland. Lower part of prostate enveloped 
in basal sections of uterus. Uterus (UT) tripartite. Upper section 
narrow, sacculated, equal in width to prostate. Basal '/n of uterus 
surrounded by a hollow, smooth-walled, white-colored chamber with 
very fine texture. Basal section of uterus expanded into a chamber 
with lamellar walls. 

Vas deferens (VD) originating from groove of prostate enfolded 
by second uterine chamber, a narrow tube passing down to peni- 
oviducal angle, then almost immediately into epiphallus (E). Latter 
a much larger muscular tube, first half swollen with strong pilasters 
on one wall, then narrowing to about 4 times diameter of vas def- 
erens. Epiphallus entering penis (P) just below attachment of penial 
retractor (PR), which is short, quite thick, and originates on dia- 
phragm. Penis rather short, apical half bulbous, tapering toward 
atrium. Internally (fig. 31d-e) with rather heavy and complex pilas- 
ters. A single large lamellar plate (PP) starts just below epiphallic 
pore, curves up and around it, then passes longitudinally down wall 
opposite insertion of penial retractor. A second longitudinal pilaster 
(PP,) arises below origin of major pilaster and tapers gradually to- 
ward atrium. A series of circular minor pilasters complete the ar- 
mament. 

Free oviduct (UV) short and muscular, bulbous, internally (fig. 
3 Id) with rugose pilasters. Spermatheca (S) of long type, expanded 
head in visceral mass above pallial cavity and cephalic aorta, stalk 
very slender. Vagina (V) long, with twisted, bilobed accessory vagi- 
nal appendix (VC). Main lobe (fig. 31b, d) with muscular walls, a few 
longitudinal pilasters, and an intricately reticulated stimulatory pad 
(VS). Secondary lobe (VC) a bulbous, tapering, thin-walled organ, 
the expanded head with many lamellar plates. Stalk of secondary 
lobe and duct of vagina with weak longitudinal pilasters. Main ac- 
cessory lobe of vaginal appendix bound to right side of tail fan muscle 
(fig. 31c, VRM) slightly less than halfway from union with right 
tentacular retractors and insertion on foot. 

Free muscle system (fig. 31c) simple. Buccal retractor (BR) at- 
taching on posterior Va of buccal mass, in U-shaped fashion, but not 
split until very tip, and joining tail fan (FR) at site of columellar 
attachment (CR). Ommatophoral retractor (TER) and rhinophoral 
retractors (TVR) unite a short distance back and almost immediately 
fuse with tail fan. Right ommatophoral retractor passing between 
penioviducal angle. Tail fan split slightly posterior to junction with 
tentacular retractors (TR). Penis base bound to body wall, accessory 
lobe of vagina bound to right part of tail fan (VRM). 

Buccal mass rather short and muscular, generative sac promi- 
nent. Esophagus entering top of buccal mass between 1 A and '/a way 
from posterior end, large and muscular, with heavy longitudinal 
ridges anteriorly, tapering and becoming thin-walled to apical end of 
pallial cavity. Stomach a very large thin-walled sac extending less 
than 1 whorl apically, along parietal-palatal margin and occupying 
much of space in visceral hump. Reflexing as intestine along lower 
inner margin of whorl, running back to apex of pallial cavity, curv- 
ing up and abutting on kidney base, looping back diagonally down- 
ward for '/2 length of stomach, then reflexing forward diagonally as 
hindgut. Hindgut not reaching parietal-palatal margin until just 
above start of pallial cavity, following margin to anus and only 
slightly expanded in lower part of pallial cavity. Salivary glands 
paired, lying on each side of esophagus, touching above and below. 
Ducts very slender, inserting on each side of posterior esophageal 
base. Extreme duct coiling and nearness of glands to buccal mass 
probably result of extreme contraction. 

Digestive gland extending from top of pallial cavity to apex of 
soft parts (which do not reach shell apex). Lower section with intesti- 
nal loops finger-like. Minor collecting tubules pass through spaces 



between branches of ovotestis clumps. Major duct empties into base 
of stomach. 

(Based on FMNH 135428. Rio Cisnes, Aysen, Chile, L. E. Pena! 
February 1961. Four dissected adults.) 

All of these taxa agree in basic structures of the 
genitalia, pallial region, radula so far as observed, and 
muscle system. Their differences relate to visceral 
hump and compaction changes in the case of Flam- 
mulina and the Otoconchinae, multiple radiations be- 
tween terrestrial and arboreal habitats (shell sculpture 
reduction, mucus pore intensification), and species rec- 
ognition factors in the terminal genitalia. The latter 
changes are many and bewildering, since 15 to 30 en- 
dodontoids from a single locality is normal in New Zea- 
land. The variations in penis interior structure illus- 
trated by Climo (1969b, figs. 25A, 28D, 29B, C, 30E; 
1970, figs. 16B, 20D, 2 1C) can be directly compared 
with the type of sympatric variations reported below. 

Continuation of the Otoconchinae as a separate 
subfamily, based on the multiple changes in structure 
resulting from visceral hump reduction, can be jus- 
tified and is accepted here. It is less strongly 
differentiated than the other subfamilies recognized in 
this report, and rigorous structural analysis may result 
in determining that the Otoconchinae is a grade in 
structure rather than a clade. 

The remaining New Zealand Charopidae are not 
amenable to subfamily distribution at this stage. 
Champa, Phenacohelix, and Flammulina agree in all 
essential details of structure. They are reasonably 
closely related. The many illustrations of Climo 
(1969a-b; 1970) do suggest possible subgroupings, but 
until details of vergic, epiphallic, and lower female 
tract structures are described and illustrated, no real 
decisions are possible. Although the New Zealand 
charopids have radiated extensively and confusingly, 
their basic structures seem rather coherent. Retention 
within an admittedly rather broad subfamily Charo- 
pinae is suggested pending detailed revisions. 

Because most of the family names used for the 
Australian charopids have been based on New Zealand 
genera, the great variation found in these taxa will not 
be discussed further. Study of Western Australian taxa 
with primitive features is in progress (Solem, in prep- 
aration C). The family name Dipnelicidae Iredale 
(1937b, pp. 22-23) will be considered in that report. 
The structural variety in the Australian taxa seems 
greater than that reported for the New Zealand 
species, but this is only a preliminary impression. 

The family Pseudocharopidae Iredale (1944, p. 
312) was proposed by the phrases "An extraordinary 
development of beautiful small shells is indicated by 
this family name. ... I was thinking of allotting them 
to the Flammulinidae, which they slightly resemble. 
... to save further confusion the above name (Pseudo- 
charopidae) is utilized." Dissection of Pseudocharopa 
lidgbirdi (Etheridge, 1889) (fig. 32a-c) shows a typical 
pallial cavity configuration (fig. 32a), with bilobed 
kidney, complete secondary ureter, and essentially un- 



SYSTEMATIC REVIEW 



69 



branched pulmonary vein. The genitalia (fig. 32c) is 
unusual only in having a single kink in the hermaph- 
roditic duct (GD), small prostate, shortened terminal 
female organs, the epiphallus shifted in relation to the 
penial retractor muscle, and the interior of the penis 
(fig. 32b) with high, smooth pilasters in the lower 
penis, rugose low pilasters in the upper penis. Because 
the shell of this species has a reduced visceral hump, 
most of the unusual features can be interpreted as sec- 
ondary. The basic features are those of the Charopinae. 




KD 




FIG. 32. Anatomy of Pseudocharopa lidgbirdi (Etheridge). Sta- 
tion 20, northeast slope of Mt. Lidgbird, 1,500 ft. elevation, south end 
Lord Howe Island. L. Price! IX-1963. FMNH 127977: a, pallial re- 
gion; b, interior of penis complex; c, genitalia. Scale lines as marked; 
b greatly enlarged. (NB). 



Amphidoxinae Thiele (1931, p. 578) is based on 
one of the Juan Fernandez Islands species, A mphidoxa 
marmorella (Pfeiffer, 1845). A single, retracted, nearly 
adult specimen (fig. 30a-b) was dissected. The 
genitalia (fig. 30d-e) has the typical charopid aspect, 
with the addition of a vaginal caecum (VC), shortened 
epiphallus (E), and internally shows the verge (PV), 
ring pilasters (PPi), and pocket stimulator (PC) seen in 
Champa (fig. 9d) and Sinployea (figs, 43e, 57c). The 
only unusual features are the absence of a secondary 
ureter (fig. 30c) and the retention of a separate pros- 
tate. Despite this, the basic structures of Amphidoxa 
and Charopa are amazingly similar. At least the 
structures of Amphidoxa show how the Charopinae 
could be derived from more generalized stocks. 

Traditionally, the South American genus 
Stephanoda has been associated with Amphidoxa. Dis- 
section of Stephanoda binneyana (Pfeiffer, 1847) 
showed profound differences. The pallial region (fig. 
31a) has the short rectal lobe found in Phenacohelix 
(fig. lla), a complete secondary ureter (KD) that opens 
(KX) (fig. 31g) just posterior of the anus (A). In the 
genitalia (fig. 31b, d-e, h-i) the ovotestis consists of 
several follicle bundles (fig. 31b, i), the talon (GT) is 
reduced (fig. 31h), the prostate-uterus is typical, but 
the slender base of the spermatheca (S) (fig. 31b) and 
fat free oviduct (UV) again agree with Phenacohelix 
(fig. lie). The tail of the animal (fig. 31f) has the typi- 
cally conservative form found in Charopa (fig. 9a) and 
all Pacific Island taxa. The most unusual features are 
in the terminal genitalia, with a prominent vaginal 
caecum (VC) (fig. 31b), long vagina (V), and short penis 
(P). Internally (fig. 31d-e) the penis has circular pilas- 
ters and two large stimulators, but the epiphallus (E) 
enters through a pore rather than a verge (fig. 31e). 
There are irregular and complex pilasters within the 
terminal female genitalia (fig. 31d). In the absence of 
the verge and altered pilaster pattern, Stephanoda 
shows considerable similarity to structures seen in 
some of the non- verge bearing charopids illustrated by 
Climo (1969b, 1970). These structures emphasize that 
the charopids are a true Southern Hemisphere group 
and that relationships are somewhat complex. 

Comments on a few other extralimital taxa are 
included under the appropriate subfamily discussions. 

In this monograph I recognize five subfamilies in 
the Charopidae: Otoconchinae Cockerell, 1893; Rota- 
discinae Baker, 1927; Charopinae Hutton, 1884; Truk- 
charopinae, new subfamily; and Semperdoninae, new 
subfamily. Of these, the Otoconchinae is least sharply 
differentiated, and the Charopinae is deliberately the 
most broadly defined, pending adequate anatomical 
revisions of the New Caledonian, New Zealand, Aus- 
tralian, South American, and South African taxa. Un- 
doubtedly the Charopinae will be subdivided further in 
the future, but whether on a tribal basis or into several 
subfamilies will depend upon the results of future in- 
vestigations. A brief diagnosis of each subfamily fol- 
lows. 



70 



SOLEM: ENDODONTOID LAND SNAILS 



Subfamily Otoconchinae Tail and body greatly elongated, vis- 
ceral hump reduced with shell a cap or fragment, head shortened in 
relation to body length. Jaw fused, arcuate. Radula typical of 
Charopinae. Pallial region modified Charopinae because of shorten- 
ing, genital system compacted by head and visceral hump reduction, 
basically Charopinae in structures. Otoconcha Hutton, 1884, and 
Maoriconcha Dell, 1952, are the included taxa. 

Subfamily Rotadiscinae Small to minute species with apical 
sculpture of short spirally arranged segments (fig. 4a-e), visible op- 
tically as spiral cords, postapical sculpture typically charopid. Jaw of 
separate plates. Radula usually typical except for ectoconal marginal 
splitting. Pallial region without to with a complete secondary ureter, 
kidney weakly to evenly bilobed. Ovotestis of one to two clumps of 
short to long follicles, talon short to long, not with circular head. 
Terminal female system simple, sometimes with accessory caecum, 
relatively thin. Epiphallus usually below penial retractor insertion, 
short, opening into penis through a short conical verge or simple 
pore, walls of penis with longitudinal pilasters. Rotadiscus Pilsbry, 

1926, Radiodiscus Pilsbry & Ferriss, 1906, Radioconus H. B. Baker, 

1927, Radiodomus H. B. Baker, 1930, Microcharopa, new genus, and 
probably some undescribed genera from Western Australia are the 
included taxa. 

Subfamily Charopinae Small to large species with apical shell 
sculpture of spiral cords, radial ribs, or secondarily smooth, postapi- 
cal sculpture typically charopid, frequently secondarily reduced. 
Apertural barriers developed in several lineages. Jaw of separate 
plates in smaller taxa, fused partly to completely in large. Radula 
generally with tricuspid laterals and marginals, but modified to 
bicuspid or unicuspid laterals and sometimes with ectoconal splitting 
on the marginals. Genitalia highly variable, epiphallus normally 
present, terminal genitalia normally thick at least in part, penis 
with or without verge, normally with complex stimulatory pilasters. 
As a temporary measure, all charopid genera not specifically as- 
signed to other subfamilies are lumped in the Charopinae. 

Subfamily Trukcharopinae Small to medium-sized species 
with apical sculpture of spiral cords, postapical sculpture strong to 
secondarily reduced. Aperture with or without barriers. Jaw and 
radula typically charopid. Genitalia unusual in lacking an epiphal- 
lus. Vas deferens passing through the penial retractor muscle before 
entering the penis either through a verge or pore. Interior of penis 
with stimulatory pilasters of varied form. Trukcharopa, 
Kubaryiellus, Russatus, Roimontis, Palikirus, Jokajdon, andPalline, 
new genera from Micronesia, are the included taxa. 

Subfamily Semperdoninae Medium-sized to large species with 
spiral apical cords, typical to greatly reduced postapical sculpture. 
Aperture with or without barriers. Jaw and radula without unusual 
features. Pallial region typical, with some variation in kidney lobe 
lengths. Genitalia unusual in the altered penis complex. Penial re- 
tractor inserting on head of epiphallus, which is a coiled double tube, 
outer wall thin muscle sheath, inner a thick glandular tube rolled 
inward on one side, opening into penis through a simple pore. Penis 
with an outer heavy muscle sheath of circular fibers that extends up 
around the lower epiphallus, with an inner tube containing fine lon- 
gitudinal pilasters above that usually coalesce into 3 glandular 
pilasters below. Semperdon, Ladronellum, and Himeroconcha, new 
genera from Micronesia, are the included taxa. 

The affinities of the subfamilies are still uncer- 
tain. The New Zealand Otoconchinae is an obvious 
sluglike secondary derivative from the New Zealand 
Charopinae. In structure it shows only minor dif- 
ferences. The Trukcharopinae from Micronesia are de- 
rivable from the typical Charopa-Sinployea stock by 
loss of the epiphallus and specialization of the penial 
surfaces. At present there are no Australian or New 
Zealand structural equivalents known, but I would not 
be surprised if relatives turn up in this area. The Sem- 
perdoninae from Micronesia have a very different 
penis structure than anything I have seen elsewhere. 



Other features of their anatomy are compatible with 
derivation from the Charopinae, and finding relatives 
in a remote area of the family range would not surprise 
me in the least. 

In contrast, the Rotadiscinae have a relict (Idaho, 
Arizona-Costa Rica, Trinidad, probably South 
America), Fiji, Western Australia (Solem, unpublished 
data) distribution. Their anatomy has a punctid aspect 
in part to the ovotestis, the pallial region is more 
primitive than that of the Charopinae, and its slender 
terminal genitalia with some accessory organs is very 
different from the Punctidae and most of the Charo- 
pinae. This group may prove to be a relatively base 
stock of the charopid radiation. 

Further phylogenetic comments are postponed 
pending completion of work on the Western Australian 
radiation and additional work on New Caledonian and 
Tasmanian taxa. Comments on relationships within 
subfamilies are given below. 

Subfamily ROTADISCINAE 

A more expanded definition and discussion is post- 
poned until completion of the Australian work. Inclu- 
sion of Microcharopa in the Rotadiscinae is based on its 
unique apical sculpture (fig. 4c-e) that is shared with 
Radiodiscus (Solem, 1977b, pp. 150-154, figs. 7-8), 
Rotadiscus, and undescribed Western Australian 
material. Microcharopa could not be dissected. In shell 
features it differs in its very small size and reduced 
whorl count, finer and more crowded radial ribs, and 
wide umbilicus. All of the New World genera are con- 
siderably larger, more coarsely ribbed and generally 
more variously umbilicated. 

Genus Microcharopa, new genus 

Shell minute, with less than 3M> tightly coiled whorls. Apex and 
spire slightly elevated, body whorl descension a little more rapid. 
Apical sculpture (fig. 4a-e) of short, twisted, open-ended ridges ar- 
ranged in 16-20 spiral rows on top of a weakly and broadly undulat- 
ing radially ridged surface. Postnuclear whorls with narrow, very 
crowded, protractively sinuated radial ribs. Microsculpture of very 
fine radials, finer spirals, and some weak secondary spiral cords. 
Umbilicus widely open, cup-shaped, regularly decoiling, margins 
weakly shouldered. Whorls almost evenly rounded, slightly com- 
pressed laterally above periphery and on basal margin. Aperture 
without barriers or heavy callus. Anatomy unknown. 

Type species. Microcharopa mimula, new species. 

Despite attempts on two different occasions to col- 
lect material on Lami Ridge and near Sigatoka, Viti 
Levu, no specimens could be found. Because only dead 
material was collected previously, the anatomy of this 
very unusual species could not be studied. Even under 
100 x magnification with the light microscope, the 
apex looks like it has only slightly wavy spiral cords, 
more frequent in number than with most species, but 
not very unusual in appearance. Scanning electron 
microscope photographs at 300 x- 10,000 x magnifica- 
tion (fig. 4a-e) show that the apical sculpture is very 
complex. Instead of continuous spirals, there are rela- 
tively short segments that are equal in width to the 



SYSTEMATIC REVIEW 



71 



vague radial undulations (fig. 4a). Each individual 
segment starts as a cord, twists up and over, then ends 
as a cuplike structure (fig. 4d-e). These are in spiral 
rows and are present except for the first quarter apical 
whorl (fig. 4c), which seems to have low spiral cords 
instead. Postnuclear sculpture (fig. 4b) shows no really 
significant difference from that found in many 
Charopidae. 

Separating Microcharopa from other Pacific gen- 
era is simple. The smallest Sinployea that has been 
dissected, S. inermis lakembana, is more than twice the 
diameter of Microcharopa, 2.36 mm. compared with 
1.07 mm., and the smallest species assigned to 
Sinployea, S. ellicensis nukulaelaeana, is 2.08 mm. in 
mean diameter, only slightly less than twice the size. 
Species of Vatusila are in the 1.65-1.88 mm. size 
range, but these have very prominent apertural bar- 
riers and much simpler apical sculpture. The only 
genus in which the shells are approximately equal in 
size is Discocharopa (p. 59). Populations of that taxon 
average about 1.37 mm. in mean diameter and have 
3V2-4 whorls, an even wider umbilicus, and, most im- 
portant, very different apical sculpture (fig. 5a-c, p. 
16). 

The peculiar apical sculpture is shared with 
Radiodiscus (see Solem, 1977b, p. 152, figs. 7-8), which 
differs in having 3 3 /4-4Mi whorls, reaches 2.0-2.25 mm. 
in diameter, and has about 21 ribs/mm, on the body 
whorl, compared with 36.5 ribs/mm, in Microcharopa. 
The narrower umbilici and larger sizes of Rotadiscus 
and Radiodomus are easy recognition features of these 
genera. 

Only the one species, Microcharopa mimula, which 
is widely distributed in the Lau Archipelago and on 
Viti Levu, is known. The name Microcharopa refers to 
the very small size of this species and its resemblance 
in shape to the historical usage of the name Charopa. 

Microcharopa mimula, new species. Figures 4a-e; 
33a-c. 

Diagnosis. Shell minute, diameter 0.95-1.13 mm. (mean 1.07 
mm.), with 3Vs-3% rather tightly coiled whorls. Apex and spire 
slightly to moderately and evenly elevated, body whorl descending 
slightly more rapidly, spire protrusion about Ve-Vs body whorl width, 
H/D ratio 0.400-0.469 (mean 0.449). Apical sculpture of 16-20 
(mean 17.8) prominent, crowded spiral cordlike structures. Postnu- 
clear sculpture of narrow, crowded, fine, sharply defined, protrac- 
tively sinuated radial ribs, 117-136 (mean 126.6) on the body whorl, 
whose interstices are 2-3 times their width. Ribs/mm. 32.3-40.5 
(mean 36.5). Microsculpture of barely visible radial riblets, 1-2 be- 
tween each pair of major ribs, equally fine spiral riblets, with occa- 
sional traces of weak secondary spiral cording. Umbilicus broadly 
open, cup-shaped, regularly decoiling, contained 2.43-3.05 times 
(mean 2.68) in the diameter, margins weakly shouldered. Whorl con- 
tours almost evenly rounded, slightly compressed laterally above 
periphery and on basal margin, aperture circular, inclined about 10 
from shell axis. 

The similarities in size and shape to Discocharopa 
are marked, but the spiral apical cording and absence 
of a deeply recessed parietal lamella immediately 
separate Microcharopa mimula from that genus. Other 
barrier-free Fijian Charopinae are at least twice the 






FIG. 33. a c, Microcharopa mimula, new species. Lami Ridge, 
west of Suva, Viti Levu, Fiji. Holotype. BPBM 178490. Scale line 
equals 0.5 mm. (MM). 



size of M. mimula at 3 l /& whorls and show numerous 
alterations in sculpture and shape. 

Description. Shell minute, with 3 1 A rather tightly coiled whorls 
that very slightly increase in width. Apex and spire slightly and 
evenly elevated, body whorl descending moderately, H/D ratio 0.467. 
Embryonic whorls slightly less than \ l k, sculpture of 18 crowded 



72 



SOLEM: ENDODONTOID LAND SNAILS 



spiral riblets, sinuated where crossing low, broadly rounded, rela- 
tively vague radial swellings. Postnuclear whorls with protractively 
sinuated, crowded, narrow, lamellar radial ribs, 136 on the body 
whorl, whose interstices are 1-2 times their width. Microsculpture 
as in diagnosis. Sutures deep, whorls shouldered above, gently and 
evenly rounded on outer margins. Umbilicus broadly open, saucer- 
shaped, regularly decoiling, contained 2.70 times in the diameter, 
margin weakly shouldered. Color very light yellow-brown. Aperture 
circular, strongly rounded near periphery and at baso-umbilical 
margin, inclined about 10 from the shell axis. Height of holotype 
0.48 mm., diameter 1.02 mm. 

Holotype. Fiji: Viti Levu, Station 47, Lami ridge, 
5 miles west of Suva, at 5-150 ft. elevation. Collected 
by Yoshio Kondo and C. M. Cooke, Jr., on July 24, 
1938. BPBM 178490. 

Range. Munia, Mothe, Wangava, Nayau, Na- 
muka, and Yangasa Levu, Lau Group, and Viti Levu, 
Fiji. 

Paratypes. Viti Levu: Near Sigatoka, '/2 mile in- 
land, under dead leaves, 50 ft. elevation (6 specimens, 
BPBM 88524, BPBM 88562); Sigatoka, near base of 
limestone cliff, Vz-lVfe miles from shore (1 specimen, 
BPBM 87930); Nayavu, upper Wainibuka River, 14 
miles inland, 190 ft. elevation (1 specimen, BPBM 
87959); Lami Ridge, 5 miles west of Suva (Station 47), 
limestone ridge, 5-150 ft. elevation (3 specimens, 
BPBM 178490). 

Lau Group: Munia, Vi mile inland (Station 65), 
hillside in copra plantation at 600-900 ft. elevation (2 
specimens, BPBM 179323); Mothe, central wooded 
peak, on leaf mold at 590 ft. elevation (4 specimens, 
BPBM 78585); Wangava, probably Station 27 (4 
specimens, ex BPBM 166994); Nayau, Nauko (Station 
43), hillside % mile inland at 250 ft. elevation (4 speci- 
mens, BPBM 167231); Namuka, Matandolo (Station 2), 
north-central point of island on hillside 500 ft. inland 
at 65-75 ft. elevation (2 specimens, ex BPBM 166614); 
Yangasa Levu, south end of island (Station 30), hill- 
side, 150 ft. inland at 40 ft. elevation (2 specimens, ex 
BPBM 167137). 

Remarks. The extremely small size, widely open 
umbilicus, and very crowded sculpture easily separate 
Microcharopa mimula from all other Fijian endodont- 
oids. It is most similar to the Melanesian-Polynesian 
Discocharopa in size and form but obviously differs in 
its spiral apical sculpture and the lack of parietal bar- 
riers. The wide distribution recorded above is undoubt- 
edly very incomplete because the minute size of this 
species would result in its discovery by only the most 
diligent collector. 



Subfamily CHAROPINAE 

A brief diagnosis of the Charopinae was given on 
p. 70. It undoubtedly will be extensively modified when 
taxa extralimital to this study are reviewed. Because 
so few genera have had details of their terminal 
genitalia described or figured, I have not attempted to 
split the Charopinae into smaller clusters of genera. I 
can emphasize that eventual generic clusters will in- 



clude taxa from highly diverse geographic areas and 
will deny many traditional associations. For example, 
the basic anatomical structures of Notodiscus Thiele, 
1931, found on Kerguelen and other Subantarctic is- 
lands (Solem, 1968c), are very close to those seen in 
Phenacohelix Suter, 1892, from New Zealand (fig. 11); 
the structures seen in Graeffedon from Samoa (fig. 87) 
are matched closely by those found in Helenoconcha 
relicta Solem, 1977, from St. Helena (Solem, 1977c) and 
differ markedly from any Austro-Zelandic taxa dis- 
sected to date; Amphidoxa Albers, 1850, from Juan 
Fernandez (fig. 30a-e) shows a combination of basic 
Charopinae and unusual features and is quite unlike 
the anatomical pattern of Stephanoda Albers, 1860 
(fig. 31a-i), from Chile, which was treated as a sub- 
genus of Amphidoxa by Pilsbry (1893-1895, pp. 
39-41), Thiele (1931, p. 575), and Zilch (1959-1960, pp. 
221-222). The last commentator on these two genera, 
Odhner (1922, pp. 230-233, figs. 8-13, pi. 9, figs. 
33-35), described the jaw and radulae of several Juan 
Fernandez species, plus an external view of the 
genitalia in "Stephanoda quadrata." Structures of the 
latter agree quite closely with the data presented here 
on Amphidoxa marmorella (Pfeiffer, 1845). Probably 
the Juan Fernandez "Amphidoxa and Stephanoda" of 
Odhner (1922) are monophyletic and best lumped 
under Amphidoxa. They are quite distinct from the 
Chilean Stephanoda. Given the above diversity and 
uncertainty, no firm allocations into clusters are pos- 
sible. 

Fortunately, the Pacific Island taxa, with three 
exceptions, appear to be a monophyletic unit. Dis- 
cocharopa Iredale, 1913, with a Philippine to Society 
Island range, Lagivala, known from Indonesia to Fiji 
and the Ellice Islands, and the endemic Graeffedon 
from Samoa and Tonga show major differences from 
the other Charopinae. The remaining genera, insofar 
as they have been dissected, belong to the same basic 
stock as the New Zealand Charopa Albers, 1860, and 
presumably will remain Charopinae regardless of fu- 
ture revisions. 

Discocharopa differs in shell microsculpture (figs. 
5a-f, 37d). The only partial knowledge of its genital 
anatomy (fig. 34a-d) makes relating it to other taxa 
difficult. The extent to which the minute shell size 
(mean diameter 1.36 mm.) has resulted in slenderiza- 
tion and/or simplification of the anatomy is unknown. 
As discussed below (p. 75), a degree of similarity to the 
structures of Phenacohelix pilula (Reeve) (fig. lla-e) 
exists, but their degree of relationship is unknown. 

Lagivala has not been dissected and has an exten- 
sive extralimital range, with species known from In- 
donesia, New Guinea, and the Bismarck Archipelago. 
The extent of its relationship to the Philippine- 
Indonesian problematic genus Beilania Preston, 1913, 
is uncertain (see p. 184). Whether Lagivala is conver- 
gent with the New Zealand Mylesia (Climo, 1978) and 
Ptychodon Ancey, 1888, will require detailed compari- 
sons that are beyond the scope of this report. It is only 



SYSTEMATIC REVIEW 



73 



possible to state that in barrier structure, distribution, 
and basic shell features, Lagivala is not similar to 
other Pacific Island genera. 

Graeffedon differs quite dramatically in its anat- 
omy (p. 200) and has very different barrier structure (p. 
17). It is clearly related to the St. Helena endemic, 
Helenoconcha, and probably will receive eventual sub- 
family separation. 

Of the remaining Pacific Island Charopinae, 
adequate anatomical data is available for Sinployea, 
Ba, and Tuimalila, fragmentary data for one species of 
Vatusila, and no data on the anatomy of the monotypic 
Maafu and Lauopa. The anatomical features of the 
Pacific Island genera agree with those of Champa (figs. 
9a-g, 10) in the basic pallial structure (except for the 
lack of a glandular extension onto the pallial roof) and 
the components of the interior terminal genitalia. The 
penes have a verge, pocket stimulator, and circular 
ridges. The lower female tracts have complex valvular 
arrangements at the free oviduct and longitudinal 
pilasters (compare figs. 9d and 57f). As pointed out 
above (pp. 27-29), the relative prominence of these pe- 
nial complex structures differs dramatically from 
species to species under conditions of sympatry, but the 
presence of the same exact structural elements in 
Charopa, Sinployea, Tuimalila, Ba, and such extralim- 
ital taxa as Amphidoxa (fig. 30e), Pilsbrycharopa (see 
Solem 1970a, fig. 2i), and Pseudocharopa (fig. 32b) 
indicate that this is a basic pattern. The lengthened 
terminal female organs of Charopa (fig. 9b) are a minor 
change of little or no phyletic significance. Charopa 
(fig. 10) differs from Pacific Island taxa most noticeably 
in the comparatively simple entrance of the vas def- 
erens into the epiphallus. This contrasts with the 
elaborate valve seen in Sinployea (fig. 57e). Despite the 
extreme compaction of its genital system, the New Zea- 
land Flammulina shows the same basic internal ele- 
ments of the penis (fig. 29e) and has a complex vas 
deferens-epiphallus junction (fig. 29i-j) equivalent to 
that seen in the Pacific Island genera. 

Without added reference dissections of extralimi- 
tal taxa, attempts to propose a phylogenetic tree for the 
Charopinae are premature. The important point here 
is that many Pacific Island genera are clusterable with 
New Zealand, New Guinea, Lord Howe Island, and 
Juan Fernandez genera. That their degree of relation- 
ship and phylogenetic order cannot yet be ascertained 
is a secondary problem. 

Just in reference to the Pacific Island taxa grouped 
here, Sinployea is most generalized, with Ba an obvi- 
ous specialization from an ancestor very similar to if 
not actually the extant species Sinployea irregularis 
(Garrett). The Tongan Tuimalila is an experiment in 
increased whorl width and gigantism also derivable 
from a Sinployea ancestor. Both Maafu and Lauopa, 
neither of which have been dissected, present prob- 
lems. Maafu shows great conchological specializations, 
but can be derived from the Sinployea inermis groups 
found in the same area. Its changes are of the same 



order of magnitude found in the Lau Archipelago En- 
dodontidae radiation of Zyzzyxdonta, Priceconcha, and 
Thaumatodon (Solem, 1976b, pp. 461-467), and this is 
interpreted as a local experiment. Lauopa, known from 
limited subfossil material, is grouped here for conveni- 
ence but conceivably could be an outlier of the Semper- 
doninae (see p. 177). The data needed to determine its 
affinities are not available. Vatusila has only a frag- 
ment of its anatomy known, but the penis structures 
(fig. 84a-b) are matched almost exactly by those seen 
in Sinployea aunuuana (fig. 53c). I am confident that 
Vatusila is correctly associated. Similarities in barrier 
size, shape, and positioning (figs. 82-83, 85) to some of 
the New Zealand Ptychodon andFectola (Climo, 1978, 
figs. 3-4, 7-8) are intriguing, but deciding whether 
this is convergence, parallelism, or genealogy is be- 
yond the scope of this review. Within Vatusila, there is 
a trend toward barrier reduction (see p. 192) and wider 
spacing of the radial sculpture. Although it would be 
tempting to suggest because of this that Vatusila 
might represent the ancestor of Sinployea, I suspect 
that they do not have a direct ancestor-descendant re- 
lationship but represent two independent colonizations 
of the Pacific Islands. 

Thus, of the Charopinae on the Pacific Islands, I 
interpret Ba, Tuimalila, and Maafu as local direct der- 
ivations from Sinployea, with Vatusila an independent 
colonization. Lauopa is of uncertain status, whereas 
Discocharopa, Lagivala, and Graeffedon also represent 
independent colonizations of the Pacific Islands. Be- 
cause there are several independent colonizations 
where external affinities are unknown, preparation of 
phylogenetic trees equivalent to those of the Endodon- 
tidae (Solem, 1976b, pp. 110-111, figs. 57-58) was not 
attempted. 

The fossil evidence is meager but intriguing. Vat- 
usila eniwetokensis (Ladd, 1958) is a Miocene species 
from Eniwetok in the Marshall Islands, whereas ex- 
tant species are found in Lau Archipelago, Tonga, 
Niue, and Vaitupu, Ellice Islands. Lagivala davidi 
(Ladd, 1968) is a Late Pleistocene or Recent fossil from 
Funafuti, Ellice Islands. Extant species are known 
from Indonesia, New Guinea, New Britain, and Viti 
Levu. The genus has not been recorded from the Sol- 
omon Islands, New Hebrides, or New Caledonia. The 
Funafuti species is very closely related to the Fijian L. 
vivus. The Marshall Islands' Miocene Vatusila is 
matched by the Miocene Cookeconcha subpacificus 
(Ladd, 1958) and the Pliocene to Pleistocene Minidonta 
inexpectans (Ladd, 1958) from Bikini, Marshall Is- 
lands. All three genera show a shrunken post-Miocene 
range correlating with the changing of the Marshall 
Islands from high islands to atolls (see Solem, 1976b, 
pp. 117-118). The distributional shift is thus inter- 
preted as range contraction, rather than the more 
simplistic assumption of migration. 

Better understanding of the Pacific Island 
Charopinae relationships requires more intensive in- 
vestigation of the Lau Archipelago, especially to obtain 



74 



SOLEM: ENDODONTOID LAND SNAILS 



anatomical material of Lauopa and Maafu, combined 
with analysis of extralimital genera. 

The generic ordering used below places two of the 
"convenience-grouped" genera first (Discocharopa) and 
last (Graeffedon), with Lagivala sited for comparative 
purposes next to Vatusila. Detailed discussions of vari- 
ation within and among genera are grouped under the 
appropriate generic discussions rather than here. 

Genus Discocharopa Iredale, 1913 
Proc. Malacol. Soc. London, 10 (6), pp. 379-380. 

Shell minute, with slightly more than 3'/2 normally coiled 
whorls. Apex and spire usually slightly to moderately elevated, body 
whorl descending more rapidly. Apical sculpture of major radial ribs 
more crowded near end of apex, with periostracal, relatively regular 
folds providing a microspiral element. Postnuclear whorls with nar- 
row, sharply defined, crowded, strongly protractively sinuated radial 
ribs. Microsculpture of very fine radial riblets with serrated edges, 
no trace of spiral sculpture. Umbilicus very widely open, saucer- 
shaped, regularly decoiling, margins rounded. Whorls flattened lat- 
erally above and below rounded periphery. Color white with a yellow 
tinge. Aperture with or without a small to very large and deeply 
recessed, medial parietal barrier. Pallial region with complete sec- 
ondary ureter, kidney bilobed, pericardial lobe longer than rectal. 
Genitalia with short bilobed ovotestis, slender talon. Terminal 
genitalia slender with subapical entrance of vas deferens, no exter- 
nally recognizable epiphallus, penis internally with longitudinal 
pilasters. 

Type species. Charopa (Discocharopa) exquisita 
Iredale, 1913, by original designation. 

The shell sculpture of Discocharopa (figs. 5a-f, 
37d) is quite different from that found in any other 
Pacific Island Charopidae. Like the Endodontidae, the 
apical whorls (fig. 5b-c) have prominent radial ribs 
that become quite crowded near the nuclear-post- 
nuclear boundary (fig. 5a), but the microsculpture is 
quite different from that seen in the Endodontidae. The 
latter have very fine, regular, "squiggly" microspirals 
that continue onto at least the early postnuclear 
whorls (Solem, 1976b, p. 38, fig. 28c-e; p. 39, fig. 29a). 
The Charopidae have fine to prominent spiral cords 
(for example Sinployea modicella, fig la-c, p. 10) with 
weak to strong (fig. 2c, p. 11) secondary radial ele- 
ments. Discocharopa has spiral sculpture that is of ir- 
regular shape, length, spt cing, and angle of orienta- 
tion. Similar apical sculpture has been found in several 
New Zealand species usually classified in Ptychodon 
(for example, P. microundulata in Solem, 1970b, pi. 58, 
figs. 4-6) but now referred to Mylesia (see Climo, 1978, 
p. 185). The postnuclear sculpture in Discocharopa 
differs from all endodontoids examined to date. All 
these have both microspiral and microradial elements, 
but Discocharopa (fig. 5b, d-f) shows no trace of micro- 
spiral elements, and the radial sculpture has unusual 
features. The major ribs (fig. 5a-b, d-f) retain rem- 
nants of microradial clustering on their apices only on 
early postnuclear whorls (a-b), whereas the latter 
whorls (d-e) appear to have a "rolled" or expanded rib 
peak. The microradials (e-f) are partly broken up into 
irregular short segments by transverse grooves. The 



latter have no counterpart in any Pacific Island species 
examined to date. 

After considerable effort over many years, it was 
possible to obtain material and partially work out the 
basic anatomy of Discocharopa. The wide range of Dis- 
cocharopa yielded hope of borrowing material to dis- 
sect, and fieldwork by L. Price in Fiji and by Price and 
Solem in Australia could have produced live material. 
No material was obtained until two specimens from the 
National Museum, New Zealand, were provided for 
study by their collector, Frank Climo. They were col- 
lected "behind the landing," Raoul Island, Kermadec 
Islands, on August 24, 1972, and thus represent near 
topotypic material of the genotype, Discocharopa ex- 
quisita (Iredale, 1913, pp. 379-380), which was de- 
scribed from "Sunday Island, Kermadec Group. Living 
under rotten wood, stones, etc." 

Both examples had completely retracted within 
their shells, and the albumen gland had crystallized, 
which made interpretation of most structures difficult. 
The main uncertainties include whether the prostate- 
uterus are combined or separated, the talon structure, 
and the position and shape of the spermathecal head. I 
suspect that the latter is bound into the aorta at the 
area where the kidney base and albumen gland touch 
the diaphragm. In neither specimen was I successful in 
separating the kidney base, albumen gland, intestines, 
and hindgut because of extensive albumen gland crys- 
tallization. Undoubtedly the spermathecal head was 
mangled and lost in the attempts to work out 
structures in this area. The exact shape, size, and re- 
lationship of the talon to the hermaphroditic duct could 
not be determined precisely. Figure 34b represents the 
best interpretation of dissector and illustrator, but 
could be incorrect. 

The ovotestis (fig. 34a, G) is unusual in that it lay 
alongside the stomach and below the main area of di- 
gestive gland. The bilobed condition is found in many 
charopids, but the anterior shift in position has not 
been seen by me in other taxa. The hermaphroditic 
duct (GD) was iridescent, again as in many charopids, 
and the albumen gland was deeply dented by loops of 
the intestine, thus having a very irregular shape. A 
swollen uterine area was clear, but the prostate tissue 
was very small and the spermathecal shaft closely 
bound to the margin. Because of partial crystallization 
in the uterine area, details could not be worked out. 
The vas deferens broke and is shown (VD) reflexed, but 
the long vagina and penis have been unfolded in order 
to show a little more detail (fig. 34c). The entrance of 
the vas deferens into the penis is subterminal, with a 
short penial retractor muscle inserted apically. Appar- 
ently there are at least two longitudinal pilasters 
within the penis, but exact structures are too small to 
be determined from available material. 

The pallial region (fig. 34d) has a closed and com- 
plete secondary ureter (KD); the kidney (K) is bilobed 
with a large section touching to partly overlapping the 



SYSTEMATIC REVIEW 



75 



GG 





FIG. 34. Anatomy of Discocharopa aperta (MbllendorfD. Forest 
behind landing, Raoul Id., Kermadec Islands. F. Climo! VIII-24-1972. 
FMNH 193764: a, ovotestis; b, detail of talon and carrefour; c, post- 
apical genitalia; d, pallial region. Scale lines as marked. (EL). 



hindgut (HG). Details of the anus (A) and external 
ureteric pore (KX) are only approximate. The apex of 
the kidney is curved over around the start of the ure- 
ter, possibly as a result of retraction into the shell. 

The larger of the two dissected specimens was 0.62 
mm. high, 1.51 mm. in diameter, H/D ratio 0.413, 
whorls 3%, umbilicus 0.43 mm. wide, D/U ratio 3.54. 
The more successful dissection was made on the 
smaller example, which was 0.58 mm. high, 1.25 mm. 
in diameter, with 3% whorls. 

All observed anatomical features are consistent 
with Discocharopa being a member of the Charopinae. 
Unfortunately the small size, contraction, and partial 
crystallization combined to prevent full study. 
Whether the slender terminal genitalia is a secondary 
specialization is uncertain. Unquestionably the fea- 
tures observed are different from Sinployea and its re- 
latives. To a certain extent there are similarities with 
Phenacohelix (fig. lie, e) in general penis shape, 
ovotestis, and pallial configuration, but there is a clear 
difference in vas deferens insertion, talon, and termi- 
nal female genitalia configurations. Of the extralimi- 
tal taxa considered here, Discocharopa has the most 
similarities to Phenacohelix, but the degree of their 
relationship remains unknown. 



Because only partial anatomical data was re- 
covered, this general discussion substitutes for a for- 
mal description. 

Originally, only the Kermadec Islands populations 
and the Tasmanian D. bassi (Legrand, 1871) were 
placed in Discocharopa. Subsequently, Iredale (1937a, 
p. 325) included several additional Australian and 
Tasmanian species, Cotton (1939, p. 176) described a 
Paralaoma as belonging to this genus, van Benthem 
Jutting (1951) named/), microdiscus from the Celebes, 
and I (Solem, 1957, 1959a) added Philippine and New 
Hebridean (fig. 36) taxa. None of the Tasmanian 
species, D. vigens (Legrand, 1871), D. bassi (Legrand, 
1871), D. lottah (Petterd, 1879), orD. mimosa (Petterd, 
1879), appear to be correctly classified in Discocharopa, 
but generic reallocation is not attempted here. Dis- 
cocharopa concinna (Hedley, 1901) and D. planor- 
bulina (Tate, 1896) from Queensland and the Northern 
Territory of Australia are synonymized with D. aperta 
(Mollendorff) from the Philippines. The latter is the 
oldest name and applies to the single species recog- 
nized. Discocharopa werneri Solem (1957, pp. 4-6, fig. 
2) from Mindanao is totally unrelated to Discocharopa. 
Its original assignment here was a desperation effort to 
avoid description of a new genus, compounded by an 
unwillingness to place it in a New Zealand genus when 
adequate comparative material was lacking. 

Combining all described species with D. aperta is 
based upon examination of all types and study of over 
300 specimens. This results in recognizing a species 
with the widest distribution of any endodontoid, ex- 
tending from the northern Philippines and Java to 
Central Australia, Kermadec Islands, and the Society 
Islands. The only approximately equivalent distribu- 
tion is that shown by Stenopylis coarctata (Mollendorff, 
1894), which has almost the same Indonesian and 
Australian range, but only reaches the Solomon Is- 
lands and has not been found in the New Hebrides, Fiji, 
Kermadec Islands, or Polynesia. Data on this species 
are summarized by Solem (1957, pp. 8-11, fig. 4). Sub- 
sequently it was reported from Bach Long Vi (Nightin- 
gale Island) in the Gulf of Tonkin (Saurin, 1960, pp. 
7-9, fig. 3, pi. 1; figs. 6a-b, 7) and was mistakenly 
described as a new species, Microphyura nightingali. 
Microphyura is a New Caledonian endemic rhytidid 
genus whose conchological appearance is convergent 
with that of Stenopylis. Hedley (in Tate, 1896, pp. 
221-222, fig. C) illustrated the jaw and radula of 
Stenopylis. Tentatively, I (Solem, 1975) classified it in 
the Helicodiscidae, an otherwise North American com- 
plex, and subsequent dissection has confirmed this as- 
signment (Solem, unpublished data). 

Discocharopa and Stenopylis have been collected 
together on several occasions, and I suspect their 
ecological requirements are very similar. The method 
of apertural narrowing is very different, with Dis- 
cocharopa using a single, deeply recessed parietal, 
whereas Stenopylis uses a combination of lip thicken- 



76 



SOLEM: ENDODONTOID LAND SNAILS 



ing and internal tubercles (Solem, 1957, p. 11, fig. 
4b-c, e). 

Differences of Discocharopa from other genera are 
gross in nature. Microcharopa from Fiji is most apt to 
be confused in size but is slightly smaller (mean diame- 
ter 1.07 mm.), has strong spiral apical sculpture (fig. 
4a-e, p. 13), proportionately larger apical whorls, and 
more evenly rounded body whorl contours. Punctum 
has a much higher spire, is distinctly smaller, has 
prominent spiral apical cords and a narrower um- 
bilicus. No Pacific Island Charopidae without promi- 
nent apertural barriers average less than 2.00 mm. in 
diameter, whereas those Lagiuala and Vatusila in the 
1.40-1.75 mm. size range have very prominent aper- 
tural barriers at or very near the lip edge. Most of these 
have marked lateral body whorl compression. Of the 
six Endodontidae averaging less than 2 mm. in diame- 
ter, only the Manu'a Group Minidonta manuaenis 
Solem 1976, occurs within the geographic range of Dis- 
cocharopa. That species has more than four whorls, a 
very narrow umbilicus, H/D ratio over 0.550, and seven 
prominent apertural barriers. The other species of 
Minidonta are Hawaiian, Mangarevan, and the Bikini 
Atoll fossil, M. inexpectans (Ladd, 1958). 

Discocharopa aperta (Mollendorff, 1888). Figures 
5a-f, 34a-d, 35a-b, 36a-c, 37a-d. 

Patula aperta Mollendorff, 1888, Nachr. Bl. deut. Malak. Gesell., 
20 (5-6), p. 89 Montalban, Rizal, Luzon, Philippines; Pilsbry, 
1892, Man. Conchol., (2) 8, pp. 80-81, pi. 37, figs. 35-37; Mol- 
lendorff, 1898, Abhl. Naturf. Ges. Gorlitz, 22, p. 88 Luzon, 
Leyte, Calamianes. 

Endodonta (Champa) planorbulina Tate, 1896, Rep. Horn Sci. 
Exped. Central Australia, (2), p. 187, pi. 17, fig. 3 Palm 
Creek, Krichauff Range, Northern Territories, Australia. 

Endodonta concinna Hedley, 1901, Proc. Linn. Soc. New South 
Wales, 25, p. 729, pi. 48, figs. 1-3 Bundaberg, Queensland, 
Australia. 

Charopa (Discocharopa) exquisita Iredale, 1913, Proc. Malacol. 
Soc. London, 10 (6), pp. 379-380, pi. XVIII, fig. 8 Sunday 
Island, Kermadec Group. 

Ptychodon celebica B. Rensch, 1932 (not Sarasin & Sarasin, 1899), 
Zool. Jahrb., Syst, 63, p. 101 Sumba; B. Rensch, 1935, Sitz.- 
Ber. Gesell. Naturf. Freunde, Berlin, 1935, p. 322 Timor. 

Pyramidula aperta (Mollendorff), Faustino, 1930, Philippine J. 
Sci. ,42(1), p. 110. 

Discocharopa planorbulina (Tate), Iredale, 1937, Australian Zool., 
8 (4), p. 325; Iredale, 1937, South Australian Nat., 18 (2), p. 25, 
pi. 1, fig. 21. 

Discocharopa concinna (Hedley), Iredale, 1937, Australian Zool., 8 
(4), p. 325. 

Charopa exquisita (Iredale), I. Rensch, 1937, Arch. f. Naturgesch., 
n. f., 6 (4), pp. 590-591 Malkong-Bach, Weiten Buch and 
Vunapope, Gazelle Peninsula, New Britain, Bismarck Ar- 
chipelago. 

Charopa (Discocharopa) microdiscus van Benthem Jutting, 1951, 
Basteria, 15 (1-2), pp. 28-29, fig. 1 South Celebes and West 
Java; van Benthem Jutting, 1952, Treubia, 21 (2), p. 398 
West Java; van Benthem Jutting, 1953, ibid., 22 (2), p. 302 
Ambon. 

Discocharopa aperta (Mollendorff), Solem, 1957, Fieldiana: Zool- 
ogy, 42 (1), pp. 3-4, fig. 1, a-d Luzon and Catanduanes, 
Philippine Islands. 

Discocharopa microdiscus van Benthem Jutting, Solem, 1958, 
Arch. f. Mollusk., 87 (1-3), p. 21. 




FIG. 35. Parietal barrier variation in Discocharopa aperta (Mol- 
lendorff): a, Tjamba Pass, near Makassar, Celebes. ZMA: b, Blue 
Lake, Sunday Island, Kermadec Islands. Paratype of Charopa ex- 
quisita Iredale, 1913. FMNH 117214. Scale line equals 1 mm. (JC). 



Charopa microdiscus van Benthem Jutting, 1958, Verh. Naturf. 

Ges. Basel, 69 (1), p. 106 Sumba, Indonesia. 
Discocharopa planulata Solem, 1959, Fieldiana: Zoology, 43 ( 1), pp. 

82-83, pi. 32, figs. 1-3 stream drift in Sarakata River Valley, 

Espiritu Santo, New Hebrides. 

Diagnosis. Shell minute, diameter 1.18-1.84 mm. (mean 1.36 
mm.), with 3*78-4(4 normally coiled whorls. Apex and spire rarely 
flat, usually slightly to moderately elevated, body whorl descending 
more rapidly to very strongly deflected (Rurutu, Austral Islands), 
spire protrusion 'Az to more than 4 body whorl width, usually about 
l k, H/D ratio 0.341-0.615 2 (mean 0.438). Apical sculpture of radial 
ribs that on lower portion of apex becomes more crowded, with barely 
visible spirals. Postnuclear whorls with prominent, narrow, sharply 
defined, strongly protractively sinuated radial ribs, 77-148 (mean 
110.6) on the body whorl, whose interstices usually are 2-4 times 
their width. Ribs/mm. 18.0-40.4 (mean 26.9). Microsculpture of very 
fine radial riblets, 3-5 between each pair of major ribs, no secondary 
spiral sculpture. Umbilicus very widely open, saucer-shaped, regu- 
larly decoiling, contained 2.00-3.21 times (mean 2.58) in the diame- 
ter, margins rounded. Sutures impressed, whorls strongly rounded 
above and on basal margin, usually very slightly to slightly com- 
pressed laterally above and below evenly rounded periphery. Aper- 
ture subcircular, moderately to strongly compressed laterally above 
periphery, less strongly compressed laterally below periphery, in- 
clined 10-30 from shell axis. Parietal wall with or without a medial 
barrier, which, when present, is usually recessed Vt to more than !4 
whorl and varies from a low cordlike ridge extending 3 Ae of a whorl to 
a very high crescentic lamella of up to % aperture height, with 
gradual anterior and posterior descensions, moderately expanded 
and weakly serrated medially when very high. Presence or absence 
on either a populational or individual basis. Color white, with at 
most a faint yellow tinge. 

In having stronger radial than spiral apical 
sculpture, a very widely open umbilicus, and at most 
only a single very deeply recessed parietal barrier, Dis- 
cocharopa aperta is immediately differentiated from 
other species of similar size. Species of Beilania and 

2 The only two specimens with H/D ratio greater than 0.500 
(0.549, 0.615) are Rurutu examples with greatly deflected body 
whorls. 



SYSTEMATIC REVIEW 



77 



Lagivala have spiral apical sculpture predominating, 
more apertural barriers that are near the lip edge (at 
least for the parietals), and generally much thicker 
body whorls with less sinuated sculpture. Mi- 
crocharopa mimula is most similar in size and shape, 
but differs in its very distinctive apical sculpture (fig. 
4), lower whorl count, lack of apertural barriers, and 
proportionately much larger apical whorls. 

Description of Patula aperta. Shell minute, with slightly less 
than 3% normally coiled whorls. Apex flat, spire slightly elevated, 
body whorl descending a little, H/D ratio 0.443. Apical whorls l'/2, 
sculpture partially eroded, but traces of radial ribbing remaining. 
Postnuclear whorls with narrow, somewhat irregular radial ribs, 
eroded over much of the shell. Microsculpture obscured by incrusta- 
tions and eroding of surface except for faint traces in umbilicus. 
Sutures deep, whorls strongly rounded above, somewhat flattened 
laterally above periphery with evenly rounded outer margins. Um- 
bilicus saucer-shaped, regularly decoiling, contained 2.51 times in 
the diameter, with evenly rounded margins. Color white, with 
periostracal remnants having a faint yellow tone. Aperture subcircu- 
lar, slightly flattened laterally above periphery, inclined about 15 
from the shell axis. No parietal barrier visible from aperture. Height 
of lectotype 0.64 mm., diameter 1.45 mm. 

Lectotype. Philippine Islands: Luzon, Montalban. 
SMF 165358. 

Description of Endodonta (Charopa) planorbulina. Shell mi- 
nute, with slightly more than 3Vi normally coiled whorls. Apex and 
spire barely protruding, last whorl not descending, H/D ratio 0.400. 
Apical whorls 1%, surface badly eroded with only faint traces of 
regularly spaced radial ribbing remaining. Postnuclear whorls with 
moderately widely spaced, protractively sinuated radial ribs, 91 on 
the body whorl, whose interstices are 2-3 times their width. Micro- 
sculpture mainly eroded, consisting, when visible, of faint radials. 
Sutures relatively deep, whorls strongly rounded above, slightly flat- 
tened laterally above periphery with evenly rounded outer margin. 
Color white. Umbilicus saucer-shaped, regularly decoiling, con- 
tained 2.70 times in the diameter with rounded margins. Aperture 
subcircular, slightly flattened laterally above periphery, inclined 
about 10 from shell axis. Height of holotype 0.61 mm., diameter 1.52 
mm. 

Holotype. Australia: Palm Creek, Krichauff 
Range, Northern Territories. SAM D.3222. 

Description of Endodonta concinna. Shell minute, with 3'/a rel- 
atively loosely coiled whorls. Apex and spire barely elevated, last 
half of body whorl descending moderately, H/D ratio 0.445. Apical 
whorls 1%, sculpture of relatively widely spaced radial ribs with an 
extremely fine microsculpture of more crowded spiral riblets. Re- 
maining whorls with narrow, relatively crowded, protractively 
sinuated radial ribs, about 102 on the body whorl, whose interstices 
are 2-4 times their width. Microsculpture barely visible, consisting 
of fine radial riblets. Sutures relatively impressed, whorls strongly 
rounded above, flattened laterally above periphery and partially on 
basal margin. Color very light yellowish white. Umbilicus saucer- 
shaped, regularly decoiling, contained 2.81 times in the diameter. 
Aperture ovate, compressed laterally above periphery and on basal 
margin, inclined about 15 from the shell axis. Height of lectotype 
0.61 mm., diameter 1.48 mm. 

Lectotype. Queensland: Bundaberg. AMS C.8970. 

Description of Charopa (Discocharopa)exquisita. Shell minute, 
with 314 normally coiled whorls that only slightly increase in width. 
Apex and spire barely emergent, body whorl descending slightly, 
H/D ratio 0.450. Apical whorls l'/2, first quarter whorl worn smooth, 
lower sculpture of prominent, narrow radial ribs, more widely spaced 
above, becoming crowded near end, with microsculpture of very fine 
spiral ribbing. Remaining whorls with narrow, strongly protrac- 
tively sinuated lamellar radial ribs, 100 on the body whorl, whose 
interstices are 3-5 times their width. Microsculpture of very faint 



radial riblets. Sutures deeply impressed, whorls evenly rounded. 
Umbilicus saucer-shaped, broadly opened, regularly decoiling, con- 
tained 2.58 times in the diameter. Color light yellow-horn without 
darker maculations. Aperture circular, flattened laterally above 
periphery, inclined about 20 from shell axis. Parietal wall with 
single, very deeply recessed barrier visible through shell. Height of 
lectotype 0.59 mm., diameter 1.32 mm. 

Lectotype. Kermadec Islands: crater slopes 
northeast of Blue Lake, Sunday Island. Collected 
under stones by W. R. B. Oliver on October 3, 1908. 
BMW. 

Description of Charopa (Discocharopa) microdiscus. Shell 
minute, with slightly more than 3% normally coiled whorls. Apex 
and spire slightly and evenly elevated, body whorl descending a little 
more rapidly, H/D ratio 0.424. Apical whorls 1%, sculpture of nar- 
row, prominent, moderately widely spaced radial ribs, becoming 
more crowded near end with a much finer, very inconspicuous mi- 
crosculpture of spiral ribs. Postnuclear whorls with fine, strongly 
protractively sinuated lamellar radial ribs, 111 on the body whorl, 
whose interstices are 2-3 times their width. Microsculpture barely 
visible under extreme magnification, consisting of very fine radial 
riblets. Color very faint yellowish white. Sutures deep, whorls 
strongly rounded above, flattened laterally above and below evenly 
rounded periphery with slightly flattened columellar margin. Um- 
bilicus very broadly open, saucer-shaped, regularly decoiling, con- 
tained 2.25 times in the diameter. Aperture subcircular, flattened 
laterally above and below periphery, inclined about 15 from the 
shell axis. No parietal barrier visible from aperture. Height of 
holotype 0.69 mm., diameter 1.63 mm. 

Holotype. South Celebes: hills of Pangkadjene, 
Makassar. ZMA. 

Range. Philippines and Java east through 
Sumba, Timor, and Biak to the Bismarck Archipelago, 
New Hebrides, and Fiji, Kermadec Islands, Manu'a 
Group in Samoa, Rurutu in the Austral Islands, 
Borabora in the Society Islands, Northern Australia as 
far south as the MacDonnell Ranges and Northern Vic- 
toria, and Bundaberg in coastal Queensland. Probably 
in most intermediate areas. 

Material. Philippine Islands: north Luzon (4 
specimens, RSME); Montalban (7 specimens SMF 
165358-60, NMWC). Catanduanes (1 specimen, 
FMNH 57193). Panglao (1 specimen, RSM). Cala- 
mianes: Koron (3 specimens, SMF 165361). Mindanao: 
Samial (5 specimens, SMF 165362). Busuanga (4 
specimens, SMF 165363, FMNH 18725). Bulacan: 
Bayabac (7 specimens, SMF 165364-5). 

Java: Koeripan, Buitzenborg (2 specimens, ZMA, 
collected February 21, 1932, by van Benthem Jutting). 

Sumba: Waikarudi (2 specimens, NHB 6092-c); 
Langgaliru (1 specimen, NHB 6092-d); Kedi (1 speci- 
men, NHB 6092-b); Mau Marru (5 specimens, NHB 
6092-a, ZMA). 

Ambon (2 specimens, ZMA, collected October 
11-13, 1949, by M. A. Lieftinck). 

South Celebes: Pangkadjene near Makassar (21 
specimens, ZMA, collected May 9 and October 14, 1948, 
by G. A. Tammes-Bolt); Palopo near Rante Pao, 23 km. 
from Makale at 700-800 m. elevation (1 specimen, 
ZMA, collected October 13, 1948, by G. A. Tammes- 
Bolt); Kalkrotsenlangs Grote, on way from Makale, 



78 



SOLEM: ENDODONTOID LAND SNAILS 



near Kalossi at 700-800 m. elevation (4 specimens, 
ZMA, collected September, 1948, by G. A. Tammes- 
Bolt). 

Timor: Araki-Vlakte (4 specimens, ZMA, collected 
April 12, 1950, by B. Polak); Niki-Niki (5 specimens, 
NHB 4169-a); Noilmina (40 specimens, ZMB, FMNH 
146025). 

Biak: Zuid-Biak Subdistrict (Station G-24) (olim 
Sorido) (1 specimen, ZMA, collected March 26, 1952). 

Aru Islands: Wokan (1 specimen, MSNG). 

Queensland: Bundaberg (9 specimens, AMS 
C.8970, FMNH 117331). 

Northern Territory: Palm Creek, Krichauff Range, 
near Hermannsburg (3 specimens, SAM D.3222); Sta- 
tion WA-130, Cycad Gorge, Palm Creek, Krichauff 
Range (7 specimens, FMNH 171558); Station WA-113, 
Glen Helen entrance to Finke Gorge, MacDonnell 
Ranges (1 specimen, FMNH 182104); Station WA-133, 
Temple Bar Gap, MacDonnell Ranges (1 specimen, 
FMNH 182119); Station WA-446, Kathleen Spring, 
Gill Range (3 specimens, FMNH 200476). 

Western Australia: Stations WA-303 and WA-305, 
0.5-1.0 km. east of Yammera Gap, Napier Range (2 
specimens, FMNH 199179, FMNH 199245); northwest 
of Stumpy's Well, southwest side Napier Range (1 
specimen, WAM 824-76); "The Tunnel," Tunnel 
Creek, Napier Range (1 specimen, WAM 1758-78); 
Stations WA-257 and WA-258, Brooking Gorge, Oscar 
Ranges (9 specimens, FMNH 199491, FMNH 199547); 
Station WA-255, near Gieke Gorge Ranger station, 
Gieke Gorge National Park, Oscar Ranges (3 speci- 
mens, FMNH 199597); Cave Spring, Bugle Gap, Law- 
ford Ranges (WAM). 

New Britain: Vunapope (12 specimens, ZMB, 
FMNH 146027); Malkong (3 specimens, ZMB). 

New Hebrides: Vila, Vate (3 specimens, BMNH 
1915.12.31.602, AMS C. 15679); Sarakata River, Es- 
piritu Santo (3 specimens, MCZ 186826, FMNH 54904, 
BPBM 212378). 

Kermadec Islands: Sunday Island (34 specimens, 
AIM, IRB, DMW 12449); under pumice on Mt. Junction 
(4 specimens, DMW 12424, collected September 18, 
1908, by W. R. B. Oliver); under stones on crater slope, 
northeast of Blue Lake (14 specimens, DMW 12423, 
FMNH 117214, collected October 3, 1908, by W. R. B. 
Oliver). 

Fiji: Viti Levu: Sigatoka, near base of limestone 
cliff, Vz-V/z miles inland (1 specimen, BPBM 87930, 
collected July 12, 1928, by H. S. Ladd); Lami Ridge 
(Station 47), 5 miles west of Suva at 5-150 ft. elevation 
(2 specimens, BPBM 178489, collected July 24, 1938, 
by Y. Kondo and C. M. Cooke, Jr.). 

Fiji: Lau Archipelago: Yangasa Levu, south end 
(Station 30), about 150 ft. inland at 40 ft. elevation (17 
specimens, BPBM 167137, collected July 26, 1934, by 
H. S. Ladd); Navutu-I-Loma (1 specimen, BPBM 
167102); northeast quarter (Station 28), 150 yd. in- 
land at 100 ft. elevation (3 specimens, BPBM 167017, 



collected July 24, 1934, by H. S. Ladd); Namuka (1 
specimen, BPBM 166668): Matandolo (Station 2), 
north-central point of island, 500 ft. inland at 65-75 ft. 
elevation (6 specimens, BPBM 166614, collected July 
7, 1934, by H. S. Ladd); Wangava (4 specimens, BPBM 
166994): northeast end (Station 27), V* mile inland at 
75 ft. elevation (3 specimens, BPBM 166965, collected 
July 22, 1934, by H. S. Ladd); Karoni (1 specimen, 
AIM, collected July 4, 1977, by Walter Cernohorsky): 
slope of peak at about 90 ft. elevation (6 specimens, 
BPBM 78603, BPBM 78606, collected August 15, 1924, 
by E. H. Bryan, Jr.); Aiwa: southwest quarter of island 
(Station 40), 50 yd. inland at 25 ft. elevation (1 speci- 
men, BPBM 167208, collected August 7, 1934, by H. S. 
Ladd); Nayau: Nauko (Station 43), Vz mile inland at 
base of limestone cliff, 250 ft. elevation (3 specimens, 
BPBM 167230, collected August 13, 1934, by H. S. 
Ladd); Vanua Mbalavu: between Valika and Mosomo 
Bay (Station 78), limestone hill about % mile inland at 
200-250 ft. elevation (4 specimens, BPBM 179582-3, 
collected August 9, 1938, by E. C. Zimmerman and Y. 
Kondo). 

Samoa: Aunuu Island off Tutuila: 10 yd. east of 
lighthouse, 200 yd. inland at 250 ft. elevation (4 speci- 
mens, BPBM 83243, collected February 4, 1926, by T. 
Dranga) (1 specimen, BPBM 171020). 

Ta'u: Siufaga (Station 76) (1 specimen, BPBM 
171074); Tavalagi Ridge (Station 75), Va mile inland at 
450 ft. elevation (27 specimens, BPBM 171066, BPBM 
171107); Lepuu Cliffs (4 specimens, BPBM 171126). 
Olosega: Olosega Village (1 specimen ex BPBM 
188720). 

Society Islands: Borabora, south slope (Station 
1093) of Pahio-Temanu ridge, 800 ft. elevation in dirt 
at cave entrance (1 specimen, BPBM 152396, collected 
October 13, 1934, by Gessler and St. John). 

Austral Islands: Rurutu: Mato Naa, bluff north of 
Moerai (Station 760), 20-30 yd. inland at 5-15 ft. ele- 
vation (6 specimens, BPBM 148292); Mato Arei (Sta- 
tion 775), cliff southeast of Moerai at 50-150 ft. eleva- 
tion (6 specimens, BPBM 148559, collected August 27, 
1934, by Y. Kondo and D. Anderson). 

Remarks. Except for collections from river drift 
or detritus sweepings, Discocharopa aperta (Mollen- 
dorff) is represented in collections by very small sam- 
ples. Partly this results from its small size, but more 
probably from the secretive habits and actual rarity. 
Iredale (1913, p. 366) reported that specimens were 
found under "a piece of black bark," later "on the 
underside of moss-covered stones deeply imbedded in 
earth on the side of the cliff," a "few . . . under dead 
nikau leaves," and "under loose dry pumice stones in 
the crater." He reported that "Only two or three were 
found at a time." No other data have been recorded 
concerning the habits of living specimens. 

At different times since 1961, I have been able to 
examine the types of all described species. Some were 
restudied a second time in order to check on details 
overlooked initially or to confirm structures. Despite 



SYSTEMATIC REVIEW 



79 



the variations outlined below, I conclude that only one 
species can be recognized. My list of material covers all 
the references cited above except for that of'Ptychodon 
celebica" from B. Rensch (1932, p. 101; 1935, p. 322). 
Broken or juvenile examples from these materials were 
seen in the SMF collection, but were not measured or 
listed. They are Discocharopa, however, and not 
Beilania philippinensis (Semper). Description of sev- 
eral species was caused by the wide geographic range, 
large shape variation, and the short diagnoses origi- 
nally presented. I am among the guilty, since Dis- 
cocharopa planulata Solem (1959a) from the New Heb- 
rides is based on rather large and widely umbilicated 
shells (fig. 36a-c). Previously (Solem, 1957, p. 4) I had 



suggested that the Philippine/), aperta and Indonesian 
D. microdiscus might be synonymous, but failed to ex- 
tend this to the other taxa. 

Data on size and shape variation are summarized 
in Table XIII. The growth pattern is such that certain 
separation of adult from subadult examples is very dif- 
ficult with fresh material and virtually impossible 
with worn, dead shells from river drift or detritus. Cal- 
culations have been restricted to means and ranges 
under these circumstances. Despite a certain partial 
mixture of subadults and adults, there is quite minor 
variation in size and shape. Specimens from the Bis- 
marck Archipelago are distinctly smaller and have 
more numerous, crowded radial ribs. Australian 



TABLE XIII. - GEOGRAPHIC VARIATION IN DISCOCHAROPA APERTA. 






NUMBER OF 
NAME SPECIMENS 


RIBS 


RIBS/MM. 


HEIGHT 


DIAMETER 


Montalban, Luzon, 


11 


102 


.0(99-105) 


23. 


7(23 


,2-24, 


0) 


0. 


58(0.49-0 


.66) 


1.38(1 


.27-1.53) 


Philippines 


Northern Luzon 


5 


104 


.7(100-110) 


24. 


6(24 


,0-25, 


4) 


0. 


59(0.53-0 


.66) 


1.31(1 


.15-1.38) 


Sumba and Celebes 


12 
25 


101 
100 


.4(78-123) 
.9(77-127) 


26. 

24. 


3(21 
4(18 


,8-31, 
,0-34. 


3) 
7) 


0. 

0. 


55(0.42-0 
56(0.48-0 


.69) 
.69) 


1.27(0 
1.34(1 


.99-1.63) 
.09-1.55) 


Timor 


Mew Britain. 


11 


133 


.0(118-148) 


35. 


3(30. 


,2-40. 


6) 


0. 


65(0.48-0 


.67) 


1.21(1 


.07-1.45) 


Bismarcks 


Krichauff Mts.. 


3 


88 


.3(84-91) 


19. 


7(19. 


2-20. 


8) 


0. 


59(0.58-0 


.61) 


1.43(1 


.38-1.51) 


N. Territories. 


Australia 


Bundaberg. 
Queensland, Australia 


5 


102 




21. 


9 






0. 


63(0.59-0 


.67) 


1.47(1 


.40-1.56) 


New Hebrides 


5 


127 


.3(125-131) 


29. 


5(29. 


2-29. 


6) 


0. 


54(0.46-0 


.63) 


1.36(1 


.25-1.43) 


Fill. Lau Group 


7 
3 
15 


135 
140 
114 


.7(125-147) 
.0(102-128) 


31. 
29. 
28. 


1(28, 
2 
0(25, 


5-35, 
4-30. 


7) 
5) 


0. 
0. 
0. 


64(0.53-0 
58(0.46-0 
64(0.49-0 


.74) 
.70) 
.72) 


1.56(1 
1.39(1 
1.32(1 


.32-1.84) 
.25-1.53) 
.07-1.45) 


Fiii. Viti Levu 


Samoa 


Rurutu. Austral Is. 


6 
1 
22 


109 
103 


.7(85-123) 


25. 
25. 


8 
1(20, 


9-30. 


6) 


0. 
0. 
0. 


70(0.63-0 
64 
57(0.49-0 


.79) 
.63) 


1.37(1 
1.43 
1.32(1 


.28-1.48) 
.23-1.41) 


Borabora. Society Is. 


Sunday Id. ., 
Kermadec Is. 



Monta. 
N. Luz. 


H/D 

0.418(0 
0.452(0 


RATIO 
.385-0 
.425-0 


.476) 
.476) 


WHORLS 
3 l/2-(3 1/4-3 3/4) 
3 3/8(3 1/8-3 1/2) 


UMBILICUS 
0.56(0.53-0.65) 
0.52(0.43-0.58) 


n/u 

2.57(2 
2.53(2 


RATIO 
.40-2.96) 
,40-2.69) 


BODY WHORL 
WIDTH SP/BWW 





Sumba 
Timor 
N. Bri. 
Krich. 
Bunda. 
N. Heb. 


0.430(0 
0.419(0 
0.444(0 
0.409(0 
0.428(0 
0.400(0 


.396-0 
. 382-0 

.402-0 
.400-0 
.404-0 
.341-0 


.487) 
.488) 
.466) 
.417) 
.445) 
.437) 


3 l/2-(3 1/8-4) 
3 1/2-O-3 7/8) 
3 3/8+(3 1/4-3 5/8) 
3 1/8(3-3 1/4) 
3 l/2+(3 1/2-3 3/4) 
3 1/2(3 1/4-3 5/8) 


0.53(0. 
0.53(0. 
0.49(0. 
0.54(0. 
0.55(0. 
0.60(0. 


39-0.72) 
39-0.61) 
43-0.59) 
51-0.56) 
51-0.64) 
53-0.66) 


2.39(2 
2.57(2 
2.48(2 
2.66(2 
2.68(2, 
2.29(2, 


,22-2 
,36-2 
,38-2 
,57-2 
,44-2 
,16-2 


.63) 
.77) 
.58) 
.71) 
.81) 
.63) 











0.38 0.130 


Lau 


0.410(0 


. 390-0 


.450) 


3 3/4+(3 


1/2-4 1/4) 


0.72(0. 


48-0.92) 


2.21(2, 


00-2 


.76) 


0.40(0.35-0.44) 0.178(0.111-0.292) 


V. Levu 


0.418(0 


. 368-0 


.457) 


3 3/8+(3 


1/4-3 5/8) 


0.46(0.44-0.48) 


3.04(2. 


81-3 


.21) 





Samoa 


0.486(0 


.429-0 


.524) 


3 l/2-(3 


1/4-3 5/8) 


0.47(0. 


39-0.61) 


2.80(2. 


59-3 


.09) 


0.39(0.34-0.41 0.171(0.091-0.286) 


Rurutu 


0.514(0 


.444-0 


.615) 


3 5/8(3 


3/8-4) 


0.51(0. 


41-0.59) 


2.71(2. 


50-3 


.12) 


0.41(0.38-0.44) 0.208(0.146-0.240) 


Borab. 


0.448 






3 1/2+ 




0.61 




2.35 






0.41 0.120 


Sunday 


0.434(0 


. 390-0 


.469) 


3 l/4+(3-3 1/2) 


0.48(0. 


46-0.56) 


2.57(2. 


42-2 


.68) 






80 



SOLEM: ENDODONTOID LAND SNAILS 




*** ^^ 





FIG. 36. a-c, Discocharopa aperta (Mollendorff) from the New 
Hebrides. River drift, Sarakata River, Espiritu Santo, New Hebrides. 
Holotype of Discocharopa planulata Solem, 1959. UMMZ 186037. 
Scale line equals 1 mm. (Solem, 1959a, pi. 32, figs. 1-3). (EJP). 

specimens are large and have fewer and more widely 
spaced radial ribs (fig. 37a-d). Lau Archipelago shells 
are very large and widely umbilicated, with the New 
Hebridean shells only slightly smaller (fig. 36a-c). 
Both have intermediate ribbing frequency and spacing. 



Shells from Viti Levu and Samoa are much more nar- 
rowly umbilicated. The higher H/D ratio and narrower 
umbilicus in the Rurutu shells results from drastic de- 
flection of the body whorl. Otherwise, differences are 
well within the range of statistical error. 

Probably more importance can be attached to var- 
iation in the parietal barrier. Kermadec shells (fig. 
35b) have a very large barrier recessed about one- 
quarter whorl and usually visible through the shell, 
but not from the aperture. Rurutu examples have a 
slightly smaller parietal that was clearly visible from 
the aperture, as did the single example from the Soci- 
ety Islands. Specimens from the Lau Archipelago had a 
parietal visible in juveniles, sometimes large and 
sometimes small, but not visible from the aperture in 
any adults. No Lau Archipelago adults had a translu- 
cent shell, but unless there was deeper recession of the 
barrier in adults, it should have been visible. No 
Australian, Samoan, Viti Levu, or New Hebridean 
shells showed any trace of a parietal. 

A number of Samoan shells were sufficiently 
translucent to allow confirmation of the parietal's ab- 
sence. Some Philippine specimens had a low to medium 
parietal, but most showed no trace of a barrier. Most 
Sumba examples had a large parietal, whereas the 
Celebes shells (fig. 35a) showed a moderately recessed, 
rather long barrier that at most extended one-third of 
the apertural width. 

It is obvious from the above paragraphs that varia- 
tions show a mosaic pattern and that the variables are 
not correlated in any obvious fashion. Widely umbili- 
cated Celebes shells have a strong parietal, whereas 
the widely umbilicated New Hebridean specimens lack 
all trace of a parietal, and the Lau Archipelago exam- 
ples have a parietal in the juvenile stage but not the 
adult. Viti Levu examples are narrowly umbilicated 
and without a parietal barrier. 

Material from the island groups covered in this 
monograph were relatively distinct from each other, 
but when specimens from extralimital areas were con- 
sidered, the differences were duplicated. Only one 
example from Borabora (BPBM 152396) was seen. Al- 
though broken and with the sculpture worn, the large 
parietal, relatively wide umbilicus and rather low 
spire are very different from the Rurutu shells. These 
have the spire markedly elevated and a correlated 
rather tight coiling pattern. In some (BPBM 148559) 
the body whorl is abruptly deflected, which greatly in- 
creases the H/D ratio and narrows the umbilicus. All 
examples had a parietal visible from the aperture. 

Samoan specimens have a raised spire, slightly in- 
creased descension of the body whorl, a relatively nar- 
row umbilicus, fairly high H/D ratio, no visible 
parietal, and, in the few translucent shells, no parietal 
visible through the shell. A subjective impression is 
that they have a slightly tighter coiling pattern than 
do the Fiji specimens. 

Lau Archipelago shells, particularly the largest 
sample (BPBM 167137) from Yangasa Levu, are very 
widely umbilicated, large shells, with a slight increase 



SYSTEMATIC REVIEW 



81 




FIG. 37. Australian Discocharopa aperta (MbllendorfD. Station WA-113, Glen Helen opening to Finke Gorge, MacDonnell Range, west of 
Alice Springs, Northern Territory. A. Solem and L. Price! 111-12-1974. FMNH 182104: a, top of shell (50x); b, side of shell (54x ); c, base of shell 
(49x); d, sculpture on body whorl (505x). 



in whorl count and distinctly more crowded radial ribs 
on the last portion of the body whorl. Visually they are 
very similar to the New Hebridean shells. All juveniles 
showed a large parietal barrier that was readily visible 
from the aperture. No adults showed any trace of a 
parietal. The few Viti Levu shells (BPBM 87930, 
BPBM 178489) are much more narrowly umbilicated 
and are shaped like the Samoan shells. They lack any 
trace of a parietal, although at least one is obviously 
subadult. 

Specimens from Rurutu, Samoa, and the Lau Ar- 
chipelago do have a characteristic appearance and if 
mixed in a tray could be separated with only slight 
difficulty. When compared with various Indonesian 
and Bismarck sets, except for the few Rurutu shells 
with extreme body whorl deflection, duplicates can be 
found for each of the Pacific Island types. Nomen- 
clatural recognition would require designation of con- 
vergent, isolated geographic races based on use of one 
or two characters and ignoring those that disagreed, or 



recognizing probably polyphyletic subspecies with in- 
termingled mosaic distributions. Neither alternative 
seems really adequate, particularly since only material 
from one population has been dissected. Eventual rec- 
ognition of races or species may be possible, but not 
until anatomical studies have been made and more 
data are available concerning the meaning of variation 
in the parietal barrier. 

Genus Sinployea, new genus 

Small to very large Charopidae with SV-j-S'/i (more in plano- 
spira) normally to rather tightly coiled whorls. Apex and spire rarely 
flat, slightly depressed (lamellicosta, tenuicostata) or barely protrud- 
ing (descendens), normally moderately protruded, sometimes ele- 
vated more than !4 body whorl width (harueyensix, vicaria, angularis, 
recursa). Body whorl descending moderately to drastically (ir- 
regularis). Apical sculpture normally of 10-12 prominent to fine spi- 
ral cords, rarely averaging less than 9 (tahitiensis, allecta) or more 
than 13 (angularis, recursa, irregularis), sometimes (peasei, av- 
anaensis) greatly reduced in prominence. Postnuclear sculpture of 
extremely fine and crowded (avanaensis) to broader and quite widely 
spaced (angularis, tenuicostata), usually protractively sinuated ra- 



82 



SOLEM: ENDODONTOID LAND SNAILS 



dial ribs, rarely (recursa) lost on lower spire and body whorl, some- 
times becoming too fine and crowded (harveyensis, irregularis) or 
irregular (rudis) to count. Microsculpture of fine radial and finer 
spiral riblets, secondary spiral cording present in % of species. Um- 
bilicus sometimes widely open (descendens, planospira), rarely 
(clinta, clausa) closed or barely perforate, normally V-shaped or U- 
shaped, with last whorl decoiling more rapidly, margins rounded. 
Shell color flammulated, light horn or reddish brown. Sutures deep, 
channeled only in proximo, whorls evenly rounded to strongly com- 
pressed laterally above periphery, rarely (canalis) with strong lat- 
eral compression, periphery rounded, angulated only in angularis. 
Aperture circular to compressedly ovate, inclined 5-35 from shell 
axis. Body color white or with gray on head and ommatophores. 
Pallial cavity with lobes of kidney subequal in length, generally a 
strip of lung roof visible between arms of ureter. Genitalia same type 
as in Champa. Penis length 0.30-0.53 shell diameter, proportion- 
ately larger only in tahitiensis and montana, internally with vergic 
papilla, muscular collar just below verge tip, and large stimulatory 
pad near middle portion. Relative size of the penial structures vari- 
able. 

Type species. Sinployea peasei, new species. 

Of the other Polynesian genera with typical 
stimulatory pad, the Viti Levu Ba has altered shell 
form (fig. 74a-c), pallial structures (fig. 75a), and is an 
experiment in whorl count reduction. The Tongan 
Tuimalila is much larger (table XXXIX), has looser 
whorl coiling (fig. 78b, e), altered sculpture, and is 
much larger. Maafu (fig. 76a-c) has not been dissected, 
but its grossly altered shell sculpture and form im- 
mediately distinguish it from any Sinployea. All of the 
above groups appear to be specialized derivatives from 
Sinployea. In contrast, the genera with apertural bar- 
riers, Lauopa, Vatusila, Lagiuala, and Graeffedon, 
probably are much more distantly related. Graeffedon 
(fig. 87c) has a very distinctive epiphallic structure. 
Only a partial specimen of Vatusila tongensis could be 
dissected (fig. 84a-b). It seems closely related to 
Sinployea. 

Intraspecific variation is large enough and 
similarities between species sufficiently numerous 
that preparation of a key to all species would have 
required multiple entries. The more practical solution 
of preparing keys to taxa from limited segments of the 
generic range has been adopted. Discussions and either 
geographic or formal dichotomous keys have been pre- 
pared for the species from Society; Cook; Samoan clus- 
ter plus Swains Island; Tonga, Ellice, Rotuma, Kusaie 
in the Carolines, and Saipan in the Marianas; Fiji; and 
Melanesia (New Hebrides, Solomon Islands, Bismarck 
Archipelago). In the individual species discussions, 
cross-references are made to morphologically similar 
forms from other areas, but the main emphasis is on 
differentiating taxa from their sympatric or neighbor- 
ing species. 

Some general discussion of variation patterns is 
presented here, although Sinployea is a relatively 
uniform group in terms of structure. 

Sinployea has the widest geographic range and 
greatest number of species recorded for any Polynesian 
endodontoid genus. I discuss 49 named species-level 
units below and mention two additional taxa rep- 
resented by material that is inadequate for description. 



Unquestionably, additional species remain to be dis- 
covered. Further collections in Tonga, Fiji, Melanesia, 
and probably on Tahiti should be especially productive. 
Dissection has confirmed placement of the New Hebri- 
dean S. euryomphala and the Caroline Island S. 
kusaieana in this genus, but inclusion of the Solomon 
Island and Bismarck Archipelago species might not be 
correct. None of them have been dissected. A record 
from Saipan in the Marianas is questionable. It is pos- 
sible that Sinployea occurs in New Guinea or even 
Wallacea, since Pilsbrycharopa kobelti (Bottger) and P. 
brunnescens (Mollendorff) might be classified in 
Sinployea. Without dissection, their position will re- 
main uncertain. Similarly, some of the New Caledo- 
nian species listed in Andrefrancia by Solem (1961, pp. 
454-463) possibly might be Sinployea, although I con- 
sider this unlikely, since several dissected Andrefran- 
cia have distinctive anatomy. Determination of the 
exact western distributional limits is beyond the scope 
of this study and would require collection of additional 
preserved material. 

The great Polynesian distribution of Sinployea 
probably can be explained on the basis of ecology. 
Many species are semiarboreal in habitat, ascending 
trees to at least 15 ft., and specimens have been found 
frequently in leaf axils or attached to loose bark on stilt 
roots. The opportunity for rare accidental transport 
during a severe storm is infinitely greater for a species 
on loose bark than for one in the dense ground litter. 
Correlated with this habitat difference is the develop- 
ment of gray markings on the ommatophores, head, 
neck, and sometimes mantle edge in those species that 
have been found on trees. No trace of gray coloration is 
seen on those species living only in ground litter or 
under logs and stones. All of the Society Islands and 
Cook Island species collected alive in this century show 
the darker markings. The condition in many Raroton- 
gan species collected on the ground by Garrett, but not 
found since the 1880s, is unknown. Of the Samoan 
species for which soft parts are known, only the Swains 
Island S. intermedia lacks gray coloration. The atoll 
environment with its thinner vegetation cover may 
have caused this species to become a litter dweller. In 
contrast, the Caroline Island S. kusaieana, Fijian S. 
inermis, S. adposita, and S. irregularis plus the New 
Hebridean S. euryomphala lack all traces of gray col- 
oration and have been recorded only from the ground 
stratum. Two Lau Archipelago species in particular 
may be semiarboreal, but they are known only from 
empty shells (S. recursa, S. angularis). 

Presently available data suggest that there are 
two main centers of speciation, Lau Archipelago (six 
species) and Rarotonga (11 species). In both areas there 
is evidence of a radiation encompassing both semiar- 
boreal and ground stratum niches. It is quite possible 
(p. 97) that the Rarotonga litter species are extinct, but 
I suspect that the Lau Archipelago habitats may be 
less disturbed. Secondary centers of speciation are 
found in Samoa (five species) and Tahiti (six species). 



SYSTEMATIC REVIEW 



83 



Elsewhere, only single species on several islands 
(Tonga) or separate species on individual islands (Sol- 
omons) have been recorded. On Tahiti, only one species 
has been collected at a station, and on Rarotonga, 
fragmentary data from recent collecting and records in 
Garrett (1881) suggest that most, if not all, of the 11 
Sinployea species were geographically isolated. In 
Samoa and the Lau Archipelago there is ample evi- 
dence of rather extensive sympatry. The above specia- 
tion pattern suggests a group of relatively recent arri- 
val that is well adapted for rare accidental transport 
over great distances, but on particular islands tends to 
break up into geographic isolates. 

In several cases it was possible to dissect sympat- 
ric species. In other situations I could study allopatric 
species populations from only a few hundred yards 
apart. In each situation there have been shifts in size 
and/or relative development of the functioning penial 
elements (verge, muscle collar, and stimulatory pad) 
that probably serve to enable species recognition and to 
maintain species isolation. Details of variation in each 
situation are given in the discussion of Tahitian 
species (p. 85), Cook Island (p. 95), Samoan (p. 118), 
and Fijian (p. 147) taxa. Except for these situations of 
sympatry or almost certain contact between species, I 
found only slight anatomical differentiation. Varia- 
tions in whorl count and body whorl capacity affect the 
relative proportions of the kidney and whether or not a 
strip of lung roof is visible between the ureter arms. 
Neither available time nor available material was suf- 
ficient to make detailed comparisons of genital propor- 
tions. No obvious differences were noted. Although 
penis length ranged from 0.8-2.2 mm., when allowance 
is made for differences in adult size by calculating the 
ratio of mean penis length/mean shell diameter of dis- 
sected specimens (table XIV), obviously there is rela- 
tively minor variation. Fourteen of 21 dissected 
Sinployea have a PL/XD ratio between 0.35 and 0.47. 1 
consider it significant that departures from this pat- 
tern are shown on the opposite ends of distribution. 
The New Hebridean S. euryomphala (Solem), Fijian S. 
irregularis (Garrett), and Society Island S. neglecta 
have small penes, with a ratio of 0.30-0.31. In contrast, 
two Tahitian species found in close proximity have 
proportionately very large penes, with ratios of 0.61 
and 0.66. Selection pressure for change would occur 
only where species came into contact or were sympat- 
ric, so that the stable genital pattern is not unexpected. 
The apparent correlation between white body color- 
litter dweller, gray head and ommatophores-tree trunk 
habitat was mentioned above. 

Conchological patterns of variation are somewhat 
more extensive, but appear mosaic in nature rather 
than geographic. Although the range of variation in 
most measured characters is rather large, the average 
pattern is relatively conservative. Table XV sum- 
marizes mean measurements, giving the minimum, 1st 
quartile, median, 3rd quartile, and maximum means 
for 50 taxa (all described material and the Borabora 



TABLE XIV. - RATIO OF PENIS LENGTH TO SHELL DIAMETER IN 
SINPLOYEA. 



Species 


Penis length 
in mm. 


X Shell 
Diameter 


PL/D 


tahitiensis 


2. 
1. 


2-2. 
4-1. 


4 
5 


3.50 
3.25 


0.66 
0.45 


lamellicosta 


modicella 


1. 

1. 


0-1. 
4-1. 


1 
6 


3.35 
2.83 


0.31 
0.53 


neglecta 


montana 


1. 


4-1. 


65 


2.50 


0.61 


atiensis 


1. 

1. 


1-1. 
1-1. 


3 
3 


2.70 
2.95 


0.44 
0.41 


andrewi 


peasei 


1. 


1-1. 


3 


3.15 


0.38 


avanaensis 


1. 


4-1. 
1.2 


6 


3.21 
2.53 


0.47 
0.47 


clista 


aunuuana 




1.0 




2.76 


0.36 


allecta allecta 


1.3-1. 
1.2-1. 
1.9-2. 


5 
6 
2 


3.13 
3.49 
4.92 


0.45 
0.40 
0.42 


intermedia 


complementaria 


vicaria vicaria 




1.3 
1.0 




2.86 
2.60 


0.45 
0.38 


kusaieana 


inermis inermis 




1 


,8-1. 
,8-1. 
.65 






2.60 
2.36 
3.29 


0.35 
0.38 
0.50 


i . lakembana 


adposita 


irregularis 





.8-1. 
0.8 





2.87 
2.64 


0.31 
0.30 


euryomphala 



adult). For the primary measurements of height, 
diameter, whorls, and rib count, the 2nd and 3rd quar- 
tiles cover from 12%-30% of the total range. Only a few 
species are strikingly distinctive in size or shape, most 
are very similar. 

Four of the seven smallest species were from the 
Solomons and Bismarcks, S. nissani, S. kuntzi, S. de- 
scendens, and S. novopommerana. The other three 
were S. ellicensis nukulaelaeana from Nukulaelae, El- 



TABLE XV. - MEAN SPECIES MEASUREMENTS IN SINPLOYEA. 



Shell Height In mm. 
Shell Diameter in mm 
H/D f.atio 
Whorl Count 

Diameter /Umbilical 
Width Ratio 

Ribs on Body Whorl 1 



MINIMUM 
1.00 
2.08 
0.463 
3 1/2 

2.73 
38.0 



1ST QUARTILE MEDIAN 3RD QUARTILE MAXIMUM 



Ribs/mm, on Body Whorl 1 3.35 
Apical Cords 8.60 



1.38 
2.50 
0.505 
3 7/8+ 

3.69 
77.1 
8.04 
9.40 



1.55 
2.84 
0.538 
4 1/8- 

4.29 
92.5 
10.8 
10.2 



1.77 
3.35 
0.573 
4 3/8- 

4.99 
112.2 
12.2 
11.8 



2.55 
4.92 
0.625 
6 5/8 

CLOSED 
225.8 
21.0 
18.5 



1. Four species have reduced ribbing and were omitted 
from this table. 



84 



SOLEM: ENDODONTOID LAND SNAILS 



lice Islands, S. godeffroyana from Viti Levu, and the 
very modified S. angularis from Namuka, Lau Ar- 
chipelago. The largest species is the Upolu Island S. 
complementaria, with nine of the 11 Rarotongan 
species comprising the other species averaging more 
than 3.60 mm. in mean diameter. Concentration of the 
smaller species in Melanesia may indicate competition 
from terrestrial helicarionids in the 2.5-3.5 mm. size 
range, whereas the development of larger Sinployea on 
Rarotonga may be the result of that island lacking ter- 
restrial helicarionids. Because the Rarotongan radia- 
tion of Sinployea probably is extinct, fieldwork to check 
these possibilities may be 50 years too late. 

Height is closely correlated with diameter in terms 
of large and small species, although H/D ratio diverges 
in special circumstances. The great increase in whorl 
count found in S. planospira (fig. 46d) has resulted in a 
lowered H/D ratio, whereas the flat spire, wide um- 
bilicus, and slightly increased whorl count in S. canalis 
(fig. 49a-c) combine to reduce the H/D ratio. In con- 
trast, the nearly closed umbilicus, strongly elevated 
spire, and thicker body whorl of the Samoan S. clausa 
(fig. 51d-f) give it the highest H/D ratio in the genus. 
Only the Tahitian S. montana, with a slightly more 
open umbilicus, equally protruded spire, but more 
sharply deflected body whorl, is similar in H/D ratio 
(fig. 41a-c). The Viti Levu S. irregularis has a flat or 
only slightly elevated spire, but a high H/D ratio is 
produced solely by an often drastic deflection of most or 
all of the body whorl (fig. 65d-f). 

Whorl count is a much less variable character. Ig- 
noring S. nissani and S. montana, for which only sub- 
adult or barely adult material was available, and S. 
planospira (fig. 46d), which is characterized by an ab- 
normally high whorl count, mean whorls range from 
3%- to 5-. Of these, only five of the very largest 
species average more than 4 l /2 whorls. There are no 
species I can cite as being experiments in whorl count 
reduction, whereas only S. planospira shows clear evi- 
dence of whorl increase. Its mean whorl count of 6% 
includes a juvenile shell and is thus quite conservative, 
yet this species averages 1% whorls more than any 
other Sinployea. 

Relative umbilical width, as measured by the D/U 
ratio, is determined by the pattern of coiling, number 
of whorls, body whorl contour, and body whorl descen- 
sion. Those with wide, cup-shaped umbilici are found 
in the Bismarcks (S. novopommerana, fig. 73c), 
Rarotonga (S. canalis, fig. 49c; S. planospira, fig. 46f), 
Ellice Islands (S. pseudovicaria, fig. 61c), whereas the 
very narrowly umbilicated forms are concentrated in 
Samoa (all the small species) and Lau (S. angularis, 
fig. 64f; S. recursa, fig. 64c; S. adposita, fig. 69f). I do 
not know the reason for this. 

Whorl contour is relatively stable, with the chan- 
neled suture of S. proxima (fig. 46a), strong lateral 
compression in S. canalis (fig. 49b), weak supra- 
peripheral sulcus in S. rudis (fig. 47e) and S. ir- 
regularis (fig. 65e), and angulated periphery in S. an- 



gularis (fig. 64e) providing the only major alterations. 
Most species vary in the narrow range from an evenly 
rounded periphery, as in S. ellicensis (fig. 62b) and S. 
vicaria (fig. 58b), to taxa with rather prominent lateral 
compression above the rounded periphery, such as S. 
recursa (fig. 64b). When specimens are compared di- 
rectly, such minor differences are obvious, but no 
quantification of these changes was attempted. 

Shell ornamentation is relatively complex and the 
variations numerous. In several species no counts of 
major radial ribs were possible. The Rarotongan S. 
rudis (fig. 47d) and the Lau Archipelago S. recursa (fig. 
64a) have widely spaced major radial ribs on the upper 
spire that become reduced to absent by the body whorl. 
In contrast, the Rarotongan S. harveyensis (fig. 48a-b) 
and Viti Levu S. irregularis (fig. 65d-e) have the ma- 
jor ribs become so fine and crowded on the body whorl 
that differentiation of ribs and growth irregularities 
becomes impossible. Specimens with few radial ribs in- 
clude the very small S. angularis (fig. 64e) and S. in- 
ermis lakembana (fig. 66e), the very large S. tenuicos- 
tata (fig. 50d), and the larger than average S. adposita 
(fig. 69e) and S. lamellicosta (fig. 40a). By far the 
greatest number of ribs, mean 225.8, is found in S. 
auanaensis (fig. 45e) from Rarotonga. Of the smaller- to 
medium-sized species, only the Fijian S. godeffroyana 
(fig. 69a) and Lau Archipelago S. navutuensis (fig. 68e) 
have actual rib counts equaling some of the larger 
Rarotongan shells. The latter three species have by far 
the highest ribs/mm, index, whereas S. adposita and S. 
tenuicostata have the most widely spaced ribs. The 
number of microradial riblets varies with the rib spac- 
ing and actual sculpture size. Development or loss of 
secondary spiral cording and relative spacing of the 
cording showed no consistent pattern that I could re- 
late to size or major sculpture. 

Apical cording is variable in prominence; the con- 
trast between the large cords inS. modicella (fig. la-b) 
and the faint ones in S. peasei (fig. 2a-b) indicate the 
extent of variation. Actual numbers of cords vary from 
a mean of 8.6 in the Samoan S. allecta allecta to a high 
of 18.5 in the Viti Levu S. irregularis. There is no sim- 
ple correlation between the number of cords and their 
relative prominence, because the cord spacing is an 
additional variable factor. It was not practical to pre- 
pare scanning electron microscope photographs of more 
than a few species. The apical cording is too fine for 
accurate measurement with a dissecting microscope, 
and the relief is too great for effective study with a 
compound microscope. Thus, no detailed analysis of 
apical sculpture spacing and cord width was possible. 
Those species averaging 8.6-9.0 apical cords were S. 
allecta allecta from Samoa, S. tahitiensis from Tahiti, 
S. decorticata from Rarotonga, S. inermis from Lau, 
and S. princei from Taveuni. Those with an enlarged 
number of apical cords are clustered geographically. 
The Society Island S. montana, S. lamellicosta, and S. 
neglecta average 12.5-13.0 cords, whereas the Lau Ar- 
chipelago S. adposita, S. inermis lakembana, S. an- 



SYSTEMATIC REVIEW 



85 



gularis, and S. recursa average 13.0-15.9 apical cords. 
The presence of both low and high counts in Lau and 
Fiji emphasizes the complexity of speciation patterns 
in that area. 

SOCIETY ISLAND Sinployea 

It is rather surprising that no species are known 
from Tahaa, Raiatea, or Maupiti. Suitable habitats 
were investigated by both Garrett and members of the 
Mangarevan Expedition. Only one specimen, too worn 
to merit description, has been found on Borabora. Pos- 
sibly Sinployea does not live on Maupiti or the Tahaa- 
Raiatea complex. Because the Society Islands repre- 
sent the eastern limit of distribution for Sinployea, 
patchiness of occurrence would not be unexpected. 

Single species are found on Huahine and Moorea; 
three species, on Tahiti. Because upland collecting on 
Tahiti has been limited to two partial transects on Mt. 
Aorai and Mt. Orofena, additional taxa may be found 
later. Sinployea neglecta (fig. 41d-f) from Huahine 
comes the closest of any species in the genus to being 
average in characteristics. Its mean measurements 
(table XVI) depart from the median Sinployea figures 
(table XV) only in having a few more crowded major 
radial ribs, more spiral apical cords, and a slightly re- 
duced number of microradial riblets. Sinployea mod- 
icella (fig. 38a-c) from Moorea is much larger, more 
widely umbilicated, and averages fewer apical cords. 
There is also a difference in penial size, with S. neg- 
lecta having a penis length of 1.4- 1.6 mm. (53% of shell 
diameter, which averages 2.83 mm.) and S. modicella 
having a penis length of 1.0-1.1 mm. (31.4% of shell 
diameter, which averages 3.35 mm.). 

All three Tahitian species were collected by the 
Mangarevan Expedition during the Mt. Aorai transect 



of the malacological assistants. None were taken dur- 
ing the Mt. Orofena journey by the expedition bot- 
anists and entomologist. Sinployea tahitiensis occurred 
in sparse numbers at several stations from 1,400-6,300 
ft. elevation and in Fautaua Valley at 250 m. eleva- 
tion. Sinployea montana was found at Station 866, a 
valley to the west of the Aorai Trail at 6,000 ft. eleva- 
tion; S. lamellicosta , at Station 870, a valley west of the 
Aorai Trail at 5,000 ft. elevation. Possibly the latter 
two were collected in the same valley, but obviously at 
different elevations. 

Sinployea montana (fig. 41a-c) is a very small, 
elevated species with quite narrow umbilicus and 
rather widely spaced radial ribs. Sinployea lamellicosta 
(fig. 40a-c) has a flat or slightly depressed spire, many 
apical spiral cords, a wide umbilicus, and very widely 
spaced radial ribs. Sinployea tahitiensis (fig. 38d-f) is 
fairly large, has a slightly protruded spire, fewer apical 
cords, more whorls, and a thicker body whorl. Penial 
differences are marked. Sinployea tahitiensis has a 
penis length of 2.2-2.4 mm. (65.8% of shell diameter), 
and the penis shows no particular modifications. In S. 
montana the penis length of 1.4-1.65 mm. is only 61% 
of shell diameter, and the vergic papilla is very large 
with the muscular collar reduced in size. Sinployea 
lamellicosta has a penis length of 1.4-1.5 mm., or 
44.6% of shell diameter, and the stimulatory pad is 
greatly enlarged (fig. 39h). Thus S. tahitiensis is 
differentiated by penis size, whereas S. lamellicosta 
and S. montana have penes of the same size, but in the 
former there is enlargement of the vergic papilla and 
in the latter, enlargement of the stimulatory pad. If the 
latter two species do have a zone of sympatry, probably 
the changes in penial structure serve to enforce species 
isolation. Further collecting is needed in the moun- 
tains of Tahiti in order to establish the ecological re- 



TABLE XVI. - RANGE OF VARIATION IN SOCIETY ISLAND SINPLOYEA. 



M AMP BUCWWiB ur 

SPECIMENS 
modicella 399 


RIBS RIBS/MM. 
112.9(89-145) 10.82(8.59-12.21) 

60.7(44-71) 5.86(4.67-8.37) 
52.4(43-71) 5.44(4.03-7.91) 


(Ferussac) 
tahitiensis 23 


lamellicosta 19 


(Garrett) 



Borabora 
sp. 



3 


63.0(57-69) 


8.04(7.43-8.65) 


172 


108.6(71-143) 


12.23(8.84-16.87) 


1 


126 


14.14 



HEIGHT DIAMETER 

1.72(1.35-2.04) 3.35(2.99-3.68) 

1.75(1.39-2.09) 3.50(2.98-4.04) 

1.58(1.35-1.74) 3.25(2.78-3.55) 

1.56(1.52-1.59) 2.50(2.45-2.55) 

1.51(1.28-1.81) 2.83(2.57-3.26) 

1.55 2.86 



H/D RATIO 
0.514(0.406-0.598) 

0.498(0.428-0.541) 
0.487(0.454-0.521) 

0.623(0.622-0.623) 
0.536(0.441-0.632) 
0.540 



APICAL 
CORDS 



SPIRE 
ELEVATION 



BODY WHORL 
WIDTH 



WHORLS UMBILICUS D/U RATIO CORDS ELEVATION UTnTH SP/BWW 

modic. 4 l/4+(4-4 5/8) 0.96(0.72-1.22) 3.48(2.95-4.28) 9.20(7-13) 0.17(0.07-0.23) 1.01(0.89-1.12) 0.166(0.074-0.226) 

tahit. 4 3/8-(4-4 5/8) 0.95(0.66-1.12) 3.68(3.38-4.50) 8.92(6-11) 0.10(0.07-0.13) 1.02(0.95-1.05) 0.104(0.091-0.133) 

lamel. 4+(3 3/4-4 3/8) 0.99(0.74-1.22) 3.26(2.89-3.87) 12.50(10-16) FLAT 0.86(0.72-0.99) FLAT OR DEPRESSED 

monta. 3 5/8+(3 5/8-3 3/4) 0.36(0.30-0.43) 7.24(5.93-8.55) 13 0.15(0.13-0.16) 0.85(0.84-0.86) 0.226(0.220-0.231) 

negle. 4 (3 5/8-4 1/4) 0.65(0.53-0.95) 4.39(3.21-5.00) 12.46(10-15) 0.14(0.10-0.23) 0.78(0.72-0.86) 0.180(0.115-0.275) 

Borab. 4 0.69 4.14 WORN 0.23 0.89 0.222 



86 



SOLEM: ENDODONTOID LAND SNAILS 



lationships and geographical ranges of these species. 
Despite the size and shape differences, the Society 
Island species share a characteristic appearance of 
very thin major ribs with periostracal lamellar exten- 
sions. Most species from other island groups have dis- 
tinctively lower and broader radial ribs. Secondary spi- 
ral cording is absent in S. montana, present only on the 
spire in S. modicella, and moderately to strongly de- 
veloped in the other species. Typical flammulated color 
patterning is seen in all but S. montana and S. 
tahitiensis, which have monochrome shell color. All 
species show a light to dark gray coloration on the head 
and neck. Sinployea modicella and S. tahitiensis have 
been collected on tree trunks or in axils of low trees, 
plus the usual log and under stone habitats. Sinployea 
neglecta was found only in the ground stratum, but at 
these stations no special investigation was made of ar- 
boreal habitats. 

Because most species are geographically isolated, 
no formal key has been prepared. The following synop- 
sis provides quick identification of known taxa: 
Moorea: S. modicella (Ferussac, 1840) 
Huahine: S. neglecta, new species 
Borabora: Sinployea sp. (probably new) 
Tahiti: S. montana, new species (diameter 
less than 2.75 mm., D/U ratio 
more than 5.00, H/D ratio more 
than 0.600) 

S. tahitiensis, new species (spire ele- 
vated, about 9 spiral apical cords, 
H/D ratio less than 0.550) 
S. lamellicosta (Garrett, 1884) (spire 
flat or depressed, about 12.5 spiral 
apical cords, H/D ratio less than 
0.550) 
Discovery of additional species is very probable. 

Sinployea modicella (Ferussac, in Deshayes, 1840), 
Figures la-e, 38a-c, 39a-b. 

Helix modicella Ferussac, 1840, Hist. Nat. Moll. terr. fluv., 1, pi. 
86, fig. 3; Deshayes, 1850, Hist. Nat. Moll. terr. fluv., 1, pp. 
90-91 "sur les hautes montagnes, Taiti" ( = Tahiti, Society Is- 
lands); Pfeiffer, 1853, Monog. helic. viv., 3, p. 92; Pfeiffer, 1859, 
Monog. helic. viv., 4, p. 86; Pfeiffer, 1876, Monog. helic. viv., 7, p. 
149. 

Pitys modicella (Ferussac), Pease, 1871, Proc. Zool. Soc. London, 
1871, p. 474. 

Patula modicella (Ferussac), Schmeltz, 1874, Cat. Mus. Godeffroy, 
5, p. 93 Huahine (?), Society Islands; Garrett, 1884, J. Acad. 
Nat. Sci., Philadelphia, 9 (1), pp. 28-29 partly. 

Helix (Patula) modicella Ferussac, Tryon, 1887, Man. Conchol., (2) 
3, p. 38, pi. 8, figs. 28-30. 

Endodonta (Charopa) modicella (Ferussac), Pilsbry, 1893, Man. 
Conchol., (2) 9, p. 35. 

Diagnosis. Shell larger than average, diameter 2.99-3.68 mm. 
(mean 3.35 mm.), with 4-4% normally coiled whorls. Apex and spire 
slightly to moderately and evenly elevated, last whorl usually de- 
scending more to sharply more rapidly, spire protrusion about '& 
body whorl width, H/D ratio 0.406-0.598 (mean 0.514). Apical 
sculpture of 7-13 (mean 9.2) prominent spiral cords. Postnuclear 
sculpture of narrow, prominent, numerous, protractively sinuated 
radial ribs, 89-145 (mean 112.9) on the body whorl, whose interstices 



are 2-5 times their width. Ribs/mm. 8.59-12.21 (mean 10.82). Mi- 
crosculpture of fine radial riblets, 3-7 between each pair of major 
ribs, very fine and crowded spiral riblets, plus weak secondary spiral 
cording on spire that fades out on body whorl. Umbilicus widely 
open, V-shaped, last whorl decoiling more rapidly, contained 2.95- 
4.28 times (mean 3.48) in the diameter, margins rounded. Whorl 
contours and apertural features without marked change from aver- 
age pattern. 

Sinployea modicella (fig. 38a-c) is most similar to 
S. neglecta (fig. 41d-f) from Huahine andS. tahitiensis 
(fig. 38d-f) from Tahiti. The former is distinctly 
smaller, has a narrower umbilicus, more and finer 
apical cords, and distinctly more prominent secondary 
spiral cording. Sinployea tahitiensis has half as many 
radial ribs and many more microradial riblets. No ex- 
tralimital species are close in size and shape. 

Description of figured specimen. Shell larger than average, 
with 4 normally coiled whorls. Apex and spire slightly and evenly 
elevated, last whorl descending much more rapidly, H/D ratio 0.480. 
Apical whorls 1%, sculpture of 13 narrow spiral ribs. Postnuclear 
whorls with narrow, prominent, protractively sinuated radial ribs, 
107 on the body whorl, whose interstices are 2-5 times their width. 
Microsculpture of fine radial riblets, 3-7 between each pair of major 
ribs, crossed by very fine and crowded spiral riblets, plus fine secon- 
dary spiral cording equal in size to microradials on spire that become 
weak to absent on body whorl. Sutures deep, whorls strongly rounded 
above, flattened laterally above periphery and somewhat flattened 
on basal margin, with evenly rounded outer margin. Color light red- 
dish horn, wide slightly darker flammulations visible above 
periphery. Umbilicus widely open, V-shaped, last whorl decoiling 
more rapidly, contained 3.63 times in the diameter, margins 
rounded. Aperture ovate, strongly flattened laterally above 
periphery, somewhat flattened on basal margin, inclined about 25 
from shell axis. Height 1.55 mm., diameter 3.22 mm. 

Range. Moorea, Society Islands. 

Material. Moorea: Opunohu Valley on tree 
trunks under bark (15 specimens, BPBM 89709); 
Faatoai Valley, 1 1 A miles inland (Stations 886, 893, 
896, 900) at 150-300 ft. elevation under dead logs and 
leaves or on trunk of hau (335 specimens, BPBM 
150331-4, BPBM 150375-9, BPBM 150416-21, BPBM 
150473-4); Maramu Valley, l l /z-l% miles inland (Sta- 
tions 919, 922) at 50-400 ft. elevation under logs and 
dead leaves (30 specimens, BPBM 150710-4, BPBM 
150754); Tepatu Valley (Stations 927, 929) at 50-800 
ft. elevation (7 specimens, BPBM 150872, BPBM 
150895-6); Station 478, above Rotonui, Belvidere Out- 
look, Uponohu Bay (1 specimen, FMNH 201054). 
"Tahiti" (12 specimens, FMNH 46531, RSM). 

Remarks. The taxon Helix modicella has been 
used as a catchall for the small, barrier-free endodon- 
toids of Polynesia and Fiji. In the absence of precise 
measurements and with the availability of only frag- 
mentary material, such a usage is understandable. Un- 
fortunately, it is difficult to fix the identity of this 
name. No type material could be located in Paris at the 
time of my study trips. All material collected on Tahiti 
in recent years differs from specimens in older collec- 
tions that were labeled "Tahiti." These shells agree 
more with the populations from lowland Moorea col- 
lected by the Mangarevan Expedition. The type figures 
have proportions that fall within the range of Moorean 
material, since the drawing is 9.4 mm. high, 18.5 mm. 









FIG. 38. a-c, Sinployea modicella (Ferussac). "Tahiti," Society Islands. FMNH 155940. d-f, Sinployea tahitiensis, new species. Station 863. 
Mt. Aorai, 4,700-5,000 ft., Tahiti, Society Islands. Holotype. BPBM 142072. Scale lines equal 1 mm. (SH). 



87 



88 



SOLEM: ENDODONTOID LAND SNAILS 



in diameter, with 4% whorls, H/D ratio 0.509, D/U 
ratio 4.02 and about 107 ribs on the body whorl. 

Because material from old collections agrees with 
the Moorean shells, I have no hesitation in restricting 
the type locality to Moorea. The figured specimen is 
labeled as coming from "Tahiti" and originating in a 
collection formed during the middle 1800s. Possibly it 
is part of the type set, but there is no evidence concern- 
ing this. In 1981, the Ecole National Superiure des 
Mines deposited material from the Deshayes collection 
in the Museum National d'Histoire Naturelle, Paris. 
Included were two lots of Helix modicella from 
"Tahiti." There were 8 and 11 shells glued to strips 
of cardboard. Through the courtesy of Dr. Simon Til- 
lier, it was possible to examine these specimens. 
The smaller lot agrees better in color and sculpture 
with the figured shell (fig. 38a-c) than the larger lot, 
but may not be conspecific. The latter specimens are 
much more brightly colored than the former and have 
more widely spaced sculpture. Almost certainly they 
are one of the other Society Island species. Without 
dismounting and extensive cleaning of each shell, de- 
termination of their species identity will not be possi- 
ble. I have not chosen a lectotype or neotype. I suspect 
that the material from the Deshayes collection was ob- 
tained subsequent to illustration of the original 
monochromatic specimen. The name Helix modicella 
Ferussac, 1840, is applied to the lowland Moorean 
populations on the basis of the original illustration. 

Variation in local populations is summarized in 
Table XVII. All these samples were taken between 50 
and 800 ft. elevation. Size and shape are practically 
identical, with only umbilical width varying signifi- 
cantly. Shells from Maramu Valley (Station 919) have 
a distinctly narrower umbilicus than those from 
Faatoai Valley (Stations 893, 896), with the small 
sample from Tepatu Valley (Station 927) intermediate 
in width. Although the difference between shells from 
Station 896 and Station 919 is statistically significant, 
with 24 degrees of freedom, t = 4.9127, I doubt that 
there is any systematic difference. 

Populations in Faatoai Valley showed considera- 
ble variation in percentage of adults. Station 893, from 
150-300 ft. elevation, had 20 of 65 (30.8%) adults; from 
Station 896 at 300 ft. elevation, there were only 19 of 
182 (10.4%) adults. Station 900, also at 300 ft. eleva- 
tion, had 11 of 73 (15.1%) adults. I doubt that sampling 
bias is involved, but there are insufficient data con- 
cerning the stations to suggest what factors might be 
responsible. 

Description of soft parts. Foot and tail slightly shorter than 
shell diameter. Sole undivided, pedal grooves prominent, uniting 
across tail, no caudal horn, middorsal groove weak. Head projecting 
in front of foot. Gonopore located slightly above and behind right 
rhinophore. Body color yellow-horn, usually with gray suffusion on 
head and neck. 

Mantle collar (MC) with tapered edge, no glandular extension 
onto mantle roof. Pneumostome shielded by a weak right mantle 
lobe. Anus (A) and external ureteric pore (KX) in typical position just 
inside pneumostome. 



Pallial region (fig. 39a) extending about % whorl apically. Lung 
roof without granulations. Kidney (K) bilobed, rectal lobe distinctly 
longer than pericardial, base tapering and extending apicad under 
intestinal loop, upper end medially indented. Length of kidney from 
anterior end of rectal lobe to base about 1.51 mm. Ureter (KD) follow- 
ing edges of kidney arms, a strip of lung roof visible between arms, 
opening next to anus. Heart (H) about half length of pericardial 
kidney arm. Principal pulmonary vein (HV) angling toward 
pneumostome, not conspicuously branched. Hindgut (HG) starting 
just above kidney apex, not tapering. 

Ovotestis (fig. 39b) a clump of several finger-like lobes lying 
above stomach apex parallel to whorl sides, imbedded in digestive 
gland. Hermaphroditic duct (GD) as in S. peasei. Albumen gland 
(GG) and talon (GT) typical. Prostate (DG) with very few and large 
acini opening into tube lying inside uterine wall. Uterus (UT) bipar- 
tite, upper chamber thinner walled and narrower, lower with thick 
glandular walls. 

Vas deferens (VD) much thicker apically, tapering to slender 
tube that remains constant in size until entering epiphallus. 
Epiphallus (E) without unusual features. Penial retractor (PR) aris- 
ing from diaphragm near apex of pallial cavity, inserting as a U- 
shaped fan partly around epiphallus-penis junction. Penis (P) exter- 
nally and internally the same as S. peasei, length about 1.00-1.10 
mm. Atrium (Y) slender, rather short. 

Free oviduct (UV), spermatheca (S), and vagina (V) without un- 
usual structures. 

(Based on BPBM 150416, 4 individuals, largest 3.49 mm. in 
diameter with 4% whorls.) 

Sinployea tahitiensis, new species. Figures 38d-f, 
39c-e. 

Diagnosis. Shell large, diameter 2.98-4.04 mm. (mean 3.50 
mm.), with 4-4% normally coiled whorls. Apex and early spire 
slightly and evenly elevated, body whorl descending more rapidly, 
spire protrusion about '& of body whorl width, H/D ratio 0.428-0.541 
(mean 0.498). Apical sculpture of 6-11 (mean 8.92) spiral cords. 
Postnuclear whorls with comparatively few, narrow, widely spaced, 
protractively sinuated radial ribs, 44-71 (mean 60.7) on the body 
whorl, whose interstices are 3-5 times their width. Ribs/mm. 4.67- 
8.37 (mean 5.86). Microsculpture of fine radial riblets, 8-12 between 
each pair of major ribs, crossed by extremely fine and crowded spiral 
riblets with widely spaced secondary spiral cords whose interstices 
are about twice those of microradials. Umbilicus broadly open, V- 
shaped, last whorl decoiling rapidly, contained 3.38-4.50 times 
(mean 3.68) in the diameter, with rounded margins. Sutures chan- 
neled, whorls strongly rounded above, not compressed on basal mar- 
gin. 

The slightly elevated spire, more widely spaced 
secondary spiral cording, fewer apical cords and 
slightly narrower umbilicus combine to separate 
Sinployea tahitiensis (fig. 38d-f) from the relatively 
similar S. lamellicosta (fig. 40a-c). Both S. modicella 
(fig. 38a-c) andS. neglecta (fig. 41d-f) have many more 
radial ribs, whereas S. montana (fig. 41a-c) is much 
smaller and has a very narrow umbilicus. 

Description. Shell large, with 4V4 loosely coiled whorls. Apex 
and first whorl and one-half flat, last whorl descending moderately, 
H/D ratio 0.478. Embryonic whorls 1%, with 10 relatively prominent 
spiral cords. Remaining whorls with protractively sinuated radial 
ribs having high periostracal lamellar extensions, 61 on the body 
whorl, whose interstices are 3-5 times their width. Microsculpture of 
fine radial riblets, 8-12 between each pair of major ribs, crossed by 
extremely fine and crowded spiral riblets. Secondary spiral cording 
equal in size to microradials, twice as widely spaced. Umbilicus 
broadly open, strongly decoiling on last whorl, V-shaped, contained 
3.47 times in the diameter. Color nearly uniform light reddish yellow 
with darker periostracal coloring on ribs. Aperture ovate, slightly 
flattened laterally above periphery, basal margin evenly rounded, 



SYSTEMATIC REVIEW 



89 



TABLE XVII. - LOCAL VARIATION IN SOCIETY ISLAND SINPLOYEA, I. 



modicella 



Sta. 919 
BPBM 150710 



Sta. 900 
BPBM 150473 



Sta. 893 
BPBM 150375 



Sta. 896 
BPBM 150416 



Sta. 927 
BPBM 150872 



tahitiensis 



Sta. 867 
BPBM 145702-3 



Sta. 862 
BPBM 145166 



NUMBER OF 
SPECIMENS 



61.56.52 
(55-68) 



1.94*0.050 

(1.78-2.07) 



1.56+0.082 

(1.38-1.78) 



H/D RATIO 



1. 


.67+0. 
(1. 


056 

42-1, 


,89) 


3.2910. 
(3. 


059 

11-3 


,51) 





.508+0, 
(0 


,0166 
,430-0 


,553) 


1 


.66+0, 
(1, 


044 
39-1 


,95) 


3.2810, 
(3, 


044 
01-3, 


54) 





,505+0 
(0, 


,0087 
,456-0, 


,551) 


1 


.75+0 
(1. 


,028 
56-2 


.02) 


3.3410 
(3, 


,026 
.15-3 


.61) 





,5230, 
(0 


,0069 
,480-0, 


,598) 


1 


.73+0 
(1 


,037 
.36-2 


.05) 


3.4010, 
(3, 


035 
,18-3 


,71) 





,508+0.0085 
(0.406-0, 


,562) 


1 


, 720. 

(1. 


,051 
,62-1 


.79) 


3.39+0 
(3, 


,090 
21-3.51) 





,509+0 
(0 


,0214 
,471-0, 


,546) 



3.74+0.110 

(3.42-4.01) 

3.2810.069 

(3.19-3.49) 



0.519+0.0065 

(0.500-0.541) 



0.47510.0173 

(0.428-0.509) 



D/U RATIO 



APICAL CORDS 






modicella 



Sta. 


919 


4 l/8+(4 1/8-4 1/4) 





.85+0 
(0 


,037 
,72-0 


.97) 


3.8910 
(3 


.106 
.54-4 


.28) 


9.00+0.69 
(7-12) 


Sta. 


900 


4 l/4-(4-4 


3/8) 





.94+0, 

(0, 


018 
,85-1 


,05) 


3.47+0 
3. 


.054 
16-3. 


77) 


9.0010.54 
(7-12) 


Sta. 


893 


4 1/4-K4-4 


1/2) 





,9810, 
(0 


017 
,86-1, 


,15) 


3.4010 
(3 


.043 
.06-3 


.78) 


9.5810.33 
(8-13) 


Sta. 


896 


4 l/4+(4-4 


5/8) 


1 


,02+0, 
(0, 


021 
86-1, 


,22) 


3.3310 
(2 


.057 
.95-3 


.79) 


9.17+0.33 
(7-12) 


Sta. 


927 


4 l/4-(4 1/8-4 1/4) 





.9310, 
(0, 


061 
86-1, 


05) 


3.6410 
(3 


.162 
.32-3.85) 



tahitiensis 



Sta. 867 4 1/2(4 3/8-4 5/8) 1.04+0.489 

(0.92-1.12) 



Sta. 862 4 l/8+(4-4 3/8) 0.9310.041 

(0.82-1.02) 



3.61+0.061 

(3.41-3.75) 



3.5510.126 

(3.38-3.92) 



9.50+0.58 
(9-10) 



7.5010.65 
(6-9) 



inclined about 25 from shell axis. Height of holotype 1.74 mm., 
diameter 3.66 mm. 

Holotype. Society Islands: Tahiti, Station 863, 
Mt. Aorai Trail at 4,700-5,500 ft. elevation. Collected 
by Yoshio Kondo and Donald Anderson on September 
14, 1934. BPBM 142072. 

Range. Interior of Tahiti at 770-6,300 ft. eleva- 
tion, Society Islands. 

Paratypes. Tahiti: Fautaua Valley (Station 20) 
at 250 m. elevation (1 specimen, BPBM 136343); right 
fork of Papenoo Valley (Station 951) at 1,400 ft. eleva- 
tion (2 specimens, BPBM 144969-70); Aorai Trail (Sta- 
tion 863) between 2nd and 3rd camps in leaf axils of 
ieie (3 specimens, BPBM 145913); ravine on west side 
(Station 862) of ridge crest, Mt. Aorai at 4,600 ft. eleva- 
tion (6 specimens, BPBM 145166-7); Aorai Trail (Sta- 
tions 865, 867) at 4,700-6,300 ft. elevation (10 speci- 
mens, BPBM 145296-7, BPBM 145702-3). 



Remarks. The flat spire and more numerous api- 
cal cords of S. lamellicosta (fig. 40a-c) are the sys- 
tematically important conchological characters sep- 
arating that species fromS. tahitiensis (fig. 38d-f). The 
former was taken by Garrett at relatively low eleva- 
tions and by the Mangarevan Expedition in a valley to 
one side of the Mt. Aorai Trail. It probably is geo- 
graphically isolated from populations of Sinployea 
tahitiensis. Sinployea montana also was collected in a 
valley to the west of the Mt. Aorai Trail, but at 1,000 ft. 
higher elevation than S. lamellicosta. The Moorean S. 
modicella (fig. 38a-c) differs primarily in having twice 
as many radial ribs on the body whorl and fewer micro- 
radial riblets. 

The penis (2.2-2.4 mm. long) is much larger than 
in either S. lamellicosta (1.4-1.5 mm. long) orS. mon- 
tana (1.4-1.65 mm. long). Internally it shows no 
differences from the typical pattern. 



K KD 




FIG. 39. Anatomy of Tahitian and Moorean Sinployea: a-b, Sinployea modicella (Ferussac). Station 896, Faatoai, Moorea. BPBM 150416. a, 
pallial region, b. genitalia; c-e, Sinployea tahitiensis, new species. Station 867, Mt. Aorai, Tahiti. BPBM 145703. c, pallial region, d, genitalia. e, 
detail of talon-carrefour region; f, genitalia of Sinployea montana, new species. Station 866, Mt. Aorai, Tahiti. BPBM 145401; g-h, Sinployea 
lamellicosta (Garrett). Station 870, Mt. Aorai, Tahiti. BPBM 145537. #, genitalia. h. detail of stimulatory pad in penis. Scale lines equal 1 mm. 
(a-d,f-g, MO;e, h, MM). 



90 



SYSTEMATIC REVIEW 



91 






Data on material from two local populations are 
presented in Table XVII. Specimens from Station 862 
were subadult. This lower H/D ratio reflects the les- 
sened body whorl descension as does the much lower 
mean height. The umbilical width is obviously less 
than in specimens of S. lamellicosta in the same size 
range. This provides a ready means of separating 
specimens where the spire protrusion is not obvious. 

Description of soft parts. Pallial region (fig. 39c) unusual only 
in having rectal and pericardia! kidney arms equal in length, about 
1.38 mm. long. Genitalia (fig. 39d) with head of talon less globular 
(fig. 39e) and a distinct shaft. Length of penis stalk, vagina, and 
atrium relatively short. Total length of penis about 2.2-2.4 mm. 
Interior of penis without unusual features. Jaw with plates partly 
fused. Radula typical, 6 laterals, with central slightly smaller. 

(Based on BPBM 145703, 3 partial individuals.) 



(Garrett, 1884). Figures 



Sinployea lamellicosta 

39g-h, 40a-c. 

Patula lamellicosta Garrett, 1884, J. Acad. Nat. Sci., Philadelphia, 

9(1), p. 30, pi. 2, fig. 11 a, b Tahiti, Society Islands. 
Helix (Patula) lamellicosta (Garrett), Tryon, 1887, Man. Conchol., 

(2) 3, p. 39, pi. 8, figs. 33-34. 
Endodonta (Thaumatodon) lamellicosta (Garrett), Pilsbry, 1893, 

Man. Conchol., (2)9, p. 27. 
Endodonta (Champa) lamellicostata (sic) (Garrett), Pilsbry, 1893, 

Man. Conchol., (2) 9, p. 35 typographical error. 

Diagnosis. Shell larger than average, diameter 2.78-3.55 mm. 
(mean 3.25 mm.), with 3%-4% relatively loosely coiled whorls. Apex 
and spire flat or slightly depressed, body whorl descending sharply, 
H/D ratio 0.454-0.521 (mean 0.487), much lower than average. Api- 
cal sculpture of 10-16 (mean 12.5) rather fine spiral cords. Postnu- 
clear sculpture of prominent, thin, protractively sinuated, widely 
spaced radial ribs, 43-71 (mean 52.4) on the body whorl, whose in- 
terstices are 4-6 times their width. Ribs/mm. 4.03-7.91 (mean 5.44). 
Microsculpture of fine radial riblets, 8-12 between each pair of major 
ribs, crossed by extremely fine and crowded spiral riblets that can be 
seen on upper spire, plus secondary spiral cording whose spacing is 
equal to the microradials. Umbilicus broadly open, wide, cup-shaped, 
regularly decoiling, contained 2.89-3.87 times (mean 3.26) in the 
diameter, margins rounded. Whorl contours and apertural features 
normal. 

Sinployea lamellicosta (fig. 40a-c) is the only Soci- 
ety Island species with a flat or slightly depressed 
spire. Sinployea tahitiensis (fig. 38d-f) is most similar, 
but has a slightly elevated spire, narrower umbilicus, 
and fewer spiral cords on the apex. All other Society 
Island species have much more crowded and numerous 
radial ribs. Extralimital species with fiat or depressed 
spires all have much more numerous and crowded ra- 
dial ribs. 

Description. Shell smaller than average, with slightly more 
than 3% loosely coiled whorls. Apex and early spire slightly de- 
pressed, body whorl descending rather rapidly, H/D ratio 0.440. Em- 
bryonic whorls P/K, sculpture partially eroded, but remnants of 11 
spiral ribs can be detected. Postnuclear whorls with protractively 
sinuated, prominent, widely spaced radial ribs, 50 on the body whorl, 
whose interstices are 4-6 times their width. Microsculpture of very 
fine radial riblets, 8-12 between each pair of major ribs, crossed by 
equal-sized secondary spiral cords that are slightly more widely 
spaced than the radial riblets, with barely visible, very fine and 
crowded spiral riblets most easily seen on upper spire. Sutures 
deeply impressed, whorls strongly rounded above, flattened laterally 
above periphery, with evenly rounded basal margin. Color pale 





FIG. 40. a-c, Sinployea lamellicosta (Garrett). Tahiti, Society 
Islands. Lectotype. BPBM 2841. Scale line equals 1 mm. (MM). 



yellow-horn with a few vague, light red flammulations. Umbilicus 
broadly open, cup-shaped, regularly decoiling, contained 2.93 times 
in the diameter, margins rounded. Aperture ovate, strongly flattened 
laterally above periphery, with gently rounded basal margin, in- 
clined about 15 from shell axis. Height of lectotype 1.19 mm., diame- 
ter 2.70 mm. 



92 



SOLEM: ENDODONTOID LAND SNAILS 



Lectotype. Society Islands: Tahiti. Collected by 
Andrew Garrett. BPBM 2841. 

Range. Tahiti, Society Islands, at low (?) to high 
elevations, probably in only one or two valleys. 

Paratype. BPBM 2841. 

Material. Tahiti (6 specimens, BPBM 2841, Zu- 
rich, NMWC); Pirae-Aorai trail (1 specimen, BPBM 
115855 Whitney Expedition); valley west of Aorai 
Trail (Station 870) at 5,000 ft. elevation (12 specimens, 
BPBM 145537-42). 

Remarks. Garrett (1884, p. 30) reported this to 
be "a scarce species, living beneath rotten wood in 
damp stations." Further collecting is needed to estab- 
lish the exact geographic relationships of S. lamel- 
licosta, S. tahitiensis, and S. montana in the Mt. Aorai 
region. 

Whereas the lectotype is the smallest shell ex- 
amined and is clearly subadult, the sculptural pres- 
ervation was excellent, and it matches in form the 
Mangarevan Expedition material. A single juvenile 
collected by the Whitney South Sea Expedition (BPBM 
115855) is tentatively referred here. The Zurich shells 
were barely adult and hence slightly higher and more 
narrowly umbilicated than the Mangarevan Expedi- 
tion specimens. Only one-third of the latter were adult. 

The flat spire and numerous apical cords are the 
main features characterizing S. lamellicosta. 

Description of soft parts. Pallial region with rectal and pericar- 
dial kidney arms almost equal in length, reaching 1.20 mm. long. No 
glandular extension onto pallial roof. Genitalia (fig. 39g) with rather 
long vagina. Penis quite thick, internal stimulatory pad (fig. 39h) 
very prominent. Length of penis about 1.41.5 mm. Glandular zone 
of spermatheca with a hardened sperm packet that was 0.45 mm. 
long, lower half cylindrical and 0.18 mm. in diameter, upper half 
expanded and spherical, diameter 0.28 mm. Radula typical, with 6 
laterals. 

(Based on BPBM 145537-8, 3 partly extracted specimens.) 

Sinployea montana, new species. Figures 39f, 41a-c. 

Diagnosis. Shell small, diameter 2.45-2.55 mm. (mean 2.50 
mm.), with 3%-3% normally coiled whorls. Apex and spire moder- 
ately and evenly elevated, last portion of body whorl starting to 
descend more rapidly, spire protrusion between V* and Vs body whorl 
width, with a high H/D ratio of 0.622-0.623 (mean 0.623). Apical 
sculpture of 13 spiral cords. Postnuclear sculpture of a few thin, 
prominent, protractively sinuated radial ribs, 57-69 (mean 63.0) on 
the body whorl, whose interstices are usually 3-4 times their width. 
Ribs/mm. 7.43-8.65 (mean 8.04). Microsculpture of fine radial rib- 
lets, 6-10 between each pair of major ribs, crossed by much finer and 
more crowded spiral riblets that become indistinct on body whorl. 
Umbilicus quite narrow, U-shaped, last whorl decoiling a little more 
rapidly, contained 5.93-8.55 times (mean 7.24) in the diameter, 
margins rounded. Basal margin of body whorl rounded, not flattened. 

The high spire and very narrow umbilicus of 
Sinployea montana (fig. 41a-c) easily separate it from 
the other Society Island species. Its rib count is the 
same as in S. tahitiensis (fig. 38d-f), but the size 
difference produces a much more crowded ribbing in S. 
montana. All extralimital species of similar size have 
higher whorl counts, lower H/D ratios, and wider 
umbilici. 



Description. Shell small, with 3% loosely coiled whorls. Apex 
and early spire moderately and evenly elevated, last whorl descend- 
ing sharply, H/D ratio 0.623. Embryonic whorls 1%, worn traces of 
about 13 relatively prominent spiral ribs remaining. Lower whorls 
with protractive, thin, radial ribs, 69 on the body whorl, whose in- 
terstices are 3-4 times their width. Microsculpture of fine radial 
riblets, 6-10 between each pair of major ribs, crossed by much finer 
and more crowded spiral riblets that become relatively indistinct on 
the body whorl. Sutures deeply impressed, whorls strongly rounded 
above, flattened laterally above periphery, evenly rounded on lower 
palatal and basal margin. Umbilicus narrowly U-shaped, last whorl 
decoiling slightly more rapidly, contained 8.55 times in the diameter, 
margins rounded. Color uniform light yellow-horn with darker 
periostracal rib extensions, no flammulations. Aperture nearly circu- 
lar, slightly flattened laterally above periphery and on basal margin, 
inclined about 10 from the shell axis. Height of holotype 1.58 mm., 
diameter 2.54 mm. 

Holotype. Society Islands: Tahiti, Station 866, 
valley west of Aorai Trail at 6,000 ft. elevation. Col- 
lected by Elwood Zimmerman, Yoshio Kondo, and 
Donald Anderson on September 15, 1934. BPBM 
145401. 

Range. Mt. Aorai, 6,000 ft. elevation, Tahiti, 
Society Islands. 

Paratypes. Tahiti: valley west of Aorai Trail 
(Station 866) at 6,000 ft. elevation (2 specimens, BPBM 
145401-2). 

Remarks. I suspect that the size of Sinployea 
montana is underrepresented by the available mate- 
rial. Two of the three examples showed only the first 
indications of gerontic growth. Probably 10%- 15% 
greater diameter is attained. 

No other Society or Cook species approaches S. 
montana in narrowness of the umbilicus. Only the Sa- 
moan S. allecta (fig. 54a-e) and the Fijian S. recursa 
(fig. 64c) could be confused on umbilical size. Sinployea 
allecta is 0.5 mm. larger and has an average of less 
than nine apical cords and a thicker body whorl. Sin- 
ployea recursa is still larger and has greatly reduced 
major sculpture. 

Description of soft parts. Pallial region typical, rectal arm 
longer than pericardial, former about 1.25 mm. long in one example. 
Apical genitalia not seen, lower portions typical. Penis 1.4-1.65 mm. 
long in two examples. Vergic papilla (fig. 39f) large and U-shaped, 
muscular bands relatively weak, stimulator typical. 

Radula with 5-6 laterals and more than 6 marginals. Central 
9 /j. long, 7 ij. wide, distinctly smaller than laterals, 1st lateral 8 n 
wide. 

(Based on BPBM 145401, 2 extracted and partly broken exam- 
ples.) 



Sinployea neglecta, new species. Figures 41d-f, 42a. 

Diagnosis. Shell of average size, diameter 2.57-3.26 mm. 
(mean 2.83 mm.), with 3%-4V4 rather tightly coiled whorls. Apex and 
spire slightly to moderately and evenly elevated, body whorl de- 
scending more rapidly, spire protrusion Ve- Vs of body whorl width, 
H/D ratio 0.441-0.632 (mean 0.536). Apical sculpture of 10-15 
(mean 12.5) relatively fine spiral cords. Postnuclear sculpture of fine, 
prominent, thin, protractively sinuated radial ribs, 71-143 (mean 
108.6) on the body whorl, whose interstices are 2-3 times their 
width. Ribs/mm. 8.84-16.87 (mean 12.23). Microsculpture of fine ra- 
dial riblets, 3-5 between each pair of major ribs, barely visible mi- 
crospiral riblets and relatively prominent secondary spiral cords. 
Umbilicus of average width, V-shaped, last whorl usually decoiling 







def 



f 




FIG. 41. a-c, Sinployea montana, new species. Station 866, Aorai Trail, 6,000 ft., Tahiti, Society Islands. Holotype. BPBM 145401: d-f, 
Sinplovea neglecta, new species. Station 1008, Tiva, Huahine, Society Islands. Holotype. BPBM 151565. Scale lines equal 1 mm. (a-c, SG; d-f. 

MM). 



93 



94 



SOLEM: ENDODONTOID LAND SNAILS 



slightly more rapidly, contained 3.21-5.00 times (mean 4.39) in the 
diameter, margins rounded. Whorl contours and apertural features 
without marked differences from average. 

The smaller size, much narrower umbilicus, and 
greater number of apical cords separate Sinployea 
neglecta (fig. 41d-f) from the Moorean S. modicella (fig. 
38a-c). Sinployea sp. from Borabora differs in the 
character of the ribbing, which is much lower and more 
rounded. Of the Cook Island species, S. andrewi (fig. 
44a) has fewer apical cords and more radial ribs, 
whereas S. atiensis (fig. 44d-e) is more elevated and 
with slightly fewer apical cords. The Tongan S. vicaria 
(fig 58a-d) has a U-shaped umbilicus, more rounded 
whorl contours, more microradial riblets, and more 
whorls. 

Description. Shell of average size, with 4 relatively loosely 
coiled whorls. Apex and spire moderately and evenly elevated, last 
whorl descending slightly more rapidly, H/D ratio 0.530. Embryonic 
whorls I'/a, sculpture of about 12 spiral cords. Postnuclear whorls 
with numerous protractively sinuated, very thin radial ribs having 
periostracal lamellate extensions, 109 on the body whorl, whose in- 
terstices are variable, but about 2-3 times their width on middle of 
spire. Microsculpture of very fine radial riblets, 3-5 between each 
pair of major ribs, with almost equally large, slightly more widely 
spaced secondary spiral cords, and barely visible microspiral riblets. 
Sutures deeply impressed, whorls strongly rounded above, flattened 
laterally with rounded periphery and somewhat flattened basal 
margin. Color light reddish horn with indistinct darker reddish 
maculations visible mainly above periphery. Umbilicus relatively 
narrow, last whorl becoming more strongly decoiled, contained 4.15 
times in the diameter. Aperture ovate, flattened laterally above 
periphery and on basal margin, inclined about 20 from the shell 
axis. Height of holotype 1.45 mm., diameter 2.73 mm. 

Holotype. Society Islands: Huahine, Station 
1008, 100 yd. inland, near Tiva at 6 ft. elevation. Col- 
lected on logs and dead leaves by Yoshio Kondo and 
Donald Anderson on October 3, 1934. BPBM 151565. 

Range. Huahine, Society Islands. 

Paratypes. Huahine (27 specimens, BPBM 6078, 
ANSP 1939, ANSP 47759, SMF 165460, ex SMF 
165351): Tiva (Station 1008) about 100 yd. inland at 6 
ft. elevation on logs and dead leaves (80 specimens, 
BPBM 151565-9); Fare (Stations 962, 963) at 100-300 
ft. elevation under stones and logs or on dead leaves (52 
specimens, BPBM 151121, BPBM 151144-9); Fare 
(Station 978) at 1,000-1,200 ft. elevation in a damp 
valley on dead fallen leaves (1 specimen, BPBM 
151284). "Tahiti" (11 specimens, RSM). 

Remarks. Garrett (1884, p. 28) lumped this 
species with S. modicella (fig 38a-c). Sinployea ne- 
glecta (fig. 41d-f) is much smaller, has more apical 
cords, a narrower umbilicus, and more prominent sec- 
ondary spiral cording. These differences require better 
optical equipment and more precise measurements 
than could be made by Garrett. Several small samples 
from his collections and two larger sets from the Man- 
garevan Expedition are compared in Table XVIII. 
There are no significant size or shape differences, ex- 
cept in regard to umbilical width. The slightly wider 
umbilici for material from Station 1008 and ANSP 
47759 is caused by a few gerontic individuals in which 
umbilical decoiling has proceeded further than usual, 



producing relatively widely open umbilici and thus dis- 
torting the means. 

Specimens from Tiva (Station 1008) were collected 
about 100 yd. inland in coastal vegetation, whereas 
those from Fare (Station 963) were found in lowland 
forest at 300 ft. elevation further inland. Percentage of 
adults was identical, with 15 of 51 (29.4%) from Station 
963 and 24 of 81 (29.6%) from Station 1008. Size and 
shape differences reached significant levels, particu- 
larly in regard to height, with 37 degrees of freedom, t 

2.3293, and D/U ratio, t 4.9127. Probably 
ecophenotypic variation is involved. 

Description of soft parts. Pallial region typical. Rectal kidney 
arm 1.45 mm. long, total cavity length 3.3 mm. Narrow strip of lung 
roof between ureter arms. 

Genitalia (fig. 42a) without unusual features, penis length 
1.4-1.6 mm. Drawing misleading in that slender shafts of penis and 
vagina folded under. Actual lengths as inS. modicella. 

(Based on BPBM 151565, 4 partial examples.) 

Sinployea sp. 

Material. Society Islands: Borabora, south slope 
of Pahio-Temanu ridge (Station 1093) at 800 ft. eleva- 
tion (1 specimen, BPBM 152395). 

Remarks. A single, probably subadult example 
from Borabora is not referable to any named form. The 
very low and crowded radial ribbing, 126 ribs on the 
body whorl, is more like some Western Polynesian 
species than any Society Island Sinployea. Measure- 
ments are given in Table XVI. Although obviously 
similar to S. neglecta in all dimensions, the character 
of the major ribbing is so distinctive that I doubt their 
identity. Both have three to six microradials between 
each pair of major ribs. Without more material and 
knowledge of the apical sculpture, description would be 
premature. 

COOK ISLAND Sinployea 

Specimens of Sinployea have been found on all the 
major Cook Islands except Mauke. Libera fratercula 
fratercula (Pease) was collected on the latter island by 
P. H. Buck, who had taken Sinployea on Atiu, 
Aitutaki, and Mangaia during the same expedition. 
Hence, there is fair probability that this is an actual 
distributional gap. 

Sinployea atiensis (Pease) lives on both Atiu and 
Aitutaki. Except for a slight increase in whorl count 
and a strongly elevated spire (fig. 44e), which increases 
the H/D ratio, this is a very average species in all 
measured characters (table XIX). Sinployea andrewi 
from Mangaia is slightly larger, has a greater number 
of more crowded major radial ribs (fig. 44a), is dis- 
tinctly less elevated, has fewer microradials, and the 
umbilicus is V-shaped, rather than U-shaped. 
Rarotonga has, or had, a remarkable radiation of 
Sinployea, including nine of the 10 largest species in 
the genus. Only the Upolu Island S. complementaria 
(Mousson) (table XXIV) reaches larger size than the 
Rarotongan species. 



SYSTEMATIC REVIEW 



95 



TABLE XVIII. - LOCAL VARIATION IN SOCIETY ISLAND SINPLOYEA, II. 



NUMBER OF 
SPECIMENS 



H/D RATIO 



lamellicosta 



Sta. 870 
BPBM 145537 



50.014.05 
(43-57) 



1.690.030 

(1.62-1.76) 



3.49*0.046 

(3.38-3.58) 



0.485+0.0120 

(0.454-0.510) 



Sta. 866 
BPBM 145401 



1.56 0.033 

(1.52-1.51) 



2.5010.050 

(2.45-2.55) 



0.62310.0005 

(0.622-0.623) 



Sta. 1008 
BPBM 151565-7 

Sta. 963 

BPBM 151144, -7 

ANSP 1939 



ANSP 47759 



BPBM 6078 





1 .tOZU 

(1 


,VL. 
.29-1 


.62) 




, /O T U 

(2 


, uz^ 
,58-2. 


98) 


u . 


3-41U 

(0, 


, uuo / 
,465-0, 


598) 





1.5510 
(1 


,033 
,39-1 


.82) 


2 


,8410.028 
(2.72-3, 


08) 


0, 


54410. 
(0, 


0096 

500-0. 


605) 


107.315.36 
(101-118) 


1.5210 
(1 


,033 
,49-1 


.59) 


2 


.8410 
(2 


,098 
65-2. 


98) 


0. 


53810 

(0 


.013 
.518-0, 


563) 


107.5112.66 
(85-143) 


1.5510 
(1 


.065 
,36-1 


.69) 


2 


.9110, 

(2 


,070 
75-3. 


08) 


0. 


54110. 
(0, 


028 
.441-0, 


594) 





1.6410 
(1 


.090 
,39-1 


,99) 


2 


,9410, 
(2. 


097 
65-3. 


28) 


0. 


55810, 
(0. 


026 
455-0. 


632) 



lamellicosta 



Sta. 870 4 l/4-(4 1/8-4 3/8) 



UMBILICUS 



1.1410.037 

(1.05-1.22) 



D/U RATIO 



APICAL CORDS 



3.0610.0851 11. 25*0. 95 

(2.89-3.29) (10-14) 






Sta. 866 3 5/8+(3 5/8-3 3/4) 

neglecta 

Sta. 1008 4-(3 5/8-4 3/8) 

Sta. 963 4+(3 7/8-4 1/4) 

ANSP 1939 4(3 3/4-4 1/8) 

ANSP 47759 4 l/8-(3 7/8-4 1/4) 

BPBM 6078 4-K3 7/8-4 1/4) 



0.36*0.066 

(0.30-0.43) 



0.6510.011 

(0.59-0.82) 



0.63*0.012 

(0.56-0.72) 



0.6010.011 

(0.59-0.63) 



0.7210.062 

(0.63-0.95) 



0.6410.053 

(0.53-0.89) 



7.24*1.31 

(5.93-8.55) 



4.2710.051 

(3.60-4.72) 



4.5310.062 

(4.00-4.94) 



4.6810.118 

(4.45-4.84) 



4.0410.222 

(3.21-4.53) 



4.6410.233 

(3.67-5.23) 



12.8910.42 

(10-15) 



12.1310.58 

(10-15) 



Sinployea peasei (fig. 45a-c) is the Rarotongan 
equivalent of S. atiensis and S. andrewi. Reduction of 
the apical sculpture (fig. 2a, p. 11) is the significant 
change, although (table XIX) there are minor size and 
shape differences from the other two species. Sinployea 
peasei is replaced in part of Avana Valley by S. av- 
anaensis (fig. 45d-f). This is a much more narrowly 
umbilicated shell with many very crowded radial ribs 
and a lower whorl count. Present data are insufficient 
to determine whether there is a zone of sympatry. 
There does seem to be ecological separation, because 
only one of 159 Sinployea peasei collected in 1964 and 
1965 was taken on the leaves of shrubs, the remainder 
under logs and stones. A few individuals of S. avanaen- 
sis were found under rotting wood, but the great major- 
ity were collected on tree trunks in moss and lichens 
from 1-15 ft. above ground level. Anatomical dif- 



ferences between the two species are slight. Sinployea 
avanaensis has the rectal kidney lobe much wider, the 
penis about 0.3 mm. longer, and the vergic papilla 
longer and narrower than in S. peasei. Study of local 
populations in Avana Valley offers an excellent oppor- 
tunity for investigation of speciation phenomena. 

The larger species show two divergent patterns 
that may represent monophyletic series, plus two 
species, S. planospira (fig. 46d-h) and S. rudis (fig. 
47d-f), that show individual specialization. 

One series of species, S. proximo, S. youngi, S. 
harveyensis, and S. canalis, retains normal or slightly 
more crowded sculpture. The smallest, S. proximo (fig. 
46a-c), has a distinctly channeled suture and rela- 
tively low spire. Garrett ( 1872, p. 230) reported it was a 
"common species lurking under stones and among rot- 



96 



SOLEM: ENDODONTOID LAND SNAILS 




pv 



FIG. 42. Anatomy of Society and Cook Island Sinployea: a, 
genitalia of Sinployea neglecta, new species. Station 1008, Tiva. 
Huahine. BPBM 151565; b, genitalia of Sinployea andreu'i. new- 
species. Arakiore Oneroa, Mangaia, Cook Islands. BPBM 97652; c-e, 
Sinployea peasei. new species. Station 26, Mt. Maungaroa, 1,400 ft. 
elevation. Rarotonga, Cook Islands. FMNH 144645. c, pallial region. 
d. penial complex, c, detail of vergic papilla and muscular collar in 
penis. Scale lines equal 1 mm. (a-b, MO; e, MM; c-d, PS). 



ten wood ... in several villages (error for valleys)." 
Later Garrett (1881, p. 386) stated that it was "Not 
uncommon in several valleys." A personally annotated 
copy of Garrett (1872) corrected "villages" to "valleys." 
Sinployea youngi (fig. 48d-f) was taken "On the ground 
in damp woods, and only noticed in a single valley" 
(Garrett, 1872, p. 229). It is characterized by its large 
size, compressed aperture, and widely open umbilicus. 
Sinployea harueyensis (fig. 48a-c) is very similar to S. 
youngi, but has a higher spire, more numerous crowded 
ribs, narrower umbilicus, and ovate aperture. It was "A 
common species found under rotten wood" (Garrett, 
1872, p. 228). Sinployea canalis (fig. 49a-c) was "found 
on the ground in damp forests, and confined to a single 
valley" (Garrett, 1872, p. 227). Although S. planospira 
(fig. 46d-h) retains typical sculpture, its great increase 
in whorl count (mean 6%), very wide umbilicus, flat- 
tened apex, plus early spire set it apart from the above 
species. Sinployea planospira is "a very rare species" 
(Garrett, 1881, p. 388). Its habitat was not defined. 

The second series differs primarily in its much 
wider spacing of the major ribs. Quite probably it is 
polyphyletic. Sinployea decorticata (fig 47a-c) has a 
higher spire and H/D ratio with between 10 and 20 
microradials between each pair of major ribs and aver- 
ages 75 ribs on the body whorl. It is confined to a single 
valley (Garrett, 1881, p. 387) where it was very abun- 
dant on the ground. Sinployea otareae (fig. 50a-c) has a 
lower spire and H/D ratio, averages about 96 ribs on 
the body whorl, and has only six to 10 microradial rib- 
lets between each pair of major ribs. It lived "under 
dead wood on the banks of a stream" in a valley about 3 
miles away from that occupied by S. decorticata. 
Sinployea tenuicostata (fig. 50d-f) is much larger, has a 
flat apex, averages about 51 ribs on the body whorl, 
and has eight to 12 rather large microradial riblets 
between each pair of majors, indicating an increase in 
microradial riblet size and spacing. It was a "somewhat 
rare species, having a wide range on the island, and 
generally found on the ground on the sides of ravines." 
Sinployea rudis (fig. 47d-f) has reasonably typical 
upper spire sculpture, but the major ribs become 
sharply reduced and irregular on the body whorl, 
whereas the lateral flattening above the periphery has 
proceeded to form a broad and shallow supraperipheral 
sulcus. It is thus an independent pattern of specializa- 
tion. Sinployea rudis was a "very abundant species . . . 
we gathered hundreds in several valleys" (Garrett, 
1881, p. 387). It was taken "on the ground in damp 
woods." 

Of the large species, four (S. proximo. S. harveyen- 
sis, S. tenuicostata, and S. rudis) were widely distrib- 
uted or found in several valleys, whereas five (S. 
youngi, S. canalis, S. planospira, S. decorticata, and S. 
otareae) were found only in a single valley. The niche of 
S. planospira was not defined, but all the other species 
were stated to occur on the ground or under rotting 
wood. Normally Garrett indicated sympatry for 
species. Thus, Mautodontha zebrina (Garrett) (see 
Solem, 1976b, p. 162) was collected in the same valley 



SYSTEMATIC REVIEW 



97 




FIG. 43. Anatomy ofSinployea avanaensis, new species. Station 32A, Avana Valley. Rarotonga, Cook Islands. FMNH 144671: a, foot length 
in relation to shell diameter; b, pallial region; c, genitalia; d, midvisceral genitalia showing uterine sections; e, interior of penis. Scale lines 
equal 1 mm. (PS). 



with S. otareae. Sinployea youngi was found in the 
same valley as S. decorticata, but Garrett did not state 
that they were sympatric. He made no other mention of 
any Sinployea being collected together. The probability 
is that these were geographically isolated. 

Use of the past tense is deliberate. Mautodontha 
unilamellata (Garrett), Thaumatodon multilamellatus 
(Garrett), Libera tumuloides (Garrett), Libera sub- 
cavernula (Tryon), Mautodontha zebrina (Garrett), and 
the nine large Sinployea were not collected in 1964 or 
1965 despite rather extensive coverage of different val- 
leys. Overlooking the highly localized species, such as 
"one-half acre . . . nearly two miles inland" (Libera 
tumuloides), would not be difficult, but the total failure 



to find any of the widely distributed species suggests 
that actual extinction has occurred. There has been no 
recent comprehensive report on the ant fauna of 
Rarotonga, but Pheidole megacephala (Fabricius), a 
species very destructive to native insects, has been on 
Rarotonga since at least 1914 (Wilson & Taylor, 1967b, 
p. 47). The same ant is common on Upolu where 
Sinployea complementaria (Mousson) still occurs, but 
that species is semiarboreal (p. 129) and the shift in 
habitat from litter to semiarboreal may have allowed 
coexistence of the ant and snail. Possibly some species 
still survive at higher elevation in isolated colonies, 
but I suspect that most of the Rarotongan endodontoids 
are extinct. 



98 



SOLEM: ENDODONTOID LAND SNAILS 



TABLE XIX. - RANGE OF VARIATION IN COOK ISLAND SINPLOYEA. 





NUMBER OF 
SPECIMENS RIBS RIBS/MM. 

atlensls 39 93.3(83-107) 11.27(10.4-12.5) 


HEIGHT DIAMETER H/D RATIO 
1.57(1.29-1.92) 2.70(2.45-3.18) 0.582(0.527-0.642) 




(Pease) 
andrewl 83 129.2(104-158) 14.01(11.63-16.09) 


1.54(1.29-1.69) 2.95(2.68-3.15) 0.519(0.472-0.568) 




pease 1 173 112.2(83-167) 11.51(8.80-14.66) 
avanaensis 138 225.8(175-290) 20.82(16.28-25.1) 


1.62(1.25-1.94) 3.15(2.97-4.01) 0.492(0.415-0.558) 
1.52(1.32-1.81) 3.21(2.86-3.82) 0.474(0.427-0.516) 




proxima 20 119.7(92-154) 10.75(8.91-12.96) 
(Carrett) 

planospira 3 128.0(108-148) 10.62(10.05-11.19) 


1.97(1.71-2.37) 3.61(3.26-4.08) 0.544(0.519-0.583) 
1.77(1.52-2.22) 3.74(3.44-4.24) 0.468(0.439-0.524) 




(Carrett) 
rudls 32 RIBS REDUCED 


2.23(1.97-2.83) 4.04(3.68-4.83) 0.553(0.500-0.642) 




(Garrett) 
decorticata 21 75.0(62-103) 5.61(4.84-6.47) 


2.36(2.04-2.98) 4.30(3.71-5.10) 0.550(0.488-0.590) 




(Garrett ) 
harveyensis 27 RIBS REDUCED 


2.39(1.92-3.01) 4.35(3.91-5.23) 0.550(0.475-0.641) 




(Garrett) 

young i 17 115.4(96-139) 8.13(7.15-9.68) 
(Garrett) 

canalls 18 119.8(98-146) 8.22(7.07-9.74) 


2.27(1.84-2.66) 4.54(3.75-5.07) 0.499(0.457-0.553) 
2.14(1.84-2.63) 4.63(4.01-5.46) 0.463(0.427-0.544) 




(Garrett) 
otareae 38 95.6(81-116) 6.48(5.72-7.73) 


2.41(2.07-2.96) 4.68(4.21-5.20) 0.516(0.470-0.570) 




(Garrett) 
tenuicostata 15 50.7(43-66) 3.46(2.87-4.23) 


2.34(1.94-2.63) 4.70(4.08-5.43) 0.499(0.437-0.537) 




(Garrett) 

WHORLS UMBILICUS n/U RATIO 
atlen. 4 3/8-(4 1/8-4 7/8) 0.63(0.56-0.82) 4.26(3.77-4.82) 


APICAL SPIRE BODY WHORL 
CORDS ELEVATION WIDTH SP/BWW 

11.13(9-14) 0.21(0.16-0.26) 0.86(0.79-0.99) 0.237(0.196-0.296) 




andre. 4 l/8+(4-4 3/8) 0.72(0.63-0.79) 4.09(3.74-4.45) 


9.19(8-11) 0.17(0.13-0.23) 0.91(0.86-0.95) 0.189(0.143-0.269) 




pease. 4 3/8-(4-5 1/8) 0.81(0.56-1.25) 3.93(3.72-4.60) 
avana. 4+(3 7/8-4 1/2) 0.69(0.53-0.99) 4.78(3.87-5.69) 


11.4(9-14) 0.18(0.10-0.30) 0.93(0.82-1.02) 0.176(0.103-0.300) 
10 ~FINE EM 0.17(0.12-0.23) 0.95(0.79-1.09) 0.181(0.113-0.264) 




proxi. 4 3/8-(4-4 7/8) 0.68(0.59-0.76) 5.35(4.48-6.16) 
piano. 6 5/8(6 1/4-7 1/4) 1.28(1.15-1.41) 2.90(2.75-2.98) 
rudls 4 l/4+(3 7/8-4 5/8) 0.83(0.69-1.12) 4.87(4.18-5.62) 


0.15(0.10-0.23) 1.17(1.05-1.32) 0.124(0.091-0.175) 
0.23 1.02 0.226 
10.02(8-12) 0.24(0.20-0.43) 1.32(1.18-1.45) 0.184(0.150-0.296) 




decor. 4 5/8-(4 3/8-5) 1.00(0.79-1.22) 4.32(3.75-5.92) 


9.00(8-10) 0.27(0.13-0.46) 1.29(1.18-1.58) 0.209(0.103-0.292) 




harve. 4 5/8+(4 1/4-5) 0.92(0.76-1.18) 4.73(3.65-5.48) 


0.36(0.20-0.59) 1.39(1.28-1.55) 0.258(0.150-0.450) 




young. 4 3/8+(3 7/8-5) 1.23(1.03-1.61) 3.73(3.15-4.39) 
canal. 5-(4 1/2-5 1/2) 1.59(1.38-1.84) 2.91(2.64-3.10) 


10.73(10-12) 0.21(0.15-0.30) 1.39(1.22-1.51) 0.149(0.116-0.196) 
10.21(8-12) 0.15(0.10-0.26) 1.53(1.41-1.74) 0.096(0.067-0.151) 




otare. 4 3/4(4 1/2-5) 1.15(0.89-1.41) 4.10(3.47-5.52) 


9.41(8-12) 0.25(0.13-0.40) 1.45(1.35-1.55) 0.171(0.093-0.261) 




tenul. 4 1/4(3 7/8-4 1/2) 0.92(0.79-1.18) 5.16(4.29-6.20) 


10.45(9-12) FLAT 1.56(1.44-1.64) FLAT 




KEY TO THE RAROTONGAN Sinployea 


6. Mean rib count over 200; mid-Avana Valley 


1 


Whorls less than 5% 2 


Sinployea auanaensis, new species 
Mean rib count less than 140; widely distributed 


?, 


Whorls more than 6V 8 Sinployea planospira (Garrett, 1881) 
Radial ribs crowded to widely spaced 4 


Sinp/oyea peasei, new species 
7. Mean ribs/mm, about 8 . . ..8 






Mean ribs/mm 37 9 


3. 


Body whorl with supraperipheral sulcus; ribs reduced to ir- 
regularity Sinp!o\ea rudis (Garrett 1872) 


8. Umbilicus widely open, mean D/U ratio about 3.00; whorls 
strongly compressed laterally 




Body whorl without sulcus; ribs very crowded 


Sinployea canalis (Garrett, 1872) 


4 


Sinployea harveyensis (Garrett, 1872) 
Suture normal 5 


Umbilicus of normal width, mean D/U ratio about 3.75; whorls 




Suture distinctly channeled (fig. 46a-b) 


Sinployea youngi (Garrett 1872) 




Sinployea proxima Garrett, 1872) 


9 Spire flat mean rib count about 50 


5 


Mean diameter less than 3.5 mm 6 


Sinployea tenuicostata (Garrett 1872) 




Mean diameter more than 4.0 mm.... . ..7 


Soire stronslv elevated; mean rib count 75-100.... ...10 



SYSTEMATIC REVIEW 



99 



10. Mean rib count about 75; 10-20 microradials between each pair 

of major ribs Sinployea decorticnta (Garrett, 1872) 

Mean rib count about 95: 6-10 microradials 

Sinployea otareae (Garrett, 1872) 

Sinployea atiensis (Pease, 1870). Figure 44d-f. 

Pithys atiensis Pease, 1870, J. de Conchyl., 18, pp. 394-395 Atiu, 
Cook Islands. 

Pitys atiensis Pease, 1871, Proc. Zool. Soc. London, 1871, pp. 453, 
474. 

Helix atiensis (Pease), PfeifTer, 1876, Monog. helic. viv., 7, p. 165. 

Patula atiensis (Pease), Garrett, 1881, J. Acad. Nat. Sci., Philadel- 
phia, 8 (4), p. 386 Atiu and Aitutaki, Cook Islands. 

Endodonta (Champa) modicella var. atiensis (Pease), Pilsbry, 
1893, Man. Conchol., (2) 9, p. 35. 

Diagnosis. Shell slightly smaller than average, diameter 
2.45-3.18 mm. (mean 2.70 mm.), with 4Vfe-4% rather tightly coiled 
whorls. Apex and spire moderately and evenly elevated, slightly 
rounded above, last whorl descending more rapidly, spire protrusion 
about 'A body whorl width, H/D ratio 0.527-0.642 (mean 0.582). Api- 
cal sculpture of 9-14 (mean 11.13) prominent spiral cords. Postnu- 
clear whorls with narrow, prominent, crowded, protractively 
sinuated radial ribs, 83-107 (mean 93.3) on the body whorl, whose 
interstices are 2-3 times their width. Ribs/mm. 10.4-12.5 (mean 
11.27). Microsculpture of fine radial riblets, 5-8 between each pair of 
major ribs, crossed by very fine and crowded spiral riblets, with a 
secondary sculpture of rather widely spaced spiral cords. Umbilicus 
rather narrow, U-shaped, regularly decoiling, contained 3.77-4.82 
(mean 4.26) in the diameter, margins rounded. Whorl and apertural 
characters not differing from average pattern. 

Sinployea atiensis (fig. 44d-f) has lower and more 
rounded radial ribs than any of the Society Island 
species. Its high spire, slightly greater whorl count, 
and finer apical cords separate it from S. neglecta (fig. 
41d-f). The related S. andrewi (fig. 44a-c) and S. 
peasei (fig. 45a-c) are distinctly less elevated and 
larger; S. andrewi has fewer and finer apical cords; S. 
peasei has the apical cords greatly reduced in promi- 
nence. 

Types. No specimens dating from Pease's ex- 
changes could be located. The possibility of specimens 
being discovered in some European museum is good, 
and I prefer not to designate a neotype. 

Range. Atiu and Aitutaki, Cook Islands. 

Material. Aitutaki (5 specimens, Zurich): Reu- 
KHI. ] '-2 mile inland at 100 ft. elevation (1 specimen, 
BPHM 95649). Atiu (13 specimens, BPBM 87407-9, 
Zurich): Matai, Vi mile inland at 50 ft. elevation (11 
specimens, BPBM 95117); below Tengatangi, I'/a miles 
inland at 150 ft. elevation (1 specimen, BPBM 94942); 
Lake Teroto, Vz mile inland at 50 ft. elevation (4 speci- 
mens, BPBM 95036); Mokoero, V 2 mile inland at 30 ft. 
elevation (2 specimens, BPBM 95141); Tarapaku track, 
Vi to '/2 mile inland at 50 ft. elevation (2 specimens, 
BPBM 94998). 

Remarks. Although Garrett (1884, p. 28) consid- 
ered this to be a synonym of the Moorean S. modicella 
(fig. 38a-c), the qualitative difference in ribbing and 
obvious changes in proportions (tables XVI, XIX) effec- 
tively separate the two species. As with the Ancey col- 
lection, Pease's types and exchanges were so widely 
scattered that I prefer not to select a type specimen at 
this time. The Aitutaki examples were barely adult, 
and hence I attach no systematic significance to the 



smaller size and lower spire shown by this set (table 

XX). 

Description of. wft parts. Partial dissection of several fragmen- 
tary examples showed no differences in structures from S. peasei. 
Penis about 1.1-1.3 mm. long, most examples flattened too badly for 
opening. One specimen with verge and stimulatory pad identical to 
S. peasei. 

(Based on BPBM 95117, several examples.) 

Sinployea andrewi, new species. Figures 42b, 44a-c. 

Diagnosis. Shell slightly larger than average, diameter 2.68- 
3.15 mm. (mean 2.95 mm.), with 4-4% normally coiled whorls. Apex 
and spire slightly to moderately and evenly elevated, body whorl 
descending more rapidly, spire protrusion less than '/s body whorl 
width, H/D ratio 0.472-0.568 (mean 0.519). Apical sculpture of 8-11 
(mean 9.19) prominent spiral cords. Postnuclear sculpture of narrow, 
rounded, crowded, protractively sinuated radial ribs, 104-158 (mean 
129.2) on the body whorl, whose interstices usually are less than 
twice their width. Ribs/mm. 11.63-16.09 (mean 14.01). Micro- 
sculpture of fine radial riblets, 3-6 between each pair of major ribs, 
with barely visible microspirals and rather crowded secondary spiral 
cording. Umbilicus open. V-shaped, last whorl decoiling more 
rapidly, contained 3.72-4.60 times (mean 4.09) in the diameter, 
margins rounded. Whorl contour and apertural features normal. 

Sinployea andrewi (fig. 44a-c) has a lower spire, 
more numerous and crowded radial ribs, and fewer api- 
cal cords than S. atiensis (fig. 44d-f). Sinployea peasei 
(fig. 45a-c) is larger, with the apical cording reduced in 
prominence. Society Island species have much finer 
and higher radial ribs. Sinployea rotumana (Smith) 
(Fig. 61d-f) has fewer, slightly more widely spaced and 
distinctly narrower radial ribs, a U-shaped umbilicus, 
and more crowded secondary spiral cording. 

Description. Shell of average size, with 4'/4 moderately loosely 
coiled whorls. Apex and spire moderately and almost evenly ele- 
vated, last whorl descending more rapidly, H/D ratio 0.516. Apical 
whorls slightly more than 1%, partly worn, sculpture of 10 spiral 
ribs. Postnuclear whorls with closely set, rounded, protractively 
sinuated radial ribs, 158 on the body whorl, whose interstices are less 
than twice their width. Microsculpture of 3-6 fine radial riblets be- 
tween each pair of major ribs, closely set and equally prominent 
secondary spiral cords, and barely visible spiral riblets. Sutures 
moderately impressed, whorls shouldered above, flattened laterally 
above rounded periphery, with slightly flattened basal margin. Color 
dark reddish yellow, with occasional lighter streaks. Umbilicus mod- 
erately open, V-shaped, last whorl decoiling more sharply, contained 
4.13 times in the diameter. Aperture subcircular, strongly flattened 
laterally above periphery, slightly flattened basally, inclined about 
30 from shell axis. Height of holotype 1.61 mm., diameter 3.12 mm. 

Holotype. Cook Islands: Mangaia, Arakiore 
Oneroa, % mile from shore at 125 ft. elevation. Col- 
lected by P. H. and M. Buck on February 16, 1930. 
BPBM 97652. 

Range. Mangaia, Cook Islands. 

Paratypes. Mangaia: Arakiore Oneroa, % mile 
from shore at 125 ft. elevation (82 specimens, BPBM 
97652-3); Oneroa, '/2 mile inland under stones at 115 ft. 
elevation (1 specimen, BPBM 97544). 

Remarks. The appearance of Sinployea andrewi 
(fig. 44a-c) is very similar to that of S. neglecta (fig. 
41d-f) from Huahine, Society Islands, and it is difficult 
to find shell characters that immediately separate in- 
dividuals of the two species. In general, S. andrewi has 
less sharply impressed sutures and wider, more 
crowded radial ribs with a lesser average number of 




FIG. 44. a-c, Sinployea andrewi, new species. Arakiore, Mangaia, Cook Islands. Holotype. BPBM 97652; d-f, Sinployea atiensis (Pease). 
Matai, Atiu, Cook Islands. BPBM 95117. Scale lines equal 1 mm. (a-c. MM; d-f. SH). 



100 



atiensis 



TABLE XX. - LOCAL VARIATION IN COOK ISLAND SINPLOYEA, I. 



NUMBER OF 
SPECIMENS 



RIBS 



HEIGHT 



DIAMETER 



H/D RATIO 



Aitutaki 
Zurich 



Atiu 

BPBM 95117 



Zurich 



BPBM 87407, 
94942, -98, 
95036.-141 



85.3+1.45 
(83-88) 



andrewi 



BPBM 96652-3 



avanaensis 



Sta. 18a' 

FMNH 144631-2 



Sta. 32a 

FMNH 144671 



24 



16 



94.42.64 

(86-102) 



233.3 



(175-290) 



1.360.038 

(1.29-1.42) 



1.57+0.034 

(1.46-1.79) 

1.550.024 

(1.49-1.62) 

1.640.059 

(1.39-1.92) 



1.54+0.018 

(1.29-1.69) 



1.54+0.031 

(1.32-1.81) 

1.47+0.062 

(1.32-1.74) 



2.510.039 

(2.45-2.58) 



2.71+0.063 

(2.55-3.18) 

2.650.058 

(2.50-2.83) 

2.750.063 

(2.55-3.01) 



2.95+0.022 

(2.68-3.15) 



3.210.060 

(2.86-3.68) 

3.220.150 

(2.89-3.82) 



0.5400.0069 

(0.527-0.551) 



0.5810.0083 

(0.543-0.616) 

0.5840.&067 

(0.573-0.609) 

0.596+0.013 

(0.539-0.642) 



0.519+0.0043 

(0.472-0.568) 



0.479+0.0062 

(0.438-0.516) 

0.459+0.0086 

(0.427-0.489) 



Zurich, AMS 



ANSP 47700 



BPBM 2317 



110.06.60 

(92-129) 

130.2+8.72 

(104-154) 



129 



(116-142) 



1.850.056 

(1.72-2.05) 

1.96*0.087 

(1.79-2.22) 

2.090.087 

(1.89-2.38) 



3.500.068 

(3.31-3.68) 

3.550.109 

(3.28-3.81) 

3.79+0.088 

(3.61-4.11) 



0.5290.0076 

(0.518-0.559) 

0.5510.0088 

(0.534-0.583) 

0.5510.0103 

(0.523-0.580) 



WHORLS 



UMBILICUS 



D/U RATIO 



APICAL CORDS 



atiensis 

Aitutaki 

Zurich 4 l/4-(4 1/8-4 1/4) 



0.59+0.010 

(0.58-0.61) 



4.21+0.012 10.670.335 

(4.19-4.23) (10-11) 



Atiu 
BPBM 4 l/4+(4 1/8-4 5/8) 



Zurich 4 l/4+(4 1/8-4 1/2) 



BPBM 



4 l/2-(4 1/4-4 7/8) 



0.640.025 

(0.56-0.82) 

0.61+0.017 

(0.56-0.66) 

0.650.019 

(0.59-0.72) 



4.16+0.090 

(3.77-4.59) 

4.360.148 

(3.87-4.70) 

4.35+0.088 

(4.05-4.82) 



11. 86 0.669 
(9-14) 



10.50.646 
(9-12) 



andrewi 

BPBM 4 1/8-K4-4 3/8) 

avanaensis 

Sta. 18a 4+(3 3/4-4 3/8) 

Sta. 32a 4 l/8+(4-4 1/2) 

proxima 

Zurich 4 l/4-(4-4 3/8) 

ANSP 4 1/4-K4-4 5/8) 

BPBM 4 l/2-(4 1/4-4 7/8) 



0.720.010 

(0.63-0.79) 



0.67+0.025 

(0.53-0.86) 

0.750.057 

(0.63-0.99) 



0.650.022 

(0.59-0.72) 

0.69+0.029 

(0.59-0.76) 

0.68+0.028 

(0.59-0.76) 



4.09+0.041 

(3.74-4.45) 



4.88+0.121 

(4.24-5.76) 

4.50+0.168 

(3.87-4.96) 



5.3510.107 

(5.00-5.55) 

5.160.246 

(4.48-5.75) 

5.530.206 

(4.96-6.16) 



9.19+0.203 
(8-11) 



101 



102 



SOLEM: ENDODONTOID LAND SNAILS 



apical ribs. The differences are sufficient that I can 
readily separate individuals on the basis of sculpture, 
but I cannot satisfactorily quantify the change. 

This species is named after Andrew Garrett, 
pioneer collector and student of Cook Island land 
shells. 

Description of soft parts. Dissection of several individuals 
showed no differences from S. peasei. Penis length was about 1.11.3 
mm., with internal sculpture typical. Partial figuring (fig. 42b) of the 
genitalia is provided. 

Radula without unusual features. 

(Based on BPBM 97652, 6 individuals). 

Sinployea peasei, new species. Figures 2a-b, 42c-e, 
45a-c. 

Patula atiensis (Pease), Garrett, 1881, J. Acad. Nat. Sci., Philadel- 
phia, 8 (4), p. 386 Rarotonga, Cook Islands (partly). 

Diagnosis. Shell rather large, diameter 2.97-4.01 mm. (mean 
3.15 mm.), with 4-5'/n normally coiled whorls. Apex and spire 
slightly and evenly elevated, last whorl descending more rapidly, 
spire protrusion a little more than Ve body whorl width, H/D ratio 
0.415- 0.558 ( mean 0.492). Apical sculpture of 9- 14 ( mean 11.4) very 
faint spiral cords (fig. 2a-b) that cannot be counted on most exam- 
ples. Postnuclear whorls with prominent, protractively sinuated, 
rounded radial ribs, 83-167 (mean 112.2) on the body whorl, whose 
interstices are usually 2-3 times their width. Ribs/mm. 8.80-14.66 
(mean 11.51). Microsculpture of fine radial riblets, 48 between each 
pair of major ribs, extremely fine spiral riblets, and irregular secon- 
dary spiral cording. Umbilicus open, V-shaped, last whorl decoiling 
more rapidly, contained 3.72-4.60 times (mean 3.93) in the diameter, 
margins rounded. Whorl contours (fig. 2a) and apertural shape typi- 
cal. 

Sinployea peasei has the apical cording (fig. 2a) 
greatly reduced in prominence, but differs only slightly 
in size and proportions from the other small Cook Is- 
land species. Sinployea modicella (Ferussac) from 
Moorea is more widely umbilicated, has finer major 
radial ribbing, and fewer, much more prominent apical 
cords (fig. Ib). 

Description. Shell large, with 4% normally coiled whorls. Apex 
and spire slightly and evenly elevated, last whorl descending more 
rapidly, H/D ratio 0.495. Apical whorls 1%, sculpture of about 12 
spiral cords, partly worn over most of surface. Postnuclear whorls 
with prominent, protractively sinuated, lamellate radial ribs, 113 on 
the body whorl, whose interstices are 2-4 times their width. Micro- 
sculpture of very fine radial riblets, 4-8 between each pair of major 
ribs, crossed by extremely fine and crowded spiral riblets, with weak 
secondary spiral cording that becomes conspicuous only in the um- 
bilicus. Sutures deep, whorls strongly rounded above, markedly flat- 
tened laterally above periphery with slightly compressed basal mar- 
gin, umbilical margin evenly rounded. Color yellow-horn, with broad 
and somewhat irregular reddish flammulations that do not fade out 
on shell base. Umbilicus broadly open, V-shaped, last whorl decoiling 
more rapidly, contained 3.62 times in the diameter. Aperture ovate, 
strongly flattened laterally above periphery, inclined about 30 from 
shell axis. Height of holotype 1.71 mm., diameter 3.45 mm. 

Holotype. Cook Islands: Rarotonga, Station 26, 
west slopes of Mt. Maungaroa at 1,400 ft. elevation. 
Collected under dead wood by Laurie Price in De- 
cember, 1964. FMNH 155947. 

Range. Widely spread over inland areas of 
Rarotonga, Cook Islands. 

Paratypes. Rarotonga (9 specimens, BPBM 2331, 
BPBM 8585): lower slopes of Mt. Maungaroa at 600 ft. 



elevation (1 specimen, BPBM 79780); Takuvaine Val- 
ley (Station 2) at 200 ft. elevation (1 specimen, FMNH 
144519); above Takuvaine stream (Station 4) at 400 ft. 
elevation (9 specimens, FMNH 144547); upper 
Takuvaine Valley (Station 5a), near Mt. Te Kou at 
1,000 ft. elevation (1 specimen, FMNH 144553); Mt. Te 
Manga, Takuvaine Valley (Station 30) at 1,200 ft. ele- 
vation (12 specimens, FMNH 144644); north slope 
(Station 3) of Mt. Maungaroa at 1,000 ft. elevation (9 
specimens, FMNH 144536); Mt. Ikurangi (Station 8) at 
800 ft. elevation (11 specimens, FMNH 144567-8); 
Taipara Valley (Station 10) at 400 ft. elevation (8 
specimens, FMNH 144578); west slope of Mt. Maun- 
garoa (Station 26) at 1,400 ft. elevation (33 specimens, 
FMNH 144645-6); Avana Valley (Station 14) at 300 ft. 
elevation (32 specimens, FMNH 144613-4); Avatiu 
Valley (Stations 11, 28) at 500-600 ft. elevation (14 
specimens, FMNH 144586, FMNH 144652); south 
slope of Mt. Te Rua Manga (Station 29) at 1,200 ft. 
elevation (9 specimens, FMNH 144654); Rutaki Valley 
(Station 13) at 400 ft. elevation (6 specimens, FMNH 
144600); Vaikapuangi Valley (Station 24), Mt. Maun- 
gatea at 900 ft. elevation (10 specimens, FMNH 
144639); summit of Mt. Te Kou (Station R-ll) at 1,930 
ft. elevation (3 specimens, FMNH 153388, FMNH 
153413). "Cook Islands" (4 specimens, BPBM 167417). 

Remarks. Garrett (1881, p. 386) recognized that 
the Rarotonga shells were slightly different from the 
Atiu and Aitutaki populations (= atiensis), but did not 
give them nomenclatural recognition. In naming this 
species after W. Harper Pease, I am recognizing his 
early encouragement of Andrew Garrett and his de- 
scription of many Pacific Island species. 

The availability of many localized sets enabled 
using this species to establish patterns of variation for 
Sinployea and to provide criteria for judging interpopu- 
lational variation. As can be seen from Table XXI, 
even with only very small numbers of adults from 
different stations, there was only minor variability. 

Description of soft parts. Foot and tail elongated, slender, not 
tapering posteriorly, rounded behind. Sole undivided, without trans- 
verse corrugations. Pedal grooves high on foot, uniting across tail, 
suprapedal distinctly weaker. No caudal foot or caudal horn. Slime 
network rectangular, more prominent on tail than head. Foot trun- 
cated anteriorly, head extended well in front. Ommatophores long. 
Gonopore slightly behind and above right rhinophore. 

Sole and tail yellow-white, head and neck with grayish suffu- 
sion. Eyespots black, stalk of ommatophores gray. 

Mantle collar (MC) rather short and thick (fig. 42c), with a wide, 
even extension of mantle glands onto lung roof. Pneumostome a 
single opening masked on each side by swellings of mantle collar. 
Anus (A) opening above, but at same level anteriorly as external 
ureteric opening (KX). A deep-grooved urinary chamber (LK) leads 
into shallow groove from anus through mantle collar. 

Pallial chamber (fig. 42c) extending '/2 whorl apically. Lung roof 
with heavy concentrations of black speckles along principal pulmo- 
nary vein (HV) and mantle glands (MG), with secondary concentra- 
tion near hindgut. Kidney (K) U-shaped, rectal lobe distinctly longer 
than pericardia!, latter about twice as long as heart (H). Rectal lobe 
of kidney lapping completely under hindgut and onto parietal wall. 
Base of kidney extended backward between loop of intestine and 
outer wall covering, rounded. Ureter (KD) sigmurethrous, U-shaped, 










a-f 





FIG. 45. a-c, Sinployea peasei, new species. Station 26, Mt. Maungaroa, 1,400 ft., Rarotonga, Cook Islands. Holotype. FMNH 155947; d-f, 
Sinployea avanaensis, new species. Station 18A, Upper Avana Valley, 600 ft, Rarotonga, Cook Islands. Holotype. FMNH 155948. Scale line 
equals 1 mm. (SH). 



103 



TABLE XXI. - LOCAL VARIATION IN SINPLOYEA PEASEI. 



NAME 



NUMBER OF 
SPECIMENS 



RIBS 



RIBS/MM. 



HEIGHT 



DIAMETER 



peasei 

FMNH 144547 
(Sta. 4) 

FMNH 144567 
(Sta. 8) 

FMNH 144664 
(Sta. 30) 

FMNH 144536 
(Sta. 3) 

FMNH 144639 
(Sta. 24) 

FMNH 144586 
(Sta. 11) 

FMNH 144652 
(Sta. 28) 

FMNH 144645 
(Sta. 26) 

FMNH 144600 
(Sta. 13) 

FMNH 144654 
(Sta. 29) 

FMNH 144578 
(Sta. 10) 

FMNH 144614 
(Sta. 14) 

BPBM 2331 



28 



26 



113.7+5.17 

(93-130) 

109.8+7.10 

(91-135) 

103.7+1.99 

(96-116) 

108.32.15 

(102-115) 

106.54.35 

(95-115) 

106.53.37 

(94-119) 

124.0+1.33 

(100-155) 

110. 1+3.07 

(83-149) 

104.7+7.70 

(96-120) 

127.215.54 

(91-167) 

121.3+7.22 

(92-156) 

114.52.00 

(88-141) 



11.57+0.478 

(9.78-13.24) 

11.08+0.366 

(10.12-11.90) 

11.13+0.186 

(10.30-11.89) 

10.970.182 

(10.39-11.52) 

L0.870.413 

(9.99-11.59) 

10.790.352 

9.58-12.12) 

11.951.23 

(8.80-14.16) 

11.55*0.309 

(9.13-14.15) 

10.49+0.774 

(9.39-11.981 

12.35+0.842 

(10.12-14.42) 

11.92+0.473 

(10.35-14.66) 

11.90+0.209 

(9.68-13.51) 



1.5010.026 

(1.38-1.58) 

1.4710.043 

(1.38-1.55) 

1.3510.022 

(1.25-1.45) 

1.52+0.058 

(1.38-1.84) 

1.4610.041 

(1.38-1.58) 

1.5510.055 

(1.38-1.74) 

1.7110.132 

(1.45-1.84) 

1.8410.020 

(1.28-1.71) 

1.5510.033 

(1.51-1.61) 

1.62+0.094 

(1.41-1.94) 

1.6310.054 

(1.41-1.84) 

1.51+0.020 

(1.32-1.71) 

1.88+0.070 

(1.69-2.12) 



3.13+0.045 

(3.03-3.36) 

3.16+0.148 

(2.86-3.85) 

2.97+0.056 

(2.66-3.16) 

3.14+0.058 

(2.93-3.32) 

3.13+0.112 

(2.89-3.42) 

3.150.033 

(2.99-3.29) 

3.340.132 

(3.09-3.65) 

3.10+0.032 

(2.76-3.42) 

3.18+0.121 

(3.03-3.321 

3.24+0.222 

(2.80-4.01) 

3.22+0.083 

(2.83-3.52) 

3.0810.030 

(2.80-3.45) 

3.6110.070 

(3.41-3.94) 



H/D RATIO 

Sta. 4 0.48010.0062 

(0.457-0.500) 



0.49410.0154 

(0.475-0.540) 

Sta. 30 0.455+0.0075 

(0.415-0.489) 

Sta. 3 0.48410.0154 

(0.440-0. 554) 

Sta. 24 n. 46910. 0051 

(0.458-0.478) 

Sta.U 0.49110.0145 

(0.442-0.546) 

Sta. 28 0.51310.0186 

(0.468-0.558) 

Sta. 26 0.49310.0049 

(0.448-0.538) 

Sta. 13 0.48710.0235 

(0.455-0.533) 

Sta. 29 0.50110.0065 

(0.484-0.521) 

Sta. 10. 0.50610.0092 

(0.463-0.535) 

Sta. 114 0.49210.0040 

(0.447-0.533) 

BPBM 2331 0.52110.0166 

(0.464-0.587) 



WHORLS 

4 3/8-(4 1/4-4 1/2) 

4 l/4-(4 1/8-4 3/8) 
4+(4-4 1/4) 

4 l/4+(4 1/8-4 1/2) 

4 3/8-(4 1/4-4 1/2) 

4 3/8-(4 1/8-4 1/2) 

4 5/8+(4 1/4-5 1/8) 

4 3/8+(4 1/8-4 3/4) 

4 3/8-(4 1/4-4 3/8) 

4 l/2-(4 1/8-5 1/8) 

4 3/8-(4 1/4-4 1/2) 

4 l/4+(4 1/8-4 1/2) 
4 5/8+(4 1/2-5) 



UMBILICUS 

0.7610.021 

(0.69-0.86) 

0.7910.045 

(0.69-0.92) 

0.73+0.025 

(0.63-0.86) 

0.8010.035 

(0.72-0.99) 

0.7810.043 

(0.69-0.86) 

0.79+0.015 

(0.72-0.86) 

0.88+0.068 

(0.76-1.05) 

0.8110.017 

(0.63-0.99) 

0.8610.038 

(0.79-0.92) 

0.8510.109 

(0.66-1.25) 

0.81+0.039 

(0.66-1.02) 

0.7910.010 

(0.66-0.89) 

0.9910.038 

(0.82-1.12) 



D/U RATIO 

4. 1110.059 

(3.92-4.38) 

4.00+0.092 

(3.75-4.18) 

4.06+0.081 

(3.54-4.29) 

3.9410.1211 

(3.26-4.21) 

4.02+0.1411 

(3.69-4.38) 

3.97+0.050 

(3.80-4.14) 

3.83+0.138 

(3.47-4.13) 

3.88+0.057 

(3.33-4.43) 

3.7210.090 

(3.61-3.83) 

3.9210.200 

(3.21-4.35) 

4.0110.109 

(3.45-4.50) 

3.90+0.046 

(3.52-4.60) 

3.6510.144 

(3.22-4.36) 



104 



SYSTEMATIC REVIEW 



105 



evenly expanded, a strip of lung roof visible between arms of ureter. 
At anterior end of rectal kidney lobe, ureter passes alongside hindgut 
on lung roof forward to pneumostome. Heart (H) fairly large, angling 
slightly from parallel to hindgut. Principal pulmonary vein (HV) 
slender, flanked by black speckling, paralleling hindgut until an- 
terior quarter of pallial cavity, then angling toward anus, finally 
reaching mantle glands. No conspicuous secondary venation. 

Ovotestis (G) imbedded in first quarter whorl above stomach- 
intestine reflexion, consisting of a single clump of varyingly split 
finger-like lobes, internally with iridescent rods. First part of her- 
maphroditic duct (GD) a very thin collecting tubule, expanding to a 
slender, opaque iridescent tube for length of stomach to middle of 
albumen gland (GG), then narrowing to a sinuous narrow tube just 
before entering talon (GT). Albumen gland small, darker in color 
than prostate (DG), cradled between intestinal loops and base of 
kidney in body cavity just above and morphologically below apex of 
pallial cavity. Head of spermatheca (S) buried between base of albu- 
men gland and head of prostate. Talon a slender tube with bulbously 
expanded head down to carrefour, hermaphroditic duct entering lat- 
erally on head of carrefour, which tapers from secondary expansion 
below apex to head of prostate-uterus. Prostate (DG) short, of large, 
white acini entering a groove in uterine wall. Uterus (UT) with 2 
chambers, upper (UT,) a very thin-walled sac partly filled with 
spongy tissue, opening through a glandular collar into a lower 
chamber, white in color, with thick glandular walls, smooth except 
for 2 pilasters extending into muscular-walled free oviduct (UV). 

Vas deferens (VD) from base of prostate at level of spermathecal 
stalk contraction, wide at first with internal pilasters, tapering down 
to a thin tube by level of penioviducal angle, passing up along penis, 
but not bound to it, until entering epiphallus (E). Latter a swollen 
muscular tube with internal longitudinal pilasters entering penis (P) 
through a small vergic papilla (PV, fig. 42e). Penial retractor (PR) 
arising on diaphragm near apex of pallial cavity, inserting in a U-fan 
around base of epiphallus. Penis proper about 1.1-1.3 mm. long, 
expanded apically with a minor constriction marking internal mus- 
cular collar, then tapering slowly to base of main glandular part, 
finally continuing as a slender tube to atrium. Internally (fig. 42d), 
penis with a cup-shaped vergic papilla (PV) surrounded by a muscu- 
lar collar of 3 circular pilasters. Below this, a huge U-shaped muscu- 
lar stimulatory pad occupies rest of penial bulge. Lower part and 
section between arms of "U" with walls longitudinally folded. 
Atrium (Y) fairly long, without distinctive internal sculpture. 

Free oviduct (UV) a short tube with smooth muscular walls ex- 
ternally, internally with smooth glandular surface, opening into va- 
gina (V) through a glandular collar. Spermatheca (S) and base of 
vagina swollen with glandular walls, tapering above to a slender 
stalk, then an oval head lying in angle between prostate and albu- 
men gland. Vagina (V) long, swollen apically with glandular walls, 
lower part a narrow tube with weak pilasters. 

Buccal mass and esophagus typical, white salivary glands unit- 
ing above esophagus. Stomach expanding just above apex of pallial 
cavity, extending % of a whorl apically, occupying parietal and most 
of palatal wall surfaces. About V-i of way from apex, stomach narrows 
from base, preparatory to stomach-intestine junction. First part of 
intestine angling down to columellar margin, running along this 
edge to apex of pallial cavity, looping up under kidney to near mid- 
point of whorl, reflexing backward for about twice its width, then 
turning up to parietal-palatal margin and forward as hindgut. Dis- 
tance between apex of pallial cavity and stomach greatly reduced, 
loops of intestine very short, hindgut only reaching parietal-palatal 
angle at base of kidney. 

Free retractor muscle system as in Champa coma. 

Enervation of penis from right cerebral ganglion. 

(Based on FMNH 144645, several whole examples.) 

Sinployea avanaensis, new species. Figures 43a-e, 
45d-f. 

Diagnosis. Shell rather large, diameter 2.86-3.82 mm. (mean 
3.21 mm.), with 3%-4!4 normally coiled whorls. Apex and spire mod- 
erately and evenly elevated, last whorl descending more rapidly, 
spire protrusion l h to l k of body whorl width, H/D ratio 0.427-0.516 



(mean 0.474). Apical sculpture of about 10-14 very fine spiral cords, 
rarely distinct enough to be tallied. Postnuclear whorls with very 
fine and crowded, protractively sinuated, narrow radial ribs, 175- 
290 (mean 225.8) on the body whorl, whose interstices are less than 
twice their width. Ribs/mm. 16.28-25.06 (mean 20.82). Micro- 
sculpture of fine radial riblets, 2-4 between each pair of major ribs, 
barely visible spiral riblets and weak secondary spiral cording visi- 
ble on upper surface and basal margins. Umbilicus narrow, V- 
shaped, last whorl decoiling more rapidly, contained 3.87-5.69 times 
(mean 4.78) in the diameter. Whorl contour typical, aperture a little 
more compressedly ovate than normal. 

The extremely fine and crowded radial ribs, dark 
coloration, and narrower umbilicus effectively sepa- 
rate Sinployea avanaensis (fig. 45d-f) from S. peasei 
(fig. 45a-c). Other Rarotonga species are much larger 
and have fewer or reduced radial ribs. Of the extralimi- 
tal species, only the much smaller S. navutuensis (fig. 
68d-f) from Lau and S. godeffroyana (fig. 69a-c) from 
Viti Levu approach the rib spacing of S. avanaensis. No 
other Sinployea has more than 175 ribs on the body 
whorl. 

Description. Shell rather large, with slightly less than 4'/4 
normally coiled whorls. Apex and spire slightly and evenly elevated, 
last whorl descending more rapidly, H/D ratio 0.510. Embryonic 
whorls 1%, sculpture of about 13 very fine spiral cords, somewhat 
worn. Postnuclear whorls with fine, narrow, very crowded, strongly 
protractively sinuated radial ribs, about 203 on the body whorl, 
whose interstices are no more than twice their width. Microsculpture 
of fine radial riblets, 2-4 between each pair of major ribs, crossed by 
barely visible spiral riblets, with a weak development of secondary 
spiral cording visible on the upper spire, becoming strong on shell 
base and in umbilicus. Sutures deep, whorls strongly rounded above, 
flatly rounded laterally above periphery, with distinctly flattened 
basal margin, umbilical margin strongly rounded. Umbilicus nar- 
row, V-shaped, last whorl decoiling much more rapidly, contained 
5.33 times in the diameter. Color dark reddish brown. Aperture com- 
pressedly ovate, slightly flattened laterally above periphery, dis- 
tinctly flattened basally, inclined about 25 from shell axis. Height of 
holotype 1.61 mm., diameter 3.16 mm. 

Holotype. Cook Islands: Rarotonga, Station 18a, 
upper Avana Valley at 600 ft. elevation. Collected by 
Laurie Price on lichens and moss-covered branches in 
December, 1964. FMNH 155948. 

Range. Upper Avana Valley at 300-1,000 ft. 
elevation, Rarotonga, Cook Islands. 

Paratypes. Rarotonga: Avana Valley between 
300 and 1,000 ft. elevation (Stations 18, 18a, 32a, R-13, 
R-16) under loose bark or fungus and in crevices on 
moss and lichen-covered tree trunks and branches up 
to 15 ft. above ground level (137 specimens, FMNH 
144631-2, FMNH 144671, FMNH 153381, FMNH 
153383-5). 

Remarks. At the lowest elevation (Station R-13, 
300 ft.) specimens were found only 1-6 ft. above ground 
level, whereas at the higher stations examples were 
taken up to 15 ft. above the ground. Probably they 
move higher, but practical problems in collecting pre- 
vented determination of upper limits. All examples 
were taken in and under moss and lichens or under 
loose bark, except for a few from Station 18 that were 
found under dead wood on the ground. Conditions were 
quite dry at the time, and the latter occurrence may 
represent a partial retreat after moisture. Size and 
shape variation is minimal (table XX). 



106 



SOLEM: ENDODONTOID LAND SNAILS 



The very reduced apical sculpture, almost uniform 
color, very crowded and numerous radial ribs, much 
narrower umbilicus, and more compressedly ovate 
aperture immediately separate S. avanaensis from the 
common S. peasei. The latter has been taken in lower 
Avana Valley (Station 14) at about 300 ft. elevation. It 
is widely distributed on other parts of Rarotonga and 
has been found on the summit of Mt. Te Kou, which 
heads both Takuvaine and Avana Valleys. The two 
species are probably allopatric, with S. peasei living 
lower in Avana than S. avanaensis. 

Description of soft parts. Foot and tail as in S. peasei, length 
slightly less than shell diameter (fig. 43a). Sole undivided, pedal 
grooves uniting above tail, suprapedal distinctly weaker. No caudal 
horn, middorsal groove weak, slime network typical. Gonopore 
slightly behind and above right rhinophore. Body color yellow-white, 
with gray-black speckling on head and neck region. Mantle collar 
with tapered edge, a weak glandular extension onto pallial roof. 
Pneumostome shielded by a weak thickening of collar to form a right 
mantle lobe. Anus (A) and external ureteric pore (KX) opening at 
angle just inside pneumostome. 

Pallial region (fig. 43b) extending apically M> whorl, lung roof 
clear, without granules. Kidney (K) bilobed, rectal lobe distinctly 
longer and broader than pericardial, former lobe crossing hindgut to 
reach parietal wall, pericardial lobe about twice as long as heart. 
Apex tapering, U-shaped. Ureter (KD) complete, opening alongside 
anus, a strip of lung roof visible between ureter arms. Heart (H) 
almost parallel to hindgut, proportionately broad. Principal pulmo- 
nary vein (KV) angling toward pneumostome, without conspicuous 
lateral branches. Hindgut iHG) starting at apex of kidney, following 
parietal-palatal margin to anus. 

Ovotestis (fig. 43c) occupying slightly less than M> whorl above 
stomach reflexion, composed of 5-6 long, single or bifurcated, 
finger-like lobes lying parallel to whorl sides, merging irregularly 
near base. Hermaphroditic duct (GD) at first a very slender tubule, 
becoming abruptly swollen below stomach apex, iridescent until ab- 
ruptly narrowing to reflex up albumen gland surface to base of talon. 
Albumen gland (GG) oval, deeply indented by head of spermatheca, 
intestinal loop, and digestive gland. Talon (GT) with circular head, 
hermaphroditic duct entering shaft just below head area, tapering to 
head of prostate-uterus. Prostate (fig. 43d) of long, finger-like acini, 
sometimes bifurcated, opening into a closed tube contained within 
uterine chamber. Uterus (UT) as in S. peasei. 

Vas deferens (VD) wide and with glandular walls at first, nar- 
rowing before penioviducal angle, becoming a slender tube passing 
alongside penis to epiphallus. Latter an enlarged muscular tube with 
longitudinal pilasters, ending in a narrow neck before a muscular 
collar leading into penis. Penial retractor (PR) arising from dia- 
phragm near pallial cavity apex, inserting in a U-shaped fan around 
base of epiphallus. Penis (P) about 1.4-1.6 mm. long, swollen on 
apical '/2-%, a slight constriction marking end of muscular collar, 
lower '/s-'/z a slender tube. Internally (fig. 43e) with a small verge 
(PV) nestled in a muscular collar generally containing two circular 
bands and followed by the typical doughnut cushion. Lower portion 
without distinctive sculpture. Atrium (Y) relatively long, equal in 
diameter to basal portion of penis, without special sculpture. 

Free oviduct (UV) tapering, internally with glandular tissue and 
weak pilasters, opening above end of glandular zone in spermathecal 
region. Spermatheca (S) with head lying next to albumen gland, 
expanded near base with strong glandular zone that ends abruptly 
below junction with free oviduct. Vagina (V) tapering sharply after 
end of glandular zone, without special structures present. 

Free muscle system as in S. peasei. Buccal retractors not split, 
esophagus typical. Stomach extending for % of a whorl, occupying 
normal position, expanding and narrowing abruptly. Intestinal loops 
compressed into 'Ae of a whorl between pallial cavity apex and 
stomach, tapered part of kidney extending well above much of loops. 
Digestive glands extending about l*/4 whorls above ovotestis apex, 



reduced to narrow strips along stomach, expanding in short area 
between stomach and kidney apex. 

(Based on FMNH 144671, Station 32a, 4 adults, largest 3.49 
mm. in diameter with 4V4+ whorls.) 

Sinployea proxima (Garrett, 1872). Figure 46a-c. 

Pitys proxima Garrett, 1872, Am. J. Conchol., 7 (4), p. 230, pi. 19. 
fig. 24 common in several valleys on Rarotonga, Cook Islands. 

Patula proxima (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 5, 
p. 93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p. 
386. 

Helix proxima (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p. 
543. 

Helix (Patula) proxima (Garrett), Tryon, 1887, Man. Conchol., (2) 
3, p. 39, pi. 8, figs. 41-43. 

Endodonta (Charopa) proxima (Garrett), Pilsbry, 1893, Man. Con- 
chol., (2)9, p. 35. 

Diagnosis. Shell relatively large, diameter 3.26-4.08 mm. 
(mean 3.61 mm.), with 44% normally coiled whorls. Apex and spire 
slightly to moderately and evenly elevated, rounded above, last 
whorl descending much more rapidly, spire protrusion about 'A body 
whorl width, H/D ratio 0.519-0.583 (mean 0.544). Apical sculpture of 
12-15 very weak spiral ribs, usually too fine or worn for accurate 
counting. Postnuclear whorls with fine, narrow, prominent, strongly 
protractively sinuated radial ribs, 92-154 (mean 119.7) on the body 
whorl, often irregularly spaced, whose interstices usually are 2-3 
times their width. Ribs/mm. 8.91-12.96 (mean 10.75). Micro- 
sculpture of fine radial riblets, 3-8 between each pair of major ribs, 
crossed by much finer and more crowded spiral riblets. Umbilicus 
narrow, U-shaped, last whorl decoiling slightly more rapidly, con- 
tained 4.48-6.16 times (mean 5.35) in the diameter, margins 
rounded. Whorl contours typical, sutures unique in distinct channel- 
ing after first postnuclear. 

Sinployea proxima (fig. 46a-c) is immediately rec- 
ognizable by its channeled suture. Its reduced apical 
sculpture and relatively narrow umbilicus separate it 
from most Rarotongan species. Sinployea avanaensis 
(fig 45d-f) is similar in umbilical size, but has a normal 
suture, much more numerous and crowded radial ribs, 
and a distinctly more elevated spire. 

Description. Shell larger than average, with 4% moderately 
tightly coiled whorls. Apex and spire moderately elevated, rounded 
above, body whorl descending distinctly more rapidly, H/D ratio 
0.557. Apical whorls 1%, sculpture eroded with traces of weak spiral 
ribs remaining. Postnuclear whorls with moderately prominent, very 
crowded, protractively sinuated radial ribs, 154 on the body whorl, 
whose interstices are about twice their width. Microsculpture of fine, 
radial riblets, 2-6 between each pair of major ribs, crossed by 
slightly finer and more crowded spiral riblets, with a secondary 
sculpture of rather widely spaced spiral cords that become less dis- 
tinct on body whorl. Sutures deeply channeled, whorls flatly rounded 
laterally above periphery and on basal margin. Color light yellow- 
horn with prominent, regular, reddish flammulations. Umbilicus 
narrow, U-shaped, slightly and regularly decoiling, contained 5.24 
times in the diameter. Aperture ovate, flattened laterally above 
periphery and on basal margin, inclined about 20 from shell axis. 
Columellar and basal wall with moderate white callus. Height of 
lectotype 2.11 mm., diameter 3.79 mm. 

Lectotype. Cook Islands: Rarotonga. Collected by 
Andrew Garrett. ANSP 47700. 

Range. Under stones and among rotting wood in 
several valleys on Rarotonga, Cook Islands. 

Paratypes.BPBM 2317, ANSP 47700. 

Material. Rarotonga (14 specimens, AMS, 
Zurich). 






a be 



def 




FIG. 46. a-c, Sinployea proximo (Garrett). Rarotonga, Cook Islands. Paratype. BPBM 2317; d-h, Sinployea planospira (Garrett). g-h, 
Details of microsculpture. Rarotonga, Cook Islands. Lectotype. Zurich. Scale lines equal 1 mm. (a-c, SH; d-h, SG). 



107 



108 



SOLEM: ENDODONTOID LAND SNAILS 



Remarks. Although Garrett (1872, p. 230; 1881, 
p. 386) reported this as a "common" or "not uncommon" 
species found in several valleys, no additional material 
has been collected. The scattered Museum sets show 
clear indication of trading bias (table XX), with the 
BPBM residue of Garrett' s personal collection contain- 
ing the largest shells, the types in the ANSP only 
slightly smaller, and the specimens that eventually 
reached Mousson (now Zurich) and Hedley (AMS) 
noticeably smaller. 

Channeling of the suture is relatively uncommon 
in the Pacific Basin endodontoids, being found 
elsewhere in the New Caledonian Andrefrancia al- 
veolus (Gassies) (see Solem, 1961, pp. 454456, fig. 12) 
and the Bismarck Lagiuala macroglyphis (Rensch) (see 
p. 188). Ignoring the sutural alteration, S. proximo is 
only a slightly enlarged, more narrowly umbilicated 
derivative from the peasei stock. 

Sinployea planospira (Garrett, 1881). Figure 46d-h. 

Patula planospira Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 

(4), p. 388 Rarotonga, Cook Islands. 
Helix (Patula) planospira (Garrett), Tryon, 1887, Man. Conchol., 

(2)3, p. 41. 
Endodonta (Champa) planospira (Garrett), Pilsbry, 1893, Man. 

Conchol., (2)9, p. 35. 

Diagnosis. Shell relatively large, diameter 3.444.24 mm. 
(mean 3.74 mm.), with 6'/4-7'/4 very tightly coiled whorls. Apex and 
early spire flat, body whorl descending rapidly, spire protrusion 
about Vs-'/4 body whorl width, H/D ratio 0.439-0.524 (mean 0.468). 
Apical sculpture eroded in available material, traces of spiral cord- 
ing visible in one umbilicus. Postnuclear whorls with narrow, prom- 
inent, protractively sinuated radial ribs. 108-148 (mean 128.0) on 
the body whorl, whose interstices are 2-4 times their width. 
Ribs/mm. 10.05-11.19 (mean 10.62). Microsculpture of fine radia! 
riblets, 5-8 between each pair of major ribs, slightly finer and more 
crowded spiral riblets, and secondary spiral cording more clearly 
visible in umbilicus. Umbilicus widely open, cup-shaped, regularly 
decoiling, contained 2.75-2.98 times (mean 2.90) in the diameter, 
margins strongly rounded. Whorls strongly rounded above, flattened 
laterally and on basal margin. 

Sinployea planospira (fig. 46d-h) has a wide um- 
bilicus and much higher whorl count than any other 
Polynesian species. Sinployea canalis (fig. 49a-c) is 
much larger, has one whorl less, and a much less ele- 
vated spire, although it is very similar in rib count and 
umbilical proportions (table XIX). 

Description. Shell larger than average, with 6'A tightly coiled 
whorls. Apex and spire flat, body whorl descending slightly, H/D 
ratio 0.442. Embryonic whorls 1%, sculpture mostly eroded, with 
traces of widely spaced spiral ribbing remaining. Postnuclear whorls 
with narrow, protractively sinuated radial ribs, 108 on the body 
whorl, whose interstices are 2-4 times their width. Microsculpture of 
very fine radial ribs, 5-8 between each pair of major ribs, crossed by 
slightly finer and more crowded spiral riblets with a secondary 
sculpture of spiral cording developed in the umbilicus. Sutures deep, 
whorls strongly rounded above, flattened laterally and on basal mar- 
gin. Umbilicus U-shaped, broadly open, regularly decoiling, con- 
tained 2.98 times in the diameter. Color light yellowish white with 
traces of irregularly spaced, broad, reddish maculations. Aperture 
subtriangular, strongly flattened laterally and on basal margin, in- 
clined about 15 from the shell axis. Height of lectotype 1.51 mm., 
diameter 3.42 mm. 

Lectotype. Cook Islands: Rarotonga. Collected by 
Andrew Garrett. Zoologisches Museum der Universitat 



Zurich, Mousson collection ex Museum Godeffroy in 
1882. 

Range. Rarotonga, Cook Islands. 

Paratypes. Rarotonga (2 specimens, Zurich). 

Remarks. Because the collections of the Museum 
Godeffroy were destroyed during World War II and 
Garrett (1881, p. 388) stated that all material of this 
species had been deposited in that collection, I had as- 
sumed that this was a lost species. The discovery of two 
shells in the Mousson collection at Zurich that had 
been obtained from the Museum Godeffroy in 1882 al- 
lowed recognition of this taxon. Although Mousson had 
attached a manuscript name "rarotongensis," I have no 
doubt as to their identity. A third shell, also obtained 
by Mousson from the Museum Godeffroy, was under 
the manuscript name "ortospirata Mousson." It is a 
gerontic individual. Unfortunately, the type specimen 
was damaged during handling, hence the figures have 
dotted lines indicating the upper whorls. 

The high whorl count is unmatched by other 
Polynesian Charopidae and is much more similar to 
the pattern observed in the Endodontidae. Traces of 
apical sculpture are sufficiently clear to leave no doubt 
as to its affinities. Only S. canalis of the Rarotongan 
species has the umbilicus as widely open. Differences 
are given in the diagnosis. 

Sinployea rudis (Garrett, 1872). Figure 47d-f. 

Pitys rudis Garrett, 1872, Am. J. Conchol., 7 (4), pp. 227-228, pi. 
19, fig. 18 several valleys on Rarotonga, Cook Islands. 

Patula rudis (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 5, p. 
93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p. 387. 

Patula sublaminata "Mousson" Schmeltz, 1874, Cat. Mus. Godef- 
froy, 5, p. 94 Nude name, tentatively synonymized by Garrett 
(1881). 

Helix rudis (Garrett), Pfeiffer, 1876, Monog. helic, viv., 7, p. 547. 

Helix (Patula) rudis (Garrett), Tryon, 1887, Man. Conchol., (2) 3, 
pp. 39-40, pi. 8, figs. 44-46. 

Endodonta (Charopa) rudis (Garrett), Pilsbry, 1893, Man. Con- 
chol., (2) 9, p. 35. 

Diagnosis. Shell large, diameter 3.68-4.83 mm. (mean 4.04 
mm.), with 378-4% normally coiled whorls. Apex flat, spire moder- 
ately and evenly elevated, last whorl descending slightly more 
rapidly, spire protrusion more than Vs body whorl width, H/D ratio 
0.500-0.642 (mean 0.553). Apical sculpture of 8-12 (mean 10.02) 
spiral cords and weak radial corrugations. Postnuclear sculpture of 
low, narrow, rather widely spaced protractive radial ribs on upper 
spire, becoming reduced and very irregular early on body whorl, 
absent near aperture. Microsculpture of fine radial riblets, crossed by 
finer and more crowded spiral riblets, reduced on body whorl. Um- 
bilicus narrow internally, V-shaped, body whorl decoiling much 
more rapidly, contained 4.18-5.62 times (mean 4.87) in the diameter, 
margins rounded. Whorls flattened and indented laterally above 
periphery, producing a distinct supraperipheral sulcus, strongly 
rounded on lower margins, aperture inclined about 35 from shell 
axis. 

In having a supraperipheral sulcus and losing 
major ribbing on the lower spire and body whorl, 
Sinployea rudis is quite distinctive among Polynesian 
species. 

Description. Shell relatively large, with slightly more than 4V4 
loosely coiled whorls. Apex and spire slightly but evenly elevated, 
body whorl descending more rapidly, H/D ratio 0.531. Apical whorls 







l-tL-, 



f 




FIG. 47. a-c, Sinployea decorticata (Garrett). Rarotonga, Cook Islands. Paratype. BPBM 2313; d-f, Sinployea rudis (Garrett). Rarotonga, 
Cook Islands. Paratype. BPBM 2320. Scale line equals 1 mm. (a-c, SH; d-f, MM). 



109 



110 



SOLEM: ENDODONTOID LAND SNAILS 



I'/a, sculpture of 11 prominent spiral ribs with a secondary micro- 
sculpture of radial and spiral riblets. Postnuclear whorls with nar- 
row, low, irregular, protractively sinuated radial ribs, becoming 
greatly reduced or absent on the body whorl. Microsculpture of fine 
and somewhat irregular radial riblets crossed by distinctly finer and 
more crowded spiral riblets that are visible only at 96 x magnifica- 
tion under strong lighting. Sutures deep, whorls shouldered above, 
flattened to indented laterally above periphery with gently rounded 
basal margin. Color greenish yellow-horn with extensive irregular 
reddish flammulations. Umbilicus narrow, V-shaped, regularly de- 
coiling, aperture contained 5.42 times in the diameter. Height of 
lectotype 2.27 mm., diameter 4.28 mm. 

Lectotype. Cook Islands: Rarotonga. Collected by 
Andrew Garrett. ANSP 47701. 

Range. Several valleys on Rarotonga, Cook Is- 
lands. 

Paratypes. BPBM 2320, ANSP. 

Material. Cook Islands (9 specimens, BPBM 
167420, FMNH 93820, FMNH 108559): Rarotonga (20 
specimens, BPBM 167421, FMNH 46267, ANSP 
47701). 

Remarks. Garrett (1881, p. 387) indicated that 
Sinployea rudis was common and that "we gathered 
hundreds in several valleys." In its flat spire, reduced 
sculpture, and very prominent supraperipheral sulcus, 
S. rudis is immediately separable from the other 
Rarotongan species. 

The size difference between the type set and 
BPBM material (table XXIII) undoubtedly reflects re- 
tention of large specimens by Garrett. 

Sinployea decorticata (Garrett, 1872). Figure 47a-c. 

Pitys decorticata Garrett, 1872, Am. J. Conchol., 7 (4), p. 228, pi. 

19, fig. 19 a mountain ravine on Rarotonga, Cook Islands. 
Patula decorticata (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 

5, p. 93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p. 

387. 
Helix decorticata (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p. 

542. 
Helix (Patula) decorticata (Garrett), Tryon, 1887, Man. Conchol., 

(2) 3, p. 40, pi. 8, figs. 47-49. 
Endodonta (Champa) decorticata (Garrett), Pilsbry, 1893, Man. 

Conchol., (2) 9, p. 35. 

Diagnosis. Shell very large, diameter 3.71-5.10 mm. (mean 
4.30 mm.), with 4%-5 normally coiled whorls. Apex and spire mod- 
erately to strongly elevated, body whorl descending more rapidly, 
spire protrusion about 1 k body whorl width, H/D ratio 0.488-0.590 
(mean 0.550). Apical sculpture of 8-10 (mean 9.0) fairly prominent 
spiral cords. Postnuclear whorls with high, thin, prominent, protrac- 
tively sinuated and widely spaced radial ribs, 62-103 (mean 75.0) on 
the body whorl, whose interstices are 3-4 times their width. 
Ribs/mm. 4.84-6.47 (mean 5.61). Microsculpture of very fine radial 
riblets, 10-20 between each pair of major ribs, crossed by even finer 
and more crowded spiral riblets. No secondary spiral cording. Um- 
bilicus narrow, U-shaped, last whorl decoiling more rapidly, con- 
tained 3.75-5.92 times (mean 4.32) in the diameter, margins 
rounded. Whorl contours and aperture typical. 

Sinployea decorticata (fig. 47a-c) is more elevated 
and has fewer ribs and double the number of microra- 
dial riblets found in S. otareae (fig. 50a-c). Sinployea 
tenuicostata (fig. 50d-f) is larger, more depressed, has 
fewer ribs, and a flat spire. Other Rarotongan species 
differ in their finer and more crowded ribbing. 



Description. Shell very large, with 4% normally coiled whorls. 
Apex and spire moderately elevated, body whorl descending a little 
more rapidly, H/D ratio 0.528. Apical whorls 1%, sculpture of 8 prom- 
inent spiral ribs, microsculpture obscured. Postnuclear whorls with 
low, rounded, protractively sinuated radial ribs, 73 on the body 
whorl, whose interstices are 3-4 times their width. Microsculpture 
occasionally visible as fine radial riblets, 10-20 between each pair of 
major ribs, crossed by much finer and more crowded spiral riblets. 
Sutures deep, whorls flattened laterally above periphery and on 
basal margin. Color mainly leached from shell with traces of reddish 
flammulations remaining. Umbilicus moderately open, U-shaped, 
slightly and regularly decoiling, contained 3.84 times in the diame- 
ter. Aperture subcircular, flattened laterally above periphery and on 
basal margin, inclined about 20 from the shell axis. Height of lec- 
totype 2.47 mm., diameter 4.67 mm. 

Lectotype. Cook Islands: Rarotonga. Collected by 
Andrew Garrett. ANSP 47754. 

Range. One mountain ravine on Rarotonga, 
Cook Islands. 

Paratypes.BPEM. 2313, ANSP. 

Material. Rarotonga (17 specimens, BPBM 2313, 
BPBM 167422, ANSP 47754, SMF 165709, FMNH, 

NMWC). 

Remarks. Although Garrett (1881, p. 387) con- 
sidered Sinployea decorticata and S. otareae to be "va- 
rieties" of one species, the differences in sculpture and 
shape (table XXIII) are sufficient to maintain specific 
separation. The two species were restricted to separate 
valleys "about three miles apart" (Garrett, 1881, p. 
387). 

Sinployea harveyensis (Garrett, 1872). Figure 48a-c. 

Pitys harveyensis Garrett, 1872, Am. J. Conchol., 7 (4), p. 228, pi. 

19, fig. 20 common on Rarotonga, Cook Islands. 
Patula harveyensis (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 

5, p. 93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p. 

387. 
Helix harveyensis (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p. 

542. 
Helix (Patula) harveyensis (Garrett), Tryon, 1887, Man. Conchol., 

(2)3, p. 40, pi. 8, figs. 51-53. 
Endodonta (Champa) harveyensis (Garrett), Pilsbry, 1893, Man. 

Conchol., (2) 9, p. 35. 

Diagnosis. Shell very large, diameter 3.91-5.23 mm. (mean 
4.35 mm.), with 414-5 normally coiled whorls. Apex and spire rather 
strongly elevated, slightly rounded above, body whorl descending 
more rapidly, spire protrusion about 14 body whorl width, H/D ratio 
0.475-0.641 (mean 0.550). Apical sculpture of fine spiral cords, 
eroded or too faint to count accurately. Postnuclear whorls with fine, 
irregular, closely spaced, protractively sinuated radial ribs, too 
crowded and irregular to count on body whorl, whose interstices are 
often less than their width near aperture. Microsculpture visible on 
upper spire as very fine radial riblets, 3- 7 between each pair of major 
ribs, crossed by slightly finer and more crowded spiral riblets. Um- 
bilicus rather narrow, U-shaped, regularly decoiling, contained 
3.65-5.48 times (mean 4.73) in the diameter, margins rounded. 
Whorls somewhat flattened laterally above periphery, very slightly 
on basal margin. 

Sinployea harveyensis (fig. 48a-c) is characterized 
by its crowded, irregular major radial ribs and elevated 
spire. Sinployea youngi (fig. 48d-f) has a wider um- 
bilicus, lower spire, more depressed shape, and less 
crowded radial ribbing (table XIX). 



SYSTEMATIC REVIEW 



111 



TABLE XXII. - LOCAL VARIATION IN COOK ISLAND SINPLOYEA, II. 



. 



NUMBER OF 
SPECIMENS 


RIBS 


HEIGHT 


DIAMETER 


H/D RATIO 


planospira 










Zurich 3 


128.0+20.02 
(108-148) 


1.77+0.226 
(1.52-2.22) 


3.7410.250 
(3.44-4.24) 


0.46810.0277 
(0.439-0.524) 


youngi 










Zurich 3 


102.0+3.05 
(96-106) 


2.0210.119 
(1.85-2.25) 


4.0610.154 
(3.77-4.30) 


0.49710.0139 
(0.476-0.523) 


BPBM 2315 4 


127.87.43 
(107-139) 


2.47+0.098 
(2.28-2.68) 


4.76+0.117 
(4.57-5.10) 


0.518+0.0135 
(0.493-0.553) 


ANSP 47703 3 


117.712.33 
(114-122) 


2.3810.066 
(2.32-2.52) 


4.9310.077 
(4.80-5.07) 


0.48310.0156 
(0.457-0.510) 


harveyensis 










ANSP 47756 10 




2 19+0 045 


4 12+Q 033 


531+0 0092 






(1.92-2.35) 

2 31+0 O''! 


(3.91-4.27) 

4 21 + 76 


(0.475-0.572) 
548+0 0065 


Zurich 4 




(2.25-2.35) 


(4.11-4.44) 


(0.530-0.560) 


BPBM 2343, 8 
BPBM 167423 


REDUCED 


2.6410.077 
(2.32-3.01) 


4.71+0.111 
(4.37-5.23) 


0.561+0.0085 
(0.530-0.604) 


canalis 










Zurich, AMS 4 


117.318.50 
(98-133) 


2.0310.068 
(1.85-2.15) 


4.5210.151 
(4.17-4.90) 


0.44910.0055 
(0.439-0.464) 


ANSP 47752 5 


122.0112.23 
(106-146) 


2.0910.055 
(1.89-2.19) 


4.62+0.156 
(4.04-4.90) 


0.45210.0103 
(0.427-0.481) 


BPBM 2328 5 


120.8+5.46 
(107-130) 


2.2810.119 
(2.02-2.65) 


4.7210.197 
(4.40-5.50) 


0.48410.0170 
(0.446-0.544) 




WHORLS 


UMBILICUS 


D/U RATIO 


APICAL CORDS 


planospira 
Zurich 


6 5/8(6 1/4-7 1/4) 


1 . 28+0.076 


2 90+0 075 








(1.15-1.41) 


(2.75-2.98) 




youngi 










Zurich 


4+(4-4 1/4) 


1.04+0.016 
(1.02-1.09) 


3.8710.098 
(3.68-4.00) 


11.3310.34 
(11-12) 


BPBM 


4 l/2-(4 3/8-4 5/8) 


1.3410.097 
(1.15-1.61) 


3.5710.190 
(3.15-4.06) 


10.2510.25 
(10-11) 


ANSP 


4 3/4+(4 1/2-5) 


1.33+0.115 
(1.12-1.51) 


3.75+0.323 
(3.33-4.39) 


10.6710.66 
(10-12) 


harveyensis 












4 1 / 2+ (4 3 / 8 4 7 / 8 ) 


89+0 031 


4 61+0 1 36 




Zurich 


4 1/2(4 I/ 44 5/8) 


(0.79-1.12) 
86+0 056 


(3.65-5.08) 

4 89+0 240 








(0.76-1.02) 


(4.32-5.48) 




BPBM 


4 7/8-(4 5/8-5 1/8) 


0.9910.039 
(0.82-1.18) 


4.77+0.156 
(4.11-5.29) 





canalis 










Zurich 


4 3/4-K4 1/2-5 1/8) 


1.5310.064 
(1.38-1.68) 


2.9310.030 
(2.87-3.00) 


10.2510.25 
(10-11) 


ANSP 


5-(4 5/8-5 1/4) 


1.6310.076 
(1.38-1.84) 


2.8310.054 
(2.64-2.96) 


9.60+0.60 
(8-11) 


BPBM 


5+(4 3/4-5 1/2) 


1.61+0.079 
(1.41-1.84) 


2.97+0.067 
(2.73-3.10) 


10.80+0.49 
(9-12) 



Description. Shell very large, with 4% normally coiled whorls. 
Apex and spire slightly elevated, body whorl descending a little more 
rapidly, H/D ratio 0.524. Apical whorls 1%, sculpture mainly eroded 
with traces of spiral ribbing remaining. Postnuclear whorls with low, 
rounded, irregularly spaced, protractively sinuated radial ribs, be- 
coming indistinct on body whorl. Microsculpture a lattice of coequal 
radial and spiral riblets. Sutures moderately impressed, whorls flat- 



tened laterally above periphery and on basal margin. Color light 
yellow-horn with widely spaced, reddish markings absent on base of 
shell. Umbilicus U-shaped, relatively narrow, regularly derailing, 
contained 4.66 times in the diameter. Aperture ovate, flattened lat- 
erally above periphery and slightly on basal margin, inclined about 
20 from the shell axis. Height of lectotype 2.17 mm., diameter 4.14 



TABLE XXIII. - LOCAL VARIATION IN COOK ISLAND SINPLOYEA, III. 



NUMBER OF 
SPECIMENS RIBS 


HEIGHT 


DIAMETER 


H/D RATIO 


rudis 




2.12+0.047 
(1.85-2.32) 

2.440.088 
(2.28-2.85) 


3.950.049 
(3.71-4. 

4.26+0.135 
(3.94-4. 


0.537+0.0085 
30) (0.500-0.575) 

0.5740.0165 
83) (0.520-0.642) 


ANSP 47701 10 

BPBM 2320, 6 
BPBM 167420 






decorticata 


89.513.54 
(76-103) 


2.410.118 
(2.19-2.98) 


4.290.170 
(3.94-5. 


0.562+0.0118 
10) (0.520-0.590) 


BPBM 2313 6 


BPBM 47754 4 
otareae 


73 


2.53+0.051 
(2.45-2.68) 

2.34+0.083 
(2.07-2.63) 


4.630.079 
(4.44-4. 

4.570.08A 
(4.38-4. 


0.5470.0090 
80) (0.528-0.566) 

0.511+0.0097 
90) (0.474-0.540) 


BPBM 167418-9 8 




BPBM 2335 7 


107.05.87 
(96-116) 


2.540.092 
(2.25-2.98) 


4.800.097 
(4.47-5. 


0.5290.0136 
23) (0.479-0.570) 


ANSP 47755 8 


92.293.76 
(81-110) 


2.450.039 
(2.32-2.65) 


4.830.065 
(4.54-5. 


0.507+0.0088 
00) (0.470-0.555) 


tenulcostata 


46.01.00 
(45-47) 


2.040.083 
(1.95-2.12) 


4.140.033 
(4.11-4. 


0.4910.0163 
17) (0.475-0.508) 


ANSP 47702 2 


Zurich 3 


47.71.20 
(46-50) 


2.33+0.040 
(2.25-2.38) 


4.550.192 
(4.20-4. 


0.5130.0133 
87) (0.489-0.535) 


BPBM 2325 5 


51.43.65 
(43-65) 


2.46+0.049 
(2.38-2.65) 


4.960.171 
(4.44-5. 


0.4990.0170 
46) (0.437-0.537) 




WHORLS 


UMBILICUS 


D/U RATIO 


APICAL CORDS 


rudis 


4 l/4-(3 7/8-4 3/8) 


0.800.016 
(0.76-0.92) 


4.900.111 
(4.18-5.42) 


10.430.31 
(9-12) 


ANSP 


BPBM 


4 3/8+(4 1/4-4 5.8) 


0.880.048 
(0.79-1.12) 


4.830.143 
(4.30-5.17) 


9.000.58 
(8-10) 


decort. 


4 5/8+(4 3/8-5) 


0.900.036 
(0.79-1.02) 


4.770.257 
(4.20-5.92) 


8.500.50 
(8-9) 


BPBM 


BPBM 
otareae 


4 3/4(4 1/2-5) 
4 5/8+(4 1/2-5) 


1.130.044 

(1.02-1.22) 

1.11+0.031 
(1.02-1.25) 


4.06+0.099 
(3.84-4.32) 

4.110.051 
(3.92-4.29) 


8 

10.000.46 
(8-12) 


BPBM 


BPBM 


4 7/8(4 3/4-5) 


1.22+0.036 
(1.09-1.35) 


3.910.061 
(3.71-4.22) 


9.290.42 

(8-11) 


ANSP 


4 3/4+(4 1/2-5) 


1.160.064 
(0.89-1.41) 


4.240.230 
(3.47-5.52) 


8.870.40 
(8-11) 


tenuico. 


4(3 7/8-4 1/8) 


0.8210.033 
(0.79-0.86) 


5.010.240 
(4.77-5.25) 


11.50+0.50 
(11-12) 


ANSP 


Zurich 


4 1/8(4-4 1/4) 


0.840.040 
(0.79-0.92) 


5.350.087 
(5.25-5.52) 


10.000.58 
(9-11) 


BPBM 


4 3/8-(4-4 1/2) 


1.000.074 
(0.82-1.18) 


5.02+0.327 
(4.29-6.20) 


10.000.32 
(9-11) 



112 






a-f 




FIG. 48. a-c, Sinployea harveyensis (Garrett). Rarotonga, Cook Islands. Lectotype. BPBM 2343; d-f, Sinployea youngi (Garrett). Rarotonga, 
Cook Islands. Paratype. BPBM 2315. Scale line equals 1 mm. (a-c, MM; d-f, SH). 



113 



114 



SOLEM: ENDODONTOID LAND SNAILS 



Lectotype. Cook Islands: Rarotonga. Collected by 
Andrew Garrett. ANSP 47756. 

Range. Common on Rarotonga, Cook Islands. 

Paratypes. BPBM 2343, ANSP. 

Material. Rarotonga (20 specimens, BPBM 
167423, ANSP 47756, FMNH 46258, Zurich). 

Remarks. Garrett did not state whether 
Sinployea harveyensis was widely distributed or re- 
stricted in its habitat "under rotten wood" (Garrett, 
1872, p. 228). Usually he specified if only a single val- 
ley was involved. I suspect that S. harveyensis had a 
wide range on the island. The very fine radial 
sculpture is similar to that seen in the Society Island 
Sinployea, but S. harveyensis lacks the prominent 
periostracal extensions that characterize the former. 
Garrett's retention of large examples is very clearly 
shown in Table XXII. 

Sinployea youngi (Garrett, 1872). Figure 48d-f. 

Pitysyoungi Garrett, 1872, Am. J. Conchol., 7 (4), p. 229, pi. 19, fig. 
22 a single valley on Rarotonga, Cook Islands. 

Patula youngi (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 5, p. 
93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p. 388. 

Helix youngi (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p. 543. 

Helix (Patula) youngi (Garrett), Tryon, 1887, Man. Conchol., (2) 3, 
p. 40, pi. 8, figs. 54-56. 

Endodonta (Champa} youngi (Garrett), Pilsbry, 1893, Man. Con- 
chol., (2) 9, p. 35. 

Diagnosis. Shell very large, diameter 3.75-5.07 mm. (mean 
4.54 mm.), with 3%-5 loosely coiled whorls. Apex and spire moder- 
ately and evenly elevated, last whorl descending more rapidly, spire 
protrusion about 'A body whorl width, H/D ratio 0.457-0.553 (mean 
0.499). Apical sculpture of 10-12 (mean 10.73) prominent spiral 
cords. Postnuclear whorls with prominent, large, protractively 
sinuated radial ribs, 96-139 (mean 115.4) on the body whorl, whose 
interstices are about twice their width. Ribs/mm. 7.15-9.68 (mean 
8.13). Microsculpture of fine radial riblets, 3-6 between each pair of 
major ribs, crossed by slightly lower and equally crowded spiral rib- 
lets. No secondary spiral cording visible. Umbilicus widely open, 
V-shaped, last whorl decoiling more rapidly, contained 3.15-4.39 
times (mean 3.73) in the diameter, margins rounded. Whorls 
strongly flattened laterally above periphery and slightly on basal 
margin. 

Sinployea youngi (fig. 48d-f) is very similar to S. 
harveyensis (fig. 48a-c) but differs in its more de- 
pressed shape, lower spire, wider umbilicus, and less 
crowded ribbing. Sinployea canalis (fig. 49a-c) has a 
much wider umbilicus, much more depressed shape, 
greater whorl count, and more laterally compressed 
whorl contour. Other species from Rarotonga differ 
most obviously in their more widely spaced ribbing or 
much smaller size. 

Description. Shell very large, with 4% relatively loosely coiled 
whorls. Apex and spire slightly and evenly elevated, body whorl 
descending more rapidly, H/D ratio 0.510. Apical whorls 1M>, 
sculpture of 10 prominent spiral cords, microsculpture obscured. 
Postnuclear whorls with prominent, rounded, protractively sinuated 
radial ribs, about 117 on the body whorl where they become slightly 
irregular, whose interstices are 3-5 times their width. Micro- 
sculpture a lattice of very fine radial riblets, 3-6 between each pair 
of major ribs, and slightly smaller, more crowded, spiral riblets. Su- 
tures moderately impressed, whorls flattened laterally above 
periphery and basally. Color dark reddish purple-brown. Umbilicus 
V-shaped, moderately open, regularly decoiling, contained 4.39 



times in the diameter. Aperture ovate, strongly flattened laterally 
above periphery and basally, inclined about 20 from the shell axis. 
Height of lectotype 2.50 mm., diameter 4.91 mm. 

Lectotype. Cook Islands: Rarotonga. Collected by 
Andrew Garrett. ANSP 47703. 

Range. One valley on Rarotonga, Cook Islands. 

Paratypes. BPBM 2315, ANSP. 

Material. Rarotonga (13 specimens, ex BPBM 
2339, FMNH 46502, Zurich). 

Remarks. Although reported as being collected 
"in one valley, the habitat of the typical decorticata" 
(Garrett, 1881, p. 388), it was not specified if the 
species were sympatric or allopatric. Sinployea decor- 
ticata has fewer and much more widely spaced radial 
ribs and a more elevated spire than does S. youngi. 
Sinployea harveyensis is more similar, but has the api- 
cal cords reduced and the radial ribbing quite crowded 
and irregular on the body whorl besides differing in 
proportions (table XXII). 

Sinployea youngi was "A somewhat rare species. 
. . . On the ground in damp woods" (Garrett, 1872, p. 
229). 

Sinployea canalis (Garrett, 1872). Figure 49a-c. 

Pitys canalis Garrett, 1872, Am. J. Conchol., 7 (4), p. 227, pi. 19, 
fig. 17 one valley on Rarotonga, Cook Islands. 

Patula canalis (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 5, p. 
93; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p. 385. 

Helix canalis (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p. 546. 

Helix (Patula) canalis (Garrett), Tryon, 1887, Man. Conchol., (2) 3, 
p. 39, pi. 8, figs. 38-40. 

Endodonta (Charopa) canalis (Garrett), Pilsbry, 1893, Man. Con- 
chol., (2) 9, p. 35. 

Diagnosis. Shell very large, diameter 4.01-5.46 mm. (mean 
4.63 mm.), with 4M>-5'/2 tightly coiled whorls. Apex and early spire 
' fiat or barely elevated, lower spire descending slightly, body whorl 
descending sharply, spire protrusion about 'Ao body whorl width, H/D 
ratio 0.427-0.544 (mean 0.463). Apical sculpture of 8-12 (mean 
10.21) prominent spiral cords. Postnuclear whorls with fine, narrow, 
rather closely spaced, protractively sinuated radial ribs, 98-146 
(mean 119.8) on the body whorl, whose interstices are 3-4 times 
their width. Ribs/mm. 7.07-9.74 (mean 8.22). Microsculpture of 
rather widely spaced radial riblets, 3-5 between each pair of major 
ribs, crossed by slightly finer and more crowded spiral riblets. Occa- 
sionally weak secondary spiral cording visible in umbilicus and su- 
tures. Umbilicus broadly open, cup-shaped, regularly decoiling, con- 
tained 2.64-3.10 times (mean 2.91) in the diameter. Sutures very 
deep, whorls shouldered above, nearly vertically compressed on outer 
margin, basal margin strongly rounded, aperture inclined about 25 
from shell axis. 

The widely open umbilicus, strongly laterally 
compressed whorls, fine ribbing, and large size char- 
acterize Sinployea canalis (fig. 49a-c). Other Cook Is- 
land species of similar size have much more widely 
spaced ribbing and narrower umbilici (S. otareae, fig. 
50a-c, and S. tenuicostata, fig. 50d-f) or normally con- 
toured whorls and a narrower umbilicus (S. youngi, fig. 
48d-f). The slightly larger Samoan S. complementaria 
(fig. 56a-f) has a much narrower umbilicus, fewer and 
more widely spaced radial ribs, and many more micro- 
radial riblets. 

Description. Shell very large, with slightly less than 5!4 rather 
tightly coiled whorls. Apex and spire slightly sunken, body whorl 



SYSTEMATIC REVIEW 



115 






FIG. 49. a-c, Sinployea canalis (Garrett). Rarotonga, Cook Is- 
lands. Paratype. BPBM 2328. Scale line equals 2 mm. (SH). 



descending moderately, H/D ratio 0.427. Apical whorls 1%, sculpture 
of 8 prominent spiral ribs with a barely visible radial microsculpture. 
Postnuclear whorls with low, rounded, relatively crowded, protrac- 
tively sinuated radial ribs, 106 on the body whorl, whose interstices 
are 2-3 times their width. Microsculpture a lattice of fine radial 
riblets, 3-5 between each pair of major ribs, crossed by slightly finer 
and more crowded spiral riblets with a secondary sculpture of spiral 
cording visible in suture and umbilicus. Sutures deep, whorls shoul- 
dered above, flattened laterally above periphery and on basal mar- 
gin. Color yellowish horn with extensive vague reddish flammula- 
tions. Umbilicus broadly open, cup-shaped, regularly decoiling, 
contained 2.96 times in the diameter. Aperture subovate, strongly 
flattened laterally above periphery and on basal margin, inclined 
about 25 from the shell axis. Height of lectotype 2.07 mm., diameter 
4.77 mm. 

Lectotype. Cook Islands: Rarotonga. Collected by 
Andrew Garrett. ANSP 47752. 

Range. One valley on Rarotonga, Cook Islands. 
Paratypes.BPBM 2328, ANSP. 

Material. Rarotonga (13 specimens, AMS, 
Zurich, SMF 165456). 

Remarks. The strong lateral compression of the 
body whorl is not duplicated by any of the other 
Polynesian species and provides a strong contrast (fig. 
49b) to the other Rarotongan species. Garrett (1872, p. 
227) recorded this as a "somewhat rare species, found 
on the ground in damp forests, and confined to a single 
valley." Measured sets (table XXII) show very little 
variation and suggest fractions of a single population. 

Sinployea otareae (Garrett, 1872). Figure 50a-c. 

Pitys otareae Garrett, 1872, Am. J. Conchol., 7 (4), pp. 228-229, pi. 

19, fig. 21 one valley on Rarotonga, Cook Islands. 
Patula otareae (Garrett), Schmeltz, 1874, Cat. Mus. Godeffroy, 5, p. 

94. 

Helix otareae (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p. 543. 
Patula decorticata var. otareae (Garrett), Garrett, 1881, J. Acad. 

Nat. Sci., Philadelphia, 8 (4), p. 387. 
Helix (Patula) decorticata var. otareae (Garrett), Tryon, 1887, Man. 

Conchol., (2) 3, p. 40, pi. 8, fig. 50. 
Endodonta (Charopa) decorticata var. otareae (Garrett), Pilsbry, 

1893, Man. Conchol., (2) 9, p. 35. 

Diagnosis. Shell very large, diameter 4.21-5.20 mm. (mean 
4.68 mm.), with 4'/2-5 relatively tightly coiled whorls. Apex and 
spire moderately and evenly elevated, last whorl descending more 
rapidly, spire protrusion more than *k body whorl width, H/D ratio 
0.470-0.570 (mean 0.516). Apical sculpture of 8-12 (mean 9.41) 
prominent spiral cords. Postnuclear whorls, with narrow, prominent, 
protractively sinuated, rather widely spaced radial ribs, 81-116 
(mean 95.6) on the body whorl, whose interstices are 2-3 times their 
width. Ribs/mm. 5.72-7.73 (mean 6.48). Microsculpture of fine, 
crowded radial riblets, 6-10 between each pair of major ribs, crossed 
by finer and more crowded spiral riblets. No secondary spiral cord- 
ing. Umbilicus open, V-shaped, regularly decoiling, contained 3.47- 
5.52 times (mean 4.10) in the diameter, margins rounded. Whorl 
contours and aperture typical. 

The presence of only six to ten microradials be- 
tween each pair of major ribs at once separates 
Sinployea otareae (fig. 50a-c) from S. decorticata (fig. 
47a-c). Individual specimens may show the size and 
shape characters of the other species, but the sculp- 
tural difference is consistent. Sinployea tenuicostata 
(fig. 50d-f) has a flat spire, fewer ribs, and a narrower 
umbilicus. 






FIG. 50. a-c, Sinployea otareae (Garrett). Rarotonga, Cook Islands. Paratype. BPBM 2335; d-f, Sinployea tenuicostata (Garrett). Rarotonga, 
Cook Islands. Paratype. BPBM 2325. Scale lines equal 1 mm. (o-c, MM; d-f, SH). 



116 



SYSTEMATIC REVIEW 



117 



Description. Shell extremely large, with 4% relatively tightly 
coiled whorls. Apex and early spire flat, later whorls descending 
gradually, body whorl descending moderately, H/D ratio 0.470. Api- 
cal whorls 1%, sculpture of 8 prominent spiral ribs with a micro- 
sculpture of radial corrugations. Postnuclear whorls with prominent, 
high, protractively sinuated radial ribs, 110 on the body whorl, 
whose interstices are 2-3 times their width. Microsculpture a lattice 
of radial riblets, 6-9 between each pair of major ribs, and slightly 
smaller, more crowded spiral riblets. Sutures deep, whorls flattened 
laterally above periphery and on basal margin. Color reddish 
yellow-brown. Umbilicus moderately open, V-shaped, regularly de- 
coiling, contained 3.98 times in the diameter. Aperture ovate, flat- 
tened laterally above periphery and basally, inclined about 20 from 
the shell axis. Height of lectotype 2.34 mm., diameter 4.97 mm. 

Lectotype. Cook Islands: Rarotonga. Collected by 
Andrew Garrett. ANSP 47755. 

Range. Under dead wood on the banks of a 
stream in a single valley on Rarotonga, Cook Islands. 

Paratypes. BPBM 2335, ANSP. 

Material. Cook Islands (2 specimens, NMWC): 
Rarotonga (30 specimens, BPBM 167418-9, FMNH 
46411, FMNH 90615). 

Remarks. Mautodontha zebrina came from the 
same unidentified valley (Garrett, 1881, p. 387). 
Sinployea otareae was abundant and "Living examples 
are uniform deep black" (Garrett, 1872, p. 229). Only 
minor size variation was observed (table XXIII). 

Sinployea tenuicostata (Garrett, 1872). Figure 
50d-f. 

Pitys tenuicostata Garrett, 1872, Am. J. Conchol., 7 (4), p. 229, pi. 

19, fig. 23 rare, but widely distributed on Rarotonga, Cook 

Islands. 
Patula tenuicostata (Garrett), Schmeltz, 1874, Cat. Mus. Godeftroy, 

5, p. 94; Garrett, 1881, J. Acad. Nat. Sci., Philadelphia, 8 (4), p. 

385. 
Helix tenuicostata (Garrett), Pfeiffer, 1876, Monog. helic. viv., 7, p. 

547. 
Helix (Patula) tenuicostata (Garrett), Tryon, 1887, Man. Conchol., 

(2)3, p. 39, pi. 8, figs. 35-37. 
Endodonta (Champa) tenuicostata (Garrett), Pilsbry, 1893, Man. 

Conchol., (2) 9, p. 35. 

Diagnosis. Shell very large, diameter 4.08-5.43 mm. (mean 
4.70 mm.), with 3%-4'/4 relatively loosely coiled whorls. Apex and 
early spire flat or barely depressed, later whorls descending slightly, 
H/D ratio 0.437-0.537 (mean 0.499). Apical sculpture of 9-12 (mean 
10.45) prominent spiral cords. Postnuclear whorls with narrow, 
widely spaced, protractively sinuated radial ribs, 43-66 (mean 50.7) 
on the body whorl, whose interstices are 3-6 times their width. 
Ribs/mm. 2.87-4.23 (mean 3.46). Microsculpture of rather large and 
widely spaced radial riblets, 8-12 between each pair of major ribs, 
crossed by much finer and more crowded spiral riblets. No secondary 
spiral cording. Umbilicus relatively narrow, U-shaped, regularly de- 
coiling, contained 4.29-6.20 times (mean 5.16) in the diameter, mar- 
gins rounded. Aperture strongly flattened laterally above periphery. 

Sinployea tenuicostata (fig. 50d-f) has few and 
widely spaced major radial ribs, a flat spire, and rela- 
tively few and larger microradial riblets. The Samoan 
S. complementaria (fig. 56a-f) has an elevated spire, 
more major radials, and much finer and more numer- 
ous microradial riblets. Sinployea otareae (fig. 50a-c) is 
the same size, but has an elevated spire, many more 
radial ribs, and a wider umbilicus. 

Description. Shell very large, with 4Vs loosely coiled whorls. 
Apex and spire fiat, last whorl descending slightly, H/D ratio 0.508. 



Apical whorls 1%, sculpture of 11 moderately prominent spiral ribs 
with a faint microsculpture of radial corrugations. Postnuclear 
whorls with narrow, prominent, rather widely spaced, protractively 
sinuated radial ribs, 47 on the body whorl, whose interstices are 3-5 
times their width. Microsculpture of fine, rather widely spaced radial 
riblets, 6-10 between each pair of major ribs, crossed by much finer 
and more crowded spiral riblets. Sutures deep, whorls strongly 
rounded above, flattened laterally above periphery and slightly on 
basal margin. Color light yellow-horn without darker maculations. 
Umbilicus narrow, U-shaped, regularly decoiling, contained 5.25 
times in the diameter. Aperture subcircular, flattened laterally 
above periphery, inclined about 25 from the shell axis. Height of 
lectotype 2.11 mm., diameter 4.15 mm. 

Lectotype. Cook Islands: Rarotonga. Collected by 
Andrew Garrett. ANSP 47702. 

Range. Widely distributed on Rarotonga, Cook 
Islands. 

Paratypes. BPBM 2325, ANSP. 

Material. Rarotonga (8 specimens, Zurich, SMF 
165748). 

Remarks. Originally described as a "somewhat 
rare species, having a wide range on the island, and 
generally found on the ground on the sides of ravines" 
(Garrett, 1872, p. 229), later Sinployea tenuicostata 
was termed "comparatively rare" (Garrett, 1881, p. 
385). Available sets showed considerable size variation 
(table XXIII), which is probably magnified by the lack 
of clear differentiation into preadult and gerontic 
growth. 

SAMOAN AND SWAINS ISLAND Sinployea 

Except for Ofu and Olosega, every island in the 
Samoan group that has been collected on has yielded at 
least one species of Sinployea. Only one endodontid, 
Minidonta manuaensis Solem (1976b, p. 130), has been 
collected on Olosega, and no specimens of this family 
have been taken on Ofu. Although only two examples 
of Sinployea clista have been taken on Tutuila, com- 
paratively little collecting has been done on that is- 
land. Aunuu Island off Tutuila has an endemic species, 
S. aunuuana, which also is recorded from Ta'u on the 
basis of a single very worn individual that may repre- 
sent an undescribed species. Sinployea clausa and S. 
allecta tauensis are restricted to Ta'u in the Manu'a 
Group. All three Ta'u species have been found at a 
single station (Station 76), and S. clausa and S. a. 
tauensis were collected together at Stations 75, 184, 
188, and 189. None of the Ta'u examples were collected 
alive. Scattered collections from Savai'i produced 
abundant material of Sinployea allecta allecta (Cox) 
from lowland localities. Much less abundant material 
of a more heavily ribbed form of this subspecies was 
collected from diverse localities and at several eleva- 
tions on Upolu. At the foot of Mt. Tafua Upolu (Station 
7), S. clista was taken together with S. a. allecta, 
whereas both S. clista and S. complementaria (Mous- 
son) occurred at the foot of Mt. Solaua (Station 18). The 
latter is the dominant species on Upolu, and I suspect it 
is significant that S. allecta (Cox) and S. complemen- 
taria (Mousson) have not been collected at the same 



118 



SOLEM: ENDODONTOID LAND SNAILS 



stations. Swains Island has an endemic species, Sin- 
ployea intermedia. 

No adult examples of Upolu S. allecta allecta were 
available for dissection, so the degree of difference from 
S. clista remains uncertain. The latter has a penis 
about 1.2 mm. long, whereas Savai'i examples of S. 
allecta have a 1.3-1.5 mm. penis with vergic papilla 
enlarged and stimulatory pad reduced in size. Sin- 
ployea complementaria is immediately differentiated 
by its very large penis of more than 2.0 mm. in length. 
Study of Upolu S. allecta would be desirable in order to 
determine the pattern of species isolation in penis 
structure. 

Compared with the Cook and Society Island 
species, Samoan Sinployea tend to have many radial 
microriblets, lack secondary spiral cording (except S. 
clausa and S. clista), tend toward a narrowed or closed 
umbilicus (except S. aunuuana), and have a flat apex 
but elevated spire (table XXIV). Sinployea aunuuana 
(fig. 52a-c) has the widest umbilicus and relatively 
crowded major radial ribbing. Its flat spire and more 



rounded body whorl ally it to the Samoan taxa. 
Sinployea clista (fig. 51a-c) and S. clausa (fig. 51d-f) 
are small species with rather widely spaced ribbing 
whose umbilici are closed or with only a narrow lateral 
crack. Sinployea clausa has the umbilicus open to the 
apex and the spire more elevated than in S. clista 
where the umbilicus is never open to the apex. Both of 
these have rather prominent secondary spiral cording. 
Sinployea allecta (fig. 54a-c) and S. intermedia (fig. 
52d-f) are larger, have a more open umbilicus, at most 
faint traces of secondary spiral cording, and rather 
widely spaced major radial ribs. Sinployea com- 
plementaria (fig. 56a-f) is the largest known species in 
the genus, has an increased whorl count, very few ra- 
dial ribs, and a moderately open umbilicus. Sinployea 
intermedia is the easternmost Polynesian species to 
lack gray pigmentation on the head and ommato- 
phores. Since Swains Island is low and has coastal 
scrub rather than dense forest, restriction of this 
species to the ground stratum and loss of pigmentation 
would not be surprising. 



TABLE XXIV. - RANGE OF VARIATION IN SAMOAN AND SWAINS ISLAND SINPLOYEA. 



NUMBER OF 
SPECIMENS 


clausa 


36 

10 
136 


clista 


aunuuana 


allecta allecta 


(Cox) 


ALL 


151 


Savaii 


86 


Upolu 


65 


allecta tauensis 


26 
46 
126 


intermedia 


complement aria 


(Mo us son) 



RIBS 



RIBS/MM. 



HEIGHT 



DIAMETER 



H/D RATIO 



86.6(67-106) 11.80(9.04-14.23) 1.51(1.25-1.97) 2.41(2.11-2.93) 0.625(0.577-0.732) 

98.3(74-115) 11.98(9.13-14.36) 1.50(1.28-1.84) 2.53(2.17-2.96) 0.591(0.555-0.636) 

112.6(92-131) 13.13(11.18-14.57) 1.40(1.25-1.71) 2.76(2.57-3.09) 0.509(0.465-0.572) 

89.1(56-121) 9.00(5.83-11.98) 1.77(1.51-2.04) 3.13(2.63-3.62) 0.571(0.511-0.622) 

75.9(56-89) 7.84(5.85-8.97) 1.75(1.51-1.94) 3.07(2.65-3.36) 0.576(0.558-0.598) 

99.3(84-121) 9.90(8.48-11.98) 1.79(1.51-2.04) 

79.8(61-101) 8.31(6.20-10.51) 1.73(1.61-1.97) 

82.1(63-96) 7.66(5.75-9.16) 2.03(1.82-2.19) 

69.1(52-95) 4.50(3.11-5.79) 2.55(2.11-3.32) 



3.17(2.63-3.62) 0.567(0.511-0.622) 

3.00(2.70-3.39) 0.577(0.542-0.616) 

3.49(3.21-3.81) 0.581(0.534-0.616) 

4.92(4.31-5.92) 0.517(0.458-0.648) 



aunuu. 
a. all. 



ALL 
Sav. 

Upo. 



compl . 



WHORLS 



UMBILICUS 



D/U RATIO 



APICAL 
CORDS 



SPIRE 
ELEVATION 



BODY WHORL 
WIDTH 



SP/BWW 



claus - 3 7/8-(3 1/2-4 3/8) 0.11(0.07-0.20) 23.1(13.7-39) 9.40(8-10) 0.19(0.16-0.23) 0.90(0.82-0.92) 0.215(0.200-0.233) 

cllst. 3 3/4+(3 1/2-4 1/4) LESS THAN 0.13 ^QW"^ 9 - 17 <S-U> 0.07(0.03-0.13) 0.98(0.86-1.12) 0.089(0.033-0.176) 
4 l/8-(4-4 1/2) 0.75(0.66-0.95) 3.69(3.24-4.05) 9.50(8-12) 0.13(0.07-0.26) 0.89(0.82-0.92) 0.125(0.074-0.286) 



4 1/8(3 5/8-4 1/2) 0.41(0.26-0.59) 7.85(5.93-9.80) 8.60(7-10) 0.14(0.07-0.23) 1.11(0.99-1.22) 0.123(0.056-0.250) 

4+(3 5/8-4 3/8) 0.39(0.33-0.43) 7.89(6.85-9.20) 8.50(8-9) 0.16(0.12-0.21) 1.09(0.99-1.15) 0.143(0.103-0.191) 

4 1/8+0 7/8-4 1/2) 0.42(0.26-0.59) 7.82(5.93-9.80) 8.70(7-10) 0.13(0.07-0.23) 1.12(1.02-1.22) 0.117(0.056-0.250) 

4-(3 3/4-4 1/8) 0.46(0.33-0.69) 6.80(4.72-9.55) UNKNOWN 0.19(0.18-0.20) 1.09(1.05-1.12) 0.172(0.162-0.188) 

4 3/8+(4 1/4-4 3/4) 0.64(0.53-0.79) 5.43(4.59-6.12) 10.57(9-13) 0.23(0.13-0.35) 1.12(1.02-1.25) 0.209(0.125-0.309) 

4 5/8+(4 3/8-5 1/4) 0.99(0.76-1.22) 4.99(4.18-6.08) 10.08(8-13) 0.22(0.07-0.46) 1.55(1.35-1.81) 0.140(0.039-0.275) 



SYSTEMATIC REVIEW 



119 



KEY TO SAMOAN AND SWAINS ISLAND Sinployea 

1. Umbilicus closed or D/U ratio averaging more than 20 2 

Umbilicus open, D/U ratio averaging much less than 10 3 

2. Spire protrusion about Vs body whorl width; umbilicus with 

apex visible Sinployea clausa, new species 

Spire protrusion averaging less than 0.10; umbilicus closed or 
at most a lateral crack Sinployea clista, new species 

3. Diameter less than 4.00 mm.; ribs/mm, averaging more than 

6.00 4 

Diameter more than 4.00 mm.; ribs/mm, averaging less than 
5.00 Sinployea complementaria (Mousson, 1865) 

4. D/U ratio averaging more than 5.00; H/D ratio averaging more 

than 0.550 5 

D/U ratio averaging about 3.75; H/D ratio about 0.510 

Sinployea aunuuana, new species 

5. Mean diameter about 3.0-3.2 mm.; mean D/U ratio more than 

6.50 6 

Mean diameter about 3.5 mm.; mean D/U ratio about 5.50; 
Swains Island Sinployea intermedia, new species 

6. Spire more protruded; umbilicus slightly wider; Ta'u, Manu'a 

Group Sinployea allecta tauensis, new subspecies 

Spire less protruded; umbilicus slightly narrower; Upolu and 
Savai'i Sinployea allecta allecta (Cox, 1870) 

Sinployea clausa, new species. Figure 51d-f. 

Diagnosis. Shell small, diameter 2.11-2.93 mm. (mean 2.41 
mm.), with 3'A-4 3 /8 normally coiled whorls. Apex and spire moder- 
ately and evenly elevated, body whorl descending much more 
rapidly, spire protrusion more than 'k body whorl width, H/D ratio 
0.577-0.732 (mean 0.625). Apical sculpture of 8-10 (mean 9.40) fine 
spiral cords. Postnuclear whorls with narrow, rounded, protractive, 
rather inconspicuous radial ribs, 67-106 (mean 86.6) on the body 
whorl, whose interstices are 2-4 times their width. Ribs/mm. 9.04- 
14.2 (mean 11.80). Microsculpture of extremely fine radial riblets, 
5- 10 between each pair of major ribs, crossed by slightly finer and 
more crowded spiral riblets. Secondary sculpture of fine, rather 
widely spaced spiral cords visible over entire shell surface. Um- 
bilicus very narrow, open to apex, about 'A constricted by basal lip, 
contained 13.7-39 times (mean 23.1) in the diameter, margins 
rounded. Whorl profile almost circular, only flattened slightly on 
upper palatal wall, basal margin rounded, aperture inclined about 5 
from shell axis. 

Sinployea clista (compare fig. 51a-c) is very closely 
related, differing primarily in its lower spire and gen- 
erally imperforate or at most very narrowly open um- 
bilicus. Other Samoan species have wider umbilici. Ex- 
tralimital species with narrow umbilici, such as Ba 
humbugi (fig. 74a-c) and Sinployea adposita (fig. 
69d-f) from Fiji, have vastly different shapes and 
sculpture. 

Description. Shell small, with 4 tightly coiled whorls. Apex 
and spire moderately and evenly elevated, last whorl descending 
much more rapidly, H/D ratio 0.613. Embryonic whorls l'/2, sculpture 
partly eroded, remnants of about 9 spiral cords. Remaining whorls 
with moderately prominent, rounded, protractive radial ribs, 89 on 
the body whorl, whose interstices are 2-4 times their width. Micro- 
sculpture of very fine radial riblets, 5-12 between each pair of major 
ribs, even finer spiral riblets and larger, relatively widely spaced, 
secondary spiral cords. Sutures deeply impressed, whorls strongly 
rounded above with only slight lateral flattening. Umbilicus barely 
perforate, partially constricted by basal lip, contained 15 times in the 
diameter. Color a uniform, very light yellow-brown. Aperture circu- 
lar with slightly expanded baso-columellar margin, slightly flat- 
tened laterally above periphery, inclined about 5 from the shell axis. 
Height of holotype 1.51 mm., diameter 2.47 mm. 

Holotype. Samoa: Ta'u, Station 188, Faleasao, 
700 ft. inland, at 12 ft. elevation. Collected by Wray 
Harris on September 24, 1937. BPBM 187736. 



Range. Ta'u, Manu'a Group, American Samoa. 

Paratypes. Ta'u: Siufaga (Stations 16, 76, 138, 
184, 185, 186, 211) at 350-600 ft. inland at 5-500 ft. 
elevation (19 specimens, BPBM 171136, BPBM 
173784, ex BPBM 171073, ex BPBM 182410, ex BPBM 
187653, BPBM 186682, BPBM 187625, BPBM 187658, 
BPBM 187705, BPBM 188085); Tavalagi, Vi2 mile in- 
land (Station 75) at 500 ft. elevation above Faleasao 
cliffs (3 specimens, BPBM 171065, ex BPBM 171106, 
ex BPBM 182095); Faleasao, 600-700 ft. inland (Sta- 
tions 185, 188-190) at 12 ft. elevations (11 specimens, 
BPBM 187665, BPBM 187728, BPBM 187736, BPBM 
187774, ex BPBM 187753, ex BPBM 187781, BPBM 
187982, BPBM 188277); Amouli, l /z mile inland (Sta- 
tion 80) at 500 ft. elevation (1 specimen, BPBM 
182150); Palapala, Vfe mile inland (Station 201) at 600 
ft. elevation (1 specimen, BPBM 187919). 

Remarks. The only available specimens were 
dead shells taken from sweepings mixed in coral rub- 
ble. A high percentage of the shells had bits of coral 
stone and sand cemented to the shell surface, and 
many were quite worn. Specimens from Faleasao did 
not differ significantly from those taken at Siufaga 
(table XXV). 

The name clausa, from the Latin meaning closed, 
is perhaps slightly misleading, because the umbilicus 
is always minutely perforate, but the name was chosen 
to emphasize its close relationship to Sinployea clista 
from Upolu and Tutuila. 

In combining a higher spire and wider umbilicus 
than is found inS. clista, S. clausa parallels (fig. 51a-f) 
the divergence seen between the subspecies of S. al- 
lecta (fig. 54a-e). Only the proportionately greater size 
of the morphologic gap between S. clista and S. clausa 
accounts for the arbitrary consideration of them as 
species rather than subspecies. 

Sinployea clista, new species. Figures 51a-c, 53d-e. 

Diagnosis. Shell small, diameter 2.17-2.96 mm. (mean 2.53 
mm.), with 3 l /2-4'/t normally coiled whorls. Apex and spire flat to 
slightly and evenly elevated, body whorl descending much more 
rapidly, spire protrusion less than Vio body whorl width, H/D ratio 
0.555-0.636 (mean 0.591). Apical sculpture of 8-11 (mean 9.17) nar- 
row spiral cords. Postnuclear whorls with low, rounded, protractively 
sinuated radial ribs, 74-115 (mean 98.3) on the body whorl, whose 
interstices are about 3-4 times their width except becoming crowded 
and indistinct on end of body whorl. Ribs/mm. 9.13-14.36 (mean 
11.98). Microsculpture of very fine radial riblets, 5-12 between each 
pair of major ribs, crossed by slightly finer spiral riblets, relatively 
crowded and inconspicuous secondary spiral cording visible on most 
of shell surface. Umbilicus either closed (50%) or with a narrow lat- 
eral opening that prevents seeing the apex, contained more than 20 
times in the diameter. Whorls strongly rounded on all margins, very 
slightly flattened laterally above periphery, aperture inclined about 
15 from shell axis. 

Sinployea clausa (fig. 51d-f) has the umbilicus 
narrowly open and the spire distinctly more elevated. 
Other minor differences are discussed below. Ba hum- 
bugi (fig. 74a-c) from Fiji has a much more elevated 
spire (mean H/D ratio 0.817) and very few and widely 
spaced major radial ribs. All other Sinployea are at 
least narrowly umbilicated. 








FIG. 51. a-c, Sinployea clista, new species. Olomoana, Tutuila, Samoa. Holotype. BPBM 84372; d-f, Sinployea clausa, new species. Station 
188, Faleasao, Ta'u, Samoa. Holotype. BPBM 187736. Scale lines equal 1 mm. (a-c, SG; d-f, MM). 



120 



TABLE XXV. - LOCAL VARIATION IN SAMOAN SINPLOYEA, I. 



NUMBER OF 
SPECIMENS 



RIBS 



HEIGHT 



DIAMETER 



H/D RATIO 



clausa 

Faleasao 

Siufaga 



10 88.115.88 I.470.063 2.3810.080 0.6150.0006 

(75-101) (1.22-1.81) (2.07-2.93) (0.577-0.666) 

11 85.614.33 1.5410.050 2.4210.050 0.63710.0122 

(67-106) (1.38-1.97) (2.20-2.70) (0.584-0.732) 



clista 



Tutulla 
BPBM 84372 

Upolu 

BPBM 185879, 
BPBM 186642 

FMNH 153177, 
BMW MF3992 



89.016.02 
(83-95) 



74 



112.712.34 

(108-115) 



1.84 



2.9310.033 0.62910.0070 

(2.89-2.96) (0.622-0.636) 



1.2910.011 2.2310.039 0.53210.0055 

(1.28-1.32) (2.17-2.30) (0.572-0.591) 

1.4810.050 2.5710.025 0.57510.0130 

(1.41-1.58) (2.53-2.63) (0.555-0.600) 



aunuuana 

BPBM 83242 



25 



1.4110.018 2.7610.024 0.50810.0050 

(1.26-1.72) (2.58-3.11) (0.465-0.556) 



allecta allecta 






Upolu 
AMS, BPBM 



Zurich 



S avail 
Sta. 38 
FMNH 153371 



18 97.115.10 1.76+0.043 3.0910.064 0.56710.0073 

(78-121) (1.39-2.02) (2.52-3.41) (0.511-0.622) 

13 101.83.72 1.8410.020 3.2610.044 0.56610.0045 

(84-121) (1.71-2.01) (3.03-3.55) (0.543-0.598) 

6 73. 411. 86 1.7510.064 3.0410.096 0.5760.0064 

(69-80) (1.51-1.94) (2.66-3.36) (0.558-0.598) 



WHORLS 



UMBILICUS 



D/U RATIO 



APICAL CORDS 



clausa 



Fale. 



Siuf. 



clista 



Tutu. 



Upolu 
BPBM 



FMNH 



3 3/4+(3 1/2-4 1/8) 0.1210.011 

(0.07-0.20) 

3 7/8+(3 1/2-4 3/8) 0.1210.012 

(0.07-0.20) 



4 1/4 



0.13 



3 5/8(3 1/2-3 3/4) 0.0810.011 

(0.07-0.10) 

3 5/8+(3 5/8-3 3/4) 



22.412.19 

(14.8-39) 

22.712.07 

(13.6-35) 



22.5 



9.2510.48 
(8-10) 



10. Oil. 
(9-11) 



29.913.3 9 

(23.3-33.5) 



8.510.50 
(8-9) 



aunuuana 



BPBM 4 l/8-(4-4 1/2) 



a. allecta 

Upolu 
AMS 4 1/8(3 7/8-4 3/8) 



Zurich 4 l/8+(4-4 3/8) 



Savaii 
Sta. 38 4-(3 5/8-4 1/4) 



0.7410.014 

(0.66-0.95) 



0.3910.015 

(0.26-0.49) 

0.4310.020 

(0.33-0.53) 



0.3910.020 

(0.33-0.43) 



3.6910.044 9.0010.55 

(3.24-4.05) (8-11) 



8.0510.250 

(5.93-9.70) 

7.7410.316 

(6.19-9.80) 



7.8910.356 

(6.85-9.20) 



8.3310.67 
(7-9) 

8.8310.48 
(7-10) 



(8-9) 



121 



122 



SOLEM: ENDODONTOID LAND SNAILS 



Description. Shell small, with 4'/4 normally coiled whorls. Apex 
and spire slightly and evenly elevated, last whorl descending much 
more rapidly, H/D ratio 0.622. Embryonic whorls IVz, sculpture of 11 
narrow spiral cords with an intrusion of vague radial corrugations on 
last quarter whorl. Postnuclear whorls with low, rounded, protrac- 
tively sinuated radial ribs, 93 on the body whorl, whose interstices 
are 2-4 times their width, becoming irregular and more crowded on 
gerontic portions of body whorl. Microsculpture of fine radial riblets, 
5-10 between each pair of major ribs, exceedingly fine spiral riblets, 
and inconspicuous secondary spiral cords. Sutures deep, whorls flatly 
rounded above periphery with broadly rounded basal margin, sloping 
to narrow umbilical chink. Umbilicus closed by expansion of inner 
portion of body whorl and reflection of umbilical lip. Color light 
yellow-brown with reddish markings. Aperture circular with slightly 
less sharply rounded supraperipheral outer margin, inclined about 
15 from shell axis. Height of holotype 1.84 mm., diameter 2.96 mm. 

Holotype. Samoa: Tutuila, Olomoana. Collected 
under stones and on dead leaves by members of the 
Samoan Expedition on March 17, 1926. BPBM 84372. 

Range. Tutuila, American Samoa, and Upolu, 
Western Samoa. 

Paratypes. Tutuila: Olomoana, under stones and 
dead leaves (1 specimen, BPBM 84372). Upolu: Mt. 
Vaea (Station 147) at 850 ft. elevation (3 specimens, 
BPBM 185879); Tatapao-Lake Lanuto'o trail at 1,250 
ft. elevation (1 specimen, BPBM 186642); foot of south- 
east peak, Mt. Tafua Upolu (Station 7) at 1,200 ft. ele- 
vation (1 specimen, FMNH 153081); foot of Mt. Solaua 
(Station 18) at 600-800 ft. elevation (2 specimens, 
FMNH 153177); Tanumalala at 800 ft. elevation (1 
specimen, BMW MF 3992, collected by R. A. Cumber 
on May 15, 1954). 

Remarks. Besides the obvious differences in 
spire protrusion and umbilical closure, Sinployea clista 
differs from S. clausa in having the aperture more 
strongly inclined and the major ribbing protractively 
sinuated rather than being just slightly inclined. The 
latter also has the secondary spiral cording more prom- 
inent and more widely spaced. 

Sinployea clista is sparsely distributed on both 
Upolu and Tutuila. Although there are obvious size, 
shape, and ribbing differences (tables XXIV- XXV), so 
little material is available that I give no systematic 
recognition to these variations. Despite intensive ef- 
forts after field recognition, no additional examples 
could be located at the Mt. Tafua Upolu and Mt. Solaua 
stations. 

The name clista is taken from the Greek for closed 
and refers to the unusual umbilicus. 

Description of soft parts. Pallial region (fig. 53e) about 3.5 mm. 
long, rectal kidney lobe 1.7 mm. long, much longer than pericardia! 
arm. Space between ureter arms wider than in most Polynesian 
species. 

Genitalia (fig. 53d) fragmentary, observed portions without un- 
usual features. Penis about 1.2 mm. long. 

(Based on BPBM 84372, 1 partial example.) 

Sinployea aunuuana, new species. Figures 52a-c, 
53a-c. 

Diagnosis. Shell slightly smaller than average, diameter 
2.57-3.09 mm. (mean 2.76 mm.), with 4-4'/2 normally coiled whorls. 
Apex and early spire flat or slightly protruding, lower spire whorls 



descending slightly, body whorl descending rapidly, spire protrusion 
about Vx body whorl width, H/D ratio 0.465-0.572 (mean 0.509). Api- 
cal sculpture of 8-12 (mean 9.50) narrow spiral cords. Postnuclear 
sculpture of very fine, narrow, rather crowded, slightly protractively 
sinuated radial ribs, 92-131 (mean 112.6) on the body whorl, whose 
interstices are 2-3 times their width. Ribs/mm. 11.18-14.57 (mean 
13.13). Microsculpture of very fine radial riblets, 5-9 between each 
pair of major ribs, crossed by very fine and crowded spiral riblets, 
with an indistinct development of secondary spiral cording occasion- 
ally visible. Umbilicus broadly open, V-shaped, regularly decoiling, 
contained 3.24-4.05 times (mean 3.69) in the diameter, margins 
rounded. Whorl contours slightly less compressed than average, 
aperture inclined about 10 from shell axis. 

The generally flat apex and early spire of 
Sinployea aunuuana (fig. 52a-c) combine with the 
wide umbilicus and crowded radial ribbing to easily 
separate it from the other Polynesian species. Of 
similar-sized species, S. neglecta (fig. 41d-f) from 
Huahine has a much more elevated spire, proportion- 
ately narrower umbilicus, and about 12.5 apical cords; 
S. atiensis (fig. 44d-f) has a higher spire and narrower 
umbilicus with broader and more widely spaced ribs. 
Sinployea kusaieana (fig. 63a-c) has a narrower body 
whorl, fewer whorls, and a proportionately smaller 
apex. 

Description. Shell slightly smaller than average, with 4'/s 
tightly coiled whorls. Apex and early spire flat, body whorl descend- 
ing rapidly, spire protrusion about 'Ao of body whorl width, H/D ratio 
0.524. Embryonic whorls 1%, sculpture of 12 moderately prominent 
spiral ribs. Postnuclear whorls with somewhat irregularly spaced, 
high, rounded radial ribs, 123 on the body whorl, whose interstices 
are usually about twice their width. Microsculpture of very fine ra- 
dial riblets, 3-6 between each pair of major ribs, crossed by barely 
visible, spiral riblets that are very crowded. Occasional traces of 
weak spiral cording. Sutures deeply impressed, whorls strongly 
rounded above, flattened laterally above periphery and slightly on 
basal margin. Umbilicus broadly V-shaped, widely open, regularly 
decoiling, contained 3.82 times in the diameter. Color uniform light 
reddish brown. Aperture circular, strongly flattened laterally above 
periphery, inclined about 10 from the shell axis. Height of holotype 
1.45 mm., diameter 2.77 mm. 

Holotype. Samoa: Aunuu Island (off Tutuila), 
Station 14, 500 yd. from shore at 200 ft. elevation. Col- 
lected by T. T. Dranga on February 4, 1926. BPBM 
83256. 

Range. Aunuu Island off Tutuila and probably 
Ta'u, Manu'a Group, American Samoa. 

Paratypes. Aunuu: 10 yd. east of lighthouse, 200 
yd. inland at 250 ft. elevation (134 specimens, BPBM 
83242). Ta'u: Siufaga swamp, 400 ft. inland (Station 
76) at 7 ft. elevation (1 specimen, ex BPBM 182444). 

Remarks. Although the size and umbilical shape 
of Sinployea aunuuana (fig. 52a-c) is more similar to 
the Cook and Society Island species, the very fine ra- 
dial riblets, wide body whorl, flat spire, and round 
whorl contours relate it to the other Samoan species. 
Sinployea allecta (fig. 54a-e) is most similar, but 
differs in its much narrower umbilicus, more elevated 
shape (table XXIV), and absence of secondary spiral 
cording. They agree in their spire shape and relative 
protrusion. 

A single dead, very worn specimen from Ta'u is 
referred here with some hesitation. In size and proper- 









abc 






FIG. 52. a-c, Sinployea aunuuana, new species. Station 14, Aunuu Island, Tutuila, Samoa. Holotype. BPBM 83256; d-f, Sinployea inter- 
media, new species. 200 yd. inland, west side Swains Island. Holotype. BPBM 186666. Scale lines equal 1 mm. (MM). 



123 



124 



SOLEM: ENDODONTOID LAND SNAILS 



-VD 




FIG. 53. Anatomy of Sinp/oyea aunuuana, S. clista, and S. intermedia from Samoa and Swains Island: a-c, Sinployea aiinuuana, new 
species. Station 14, Aunuu, Tutuila, Samoa. BPBM 83256. a, pallial region, b, genitalia. c, interior of penis with muscular collar not slit open: 
d-e, Sinployea clista, new species. Olomoana, Tutuila, Samoa. BPBM 84372. d, terminal genitalia. e, pallial region; f-h, Sinployea intermedia. 
new species. West side of Swains Island. BPBM 186666. f, pallial region. #, genitalia. h, penis interior. Scale lines equal 1 mm. (MO). 



tions it falls within the midrange of variation, height 
1.35 mm., diameter 2.63 mm., H/D ratio 0.512, whorls 
4Vs, D/U ratio 3.64. The spire is slightly more elevated 
than normal, and this may be misidentified. If so, it 
represents an undescribed species. 

Description of soft parts. Pallial region (fig. 53a) about 3.3 mm. 
long, rectal kidney lobe much longer than pericardia!, about 1.35 
mm. long. 

Genitalia (fig. 53b) without unusual features. Penis length 
about 1.0 mm., internally (fig. 53c) penis with verge, collar, and 
stimulatory pad. Latter drawing with collar shown intact, verge not 
visible. 

(Based on BPBM 83256, 1 example.) 

Sinployea allecta (Cox, 1870). 

Material from Ta'u, Upolu, and Savai'i is suffi- 
ciently similar in size, shape, and sculpture (table 
XXV) to be considered conspecific. The Swains Island 
S. intermedia (fig. 52d-f) is conspicuously larger, has a 
wider umbilicus, and averages two more apical spiral 
cords. Sinployea aunuuana (fig. 52a-c) has the um- 
bilicus much more widely open (mean D/U ratio 3.69 as 



compared to more than 6.5 in allecta) and possesses 
relatively conspicuous secondary spiral cording. 
Sinployea complementaria (fig. 56a-f) is much, much 
larger (table XXIV) and has more widely spaced radial 
ribs. 

Taxonomic treatment of the S. allecta material is 
partly arbitrary. The populations on Ta'u are subspeci- 
fically distinct, differing significantly in having both a 
higher spire and a wider umbilicus. Usually a lower 
spire means a wider umbilicus, and a higher spire ac- 
companies a narrower umbilicus. Hence, the shifts in 
the Ta'u populations are separate rather than being 
linked. Subspecific status has been chosen because the 
similarities are many. Populations on Savai'i and 
Upolu show no significant differences in size or shape, 
but do have markedly distinctive rib counts and spac- 
ing (table XXV). The Upolu material comes from the 
1860-1870 period, whereas the Savai'i material was 
collected in 1965. As discussed below, only subadult S. 
allecta were taken on Upolu in 1965. The significance 
of this sculpture variation remains uncertain. In treat- 



SYSTEMATIC REVIEW 



125 



ing the Savai'i and Upolu populations as belonging to 
one subspecies, I am deferring judgment as to their 
exact relationship. New collections of adults from 
Upolu are needed to settle this problem. 

Sinployea allecta allecta (Cox, 1870). Figures 54a-c, 
55a-e. 

Helix allecta Cox, 1870, Proc. Zool. Soc. London, 1870, p. 81 found 
on the mountains under decayed wood, Upolu, Samoa; Pfeiffer, 
1876, Monog. helic. viv., 7, p. 162. 

Champa allecta (Cox), Tryon, 1886, Man. Conchol., (2) 2, p. 210 
name only. 

Patula allecta (Cox), Garrett, 1887, Proc. Acad. Nat. Sci., Philadel- 
phia, 1887, p. 130. 

Diagnosis. Shell larger than average, diameter 2.63-3.62 mm. 
(mean 3.13 mm.), with 3%-4'/2 rather tightly coiled whorls. Apex and 
early spire usually flat, sometimes slightly and evenly elevated, 
lower spire descending slightly, body whorl descending moderately, 
spire protrusion about VK body whorl width, H/D ratio 0.511-0.622 
(mean 0.571). Apical sculpture of 7- 10 (mean 8.60) rather fine spiral 
cords. Postnuclear whorls with narrow, rounded, rather prominent, 
and protractively sinuated radial ribs that are variable in number 
and spacing, 56121 (mean 89.1) on the body whorl, whose in- 
terstices are 2-5 times their width. Ribs/mm. 5.83-11.98 (mean 
9.00). Microsculpture of exceedingly fine radial riblets, 4-12 between 
each pair of major ribs, crossed by slightly finer and more crowded 
spiral riblets. No secondary spiral cording visible at 96 x magnifica- 
tion. Umbilicus narrow, U-shaped, last whorl decoiling more rapidly, 
slightly constricted by columellar lip expansion, contained 5.93-9.80 
times (mean 7.85) in the diameter, margins rounded. Whorls almost 
evenly rounded on outer margins, slightly flattened laterally above 
periphery, aperture inclined about 15 from shell axis. 

The narrower umbilicus and lower spire protru- 
sion readily separate Sinployea allecta allecta (fig. 
54a-c) from S. a. tauensis (fig. 54d-f). Sinployea inter- 
media (fig. 52d-f) is larger and has a higher spire and 
much wider umbilicus. Other Samoan species differ 
radically in size and umbilical proportions. Of ex- 
tralimital species with narrow umbilici, the Lau Ar- 
chipelago S. angularis (fig. 64d-f) and S. recursa (fig. 
64a-c) both have reduced radial ribbing; the Tahitian 
S. montana (fig. 41a-c) is much smaller. 

Description. Shell smaller than average with 3% loosely coiled 
whorls. Apex slightly protruding, spire flat, last whorl descending a 
little, H/D ratio 0.554. Body whorl quite obese, sutures moderately 
deeply impressed, whorls strongly rounded above and on outer mar- 
gins. Embryonic whorls 1%, sculpture of 9 relatively inconspicuous 
spiral ribs. Postnuclear whorls with moderately prominent, rounded, 
closely spaced, protractively sinuated radial ribs, 86 on the body 
whorl, whose interstices are usually slightly less than twice their 
width. Microsculpture of 4-8 very fine radial riblets between each 
pair of major ribs, and even finer and more crowded spiral riblets. 
Body whorl evenly rounded on outer margin with slightly protruded 
basal lip coming to the evenly rounded umbilical margin. Umbilicus 
partially constricted by inward expansion of columellar lip, narrowly 
U-shaped, regularly decoiling, contained 5.93 times in the diameter. 
Color yellowish white with relatively numerous reddish flammula- 
tions. Aperture circular with protruding basal margin, inclined 
about 15 from shell axis. Height of lectotype 1.53 mm., diameter 
2.73 mm. 

Lectotype. Samoa: Upolu. Collected on the moun- 
tains under decayed wood. AMS C. 63457. 

Range. Upolu and Savai'i, Western Samoa. 

Paratypes.AMS C. 63458. 

Material. Upolu (60 specimens, BPBM 106246, 



BPBM 115349, BPBM 167430, AMS C.34756, AMS 
C.63457-8, Zurich, FMNH 117267, FMNH 160001, 
SMF 165706, NMWC): Togitogiga forest area (Station 
10) at 50 ft. elevation (1 specimen, FMNH 153062); foot 
of southeast peak (Station 7), Mt. Tafua Upolu at 1,200 
ft. elevation (3 specimens, FMNH 153082); gully to foot 
of Mt. Siga'ele (Station 23) at 2,300 ft. elevation (1 
specimen, FMNH 153426). 

Savai'i: heavy forest about 5 miles (Station 30) 
northwest of Vailoa at 600 ft. elevation in heavy forest 
(20 specimens, FMNH 153039); about Vz mile inland, 5 
miles east of Vailoa (Station 38) on road to Salelologa 
wharf in open bush (66 specimens, FMNH 153371, 
FMNH 153091). 

Remarks. The type specimens are all subadult, 
and Cox's whorl count of 4'/2-5 is in error. In older 
collections, specimens of this species are usually 
labeled as S. complementaria. Numerous specimens 
were collected by Graeffe in the early 1870s and are 
rather widely distributed in museum collections. This 
was after Mousson had published on Samoan material. 
Hence, they were not mentioned in the previous litera- 
ture. 

Although the material in historical collections and 
the specimens obtained by Solem and Price in 1965 
agree almost exactly in dimensions and shape (table 
XXV), there is a marked dichotomy in ribbing. The 
specimens collected by Cox and Graeffe in the 1860s 
and 1870s on Upolu have a mean of 99.3 ribs on the 
body whorl and 9.90 ribs/mm. Those from Savai'i (ta- 
bles XXIV- XXV) average only 75.9 ribs on the body 
whorl and 7.84 ribs/mm. Unfortunately, only subadult 
material was obtained on Upolu in 1965. The specimen 
from Togitogiga (Station 10) has the typical Upolu rib- 
bing, but those from Mt. Tafua Upolu (Station 7) and 
near Mt. Siga'ele (Station 23) have the more widely 
spaced ribbing found in the Savai'i examples. 

Description of the soft parts. Foot and tail longer than shell 
diameter, bluntly rounded anteriorly, slightly tapering posteriorly. 
Sole undivided, pedal grooves prominent, suprapedal larger; no 
caudal horn or middorsal groove present. Head projecting slightly in 
front of foot. Ommatophores typical. Gonopore in normal position. 

Body color yellow-white, with a grayish suffusion on neck, head, 
and back of tail. 

Mantle collar (fig. 55d) without glandular extension onto pallial 
roof, edge swollen and providing a flap to narrow pneumostome. 
Anus (A) and external ureteric pore (KX) typical. 

Pallial region extending % whorl apically. Lung roof clear, with- 
out granulations. Kidney (K) with lobes subequal, rectal lobe 1.45 
mm. long, clearly overlapping hindgut. Base slightly indented, inner 
side dished by organs above pallial cavity. Ureter (KD) reflexed with 
very little roof space between arms (fig. 55a-b), narrow, widening 
slightly along hindgut. Heart (H) more than half length of kidney, 
angled slightly from hindgut. Principal pulmonary vein (HV) simple, 
unbranched, angling toward pneumostome. Hindgut (HG) arising 
just above kidney, not tapering. 

Ovotestis with one or two clumps of palmately clavate acini 
lying parallel to whorl sides. Hermaphroditic duct (fig. 55c, GD) nar- 
row at first, expanded and iridescent, sharply reflexed and narrowed 
at apex of albumen gland, following surface of gland to base of talon 
head, entering latter basally. Albumen gland (GG) elongately ovate, 
surface indented by intestinal loops and spermathecal head. Talon 
(GT) with ovately globular, iridescent head partly buried in albumen 








FIG. 54. a-c, Sinployea allecta allecta (Cox), Upolu, Samoa. BPBM 167430; d-e, Sinployea allecta tauensis, new subspecies. Station 188, 
Faleasao, Ta'u, Samoa. Holotype. BPBM 187753. Scale line equals 1 mm. (MM). 



126 



HV 




SYSTEMATIC REVIEW 



127 



PP 



FIG. 55. Anatomy of Sinployea allecta allecta (Cox). Station 38, 
Vailoa, Savai'i, Samoa. FMNH 153091. a, external view of pallial 
region; b, internal view of pallial region; c, genitalia; d, mantle 
collar at parietal-palatal angle showing lobe development; e, interior 
of penis. Scale lines equal 1 mm. (MM). 



gland, shaft very slender, joined by hermaphroditic duct just below 
head expansion. Prostate (DG) of few large acini opening into a 
groove on upper uterine chamber wall. Uterus (UT) bipartite, upper 
chamber very thin-walled, lower with thick, glandular surface. 

Vas deferens (VD) reflexing up from penioviducal angle, enter- 
ing epiphallus (E) through a "valve" pilaster (fig. 55e). Epiphallus 
more than half the length of penis, entering penis apically. Penial 
retractor (PR) arising on diaphragm, inserting in "U" around 
epiphallus-penis junction on side opposite epiphallic ascension. Penis 
(P) 1.3-1.5 mm. long, not tapering basally. Internally (fig. 55e) with 
vergic papilla enlarged and elongated, fastened to muscular collar on 
one side, opening to epiphallus occupying small portion of length. 
Main stimulatory pad lower and less sharply outlined than in most 
species. Atrium (Y) narrow and relatively short, without clear pilas- 
ter arrangement. 

Free oviduct (UV) with thick muscular walls, internally with 
very narrow channel. Spermatheca (S) with expanded head lying 
next to albumen gland, expanded basally with same pilaster ar- 
rangement found in S. complernentaria. Vagina (V) as in S. com- 
plementaria. 

Free muscle system without unusual features. 

Buccal mass elongately oval, small generative sac. Buccal re- 
tractors not split, inserting on lower posterior margin of buccal mass. 
Esophagus a narrow tube extending just apicad of pallial cavity. 
Stomach extending % whorl apically, anterior expansion very sud- 
den. Intestinal looping normal, compressed into less than V whorl. 

Salivary glands uniting posteriorly above esophagus. 

(Based on FMNH 153091, 5 adult, expanded examples, 3.16- 
3.29 mm. in diameter, with 4'/8-4% whorls.) 

Sinployea allecta tauensis, new subspecies. Figure 
54d-e. 

Diagnosis. Shell slightly larger than average, diameter 2.70- 
3.39 mm. (mean 3.00 mm.), with 3%-4'/ somewhat tightly coiled 
whorls. Apex and spire noticeably and evenly elevated, body whorl 
descending much more rapidly, spire protrusion more than '& body 
whorl width, H/D ratio 0.541-0.616 (mean 0.577). Apical whorls 
worn in all material, traces of fine spiral cords visible in most. Post- 
nuclear whorls with fine, low, rounded, protractively sinuated radial 
ribs, 61-101 (mean 79.8) on the body whorl, whose interstices are 
2-4 times their width. Ribs/mm. 6.20-10.51 (mean 8.31). Micro- 
sculpture of extremely fine radial riblets, 6-12 between each pair of 
major ribs, crossed by even finer and more crowded spiral riblets. No 
secondary spiral cording. Umbilicus narrow, V-shaped, last whorl 
decoiling more rapidly, contained 4.72-9.55 times (mean 6.80) in the 
diameter, margins rounded. Whorls almost evenly rounded, not or 
only slightly flattened laterally above periphery. 

The nominate race of Sinployea allecta (fig. 54b-c) 
has a narrower umbilicus and slightly lower spire than 
S. a. tauensis (fig. 54e-f). The only Polynesian species 
apt to be confused with this is the Cook Island S. av- 
anaensis (fig. 45d-f), which is much less elevated and 
has a wider umbilicus. 

Description. Shell rather large, with slightly more than 4 
rather tightly coiled whorls. Apex and spire slightly and evenly ele- 
vated, last whorl descending more rapidly, H/D ratio 0.545. Em- 
bryonic whorls 1%, sculpture of fine spiral cords visible in suture, 
eroded over most of surface. Postnuclear whorls with prominent, 
rounded, variably spaced, protractively sinuated radial ribs, 76 on 
the body whorl, whose interstices are 2-4 times their width, and 
which become very irregular near aperture. Microsculpture of ex- 
tremely fine radial riblets, 5-12 between each pair of major ribs, 
crossed by slightly finer and more crowded spiral riblets. No secon- 
dary spiral cording visible at 96 x magnification. Sutures deep, 
whorls evenly rounded on outer margins. Umbilicus narrow, V- 
shaped, last whorl decoiling more rapidly, contained 5.94 times in 
the diameter, slightly narrowed by columellar lip expansion. Color 
reddish horn on spire, last whorl separating into basal zones and 



128 



SOLEM: ENDODONTOID LAND SNAILS 



broad, irregular, sinuated reddish flammulations that coalesce on 
shell base. Aperture almost circular, with evenly rounded outer 
margins, inclined about 20 from shell axis. Height of holotype 1.81 
mm., diameter 3.32 mm. 

Holotype. Samoa: Ta'u, Station 188, Faleasao, 
700 ft. inland at 12 ft. elevation. Collected by Wray 
Harris on September 24, 1937. BPBM 187753. 

Range. Ta'u, Manu'a Group, American Samoa. 

Paratypes. Ta'u: Talavagi ridge and summit 
(Station 75) from I fi2-Vs mile inland at 450-500 ft. ele- 
vation (4 specimens, BPBM 171106, BPBM 181998, 
BPBM 182095); Siufaga (Stations 76, 184), 250-600 ft. 
inland at 400-600 ft. elevation (12 specimens, BPBM 
171073, BPBM 182055, BPBM 182400, BPBM 182410, 
BPBM 182444, BPBM 187635, BPBM 187640, BPBM 



187653); Maia (Station 87), l /iz mile inland at 75 ft. 
elevation (1 specimen, BPBM 182364); Faleasao (Sta- 
tions 188, 189), 600-700 ft. inland at 12 ft. elevation (8 
specimens, BPBM 187766, BPBM 187781, BPBM 
187788, BPBM 188357, ex BPBM 187736, ex BPBM 
188277). 

Remarks. All known material consists of dead 
specimens sorted from rubble sweepings. Normally a 
higher spire and narrower umbilicus are correlated 
changes, so that the more protruded spire and wider 
umbilicus of Sinployea allecta tauensis, compared with 
the nominate race, are doubly significant. Rib spacing 
and counts in S. a. tauensis are very similar to those 
found in the Savai'i population and rather different 
from the Upolu examples (table XXVI). 



TABLE XXVI. - LOCAL VARIATION IN SAMOAN SINPLOYEA, II. 



NUMBER OF 
SPECIMENS 

allecta tauensis 


RIBS 

77.34.18 
(66-95) 


HEIGHT 

1.760.048 
(1.61-1.97) 


DIAMETER 

3.050.076 
(2.76-3.32) 


H/D RATIO 

0.577+0.0096 
(0.542-0.606) 


Faleasao 


Siufa 


ga 


4 


88.77.89 
(74-101) 


1.72+0 
(1 


.059 
.61-1.88) 


3.040 
(2 


.141 
.70-3.39) 


0.5660 

(0 


.0120 
.543-0 


.598) 


intermedia 


BPBM 


186666 14 


81.74.93 
(65-96) 


2.030 
(1 


.032 
.82-2.19) 


3.490 
(3 


.042 
.21-3.81) 


0.581+0 
(0 


.0062 
.534-0 


.616) 


complement aria 


Sta. 
FMNH 


39, 2000' 8 
153386, -419 


66.3+1.67 
(59-73) 


2.330 
(2 


,062 
,11-2.57) 


4.71+0 
(4 


.150 
.31-5.20) 


0.497+0 

(0 


.0113 
.458-0 


,543) 


Sta. 
FMNH 
-168 


18, 600-800' 37 
153016, -24, 
,-176, -557 


69.9tl.24 

(54-87) 


2.430 
(2 


.025 
11-2.70) 


4.81+0 
(4, 


.038 
,34-5.23) 


0.5050 
(0 


.0040 
.464-0 


.563) 


Sta. 
FMNH 


26, 2450-2500' 8 
153047, -99 


62.9+2.40 
(58-78) 


2.73+0.070 
(2.53-3.03) 


5.110. 
(4 


,124 
.57-5.53) 


0.534+.0, 
(0, 


,0117 
473-0, 


561) 


Sta. 

FMNH 


24, 2300' 3 
153619 


53.01.00 
(52-55) 


2.820. 
(2. 


105 
63-2.99) 


5.210, 
(5, 


090 
03-5.33) 


0.54110, 
(0, 


0110 
523-0. 


562) 


Sta. 13 
FMNH 


,750' 5 
153380, -408 


79.8+2.98 
(73-88) 


2.83+0. 
(2. 


119 
47-3.09) 


5.32+0, 
(4, 


217 
57-5.92) 


0.532+0. 
(0. 


0124 
506-0. 


577) 



WHORLS 
all.tau 

Faleasao 4+(3 7/8-4 1/4) 



Siufaga 



4-(3 3/4-4 1/8) 



TABLE XXVI, CONTINUED 
UMBILICUS 



0.49+0.047 

(0.36-0.69) 



0.44+0.049 

(0.33-0.56) 



D/U RATIO 



6.45+0.515 

(5.53-8.45) 

7.140.483 

(6.06-8.20) 



APICAL CORDS 



BPBM 

compleme . 
Sta. 39 



Sta. 26 
Sta. 24 
Sta. 13 



4 3/8+(4 1/4-4 3/4) 0.64+0.019 

(0.53-0.79) 



4 5/8-(4 1/2-4 7/8) 0.93+0.046 

(0.76-1.18) 



Sta. 18 4 5/8+(4 3/8-5) 



0.99+0.016 

(0.82-1.15) 



4 7/8-(4 1/2-5 1/4) 1.01+0.052 

(0.82-1.22) 



5-(4 3/4-5) 

4 7/8-(4 5/8-5) 



0.99 



1.070.662 

(0.92-1.18) 



5.43+0.123 10.10.31 

(4.59-6.12) (8-12) 



5.13+0.149 

(4.61-6.00) 



4.890.060 9.82+0.48 

(4.18-5.77) (8-13) 



5.14+0.202 13 

(4.22-6.08) 



5.28+0.160 

(5.10-5.40) 



4.9710.109 

(4.61-5.29) 



SYSTEMATIC REVIEW 



129 



Sinployea complementaria (Mousson, 1865). Fig- 
ures 56a-f, 57a-f. 

Patula complementaria Mousson, 1865, J. de Conchyl., 13, pp. 
168-169, 431, pi. 14, fig. 5 Upolu, Samoa; Mousson, 1869, J. de 
Conchyl., 17, p. 333; Garrett, 1887, Proc. Acad. Nat. Sci., 
Philadelphia, 1887, pp. 129-130 behind Apia, Upolu, Samoa. 

Helix complementaria (Mousson), Pfeiffer, 1868, Monog. helic. viv., 
5, p. 157; Pfeiffer, 1876, Monog. helic. viv., 7, p. 164. 

Pitys complementaria (Mousson), Pease, 1871, Proc. Zool. Soc. Lon- 
don, 1871, p. 474. 

Helix (Patula) complementaria (Mousson), Tryon, 1887, Man. 
Conchol., (2) 3, pp. 40-41, pi. 8, fig. 57. 

Endodonta (Champa) complementaria (Mousson), Pilsbry, 1894, 
Man. Conchol., (2) 9, p. 35. 

Diagnosis. Shell extremely large, diameter 4.31-5.92 mm. 
(mean 4.92 mm.), with 4% 5!4 normally coiled whorls. Apex and 
early spire barely to strongly elevated, last whorl usually descending 
much more rapidly, spire protrusion quite variable, usually about l h 
body whorl width, H/D ratio 0.458-0.648 (mean 0.517). Apical 
sculpture of 8-13 (mean 10.08) fine spiral cords, frequently eroded. 
Postnuclear whorls with high, narrow, prominent, strongly protrac- 
tively sinuated radial ribs, 52-95 (mean 69.1) on the body whorl, 
whose interstices are 3-7 times their width. Ribs/mm. 3.11-5.79 
(mean 4.50). Microsculpture of extremely fine radial riblets, 9 to 
more than 20 between each pair of major ribs, crossed by slightly 
finer and more crowded spiral riblets. No secondary spiral cording. 
Umbilicus open, V-shaped, last whorl decoiling more rapidly, con- 
tained 4.18-6.08 times (mean 4.99) in the diameter, margins 
rounded. Whorl contour slightly more rounded than average, flat- 
tened laterally above periphery, slightly on basal margin, aperture 
inclined about 20 from shell axis. 

The extremely large size, comparatively few radial 
ribs, and a high number of microradial riblets effec- 
tively separate even juvenile examples of Sinployea 
complementaria (fig. 56a-f) from the other Samoan 
species. Of the larger Rarotongan species, only S. 
tenuicostata (fig. 50d-f), which has a flat spire, fewer 
and larger microradial riblets plus never more than 4'/2 
whorls, might be confused. The Tongan Tuimalila (fig. 
78a-f) are even larger, but the looser whorl coiling and 
reduced apical sculpture of that genus readily sepa- 
rates them. 

Description. Shell very large, with slightly more than 4V4 nor- 
mally coiled whorls. Apex barely emergent, whorls of spire descend- 
ing rapidly, H/D ratio 0.648. Apical whorls l'/2, sculpture eroded. 
Postnuclear whorls with narrow, moderately widely spaced, protrac- 
tively sinuated radial ribs, 73 on the body whorl, whose interstices 
are 3-5 times their width. Microsculpture a lattice of very fine and 
crowded radial riblets crossed by distinctly finer and slightly more 
crowded spiral riblets. No secondary spiral cording. Sutures moder- 
ately impressed, whorls slightly shouldered above, strongly flattened 
laterally above periphery and on basal margin. Umbilicus V-shaped, 
narrow, partially closed by reflection of basal lip, contained 6.00 
times in the diameter. Color light yellow-brown with irregular red- 
dish flammulations, generally parallel to the line of growth. Aper- 
ture subcircular, flattened laterally above periphery and on basal 
margin, inclined about 15 from the shell axis. Edge of lip broken. 
Height of lectotype 3.32 mm., diameter 5.14 mm. 

Lectotype. Samoa: Upolu. Collected by Graeffe. 
Zoologisches Museum der Universitat Zurich. 

Range. Upolu, Western Samoa. 

Paratypes. Zurich. 

Material. Samoa (2 specimens, BPBM 161): 
Upolu (14 specimens, Zurich, IRB); foot of Mt. Solaua 
(Station 18) at 600-800 ft. elevation on stilts ofFicus 
or under fallen bark and rotting wood (59 specimens, 
FMNH 153016, FMNH 153024, FMNH 153165, 



FMNH 153168, FMNH 153176, FMNH 153557); 
Fagaloa Pass (Station 13) at 750 ft. elevation along 
ridge crest under rotting wood (6 specimens, FMNH 
153380, FMNH 153408); Le Mafa Pass at 900 ft. eleva- 
tion under birdnest fern (2 specimens, DMW M-4002, 
collected by R. A. Cumber on February 10, 1955); top of 
range (Station 39), Mt. Solaua at 2,000 ft. elevation in 
heavy forest (18 specimens, FMNH 153386, FMNH 
153419); Afiamalu, 6 miles inland at 2,100 ft. elevation 
(3 specimens, BPBM 186029, BPBM 186214); % mile 
above Afiamalu seismographic station (Stations 2, 8) 
in secondary forest (2 specimens, FMNH 153050, 
FMNH 153089); south slope, Mt. Siga'ele (Station 24) 
at 2,300 ft. elevation in heavy forest (3 specimens, 
FMNH 153619); Afiamalu-Lake Lanuto'o track (Sta- 
tion 26) at 2,450-2,500 ft. elevation under rotting wood 
in heavy forest (9 specimens, FMNH 153047, FMNH 
153099); rim of Lake Lanuto'o crater (Station 19) at 
2,500 ft. elevation under dead leaves and rotting wood 
(18 juvenile specimens, FMNH 153133, FMNH 
153424). 

Remarks. The lectotype is the highest specimen 
seen, but closely matches the type figures and has the 
characteristic sculpture well preserved. Spire elevation 
in the juvenile example (fig. 56e) is slightly less than 
average. 

Samples collected in 1965 showed rather consider- 
able variation, some of which may be environmentally 
correlated. The very large shells (table XXVI) from 
Fagaloa Pass come from an area of very high rainfall 
that also is continually swept by a moist sea breeze. 
Hence humidity would remain continually high, and 
conditions for growth would be exceptionally favorable. 
Specimens from Mt. Siga'ele (Station 24) were taken 
under litter in heavy fog forest. Their low rib count and 
wide rib spacing may reflect the constancy of this 
niche. Only juvenile examples were taken on the rim of 
Lanuto'o crater (Station 19). Almost all examples from 
Station 18 at the foot of Mt. Solaua were collected 
around the base of a single huge Ficus that was iso- 
lated from the forested slopes by a partly overgrown 
banana patch. Specimens of Sinployea complementaria 
were most frequent under starting bark on stilt roots 
from 2-5 ft. above ground level. A number were taken 
from under accumulated debris in the center of this 
tree, and only a few under fallen branches lying in 
grass beneath the canopy spread. None were found in 
the banana patch or lower forest reaches. Collecting 
1,200 ft. higher on top of Mt. Solaua (Station 39) pro- 
duced specimens essentially identical in measure- 
ments (table XXVI) and ribbing. 

Without much more material, the question of 
geographic variation cannot be settled. The size 
difference between the Mt. Solaua and Fagaloa or 
Siga'ele specimens is large and significant, but there is 
no obvious geographic pattern. The occurrence of S. 
complementaria is strictly colonial, the specimens from 
Mt. Solaua, Fagaloa, and the Lake Lanuto'o areas hav- 
ing been obtained in a fraction of the collecting time 
devoted to the Afiamalu area with its two examples. 
Sinployea complementaria was found in both pro- 




a 





FIG. 56. Sinployea cornplementaria (Mousson). a-c, Upolu, Samoa. Lectotype of Patula complementaria Mousson, 1865. Zoologisches 
Museum Universitat Zurich; d-f, Afiamalu, 2,100 ft., Upolu, Samoa. Juvenile. BPBM 186214. Scale lines equal 1 mm. (SG). 



130 



SYSTEMATIC REVIEW 



131 



foundly disturbed (Fagaloa and foot of Mt. Solaua) and 
undisturbed vegetation zones. 

Description of soft parts. Foot and tail equal in length to shell 
diameter, slightly tapering posteriorly, truncated anteriorly. Sole 
undivided, pedal grooves high on foot, suprapedal weaker, no caudal 
horn or middorsal groove. Slime network typical. Gonopore a long, 
vertical slit behind right rhinophore and below right ommatophore. 

Body color yellow-white, with light gray speckling on head and 
neck, ommatophores not darker. 

Mantle collar (MC) swollen and rolled, no glandular extension 
onto pallial roof, a dark gray streak on expanded portion. Pneumo- 
stome masked by expansion of collar, no mantle lobe development. 
Anus (A) and external ureteric pore (KX) opening beside each other 
just inside pneumostome. 

Pallial region (fig. 57a) extending % whorl apically. Lung roof 
with black specklings along edges of pulmonary vein, otherwise 
clear. Kidney (K) bilobed, 2.57 mm. long, rectal arm slightly longer 
than pericardial and crossing hindgut onto parietal wall. Ureter 
(KD) typical, a narrow strip of lung roof visible between arms. Heart 
(H) slightly more than Vs length of kidney, angled to hindgut. Princi- 
pal pulmonary vein (HV) unbranched, fading out just short of mantle 
collar. Hindgut (HG) deflecting from parietal-palatal margin just 
before end of kidney. 

Ovotestis (fig. 57b, G) a single clump of palmately clavate al- 
veoli, lying parallel to whorl sides, occupying Vs whorl above stomach 
apex. Hermaphroditic duct (GD) slender at first, expanded and 
iridescent in middle, markedly kinked just before albumen gland, 
embedded and narrow before talon. Albumen gland (GG) with rather 
large acini, surface indented by intestinal loops and spermathecal 
head. Talon (fig. 57d, GT) with globular head, hermaphroditic duct 
inserting at base. Carrefour imbedded, a slightly expanded tube from 
neck of talon, opening into prostate-uterus. Prostate (DG) of large 
acini opening into groove on side of uterus, acini complexly folded. 
Uterus (UT) bipartite, upper chamber with very thin walls. 

Vas deferens (VD) a broad tube with heavily glandular walls to 
penioviducal angle, where it narrows abruptly to ascend along side of 
penis. Epiphallus (E) large, twisted, entering head of penis laterally, 
with a peculiar Y-shaped valve (fig. 57e) at entrance of vas deferens, 
lower portion with glandular pilasters, entering penis laterally on 
head (fig. 57c). Penial retractor (PR) short, thick, arising on dia- 
phragm, inserting onto head of penis. Penis (P) about 1.75-1.85 mm. 
long, club-shaped, gradually tapering to atrium, internally (fig. 57c) 
with large verge (PV) through which epiphallic pore (EP) enters 
medially, a reduced sphincter pilaster and a moderately large, typi- 
cal pocket pilaster. Atrium (Y) of average length, with weak lon- 
gitudinal pilasters. 

Free oviduct (UV) a long, thick muscular tube with an apical 
muscle attaching to right tentactular retractor, internally (fig. 57f) 
with heavy glandular walls ending in a hemispherical protrusion 
with subcentral pore (UVO) just at junction with spermathecal base, 
externally smoothly muscled. Spermatheca (S) with head lying next 
to albumen gland, slender shaft bound to diaphragm tissue to base of 
uterus, then grossly expanded to vagina level, with complex pilasters 
forming a circular ring at area of vagina, with a flap obviously capa- 
ble of closing the spermathecal opening (SO). Vagina (V) quite 
thin-walled, with weak internal pilasters. 

Free muscle system typical, muscle from free oviduct quite 
prominent. Right ommatophoral retractor passing through peni- 
oviducal angle. 

Buccal mass and esophagus without unusual features. Stomach 
occupying a little more than % whorl, shape and expansion typical. 
Intestinal looping occupying less than Vs whorl above pallial cavity 
apex. 

Salivary glands fused across top of esophagus. 

Central tooth of radula distinctly smaller than 1st lateral, 
tricuspid with mesocone extending anterior to basal plate edge. Lat- 
erals 4-5, with endocone increasing regularly, transitions to margi- 
nals marked by shift in support size and increasing size of endocones. 



Marginals 6-7, outer ones with endocone and mesocone subequal, 
ectocone always much smaller, no evidence of ectoconal splitting. 

(Based on FMNH 153165, 4 whole individuals, 4.9-5.1 mm. in 
diameter.) 



Sinployea intermedia, new species. Figures 52d-f, 
53f-h. 

Diagnosis. Shell large, diameter 3.21-3.81 mm. (mean 3.49 
mm.), with 4'/4-4% rather tightly coiled whorls. Apex and spire mod- 
erately and evenly elevated, last whorl descending more rapidly, 
spire protrusion about '& body whorl width, H/D ratio 0.534-0.616 
(mean 0.581). Apical sculpture of 9-13 (mean 10.57) very fine spiral 
cords. Postnuclear whorls with low, rounded, prominent, protrac- 
tively sinuated radial ribs, 63-96 (mean 82.1) on the body whorl, 
whose interstices are usually 3-5 times their width. Ribs/mm. 
5.75-9.16 (mean 7.66). Microsculpture of fine radial riblets, 5-10 
between each pair of major ribs, crossed by slightly finer and more 
crowded spiral riblets. No secondary spiral cording. Umbilicus nar- 
row, U-shaped, last whorl decoiling more rapidly, contained 4.59- 
6.12 times (mean 5.43) in the diameter, margins rounded. Whorls 
only slightly flattened laterally above periphery, strongly rounded 
on basal margin. 

Sinployea intermedia (fig. 52d-f) has a distinctly 
more protruded spire, wider umbilicus, slightly greater 
number of apical cords and is larger than any of the S. 
allecta (fig. 54a-e) races. Society and Cook Island 
species of the same size are more widely umbilicated 
and have quite different ribbing structure. 

Description. Shell larger than average, with slightly more 
than 4'/2 moderately tightly coiled whorls. Apex and spire moder- 
ately and evenly elevated, body whorl descending more rapidly, H/D 
ratio 0.592. Embryonic whorls l'/2, sculpture of 8 relatively weak 
spiral ribs with irregularly radially ribbed last quarter whorl. Re- 
maining whorls with low, rounded, moderately widely spaced, pro- 
tractively sinuated radial ribs, 90 on pregerontic portion of body 
whorl, whose interstices are about 2-3 times their width. Micro- 
sculpture of extremely fine microradial riblets, 5- 10 between each 
pair of major ribs, and smaller microspiral riblets forming a latticed 
pattern. Sutures relatively deep, whorls strongly rounded above and 
on basal margin, body whorl slightly deflected at aperture. Um- 
bilicus narrowly U-shaped, becoming more widely open on last 
whorl, contained 4.91 times in the diameter. Color light yellowish 
white with numerous irregular reddish flammulations. Aperture cir- 
cular, slightly compressed laterally above periphery by deflection of 
lip, inclined about 15 from the shell axis. Height of holotype 2.11 
mm., diameter 3.56 mm. 

Holotype. Swains Island: 200 yd. inland on west 
side at 20 ft. elevation. Collected under coconut husks 
and fallen limbs by Elwood C. Zimmerman on August 
21, 1940. BPBM 186666. 

Range. Swains Island, American Samoa. 

Paratypes. Swains Island: west side, 200-500 yd. 
inland at 20 ft. elevation (45 specimens, BPBM 
186666-8, BPBM 186679). 

Remarks. Differences from Sinployea allecta are 
adequately covered in the diagnosis above. Although 
closely related to that species, I have no doubt that 
Sinployea intermedia is distinct. Presence of an en- 
demic species on Swains Island, a very isolated but 
heavily vegetated atoll, was unexpected. Swains Island 
is less than 20 ft. high and not over IVz mile in greatest 
dimension (Bryan, 1942, p. 83). 



132 



SOLEM: ENDODONTOID LAND SNAILS 




VD 



UVO 



FIG. 57. Anatomy of the Samoan Sinployea complementaria (Mousson). Station 18, Mt. Solaua, Upolu, Samoa. FMNH 153165: a, pallial 
region; b, genitalia; c, interior of penis; d, detail of talon-carrefour area; e, junction of vas deferens and epiphallus; f, interior of spermatheca and 
free oviduct junction. Scale lines equal 1 mm. (SH). 



Description of soft parts. Foot and tail shorter than shell 
diameter. Sole undivided, slightly tapering posteriorly. Pedal 
grooves high on sides of foot, prominent. 

Body color light yellow-white, no grayish markings. 

Mantle collar (MC) thick and swollen in available material, no 
glandular extension onto mantle roof. 

Pallial region (fig. 530 extended length about '/a whorl. Lung 
roof clear of granulations. Kidney (K) bilobed, rectal lobe longer than 
pericardial, greatest length 1.65-2.00 mm., differing from S. allecta 



in having less overlap of hindgut and greater lung roof visible be- 
tween arms of ureter (KD). Heart (H) and principal pulmonary vein 
(HV) typical. Hindgut (HG) following parietal-palatal margin for 
moderate distance above kidney apex. 

Ovotestis (fig. 53g, G) composed of one or two acinar clumps in 
typical position. Hermaphroditic duct (GD), albumen gland (GG), 
talon (GT), prostate (DG), and uterus (UT) without unusual features. 

Vas deferens (VD) and epiphallus (E) as in Sinployea allecta. 
Penial retractor (PR) as in S. allecta. Penis (P) 1.2-1.6 mm. long, 



SYSTEMATIC REVIEW 



133 



tapering more on basal portion than inS. allecta, externally strongly 
swollen on upper half. Internally (fig. 53h) with typical stimulatory 
pad, vergic papilla as in S. allecta. Atrium (Y) very short, without 
unusual features. 

Free oviduct (UV), spermatheca (S), and vagina (V) typical; lat- 
ter a trifle longer than in S. allecta. 

(Based on BPBM 186666, several partial examples.) 

WESTERN POLYNESIAN AND MICRONESIAN Sinployea 

Whereas the Sinployea from Tonga, Hoorn Islands, 
Ellice Islands, and Rotuma show many similarities, in- 
clusion of the single species from Kusaie in the 
Caroline Islands and a possibly introduced form from 
Saipan in the Marianas is merely for convenience in 
reducing the number of subheadings. Sinployea 
kusaieana (fig. 63a-c) lacks gray coloration on the 
head and ommatophores, has monochrome shell colora- 
tion, a less protruded spire, fewer and more loosely 
coiled whorls, and the umbilicus wider and V-shaped. 
Two specimens collected on Saipan could represent an 
introduced population, a mislabeled set, or another un- 
described Micronesian species. 

Of the remaining species, only Sinployea 
pseudovicaria (fig. 61a-c) with its low H/D ratio and 



wide umbilicus (table XXVII) is distinctive. The Tahi- 
tian S. lamellicosta (Garrett) (fig. 40a-c) is virtually 
identical in size and shape, but has a depressed spire, 
averages less than half as many ribs, and has slightly 
fewer whorls at a given diameter. Other widely umbili- 
cated Sinployea are either much bigger (S. canalis, fig. 
49a-c), have many more whorls (S. planospira, fig. 
46d), or are much smaller (S. nissani, S. navutuensis). 
Sinployea rotumana (E. A. Smith) (fig. 61d-f) perhaps 
comes closest of any species to being median in its fea- 
tures. Only the slightly more numerous and crowded 
radial ribs depart from the median measurements out- 
lined in Table XV. Sinployea ellicensis (fig. 62a-f) is a 
rather small, normally elevated species with crowded 
radial ribbing. Sinployea vicaria (Mousson) (fig. 58a-d) 
is larger and with a rather elevated spire. Both of these 
species have less marked lateral whorl flattening than 
species from Eastern Polynesia. Although measure- 
ments overlap, the round apertures and generally ele- 
vated spires would allow immediate recognition in 
mixed sets. 

Because none of the species are sympatric, no for- 
mal key has been prepared. The following geographic 
notes will locate material of known taxa: 



TABLE XXVII. - RANGE OF VARIATION IN WESTERN POLYNESIAN AND MICRONESIAN SINPLOYEA. 



NAME 

ellicensis 
ellicensis 

ellicensis 

nukulaelaeana 

vicaria 
vicaria 
(Mousson) 

vicaria 

paucicosta 

rotumana 
(Smith) 

pseudovicaria 
Vaitupu 

Eua (?) 
kusaieana 
sp. (Saipan) 



NUMBER OF 
SPECIMENS 



326 

87 
9 

21 
7 

61 
2 



RIBS RIBS/MM. HEIGHT DIAMETER H/D RATIO 

100.3(79-125) 13.23(10.55-16.69) 1.36(1.13-1.68) 2.47(2.20-2.81) 0.549(0.486-0.600) 

84.5(72-102) 13.02(11.06-15.20) 1.11(1.06-1.16) 2.08(1.99-2.20) 0.533(0.496-0.561) 

108.5(77-145) 12.65(8.72-15.91) 1.51(1.17-1.89) 2.73(2.14-3.15) 0.552(0.469-0.633) 

77.1(51.98) 9.80(6.71-11.44) 1.45(1.18-1.78) 2.49(2.22-2.76) 0.585(0.500-0.659) 

105.2(84-144) 11.74(9.79-14.15) 1.55(1.25-1.81) 2.84(2.60-3.24) 0.547(0.481-0.607) 

108.1(82-127) 11.39(9.56-15.17) 1.57(1.41-1.74) 3.24(3.03-3.52) 0.486(0.454-0.525) 

94.8(81-113) 8.93(8.43-9.77) 1.61(1.45-1.74) 3.37(3.06-3.68) 0.479(0.455-0.509) 

91.1(84-103) 11.25(10.27-13.12) 1.32(1.19-1.49) 2.60(2.52-2.75) 0.506(0.462-0.542) 

85.0(82-88) 11.44(11.20-11.67) 1.41(1.38-1.45) 2.37(2.34-2.40) 0.598(0.592-0.603) 



APICAL 
CORDS 



SPIRE 
ELEVATION 



BODY WHORL 
WIDTH 



e.ell. 



e.nuk. 



v. pau. 

rotum. 
pseud. 
Vait. 

Eua 
kusai. 
sp. 



WHORLS UMBILICUS D/U RATIO CORDS ELEVATION WIDTH SP/BWW 

4 1/8(4-4 1/2) 0.56(0.43-0.76) 4.39(3.48-5.46) 9.42(8-12) 0.15(0.10-0.23) 0.82(0.72-0.92) 0.183(0.125-0.259) 

3 7/8+(3 5/8-4) 0.48(0.44-0.59) 4.31(4.13-4.66) 11.67(11-13) 0.11(0.10-0.16) 0.68(0.64-0.72) 0.166(0.136-0.250) 

4 1/4(3 3/4-4 3/4) 0.58(0.41-0.75) 4.47(3.88-5.80) 10.63(9-12) 0.21(0.10-0.38) 0.86(0.69-1.02) 0.251(0.111-0.417) 
4 1/2(4 1/4-5) 0.69(0.49-0.72) 4.14(3.62-4.64) 11.50(10-12) 0.21(0.10-0.33) 0.79(0.69-0.89) 0.268(0.125-0.385) 
4 1/4(4-4 5/8) 0.66(0.58-0.81) 4.32(3.93-4.86) 9.67(8-11) 0.17(0.07-0.25) 0.95(0.84-1.04) 0.175(0.073-0.268) 

4 3/8+(4 1/4-4 5/8) 1.03(0.79-1.22) 3.17(2.89-4.05) 11.74(10-13) 0.10(0.07-0.13) 0.86(0.79-0.95) 0.121(0.083-0.160) 

4 1/2(4 1/4-4 3/4) 1.08(0.99-1.25) 3.13(2.95-3.30) 

3 7/8+(3 3/4-4 1/8) 0.74(0.69-0.79) 3.51(3.21-3.67) 9.20(8-10) 0.12(0.10-0.16) 0.79(0.72-0.86) 0.150(0.125-0.208) 

3 7/8+ 0.53(0.49-0.56) 4.52(4.30-4.73) 



134 



SOLEM: ENDODONTOID LAND SNAILS 



Tonga and Hoorn Islands: S. vicaria (Mousson, 
1871) 
Sinployea vicaria paucicosta, new subspecies 

on southern Vava'u 

Rotuma Island: S. rotumana (E. A. Smith, 1897) 
Ellice Islands 

Vaitupu: S. pseudovicaria, new species 

(widely umbilicated) 

Funafuti: S. ellicensis ellicensis, new sub- 
species 
Nukulaelae: S. ellicensis nukulaelaeana, new 

subspecies 
Caroline Islands 

Kusaie: S. kusaieana, new species 
Mariana Islands 

Saipan: Sinployea sp. 

Sinployea vicaria (Mousson, 1871). 

Several small samples from scattered islands of 
the Tonga and Hoorn group collected by Graeffe in the 
1860s and 1870s, plus material from Vava'u, Ton- 
gatapu, and Eua collected by L. Price in 1966 agree 
closely in proportions (table XXVIII). Four samples 
from Vava'u have a distinctive pattern of more widely 
spaced ribbing and are separated as a subspecies, 
Sinployea vicaria paucicosta (fig. 58d). Size differences 
among the remaining samples may be attributed to 
sample bias and/or ecophenotypic dwarfing (p. 108). 
They are grouped as a single subspecies. It is quite 
possible that more ample material, particularly from 
Futuna, will cause a revision in this judgment. 

General appearance and sculpture are much more 
similar to the smaller Cook Island Sinployea, S. atien- 
sis, S. andrewi, and S. peasei, than to any of the Sa- 
moan or Fijian species. 

Sinployea vicaria vicaria (Mousson, 1871). Figures 
58a-c, 60a-d. 

Patula (Patula) vicaria Mousson, 1871, J. de Conchyl., 19, pp. 
11-12, pi. 3, fig. 2 Futuna, Hoorn Group; Hufena on Hapai 
(probably is Uoleva Id., Haapai Group) and Vavao, Tonga Is- 
lands. 

Helix vicaria (Mousson), Pfeiffer, 1876, Monog. helic. viv., 7, p. 187. 

Helix modicella (Ferussac), Tryon, 1887, Man. Conchol., (2) 3, p. 
38, pi. 8, figs. 31-32. 

Rhytida (Ouagapia) vicaria (Mousson), Mollendorff, 1903, Syst. 
Conchyl. Cab., I, 12b, pp. 83-84, pi. 13, figs. 16-17. 

Diagnosis. Shell variable in size, generally slightly smaller 
than average, diameter 2.14-3.15 mm. (mean 2.73 mm.), with 
3%-4% rather tightly coiled whorls. Apex and spire markedly and 
almost evenly elevated, last whorl descending moderately to much 
more rapidly, spire protrusion averaging V* body whorl width, H/D 
ratio 0.469-0.633 (mean 0.552). Apical sculpture of 9-12 (mean 
10.63) very fine spiral cords. Postnuclear whorls with narrow, 
rounded, crowded, rather strongly protractively sinuated radial ribs, 
77-145 (mean 108.5) on the body whorl, whose interstices are 1-3 
times their width. Ribs/mm. 8.72-15.91 (mean 12.65). Micro- 
sculpture of very fine radial riblets, 5-8 between each pair of major 
ribs, barely visible microspiral riblets, and rather closely spaced sec- 
ondary spiral cords equal in size to microradial riblets. Umbilicus 
rather narrow, U-shaped, last whorl decoiling more rapidly, con- 
tained 3.88-5.80 times (mean 4.47) in the diameter, margins 
rounded. Whorls less strongly flattened laterally than in Society or 
Cook Island species. 



Sinployea vicaria (table XXVII) is very similar to 
S. atiensis (table XIX) in all measurements, differing 
only slightly in ribbing and umbilical coiling. In S. 
atiensis (fig. 44d-f) the umbilicus decoils regularly, the 
secondary spiral cording is noticeably wider spaced, 
and the whorl contour is more strongly flattened later- 
ally. Sinployea vicaria (fig. 58a-c) usually has a more 
rapid decoiling of the umbilicus and slightly more 
crowded ribbing. Sinployea pseudovicaria (fig. 61a-c) 
is immediately separable by its nearly flat spire and 
much more widely open umbilicus. Sinployea rotu- 
mana (fig. 61d-f) has the body whorl strongly flattened 
laterally and more crowded secondary spiral cording. 
Sinployea neglecta (fig. 41d-f) has finer ribs, fewer 
microradial riblets, a V-shaped umbilicus, and greater 
lateral whorl flattening. 

Description. Shell of average size, with 4V* normally coiled 
whorls, apex barely emergent, whorls of spire descending moder- 
ately, last whorl more rapidly, H/D ratio 0.538. Apical whorls 1%, 
sculpture of 11 fine, moderately widely spaced spiral ribs with a faint 
microsculpture of radial ribs and a secondary intrusion of radial 
growth wrinkles near the end. Postnuclear whorls with prominent, 
narrow, protractively sinuated radial ribs, 82 on the body whorl, 
whose interstices are 2-4 times their width. Microsculpture of very 
fine radial riblets, 4-8 between each pair of major ribs, crossed by 
barely visible spiral riblets with a prominent secondary sculpture of 
closely spaced spiral cording. Sutures moderately impressed, whorls 
slightly flattened laterally above and below periphery with evenly 
rounded outer margins. Color light yellowish white with irregular 
somewhat vague reddish flammulations. Umbilicus V-shaped, mod- 
erately open, last whorl decoiling a little more rapidly, contained 
5.05 times in the diameter. Aperture subcircular, somewhat flat- 
tened basally, inclined about 20 from shell axis. Height of lectotype 
1.61 mm., diameter 3.00 mm. 

Lectotype. Tonga: Vava'u. Collected by E. 
Graeffe. Zoologisches Institut der Universitat, Zurich. 

Range. Futuna, Hoorn Islands; Vava'u, Tonga- 
tapu, Eua, and Ha'apai Group, Tonga Islands. 

Material. Tonga: Vava'u (3 specimens, Zurich): 
Utula'aina (Station T-13) near Holonga in heavy forest 
at 10-100 ft. elevation (265 specimens, FMNH 
152372-3); 200 yd. inland on cliffs at 200-350 ft. eleva- 
tion, north side of island (4 specimens, BPBM 87859). 
Ha'apai Group, Hufena (= Uoleva) (4 specimens, 
Zurich). Eua (6 specimens, Zurich, FMNH 150773, 
BMNH 93.9.25.81): heavy primary forest near south 
end (Stations T-21, T-22) at 850-1,000 ft. elevation (5 
specimens, FMNH 152519, FMNH 152379). Ton- 
gatapu: Manima Id., 4 miles northeast of Nuku'alofa 
(Station T-4) in second growth on a coral islet (1 speci- 
men, FMNH 152397); about 11 miles northwest of 
Nuku'alofa, 200 yd. inland (Station T-26) in coastal 
scrub (32 specimens, FMNH 152312, FMNH 152461). 
Fukave, near Tonga tapu (1 specimen, BPBM 53524). 

Hoorn Islands: Futuna (6 specimens, Zurich, 
FMNH 150772). 

Remarks. Recorded variation between sets of 
Sinployea vicaria (table XXVIII) is quite large, with a 
spread of 0.75 mm. between the smallest in diameter 
(recent collection from Eua) and the largest (type series 
from Vava'u). Two factors reduce the possibility that 
this has any systematic significance. The specimens in 
the Mousson collection at Zurich were selected by 



SYSTEMATIC REVIEW 



135 



TABLE XXVIII. - LOCAL VARIATION IN SINPLOYEA VICARIA. 



vicarla vlcarla 


NUMBER OF 
SPECIMENS 

3 


RIBS 

100.39.21 
(82-111) 


RIBS/MM. 

10.5410.920 
(8.72-11.61) 


HEIGHT 

1.7210.083 
(1.62-1.89) 


DIAMETER 

3.0510.051 
(2.98-3.15) 





H/D 

.56610 
(0 


RATIO 

.034 
.526-0.633) 


Vavau 
Zurich 


Hufena, Haapaii 
Zurich 


4 


101.02.68 
(95-108) 


11.9910. 
(11 


307 
.49-12.82) 


1.6010. 
(1. 


045 
49-1.71) 


2.7010 
(2 


.035 
.65-2.80) 


0.593*0 
(0, 


,0115 
.563-0.612) 


Tongatapu 15 
Sta. T-26 
FMNH 152312, -461 


118.33.87 
(96-145) 


13.340. 
(10 


369 
.59-15.76) 


1.5310, 
(1, 


039 
30-1.78) 


2.8210 
(2 


.050 
.52-3.09) 





.54110 

(0, 


.0087 
,469-0.596) 


Fukave, Tongatapu 
BPBM 53524 


1 


128 


14.40 




1.51 




2.83 







.535 




Eua 
Zurich 


5 


105.012.98 
(96-113) 


11.8410. 

(10 


381 
.80-13.17) 


1.5010. 

(1. 


034 
.42-1.62) 


2.8410.049 
(2.75-3.03) 





.52710.0063 
(0.511-0.547) 


Sta. T-21 
FMNH 152519 


4 


92.011.53 
(90-95) 


12.7210. 
(11 


683 
.36-13.42) 


1.2510.062 
(1.17-1.43) 


2.3010 

(2 


.088 
.14-2.55) 





.546+0 
(0, 


,0166 
,518-0.585) 


BMNH 93.9.25.81 


1 


143 


15.91 




1.74 




2.86 







.609 




Put una 
Zurich 


5 


96.2*6.40 
(77-111) 


12.2010. 
(11 


442 
.03-13.69) 


1.4210 
(1 


.055 
.33-1.61) 


2.5010 
(2 


.094 
.22-2.80) 





.56910 
(0 


.0100 
.546-0.600) 


vicaria paucicosta 
























Vavau 
Sta. T-8 
FMNH 152488 


10 


78.015.85 
(51-98) 


9.3910. 
(6. 


590 
71-11.42) 


1.5110 
(1 


.041 
,27-1.78) 


2.6310 
(2 


.038 
.42-2.76) 





.57410 
(0 


.012 
.500-0.643) 


Sta. T-9 
FMNH 152414 


28 


76.811.68 
(67-89) 


9.9510.210 
(8.54-11.44) 


1.4310 
(1. 


,018 
,18-1.63) 


2.4410 
(2 


.021 
.22-2.62) 





.58910 
(0 


.0062 
.518-0.659) 


Sta. T-13 
FMNH 152372-3 


60 


89.6*1.20 
(66-108) 


10.39-10. 
(8. 


131 
08-12.59) 


1.48+0 

a 


,016 
,25-1.81) 


2.76+0.025 
(2.43-3.35) 





.536+0.0032 
(0.467-0.588) 






WHORLS 
v. vicaria 

Vavau 4 3/8+(4 1/4-4 3/4) 

Hufena 4 3/8+(4 1/4-4 5/8) 

Tongatapu 4 1/4-OV-4 1/2) 

Fukave 4 1/8 

Eua 

Zurich 4 l/8+(4-4 1/4) 

Sta.T-21 4-(3 3/4-4 1/8) 

BMNH 41/4 

Futuna 4 l/4+(4-4 1/2) 

v.paucicos. 

Sta. T-8 4 l/2+(4 1/2-5) 

Sta. T-9 4 3/8(4 1/4-4 5/8) 

Sta. T-13 4 l/4-(3 3/4-4 5/8) 



D/U RATIO 



SPIRE 
APICAL CORDS PROTRUSION 



SP/BWW 



0.6510 
(0 


.040 
,59-0.72) 


4.7110 
(4 


.312 
.09-5. 


05) 


11 


0.2510 
CO 


.067 
.16-0.38) 


0.9710 
(0 


.025 
.94-1.02) 


0.257*0 
(0 


.074 
.161-0.404) 


0.6210, 
(0, 


,024 
,56-0.67) 


4.3310 
(4 


.130 
.13-4. 


71) 


10.33*0.883 
(9-12) 


0.280 
(0 


.032 
.20-0.35) 


0.85*0 
(0 


.022 
.81-0.89) 


0.33510 

(0 


.038 
.222-0.389) 


0.66+0 
(0 


,017 
,51-0.76) 


4.28+0 
(3 


.092 
.88-5. 


26) 


11.20.374 
(10-12) 


0.23*0 
(0 


.014 
.16-0.33) 


0.86*0.013 
(0.77-0.92) 


0.26910 
(0 


.016 
.189-0.417) 


0.66 




4.30 









0.20 




0.82 




0.240 




0.63*0 
(0 


,024 
.56-0.61) 


4.5210 
(4 


.116 
.20-4. 


88) 


11.0010.578 
(10-12) 


0.150 
(0 


.078 
.10-0.21) 


0.9110 
(0 


.021 
.87-0.99) 


0.167*0 
(0 


.026 
.111-0.241) 


0.4710.037 
(0.41-0.57) 

0.49 

0.38+0.038 
(0.44-0.66) 


4.9110 
(4 

5.80 

4.3410 
(4 


.224 
.43-5. 

.172 
.03-5. 


48) 
00) 


9.7510.479 
(9-11) 


0.1710 
(0 


.012 
,15-0.20) 


0.72*0 
(0 


.022 
.69-0.79) 


0.22810 
(0 


.0172 
.208-0.279) 


0.19*0 
(0 


.022 
.13-0.26) 


0.84*0 
(0 


.009 
.82-0.87) 


0.23110 
(0 


.0199 
.157-0.302) 


0.6510.014 
(0.59-0.72) 


4.06*0 
(3 


.089 
.62-4. 


54) 


11.2510.479 
(10-12) 


0.23*0 
(0, 


.020 
.15-0.33) 


0.8310 
(0 


,017 
,76-0.89) 


0.27610 
(0, 


,0243 
,196-0.385) 


0.59+0 
(0 


.008 
.49-0.67) 


4.17+0 
(3 


.042 
.73-4. 


64) 


11.67*0.211 
(11-12) 


0.21*0.010 
(0.10-0.33) 


0.7710. 
(0, 


,007 
,69-0.86) 


0.265*0, 
(0, 


,130 
,125-0.360) 


0.5910.008 
(0.46-0.72) 


4.7110.054 
(3.90-5. 


71) 





0.16*0, 
(0, 


.006 
07-0.26) 


0.9110, 

(0, 


008 
79-1.08) 


0.18110, 
(0, 


0071 
080-0.286) 



Mousson from field collections that were then depos- 
ited in the Godeffroy collection. Presumably these 
large specimens can be equated with the retention by 
Garrett of larger examples in his collection (for exam- 
ple see S. proximo, p. 108). Hence, a major part of the 
larger size shown by the Mousson Eua Island and 
Vava'u material could result from biased sampling. 
Second, the material collected by Price in 1966 was 
taken after an extended period of drought. Quite possi- 
bly, ecophenotypic dwarfing was involved in the rela- 



tively small size of these shells. Since the proportions, 
whorl counts, and ribbing are essentially identical, I 
am ignoring the size differences and consider them 
conspecific. Inclusion of the Futuna shells is less cer- 
tain, but their proportions and ribbing match the Ton- 
gan examples. 

Comparison of measurements in Tables XIX and 
XXVII demonstrates that Sinployea vicaria and S. 
atiensis are extremely similar in size and proportions. 
Direct comparisons of specimens show that in S. vi- 



136 



SOLEM: ENDODONTOID LAND SNAILS 







FIG. 58. a-c, Sinployea vicaria vicaria (Mousson). Vava'u, Tonga. Lectotype. Zoologisches Institut der Universitat Zurich; d, Sinployea 
vicaria paucicosta, new subspecies. Station T-9, Mo'ungalafa, Vava'u, Tonga. Holotype. FMNH 158449. Scale lines equal 1 mm. (SH). 



caria the secondary spiral cording is much more 
crowded, the apical cords reduced in prominence, the 
body whorl less distinctly flattened laterally above the 
periphery, and the last whorl of the umbilicus decoils 
slightly more rapidly. On the average, specimens of S. 
vicaria have about one more rib/mm, and perhaps 15 
more ribs on the body whorl, but overlap is complete 
and these characters cannot be used to separate the 
taxa. The similarities are considered to be convergent 
and caused by the limited patterns of coiling available 
to these snails. 

Mousson's original measurements of 3 mm. in 
diameter and 4 l /2 whorls are most closely matched by 
the specimen from Vava'u selected as lectotype. It is 
atypical in having an umbilicus that is slightly nar- 
rower than usual because of regular last whorl decoil- 
ing (fig. 58c). 

Description of soft parts. Foot and tail slightly shorter than 
shell diameter, truncated anteriorly, bluntly rounded, and slightly 
tapering posteriorly. Sole undivided, pedal grooves prominent, high 
on foot, no caudal horn or middorsal groove developed. Slime net- 
work typical. Head projecting in front of foot, ommatophores long, 
dark in color. Gonopore below right ommatophore and behind right 
rhinophore, a vertical, short slit in body wall. 

Body color yellow-white on sides and sole of foot, back of head 
and neck, ommatophores, and inner part of mantle collar heavily to 
moderately speckled with gray. 



Mantle collar (MC) wide, white on outer edge, heavily marked 
with gray on inner edge, no glandular extension onto pallial roof. 
Anus (A) opening just anterior to external ureteric pore (KX). 

Pallial region (fig. 60a) extending slightly less than Vfe whorl 
apically. Lung roof clear, without granulations. Kidney (K) about 
1.45 mm. long, rectal lobe slightly longer than pericardial. Rectal 
arm crossing hindgut and extending well down onto parietal wall. 
Ureter (KD) typical, a strip of lung roof visible between arms. Heart 
(H) nearly parallel to hindgut, % length of pericardial kidney lobe. 
Principal pulmonary vein (HV) narrow, unbranched, fading out just 
before mantle collar. Hindgut (HG) extending about 0.3 mm. above 
kidney apex before starting to deflect downward. 

Ovotestis (fig. 60b) a single clump of palmately clavate alveoli 
above stomach apex, lying parallel to plane of coiling, sharply trun- 
cate anteriorly, soft parts extending one full whorl above ovotestis 
apex. Hermaphroditic duct (GD) typical. Albumen gland, talon, pros- 
tate, and uterus without unusual features. 

Vas deferens (fig. 60c, VD) typical, epiphallus (E) more than half 
length of penis, entering penis apically through a longitudinal 
epiphallic pore (EP). Penial retractor (PR) short, attached in U-fan to 
head of penis, arising from diaphragm. Penis (P) about 1.4 mm. long, 
lower third very narrow, internally (fig. 60d) with distinct collar 
(PVS) around prominent vergic papilla (PV). Normal circular muscle 
collar reduced to a low ridge, main stimulatory pad quite prominent. 
Atrium (Y) long, medially swollen. 

Free oviduct (UV) enlarging rapidly in diameter just before 
junction with spermatheca. Spermatheca (S) and vagina (V) typical. 

(Based on FMNH 152373, several expanded adults, 2.73-2.93 
mm. in diameter, with 4'/8-4V4 whorls.) 



SYSTEMATIC REVIEW 



137 




FIG. 59. Shell sculpture details of Sinployea uicaria vicaria (Mous- 
son). Station T-13, Vava'u, Tonga. FMNH 152373: a, sculpture of 
body whorl (535x); b, fracture section through body whorl showing 
periostracal (upper) and calcareous (lower) elements of sculpture 

(950x). 



Sinployea vicaria paucicosta, new subspecies. Fig- 
ure 58d. 

Diagnosis. Shell small, diameter 2.22-2.76 mm. (mean 2.49 
mm.), with 4V4-5 rather tightly coiled whorls. Apex and first post- 
nuclear whorl barely emergent, lower whorls descending more 
rapidly, body whorl much more rapidly, spire protrusion slightly 
more than Vt body whorl width, H/D ratio 0.500-0.659 (mean 0.585). 
Apical sculpture of 10-12 (mean 11.5) fine spiral cords. Postnuclear 
whorls with prominent, narrow, rather widely spaced, protractively 
sinuated radial ribs, 51-98 (mean 77.1) on the body whorl, whose 
interstices are 5-8 times their width. Ribs/mm. 6.71-11.44 (mean 
9.80). Microsculpture of very fine radial riblets, 812 between each 
pair of major ribs, barely visible microspiral riblets, and prominent, 
relatively crowded secondary spiral cords. Umbilicus open, U- 
shaped, last whorl decoiling more rapidly, contained 3.62-4.64 times 
(mean 4.14) in the diameter, margins rounded. Whorl contour and 
sutures as in the nominate race. 

Sinployea vicaria paucicosta (fig. 58d) differs from 
the nominate race in its more sharply defined and more 
widely spaced ribbing. The smaller size, slightly 
greater H/D ratio, wider umbilicus, and average of one 
more apical cord are of uncertain significance. 
Sinployea ellicensis (fig. 62a-0 has much more 



crowded ribbing and fewer whorls, whereas S. 
pseudouicaria (fig. 61a-c) is widely umbilicated and 
much larger. 

Description. Shell small, with 4% rather tightly coiled whorls. 
Apex barely elevated, later whorls descending much more rapidly, 
H/D ratio 0.595. Apical whorls and early spire with sculpture eroded. 
Postnuclear whorls with narrow, prominent, strongly protractively 
sinuated radial ribs, 69 on the body whorl, whose interstices are 4-6 
times their width. Microsculpture of very fine radial riblets, 6-12 
between each pair of major ribs, crossed by barely visible spiral rib- 
lets. Secondary spiral cording prominent and relatively crowded. Su- 
tures impressed, whorls strongly rounded above, slightly flattened 
laterally above periphery and on basal margin. Color light yellow- 
horn, with broad, irregular, reddish flammulations that fade out on 
shell base. Umbilicus open, U-shaped, last whorl decoiling more 
rapidly, contained 4.00 times in the diameter. Aperture subcircular, 
slightly flattened laterally above periphery, inclined about 20 from 
shell axis. Height of holotype 1.45 mm., diameter 2.43 mm. 

Holotype. Tonga: Vava'u, Station T-9, Mo'un- 
galafa, near Tu'anuku, about 14 miles southwest of 
Neiafu, at 700 ft. elevation. Collected dead in leaf 
mould from patches of heavy forest by Laurie Price on 
January 3, 1966. FMNH 158449. 

Range. Southern part of Vava'u, Tonga Islands. 

Paratypes. Vava'u: near Toulo (Station T-8), 2 
miles south of Neiafu in leaf mould in coral rubble 
under trees above strandline (16 specimens, FMNH 
152488); Mo'ungalafa, near Tu'anuku (Station T-9), 
about 14 miles southwest of Neiafu in leaf mould from 
heavy forest at 700 ft. elevation (69 specimens, FMNH 
152414); Muitola, 15 miles west of Neiafu (Station 
T-10) at 400 ft. elevation in heavy forest (3 specimens, 
FMNH 158455); cliffs at end of peninsula past Makave 
Village (Station T-12), 2 x /2 miles southeast of Neiafu at 
200 ft. elevation from leaf mould (1 specimen, FMNH 
152360). 

Remarks. The four localities for Sinployea ui- 
caria paucicosta are all on the south and southeast 
coasts of Vava'u. Material of the nominate race is 
known on Vava'u from the original collection of 
Graeffe in the 1860s, 265 shells from near Holonga 
(Station T-13) on the north tip, and four juvenile 
specimens from the north side (BPBM 87860) collected 
at 200-350 ft. elevation by J. E. Hoffmeister on July 
10, 1928. It is probable that they are geographic races, 
but the possibility does exist that the variation is 
ecophenotypic or that they are separate species. Unfor- 
tunately, weather conditions were so dry that no living 
material could be obtained of S. v. paucicosta (see also 
p. 183). 

Material from Stations T-8 and T-9 differs slightly 
in size (table XXVIII) but is almost identical in propor- 
tions and ribbing. Three examples from Station T-10 
are subadult. The single shell from Station T-12 is 
badly worn and quite large, diameter 3.13 mm. One 
portion of the shell contains traces of widely spaced 
ribbing in the suture. Although it has been referred to 
this subspecies, the measurements have not been used 
in figuring means and ranges because it might belong 
to the nominate race. Further collecting is needed in 
order to establish the identity of this population. 



138 



SOLEM: ENDODONTOID LAND SNAILS 



The name paucicosta refers to the reduced number 
of major radial ribs that characterize this taxon. 

Sinployea rotumana (E. A. Smith, 1897). Figure 
61d-f. 

Champa rotumana E. A. Smith, 1897, Ann. Mag. Nat. Hist., (6) 20, 
p. 520 Rotuma Island; Hedley, 1899, Mem. Australian Mus., 3 
(7), p. 488 as a synonym of "Charopa modicella" I =Sinployea 
ellicensis) from Funafuti. 

Diagnosis. Shell of average size, diameter 2.60-3.24 mm. 
(mean 2.84 mm.) with 4-4% normally coiled whorls. Apex and spire 
slightly to moderately and evenly elevated, last whorl descending 
more rapidly, spire protrusion about Vs-'A body whorl width, H/D 
ratio 0.481-0.607 (mean 0.547). Apical sculpture of 8-11 (mean 9.67) 
fine spiral cords. Postnuclear sculpture of low, rounded, rather 
crowded, strongly protractively sinuated radial ribs, 84-144 (mean 
105.2) on the body whorl, whose interstices are l'/i-3 times their 
width. Ribs/mm. 9.79-14.15 (mean 1L74). Microsculpture of fine ra- 



dial riblets, 4-7 between each pair of major ribs, slightly finer and 
more crowded spiral riblets, with secondary spiral cording that be- 
comes extremely fine on lower spire and very crowded. Umbilicus 
open, U-shaped, last whorl decoiling slightly more rapidly, contained 
3.93-4.86 times (mean 4.32) in the diameter, margins rounded. Su- 
tures deep, whorls rounded to strongly compressed laterally above 
periphery, aperture inclined about 20 from shell axis. 

Sinployea rotumana (fig. 61d-f) has the major rib- 
bing lower and less prominent, the secondary spiral 
cording much more crowded, and the body whorl more 
strongly flattened laterally than in S. vicaria (fig. 
58a-c). Sinployea atiensis (fig. 44d-f) from the Cook 
Islands has fewer major radial ribs, more apical cords, 
and much more widely spaced secondary spiral cord- 
ing. There are also subtle differences in umbilical and 
whorl contours. 

Description. Shell of average size, with a little more than 4 
normally coiled whorls. Apex and early spire barely elevated, body 








LM 





PR 



PVS 



EP 



FIG. 60. Anatomy of the Tongan Sinployea vicaria vicaria (Mousson). Station T-13, near Holonga, Vava'u, Tonga Islands. FMNH 152373: a, 
pallial region; b, apical genitalia; c, terminal genitalia; d, interior of penis. Scale lines equal 1 mm. (CW). 




a 




^^^_ ..'-J"/,ti^ 





FIG. 61. a-c, Sinployea pseudovicaria, new species. Vaitupu, Ellice Islands. Holotype. BPBM 189682; d-f, Sinployea rotumana (E. A. 
Smith). Rotuma Island. BPBM 187049. Scale lines equal 1 mm. (SH). 



139 



140 



SOLEM: ENDODONTOID LAND SNAILS 



whorl descending a little more rapidly, H/D ratio 0.497. Apical 
whorls 1%, sculpture of 10 prominent spiral cords, with a very faint 
secondary sculpture of radial riblets on last quarter whorl. Postnu- 
clear whorls with prominent, rounded, low, slightly protractively 
sinuated radial ribs, 87 on the body whorl, whose interstices are 2-3 
times their width. Microsculpture of fine radial riblets crossed by 
exceedingly fine spiral riblets with a secondary sculpture of minor, 
very crowded spiral cords. Sutures deep, whorls strongly rounded 
above, almost evenly rounded on outer and basal margins. Color 
light yellowish horn with broad, irregular reddish flammulations 
coalescing on base of shell. Umbilicus narrow, U-shaped, regularly 
decoiling, contained 4.22 times in the diameter. Aperture subcircu- 
lar, slightly flattened laterally above periphery, inclined about 20 
from the shell axis. Height of lectotype 1.38 mm., diameter 2.78 mm. 

Lectotype. Rotuma Island. Collected by J. Stan- 
ley Gardiner. BMNH 77.11.8.57. 

Range. Rotuma Island, 300 miles northwest of 
Fiji. 

Parades. BMNH 77.11.8.58-63. 

Material. Rotuma (1 specimen, BPBM 187049). 

Remarks. The single specimen collected by 
Harold St. John in 1938 (BPBM 187049) has the body 
whorl more compressed laterally above the periphery, 
but otherwise compares well with the type set. 
Differences from Sinployea vicaria and S. atiensis are 
covered in the diagnosis above. 

Sinployea ellicensis, new species. 

Materials from Funafuti and Nukulaelae collected 
by Graeffe in 1868 and Charles Hedley in 1899 differ 
from the Tongan S. vicaria (fig. 58a-d) in their 
stronger ribbing and nearly circular aperture. Species 
of similar size from Eastern Polynesia have the aper- 
ture strongly flattened laterally above the periphery 
and usually show large differences in sculpture and/or 
shape. Species from Fiji, such as S. godeffroyana (fig. 
69a-c) and S. princei (fig. 65a-c), differ in ribbing and 
higher spire (latter) or umbilical shape and much finer 
ribbing (former). 

Recognition of two subspecies is done with some 
hesitation because only a single collection has been 
made on Nukulaelae. Specimens from this island are 
much smaller and have more apical cords than the 
examples from Funafuti. Although an equally great 
size difference is found between populations of 
Sinployea vicaria on Tongatapu and Eua Islands (table 
XXVIII), the distinct difference in island topography 
between the latter islands suggests that the distinc- 
tions may be environmentally correlated. Both 
Funafuti and Nukulaelae are atolls. Presumably there 
is no major habitat difference involved. 

Sinployea ellicensis ellicensis, new species and sub- 
species. Figure 62a-c. 

Patula vicaria Mousson, 1873 (not Mousson, 1871), J. de Conchyl., 
21, p. 104 Funafuti, Ellice Islands. 

Champa modicella Hedley, 1899 (not Ferussac, 1840), Mem. Aus- 
tralian Mus., 3 (7), p. 488 Funafuti, Ellice Islands. 

Diagnosis. Shell small, diameter 2.20-2.81 mm. (mean 2.47 
mm.), with 4-4M> rather tightly coiled whorls. Apex and spire 
slightly to moderately and evenly elevated, body whorl descending 



more rapidly, spire protrusion about Vs-Vs body whorl width, H/D 
ratio 0.496-0.561 (mean 0.549). Apical sculpture of 8-12 (mean 9.42) 
prominent spiral cords. Postnuclear whorls with prominent, 
rounded, crowded, slightly protractively sinuated radial ribs, 79-125 
(mean 100.3) on the body whorl, whose interstices are usually less 
than twice their width. Ribs/mm. 10.55-16.69 (mean 13.23). Micro- 
sculpture of fine radial riblets, 3-5 between each pair of major ribs, 
barely visible and extremely crowded spiral riblets, with a secondary 
microsculpture of spiral cords slightly larger and more widely spaced 
than the microradials. Umbilicus open, U-shaped, last whorl decoil- 
ing more rapidly, contained 3.48-5.46 times (mean 4.39) in the 
diameter, margins rounded. Whorls strongly and evenly rounded, 
aperture circular, not or only slightly flattened laterally above 
periphery, inclined about 20 from shell axis. 

Sinployea ellicensis (fig. 62a-c) differs from the 
Tongan S. vicaria (fig. 58a-c) in its smaller size, lower 
spire protrusion, larger radial ribs, and more rounded 
whorl contours. Sinployea ellicensis nukulaelaeana 
(fig. 62d-f) is much smaller and has more apical cords. 
Sinployea princei (fig. 65a-c) has more whorls, mono- 
chromatic coloration, and only traces of secondary spi- 
ral cording. 

Description. Shell smaller than average, with 4% normally 
coiled whorls. Apex and spire moderately and evenly elevated, body 
whorl descending much more rapidly, H/D ratio 0.556. Apical whorls 
1%, sculpture of 10 prominent, relatively widely spaced spiral cords. 
Postnuclear whorls with rounded, relatively crowded, protractively 
sinuated radial ribs, 119 on the body whorl, whose interstices are 2-3 
times their width. Microsculpture of fine radial riblets, 3-6 between 
each pair of major ribs, crossed by barely visible spiral riblets and 
rather crowded secondary spiral cords. Sutures deeply impressed, 
whorls strongly rounded above, very slightly flattened laterally 
above periphery. Umbilicus narrowly U-shaped, last whorl decoiling 
more rapidly, contained 3.89 times in the diameter. Color light yel- 
lowish horn with vague, irregular reddish flammulations, fainter on 
base of shell. Aperture circular, slightly flattened laterally above 
periphery, inclined about 10 from the shell axis. Height of holotype 
1.56 mm., diameter 2.81 mm. 

Holotype. Ellice Islands: Funafuti. Collected by 
Charles Hedley. AMS C.6124. 

Range. Funafuti, Ellice Islands. 

Paratypes. Ellice Islands (3 specimens, ex BPBM 
1667): Funafuti (102 specimens, AMS C.6124, Zurich, 
DMW MF-8810, FMNH 116981, FMNH 158451). 

Remarks. Adult specimens in the Mousson col- 
lection are distinctly higher and slightly larger (table 
XXIX) than those collected by Hedley (AMS C.6124). 
Probably this is another example of collection bias to- 
ward retention of larger individuals. Differences from 
Sinployea vicaria are covered in the diagnosis above. 
Although direct comparison of examples leaves no 
doubt that they are distinct, measurements do overlap 
extensively. Unlocalized specimens would be hard to 
identify without direct comparisons. Measurements of 
S. princei (Liardet) are almost identical (table XXX), 
but there is a distinct difference in coiling pattern. 

Sinployea ellicensis nukulaelaeana, new sub- 
species. Figure 62d-f. 

Diagnosis. Shell very small, diameter 1.99-2.20 mm. (mean 
2.08 mm.), with 3%-4 normally coiled whorls. Apex and spire 
slightly and evenly elevated, last whorl descending more rapidly, 
spire protrusion about Ve body whorl width, H/D ratio 0.496-0.561 
(mean 0.533). Apical sculpture of 11-13 (mean 11.67) prominent spi 




a 







FIG. 62. a-c, Sinployea ellicensis ellicensis, new species and subspecies. Funafuti, Ellice Islands. Holotype. AMS C6124; d-f, Sinployea 
ellicensis nukulaelaeana, new subspecies. Nukulaelae, Ellice Islands. Holotype. AMS C6123. Scale lines equal 1 mm. (SH). 



141 



142 



SOLEM: ENDODONTOID LAND SNAILS 



TABLE XXIX. - LOCAL VARIATION IN WESTERN POLYNESIAN SINPLOYEA. 





NUMBER OF 
SPECIMENS RIBS 


RIBS/MM. 


ellicensls 






ellicensis 






Zurich 


10 104.1*3.16 
(91-125) 


13.380.474 
(11.02-16.71) 


AMS C6124 


21 98.12.33 
(79-119) 


13.230.238 
(10.55-16.69) 


ellicensis 






nukulaelaeana 






AMS C6123 


6 84.5*4.85 
(72-102) 


13.02+0.734 
(11.08-15.20) 


Vaitupu 
BPBM 1667 


5 113.54.51 
(109-118) 


11.15*0.156 
(10.99-11.30) 


Zurich 


3 111.3*4.34 
(107-120) 


10.81*0.020 
(10.79-10.85) 


Eua 
BPBM 7983 


2 87.0*6.02 
(81-93) 


8.540.110 
(8.43-8.65) 


Zurich 


3 100.017.51 
(87-113) 


9.19*0.371 
(8.50-9.77) 


rotumana 






BMNH 77.11.8.57-63 


8 104.0*7.01 
(84-144) 


11.74+0.519 
(9.79-14.15) 



H/D RATIO 



1.42*0.023 2.50*0.028 0.570*0.0066 

(1.29-1.51) (2.40-2.65) (0.534-0.596) 



1.29*0.022 2.42+0.033 0.5330.0061 

(1.14-1.57) (2.22-2.83) (0.486-0.592) 



1.11+0.016 2.08+0.032 0. 53310-.0102 

(1.06-1.16) (1.99-2.20) (0.496-0.561) 



1.55*0.066 3.18+0.050 0.487+0.0152 

(1.42-1.76) (3.05-3.34) (0.454-0.525) 

1.58+0.029 3.30+0.121 0.479+0.0121 

(1.52-1.62) (3.18-3.54) (0.458-0.500) 



1.53+0.082 



3.24+0.181 



(1.45-1.61) 



(3.06-3.42) 



1.67+0.048 3.45+0.125 

(1.58-1.74) (3.26-3.68) 



1.560.063 



2.86+0.066 



(1.26-1.82) 



(2.62-3.26) 



0.472+0.0001 

(0.471-0.473) 

0.4830.0157 

(0.455-0.509) 



0.544+0.0138 

(0.481-0.607) 



WHORLS 
e. ellicensis 

Zurich 4 3/8-(4 1/4-4 1/2) 

AMS C6124 4 l/8+(4-4 1/2) 

e.nukulaelae. 

AMS C6123 3 7/8-(3 5/8-4) 



pseudovicarla 

Vaitupu 4 
BPBM 1667 



Eua 4 
BPBM 7983 



Zurich 4 



3/8(4 1/4-4 1/2) 
l/2+(4 1/2-4 5/8) 
3/8(4 1/4-4 1/2) 
l/2+(4 1/2-4 3/4) 



rotumana 

BMNH 4 l/4-(4-4 5/8) 



UMBILICUS D/U RATIO APICAL CORDS 



0.590.022 4.24+0.117 10.50.77 

(0.49-0.76) (3.48-4.86) (9-12) 

0.5510.016 4.43+0.081 9.42+0.21 

(0.43-0.72) (3.77-5.46) (8-11) 



0.48+0.013 4.31+0.086 11.7+0.34 

(0.44-0.53) (4.13-4.66) (11-13) 



0.95+0.044 3.35+0.197 11.7+0.33 

(0.79-1.05) (2.94-4.05) (11-12) 

1.12+0.050 2.93+0.034 11 

(1.05-1.22) (2.89-3.00) 

1.0510.066 3.08+0.020 

(0.99-1.12) (3.06-3.10) 

1.100.079 3.1710.109 9.510.50 
(0.99-1.25) (2.95-3.30) (9-10) 



0.65+0.025 4.3510.117 9.67+0.88 
(0.58-0.81) (3.93-4.86) (8-11) 



ral cords. Postnuclear sculpture of narrow, prominent, crowded, pro- 
tractively sinuated radial ribs, 75-102 (mean 84.5) on the body 
whorl, whose interstices are l'/2-3 times their width, often with 
periostracal extensions. Ribs/mm. 11.06-15.20 (mean 13.02). Micro- 
sculpture of fine radial riblets, 3-6 between each pair of major ribs, 
crossed by barely visible spiral riblets, with secondary spiral cording 
that is rather crowded and equal in spacing to microradials. Um- 
bilicus open, U-shaped, last whorl decoiling more rapidly, contained 
4.13-4.66 times (mean 4.31) in the diameter, margins rounded. 
Aperture and whorls circular, slightly compressed laterally above 
periphery. 



Sinployea ellicensis nukulaelaeana differs from the 
nominate race in possessing more apical spiral cords 
and in being much smaller. 

Description. Shell very small, with slightly less than 4 nor- 
mally coiled whorls. Apex and spire moderately and evenly elevated, 
body whorl descending rapidly, H/D ratio 0.547. Apical whorls 1%, 
sculpture of 12 prominent spiral cords. Postnuclear whorls with 
prominent, narrow, protractively sinuated, crowded, radial ribs, 72 
on the body whorl, whose interstices are l'/2-3 times their width. 
Microsculpture of fine radial riblets, 3-6 between each pair of major 



SYSTEMATIC REVIEW 



143 



ribs, barely visible spiral riblets, with relatively prominent and 
crowded secondary spiral cording. Sutures deep, whorls strongly 
rounded above, very slightly compressed laterally above periphery 
and on basal margin, with evenly rounded outer margin. Color light 
yellow-horn, with vague and irregular reddish flammulations that 
tend to coalesce on shell base. Umbilicus open, U-shaped, last whorl 
decoiling slightly more rapidly, contained 4.66 times in the diameter, 
margins rounded. Aperture circular, slightly flattened laterally 
above periphery and on basal margin, inclined about 15 from shell 
axis. Height of holotype 1.14 mm., diameter 2.07 mm. 

Holotype. Ellice Islands: Nukulaelae. AMS 
C.6123. 

Range. Nukulaelae, Ellice Islands. 

Paratypes. Nukulaelae (7 specimens, AMS 
C.6123, FMNH 158452). 

Remarks. Size differences between the races of 
Sinployea ellicensis (table XXIX) equal those found be- 
tween the Tongatapu and Eua Island S. vicaria col- 
lected in 1965 (table XXVIII). Subspecific recognition 
in this case is based on the difference in apical cord 
count and absence of major topographic differences be- 
tween Funafuti and Nukulaelae. There are no consis- 
tent sculptural or shape differences between the popu- 
lations. 



contained 2.76 times in the diameter. Aperture circular, slightly flat- 
tened above periphery, inclined about 25 from shell axis. Height of 
holotype 1.25 mm., diameter 2.73 mm. 

Holotype. Ellice Islands: Vaitupu. Collected by L. 
Isaacs in May, 1941. BPBM 189682. 

Range. Vaitupu, Ellice Islands. 

Paratypes. Ellice Islands (5 specimens, BPBM 
1667): Vaitupu (8 specimens, BPBM 106246, BPBM 
189682, Zurich, FMNH 158450). 

Material. "Eua" (error) (7 specimens, BPBM 
7983 ex Mus. Godeffroy, FMNH 116980, Zurich ex 
Mus. Godeffroy). 

Remarks. Although the holotype is subadult, it 
comes from the only set whose locality data can be 
accepted without question. The specimens from the 
Museum Godeffroy labeled "Eua" are essentially iden- 
tical to the Vaitupu shells (table XXIX) and probably 
were accidentally mislabeled. Nothing at all similar 
has been recorded from Eua subsequently. 

In view of the confusion between this species and 
vicaria in the older literature, I have chosen to use the 
name pseudovicaria. 



Sinployea pseudovicaria, new species. Figure 61a-c. 

Patula vicaria var. major Schmeltz, 1869, Cat. Mus. Godeffroy, 4, 
p. 72 Vaitupu, Ellice Islands (nude name); Schmeltz, 1874, 
Cat. Mus. Godeffroy, 5, p. 93 (nude name). 

Patula modicella Mousson, 1873 (not Ferussac, 1840), J. de 
Conchyl., 21, pp. 104-105 Vaitupu, Ellice Islands. 

Diagnosis. Shell relatively large, diameter 3.03-3.52 mm. 
(mean 3.24 mm.), with 4'/4-4% normally coiled whorls. Apex and 
early spire flat, lower spire descending slightly, body whorl descend- 
ing moderately, spire protrusion about Vs body whorl width, H/D 
ratio 0.454-0.525 (mean 0.486). Apical sculpture of 10-13 (mean 
11.75) fine spiral cords. Postnuclear whorls with fine, prominent, 
protractively sinuated radial ribs, 82-127 (mean 108.1) on the body 
whorl, whose interstices are 2-4 times their width. Ribs/mm. 9.56- 
15.15 (mean 11.33). Microsculpture of prominent radial riblets, 3-6 
between each pair of major ribs, crossed by extremely fine and 
crowded spiral riblets. No secondary spiral cording present. Um- 
bilicus broadly open, regularly decoiling, cup-shaped, contained 
2.89-4.05 times (mean 3.17) in the diameter, margins rounded. 
Whorls with subcircular outline, very slightly flattened laterally 
above periphery, inclined about 25 from shell axis. 

The flat spire, widely open umbilicus, and larger 
size immediately separate Sinployea pseudovicaria 
from the other Ellice and Tongan species. Sinployea 
lamellicosta (fig. 40a-c) from Tahiti has an equally 
wide umbilicus and is nearly identical in size and pro- 
portions, but has comparatively few radial ribs, a de- 
pressed or flat spire, and fewer, more loosely coiled 
whorls. 

Description. Shell rather small, with 4V4 relatively loosely 
coiled whorls. Apex and early spire flat, last whorl barely descend- 
ing, H/D ratio 0.458. Apical whorls 1%, sculpture of 13 moderately 
prominent spiral cords. Postnuclear whorls with prominent, 
rounded, protractively sinuated radial ribs, 82 on the body whorl, 
whose interstices are 2-4 times their width. Microsculpture of fine, 
crowded radial riblets with much finer and more crowded spiral rib- 
lets. Sutures deeply impressed, whorls slightly shouldered above, 
body whorl slightly flattened laterally above periphery. Color par- 
tially leached from shell with dark, reddish brown flammulations 
remaining. Umbilicus broadly open, U-shaped, regularly decoiling, 



Sinployea kusaieana, new species. Figures 63a-c, 
67h. 

Diagnosis. Shell smaller than average, diameter 2.52-2.75 
mm. (mean 2.60 mm.) with 3%-4'/8 normally coiled whorls. Apex and 
spire almost flat, slightly and evenly to moderately elevated, last 
whorl descending somewhat to much more rapidly, spire protrusion 
about V? body whorl width, H/D ratio 0.462-0.542 (mean 0.506). Api- 
cal sculpture of 8-10 (mean 9.20) fine, rather inconspicuous spiral 
cords, whose interstices are 3-4 times their width. Postnuclear 
sculpture of prominent, strongly protractively sinuated, rather 
crowded, V-shaped radial ribs, 84-103 (mean 91.1) on the body 
whorl, whose interstices are 3-5 times their width. Microsculpture of 
very fine radial riblets, 5-10 between each pair of major ribs, crossed 
by slightly finer and more crowded spiral riblets, with secondary 
spiral cording slightly more prominent than microradials, whose in- 
terstices are about 3 times their width. Umbilicus broadly open, V- 
shaped, last whorl usually decoiling more rapidly, contained 3.21- 
3.67 times (mean 3.51) in the diameter, margins rounded. Suture 
deep, whorls strongly rounded above, flattened laterally above 
periphery and weakly flattened in columellar margin. Aperture sub- 
circular, flattened laterally above periphery, inclined about 20 from 
shell axis. Color reddish yellow-horn, without darker markings. 

Sinployea kusaieana (fig. 63a-c) differs from the 
other barrier-free Caroline Island species by its posses- 
sion of an epiphallus, vergic papilla, and Sinployea- 
type stimulatory pad in the penis, plus having the kid- 
ney arms equal in length. Trukcharopa trukana (fig. 
63d- f) is dark reddish brown in color and has an aver- 
age of 150 major ribs on the body whorl but is virtually 
identical in many measurements (pp. 209-210). Rus- 
satus nigrescens (fig. 89d-f) is much more loosely coiled 
and very much larger in size with fewer whorls and a 
very narrow umbilicus. Kubaryiellus kubaryi (fig. 
89a-c) has a depressed apex and is much larger. The 
most similar Polynesian species areS. navutuensis (fig. 
68d-f) from the Lau Archipelago, which has 127-147 
ribs on the body whorl and averages one-quarter whorl 
less, and S. rotumana (fig. 61d-f) and S. vicaria (fig. 







a-f 




Fig. 63. a-c, Sinployea kusaieana, new species. Station 44, Lele Islet, Kusaie, Caroline Islands. Holotype. BPBM 155874; d-f, Trukcharopa 
trukana, new species. Station A5, Mt. Urupot, Tol Islet, Truk, Caroline Islands. Holotype. BPBM 189318. Scale line equals 1 mm. (a-c, e, SH; d, f, 

MM). 



144 



SYSTEMATIC REVIEW 



145 



58a-c), which have higher spires and narrower um- 
bilici. 

Description. Shell smaller than average, with 4 moderately 
tightly coiled whorls. Apex flat, spire and body whorl descending 
slightly, H/D ratio 0.481. Embryonic whorls l'/2, sculpture of 9 nar- 
row, relatively low spiral ribs. Remaining whorls with sculpture of 
crowded, rounded, protractively sinuated radial ribs, 84 on the body 
whorl, whose interstices are 2-3 times their width. Microsculpture of 
fine radial riblets, 5-10 between each pair of major ribs, crossed by 
finer, slightly more crowded spiral ribs, with more prominent, 
rounded, more widely spaced spiral cords. Sutures deeply impressed, 
whorls sharply rounded above, somewhat flattened laterally with 
slightly flattened columellar margin. Umbilicus broadly V-shaped, 
regularly decoiling, contained 3.44 times in the diameter. Color light 
reddish yellow-horn without traces of flammulations. Aperture sub- 
circular, strongly flattened laterally above periphery with slightly 
flattened columellar margin, inclined about 15 from the shell axis. 
Height of holotype 1.25 mm., diameter 2.60 mm. 

Holotype. Caroline Islands: Kusaie, Station 44, 
Lele Islet, under stones and logs at 3 ft. elevation. Col- 
lected by S. Ito and Yoshio Kondo on January 21, 1936. 
BPBM 155874. 

Range. Kusaie, Caroline Islands. 

Paratypes. Kusaie: Lele Islet (Station 44) under 
stones and logs at 3 ft. elevation (59 specimens, BPBM 
155874-7); Mot District (Station 66) 300 yd. west of 
Kefwas, 10 yd. inland at 4-5 ft. elevation (1 specimen, 
BPBM 156179). 

Remarks. Both of the known localities on Kusaie 
are lowland elevations under disturbed conditions. 
Quite possibly Sinployea kusaieana will be found to 
occur on other of the Eastern Caroline Islands. 

Description of soft parts. Foot and tail equal to shell diameter, 
tapering posteriorly. Sole and pedal grooves typical. Gonopore in 
normal position. 

Body color light yellow-white, without darker markings. 

Pallial region extending % whorl apically, without unusual fea- 
tures. Lung roof clear, no granulations. Kidney typically bilobed, 
maximum length 1.0-1.3 mm., lobes equal in length. Ureter typical, 
narrow strip of lung roof visible between arms. 

Ovotestis, hermaphroditic duct, albumen gland, talon, prostate 
(fig. 67h, DG), and uterus typical, no unusual features. 

Vas deferens (VD) not expanded in any area, entering epiphallus 
through usual pore. Epiphallus (E) about half as long as penis, slen- 
der, only weakly expanded. Penial retractor (PR) short, arising from 
diaphragm, inserting on head of penis lateral to epiphallic junction. 
Penis (P) about 1 mm. long, quite slender, upper half weakly ex- 
panded, internally with typical vergic papilla and small stimulatory 
pad. Atrium (Y) long and slender. 

Free oviduct (UV) typical. Spermatheca (S) with basal portion of 
shaft less expanded than usual. Vagina (V) tapering to atrium. 

(Based on BPBM 155874, 4 whole and several partial speci- 
mens.) 

Sinployea sp. 

Material. Marianas: Saipan, north edge of Gara- 
pan at base of a century plant (2 specimens, BPBM 
190043). 

Remarks. The two dead, somewhat worn shells 
collected by Harold W. Harry in October 1944 are not 
referable to any described species. They are most simi- 
lar to Sinployea atiensis (Pease) from the Cook Islands 
and S. ellicensis from Funafuti, but are smaller and 



with fewer whorls than the former and higher and with 
fewer ribs than the latter. The coiling pattern is looser 
than in either species. 

Because they were collected under an introduced 
plant in an urban area, the possibility exists that they 
were introduced on plants from Polynesia. Without 
additional information, I am not willing to accept this 
as a valid range extension and include the specimens 
here only to call attention to the need for making more 
collections in the Northern Marianas. Measurements 
of the shells have been included in Table XXVII for 
convenient reference. 

FIJIAN Sinployea 

There is an extraordinary diversity of Sinployea 
and derivative genera in Fiji and the Lau Archipelago 
(table XXX). Two species, S. irregularis (Garrett) and 
S. godeffroyana, are known from Viti Levu; the former 
also lives on Ovalau; and one species, S. princei (Liar- 
det), is known from Taveuni and possibly Kandavu. 
The absence of any records from relatively large is- 
lands such as Vanua Levu, Koro, Ngau, Moala, and 
Thithia probably are artifacts of collecting activity. I 
know of no significant collections of land mollusks from 
any of the above islands. Undoubtedly, most of them 
will be found to have Sinployea present. Six species of 
Sinployea are known from the Lau Group, one of them, 
S. inermis (Mousson), with three geographic races. 
Genera that are probably derived directly from 
Sinployea include Maafu thaumasius, which is re- 
stricted to Nayau in the Lau Group, and Ba humbugi 
on Viti Levu. The former agrees in shell micro- 
sculpture with Sinployea inermis, although radically 
altered in shell form and size (fig. 76a-c), whereas the 
latter (fig. 74a-c) can be derived without difficulty 
from S. irregularis. I am less certain of the probable 
derivation for the Vanua Mbalavu Lauopa 
mbalauuana (fig. 77a-c). 

In Polynesia it is very unusual to have any sym- 
patric occurrences of Sinployea or even for two species 
to live in the same valley, but there is extensive sym- 
patry in the southern Lau Archipelago. The northern 
Lau islands of Kimbombo, Vanua Mbalavu, Munia, 
and Mango have only single species records, either S. 
inermis or S. adposita. On Nayau, S. adposita and S. 
lauensis were taken at the same station, whereas on 
Aiwa, Tuvutha, and Lakemba both S. adposita and S. 
inermis were found together. On Oneata, both S. 
navutuensis and S. adposita were collected, but not at 
the same stations. Wangava had large numbers of S. 
recursa and a few S. lauensis taken together, whereas 
on Navutu-i-Loma H. S. Ladd collected S. inermis, S. 
lauensis, and S. navutuensis together at two different 
stations. Finally, on Yangasa Levu large numbers of S. 
inermis, S. adposita, and S. lauensis were sympatric. 
Unfortunately most of these specimens were dead indi- 
viduals collected during the July to August dry season. 
Soft parts were available only for S. inermis and S. 
adposita. Dissection of specimens from Station 97, 



146 



SOLEM: ENDODONTOID LAND SNAILS 



TABLE XXX. - RANGE OF VARIATION IN FIJIAN SINPLOYEA. 





NUMBER OF 


NAME 


SPECIMENS 


angularis 


188 


re curs a 


108 


prince! 




(Liardet) 




Taveuni 


9 


Kandavu 


U 


inermis inermis 


197 


(Mousson) 




inermis 




meridionalis 


169 


inermis 




lakemhana 


14 


lauensis 


82 


navutuensis 


29 


jtdjjp ^s i t a 


176 


irregularis 


78 


(Garrett) 



godef f royana 



RIBS RIBS/MM. HEIGHT DIAMETER H/D RATIO 

38.0(36-42) 5.67(5.28-6.16) 1.21(1.08-1.29) 2.11(2.02-2.27) 0.574(0.512-0.615) 

REDUCED 1.66(1.51-1.84) 2.83(2.60-3.09) 0.587(0.562-0.634) 

91.1(80-102) 12.06(11.84-13.80) 1.34(1.26-1.57) 2.43(2.35-2.68) 0.551(0.524-0.586) 

96.3(84-119) 13.10(11.4-14.67) 1.27(1.17-1.45) 2.33(2.02-2.58) 0.545(0.521-0.577) 

80.9(65-104) 9.82(8.39-11.75) 1.46(1.18-1.84) 2.60(2.24-3.06) 0.562(0.500-0.629) 

72.2(53-104) 8.37(6.75-11.70) 1.64(1.32-1.94) 2.88(2.50-3.22) 0.566(0.505-0.655) 

51.3(38-62) 7.01(4.67-8.20) 1.37(1.10-1.64) 2.36(2.14-2.86) 0.583(0.508-0.634) 

103.9(95-119) 11.53(10-93-14.46) 1.47(1.18-1.78) 2.76(2.40-3.13) 0.530(0.493-0.581) 

136.7(121-147) 17.36(16.94-17.62) 1.32(1.16-1.49) 2.66(2.58-2.78) 0.498(0.499-0.566) 

53.3(36-74) 5.28(3.35-7.23) 1.81(1.58-2.20) 3.29(2.70-3.62) 0.603(0.538-0.687) 

IRREGULAR 1.77(1.51-2.30) 2.87(2.50-3.52) 0.614(0.528-0.764) 

117.7(85-152) 28.36(20.99-38.03) 1.26(1.19-1.29) 2.12(1.95-2.32) 0.595(0.577-0.619) 





WHORLS 


UMBILICUS 


D/U 


RATIO 


APICAL 
CORDS 


SPIRE 
ELEVATION 


BODY WHORL 
WIDTH 


SP/BWW 


angul. 


3 7/8+O 3/4-4) 


0.35(0.23-0 


,40) 6.14(5 


.21-8.71) 


14.6(13-16) 


0.19(0.16-0.23) 


0.62(0. 


59-0.66) 


0.316(0. 


250-0.389) 


recur. 


3 7/8-O 1/2-4 1/8) 


0.43(0.33-0 


,53) 6.63(5 


.75-7.90) 


15.9(14-18) 


0.23(0. 


20-0.30) 


0.86(0. 


81-0.91) 


0.264(0. 


222-0.346) 


princ. 


3 3/4+(3 1/2-4 1/8) 


0.57(0.49-0. 


63) 4.28(3 


.74-4.83) 


9(8-10) 


0.12(0. 


08-0.23) 


0.81(0. 


70-0.89) 


0.151(0. 


095-0.259) 


Tav. 


Kan. 


3 5/8+(3 1/2-4) 


0.46(0.39-0, 


53) 4.76(4 


.14-5.19) 


NOT 
AVAILABLE 


0.13(0. 


10-0.18) 


0.75(0. 


66-0.86) 


0.179(0. 


130-0.250) 


Line. 


4-(3 1/2-4 5/8) 


0.62(0.49-0. 


82) 4.17(3 


.47-4.69) 


8.96(6-11) 


0.16(0. 


10-0.25) 


0.81(0. 


72-0.92) 


9.192(0. 


115-0.280) 


i.mer. 


4 l/4+(3 7/8-4 5/8) 


0.82(0.69-0, 


99) 3.51(3 


.07-3.81) 


11.6(9-14) 


0.20(0. 


13-0.26) 


0.96(0. 


82-1.05) 


0.207(0. 


140-0.285) 


l.lak. 


4-t-O 3/4-4 1/2) 


0.45(0.33-0.76) 5.54(3 


.78-7.10) 


13.6(11-16) 


0.19(0. 


13-0.23) 


0.82(0. 


72-0.89) 


0.226(0. 


153-0.259) 


lauen. 


3 7/8(3 1/2-4 1/4) 


0.65(0.53-0. 


79) 4.24(3 


.88-4.65) 


9.6(9-11) 


0.10(0. 


03-0.16) 


0.91(0. 


82-1.09) 


0.113(0. 


038-0.161) 


navut. 


4 1/8(4-4 3/8) 
4 3/8+(4-4 7/8) 


0.78(0.76-0, 
0.28(0.17-0, 


82) 3.38(3 
46)11.9(7. 


.25-3.52) 
85-17.0) 


10.3(9-11) 
13.0(12-14) 


0.09(0. 
0.24(0. 


07-0.13) 
13-0.33) 


0.74(0. 
1.04(0. 


66-0.79) 
86-1.18) 


0.125(0. 
0.227(0. 


100-0.167) 
133-0.275) 


adpos. 


irreg. 


3 3/4+(3 1/4-4 3/8) 


0.64(0.39-0, 


92) 4.54(3 


.79-6.83) 


13.5(16-20) 


0.20(0. 


10-0.46) 


0.96(0. 


86-1.09) 


0.201(0. 


100-0.424) 


godef. 


3 3/4+(3 3/4-4) 


0.42(0.33-0, 


56) 5.26(4 


.11-6.30) 


9.7(8-11) 


0.12 




0.81 




0.143 





Toumba, Lakemba, Lau, showed that S. adposita had a 
longer penis with very small stimulatory pad and nor- 
mal vergic papilla, whereas S. inermis had a shorter 
penis with very large stimulatory pad and small vergic 
papilla. On Viti Levu, S. godeffroyana and S. ir- 
regularis have been collected at the same station, and I 
have found S. irregularis and Ba humbugi under the 
same piece of bark. 

Evidence from the white body color suggests that 
S. irregularis, S. inermis, and S. adposita are strictly 
terrestrial. Generally, semiarboreal species have gray 
color on the ommatophores and neck, sometimes also 
on the mantle collar edge. Two species, S. angularis 
and S. recursa, have greatly reduced sculpture, and I 
suspect that they may be semiarboreal or arboreal 
species. Unfortunately, they are known only from dead 
examples. 



Patterns of distribution are relatively simple. 
Sinployea irregularis and S. godeffroyana are widely 
distributed on forested areas of Viti Levu. Sinployea 
princei was described from Taveuni, and essentially 
identical specimens coming from the Rolle collection 
are labeled Kandavu. Sinployea inermis and S. ad- 
posita are widely distributed in the Lau Archipelago, 
with the former having a slightly greater range. 
Sinployea recursa is known only from Wangava, and S. 
angularis, only from Namuka. Sinployea navutuensis 
is recorded from Navutu-i-Loma, and a single shell 
from Oneata has been referred here with some hesita- 
tion. Sinployea lauensis has a fairly wide range in 
southern Lau, having been found on Nayau, Wangava, 
Namuka, Navutu-i-Loma, and Yangasa Levu. 

Many of the Lau Group have never been collected 
for land mollusks. Generally, only a single or at most 



SYSTEMATIC REVIEW 



147 



two stations made in a hurried visit of one to two days 
account for the collecting done to date on most islands. 
Obviously there are splendid opportunities for further 
field work. Particularly important will be efforts to 
work out patterns of species isolation between sympat- 
ric taxa. 

Sinployea inermis (Mousson) from the Lau Ar- 
chipelago, S. princei (Liardet) from Taveuni and Kan- 
davu, and S. irregularis (Garrett) from Viti Levu are 
the generalized Fijian species. Sinployea inermis is one 
of the most "average" Sinployea, with the monochrome 
coloration, details of sculpture, and proportions (fig. 
66a-c) differentiating it from the Tongan S. vicaria 
(Mousson) (fig. 58a-c) and the Ellice Island S. ellicen- 
sis (fig. 62a-c). Subspecies of S. inermis depart further 
from the typical pattern, but are still generalized in 
shape and form. Sinployea princei (fig. 65a-c) has a 
weakly flammulated color pattern, much more crowded 
radial ribbing, and reduced secondary spiral cording, 
but is very similar to S. inermis. Greater differences 
are shown by S. irregularis (fig. 65d-f). The spire is 
flat, sutures deep, apical sculpture much finer and 
more numerous, ribbing very irregular, body whorl 
often abruptly descending, and the size is slightly 
larger. 

Several species show minor character changes. 
Sinployea adposita (Mousson) (fig. 69d-f) can be de- 
rived from S. inermis by increase in size, narrowing of 
the umbilicus, and increase in rib spacing. The two 
species are sympatric and show character displacement 
in penial structure, with S. inermis (fig. 67e-f) having 
the stimulatory pad and muscular collar proportion- 
ately larger than usual and the vergic papilla shorter, 
whereas in S. adposita (fig. 67g) the vergic papilla is 
quite large. Sinployea lauensis (fig. 68a-c) has more 
crowded ribbing, a less protruded spire, fatter body 
whorl, and more rapidly decoiling umbilicus. It has not 
been dissected. Sinployea navutuensis (fig. 68d-f) has 
much more crowded radial ribbing, a wider umbilicus, 
nearly flat apex, and lower H/D ratio. Two other Lau 
Archipelago species are more strongly differentiated. 
The slightly larger S. recursa (fig. 64a-c) has a 
strongly flammulated color pattern and the sculpture 
greatly reduced. No live material is known, but I sus- 
pect this may be a more arboreal species than most 
Sinployea. Sinployea angularis (fig. 64d-f) is reduced 
in size, has widely spaced radial ribbing, a narrow um- 
bilicus, angulated periphery, and pale monochrome 
coloration. It has perhaps the most modified shell of 
any Sinployea. Possibly generic separation is war- 
ranted, but without study of the soft parts, I prefer a 
conservative treatment. 

On Viti Levu, the monotypic genus Ba (p. 172) can 
be derived from Sinployea irregularis by a drastic 
change in coiling pattern, reduction in whorl count, 
closure of the umbilicus, and decrease in number of 
spiral apical cords. Anatomical changes are covered 
under the description of that species. The other Viti 
Levu species, Sinployea godeffroyana (fig. 69a-c), is 



characterized by its small size, proportionately thick 
body whorl, and extremely crowded radial ribbing 
(mean ribs/mm. 28.4). It has not been dissected. 

Undoubtedly there are additional species of 
Sinployea in both Fiji and the Lau Archipelago. Use of 
the following key can serve only as a general guide to 
affinities when more collections become available. 

KEY TO THE FIJI AND LAU Sinployea 

1. Main islands of Fiji 2 

Lau Archipelago 4 

2. Ribbing irregular; apical cords more than 15; mean diameter 

more than 2.7 mm Sinployea irregularis (Garrett, 1887) 

Ribbing regular; apical cord average 10; mean diameter less 
than 2.5 mm 3 

3. Viti Levu; mean diameter about 2.1 mm.; ribs/mm, more than 

20 Sinployea godeffroyana, new species 

Taveuni and Kandavu; mean diameter about 2.3-2.4 mm.; 
ribs/mm, less than 15 Sinployea princei (Liardet, 1876) 

4. Mean diameter less than 3.0 mm.; mean D/U ratio less than 

7 5 

Mean diameter more than 3.1 mm.; mean D/U ratio more than 
10 Sinployea adposita (Mousson, 1870) 

5. Body whorl not angulated; mean diameter more than 2.3 

mm 6 

Body whorl angulated; mean diameter about 2.1 mm.; 
Namuka Sinployea angularis, new species 

6. Prominent major ribbing continuing to end of body whorl 7 

Major radial ribbing widely spaced on spire, becoming reduced, 

then absent on body whorl Sinployea recursa, new species 

7. Mean rib count 100-150 8 

Mean rib count 50-95 9 

8. Mean D/U ratio less than 3.5; ribs/mm, more than 16; mean 

body whorl width about 0.75 mm 

Sinployea navutuensis, new species 
Mean D/U ratio more than 4.0; ribs/mm, less than 15; mean 

body whorl width about 0.90 mm 

Sinployea lauensis, new species 

9. Mean D/U ratio usually much more than 3.8; Central and 

Northern Lau 10 

Mean D/U ratio less than 3.6; Southern Lau 

Sinployea inermis meridionalis, new subspecies 
10. Mean rib count less than 60; mean D/U ratio about 5.50; mean 

apical cords about 13; Lakemba 

Sinployea inermis lakembana, new subspecies 

Mean rib count more than 70; mean D/U ratio less than 4.50; 

mean apical cords about 9; Mango and Vanua 

Mbalavu Sinployea inermis inermis (Mousson, 1870) 



Sinployea angularis, new species. Figure 64d-f. 

Diagnosis. Shell very small, diameter 2.02-2.27 mm. (mean 
2.11 mm.), with 3%-4 normally coiled whorls. Apex and spire mod- 
erately and usually evenly elevated, sometimes rounded above, body 
whorl descending a little more rapidly, spire protrusion almost Va 
body whorl width, H/D ratio 0.512-0.615 (mean 0.574). Apical 
sculpture of 13-16 (mean 14.6) narrow and rather inconspicuous 
spiral cords. Postnuclear sculpture of low, rounded, widely spaced, 
strongly protractively sinuated radial ribs, 36-42 (mean 38.0) on the 
body whorl, whose interstices are 4-7 times their width. Ribs/mm. 
5.28-6.16 (mean 5.67). Microsculpture of extremely fine radial rib- 
lets, more than 10 between each pair of major ribs, crossed by equally 
fine spiral riblets, with narrow, rather crowded secondary spiral 
cording, becoming broader and less distinct on body whorl. All micro- 
sculpture barely visible at 96 x magnification. Umbilicus narrow, 
U-shaped, last whorl decoiling more rapidly, contained 5.21-8.71 
times (mean 6.14) in the diameter, margins rounded. Sutures im- 
pressed, whorls strongly rounded above, becoming strongly flattened 
laterally above obtusely angulated periphery, evenly rounded on 







FIG. 64. a-c, Sinployea recursa, new species. Station 27, Wangava, Lau, Fiji. Holotype. BPBM 166967; d-f, Sinployea angularis, new 
species. Station 3, Namuka, Lau, Fiji. Holotype. BPBM 166616. Scale lines equal 1 mm. (MM). 



148 



SYSTEMATIC REVIEW 



149 



outer and lower margins. Aperture inclined about 25 from shell 
axis. 

Although both Sinployea irregularis from Viti 
Levu (fig. 65e) and S. rudis from Rarotonga (fig. 47e) 
give the appearance of having an angulated periphery 
because of supraperipheral flattening, only S. an- 
gularis (fig. 64e) of Polynesian-Fijian Charopidae has 
the periphery itself actually angulated. Sinployea ad- 
posita (fig. 69d-f) agrees in rib spacing, but is much, 
much larger and has a very narrow umbilicus. Sin- 
ployea inermis (fig. 66a-e) is much larger, with a less 
protruded spire, rounded periphery, and usually much 
more numerous and crowded ribbing. 

Description. Shell very small, with 4 moderately tightly coiled 
whorls. Apex and spire distinctly and evenly elevated, last part of 
body whorl slightly deflected below periphery of penultimate, H/D 
ratio 0.591. Embryonic whorls 1%, sculpture mainly eroded, with 
traces of fine spiral cords remaining. Postnuclear sculpture of low, 
rounded, protractively sinuated radial ribs, 36 on the body whorl, 
whose interstices are 4-7 times their width. Microsculpture of ex- 
tremely fine radial riblets, crossed by much finer, more crowded spi- 
ral riblets and narrow, rather vague secondary spiral cords. Sutures 
relatively shallow on body whorl, slightly indented. Whorls with 
angulated periphery, somewhat flattened above and slightly flat- 
tened below with gently rounded basal margin. Umbilicus narrow, 
U-shaped, last whorl decoiling more rapidly, contained 5.50 times in 
the diameter. Color leached from shell. Aperture ovate with angu- 
lated periphery, inclined about 25 from the shell axis. Height of 
holotype 1.28 mm., diameter 2.17 mm. 

Holotype. Fiji: Lau Group, Namuka, Station 3, 
500 ft. inland from Matandolo on north central part of 
island at 65-75 ft. elevation. Collected dead between 
rocks by H. S. Ladd on July 7, 1934. BPBM 166616. 

Range. Namuka, Lau Group, Fiji. 

Paratypes. Namuka: 500 ft. inland from Matan- 
dolo (Station 3) on north central part of island at 65-75 
ft. elevation (187 specimens, BPBM 166616, BPBM 
166642, BPBM 166669). 

Remarks. Only 14 (7.4%) of 188 Sinployea an- 
gularis were whole adults. The remainder were 
juveniles or badly broken specimens. Although an an- 
gulated periphery is extremely common in the En- 
dodontidae and many of the New Caledonian and 
Austro-Zelandic endodontoid taxa, it is found other- 
wise among Pacific Island Charopidae only in the 
Marianas Himeroconcha quadrasi and H. fusca (fig. 
105b, e). Hence the choice of angularis as specific name 
for this species. 

Its elevated spire and angulated periphery sepa- 
rate it from any other Fijian species in appearance, but 
the pattern of sculpture is a modification of the same 
type seen in S. inermis. Sinployea angularis is much 
smaller than examples of S. inermis, coming closest in 
size to the Viti Levu S. godeffroyana. The latter has 
very numerous and crowded radial ribbing, plus the 
spire is much less protruded. 

Possibly this is an arboreal species and should be 
generically separated from Sinployea. Without dissec- 
tion and ecological data, I prefer to retain it in 
Sinployea. In size and shape it is the most drastically 
altered species. 



Sinployea recursa, new species. Figure 64a-c. 

Diagnosis. Shell of average size, diameter 2.60-3.09 mm. 
(mean 2.83 mm.), with 3 l /2-4'/8 normally coiled whorls. Apex and 
spire moderately to strongly elevated, body whorl descending more 
rapidly, spire protrusion more than Vi body whorl width, H/D ratio 
0.562-0.634 (mean 0.587). Apical sculpture of 14-18 (mean 15. 9) fine 
spiral cords, often partly to completely eroded. Postnuclear sculpture 
of indistinct, low, broadly rounded, widely spaced radial ribs on the 
upper spire that become greatly reduced to absent by early part of 
body whorl. Vague traces of microreticulations on the upper spire 
become replaced by fine and crowded secondary spiral cords on body 
whorl. All microsculpture only visible under high magnification with 
oblique lighting. Umbilicus narrow, V-shaped, slightly and regularly 
decoiling, contained 5.75-7.90 times (mean 6.63) in the diameter, 
margins rounded. Sutures impressed, whorls strongly rounded 
above, compressed laterally above periphery, with slightly com- 
pressed basal margin. Color light yellow-horn, with irregular red- 
dish flammulations that become narrower on shell base. 

In spire elevation and general appearance, Sin- 
ployea recursa shows many similarities to S. angularis. 
The latter (fig. 64d-f) is much smaller, has an angu- 
lated periphery, and much more prominent radial rib- 
bing. No other Polynesian or Micronesian Charopidae 
has the ribbing so greatly reduced, although this is 
frequent in New Caledonian, Australian, and New 
Zealand taxa and occurs sporadically in the Polynesian 
Endodontidae. 

Description. Shell of average size, with slightly less than 3% 
relatively loosely coiled whorls. Apex and spire evenly elevated, last 
whorl descending a little more rapidly, H/D ratio 0.582. Embryonic 
whorls 1!4, sculpture of about 15 fine spiral ribs, partially eroded. 
Postnuclear whorls with a few widely spaced, low, inconspicuous 
radial ribs on the upper spire, becoming reduced over most of the 
body whorl but reappearing on last quarter. Microsculpture of fine 
radial growth striae and very low, inconspicuous secondary spiral 
cording. Sutures relatively shallow, whorls gently rounded above, 
sloping flatly down to more strongly rounded periphery and slightly 
flattened basal lip, contained 7.82 times in the diameter. Color light 
yellowish white with vague irregular yellow-red or reddish brown 
flammulations. Aperture ovate, slightly flattened basally, inclined 
about 20 from the shell axis. Height of holotype 1.64 mm., diameter 
2.83 mm. 

Holotype. Fiji: Lau Group, Wangava, Station 27, 
northeast end of island on an inland ridge, % mile from 
shore, at 75 ft. elevation. Collected in soil under leaves 
and between rocks by H. S. Ladd on July 22, 1934. 
BPBM 166967. 

Range. Wangava, Lau Group, Fiji. 

Paratypes. Wangava: northeast end of island 
(Station 27), % mile from shore on ridge at 75 ft. eleva- 
tion (107 specimens, BPBM 166967, BPBM 166969, 
BPBM 166996). 

Remarks. The similarity of Sinployea recursa to 
the Austro-Zelandic flammulinid taxa with reduced 
ribbing is striking, but the probabilities are that this is 
convergent. Sinployea angularis is intermediate in 
sculpture between S. recursa and the Fijian charopids 
with typical sculpture, so that I have no hesitation in 
considering it to be a local development. No soft parts 
were available for dissection. 

No other Charopidae covered in this monograph 
has such sculptural reduction. Although many species 
have the ribbing extremely irregular, such Fijian 



150 



SOLEM: ENDODONTOID LAND SNAILS 



shells as S. irregularis (fig. 65d-f) and Ba humbugi 
(fig. 74a-c) being good examples, the almost complete 
loss of ribbing does not occur. This condition is more 
common in the Endodontidae, where a number of En- 
dodonta, several Nesodiscus, one Nesophila, and one 
Libera have undergone such reduction. In all of these 
cases, the shells are quite large, and sculptural reduc- 
tion is associated with size increase. The name recursa 
refers to the reduced shell sculpture on the lower 
whorls in particular, indicating a return to simplified 
shell structure. 

I suspect it may be an arboreal or semiarboreal 
species. 

Sinployea princei (Liardet, 1876). Figure 65a-c. 

Helix princei Liardet, 1876, Proc. Zool. Soc. London, 1876, p. 100, 
pi. 5, fig. 4, a Taveuni Island, Fiji Islands. 

Patula princei (Liardet), Garrett, 1887, Proc. Zool. Soc. London, 
1887, p. 179. 

Helix (Patula) princei Liardet, Tryon, 1887, Man. Conchol., (2) 3, p. 
27, pi. 5, fig. 83a. 

Endodonta (Charopa) princei (Liardet), Pilsbry, 1893, Man. Con- 
chol., (2)9, p. 35. 

Champa princei (Liardet), Gude, 1913, Proc. Malacol. Soc. London, 
10, (5), p. 330; Germain, 1932, Ann. Inst. Ocean., 12 (2), p. 45. 

Diagnosis. Shell small, diameter 2.35-2.68 mm. (mean 2.43 
mm.), with 3Vz-4Vn rather tightly coiled whorls. Apex and spire mod- 
erately and evenly elevated, last whorl descending more rapidly, 
spire protrusion more than Ve body whorl width, H/D ratio 0.524- 
0.586 (mean 0.551). Apical sculpture of 8-10 (mean 9.0) fine and 
relatively low spiral cords. Postnuclear sculpture of high, prominent, 
thin, protractively sinuated radial ribs, 80-102 (mean 91.1) on the 
body whorl, whose interstices are l'/2-3 times their width. Ribs/mm. 
11.85-13.80 (mean 12.06). Microsculpture of fine radial riblets, 3-6 
between each pair of major ribs, crossed by extremely fine and 
crowded spiral riblets that are barely visible under 96x magnifica- 
tion. Secondary spiral cording occasionally visible near periphery 
and shell base. Umbilicus open, U-shaped, last whorl decoiling more 
rapidly, contained 3.74-4.83 times (mean 4.28) in the diameter, 
margins rounded. Whorls very strongly rounded, aperture circular, 
sometimes slightly flattened laterally above periphery, inclined 
about 20 from shell axis. Color reddish horn, no flammulations. 

Sinployea princei is very similar to S. inermis in- 
ermis (fig. 66a-c), differing primarily in its much more 
crowded radial ribbing, greatly reduced secondary spi- 
ral cording, and slightly smaller size. Sinployea lauen- 
sis (fig. 68a-c) is larger, flatter, and has a distinctly 
less protruding spire. Other Fijian species differ 
greatly in rib spacing and count. Sinployea ellicensis 
(fig. 62a-f) has a flammulated color pattern, much 
more prominent secondary spiral cording, and aver- 
ages three-eighths of a whorl less. 

Description. Shell small, with 3% moderately tightly coiled 
whorls. Apex and spire slightly and evenly elevated, body whorl 
descending more rapidly, H/D ratio 0.545. Apical whorls 1%, 
sculpture of 8 narrow, widely spaced spiral ribs with faint traces of 
weaker radial riblets on last eighth whorl. Postnuclear whorls with 
narrow, lamellar, somewhat crowded, slightly protractively sinuated 
radial ribs, 102 on the body whorl, whose interstices are 2-4 times 
their width. Microsculpture of weak radial and finer spiral riblets 
with a secondary sculpture of weak, widely spaced spiral cords visi- 
ble only on periphery and base of body whorl. Sutures deep, whorls 
strongly rounded above, evenly rounded on outer and basal margin. 
Color light reddish yellow-horn without darker markings. Umbilicus 
narrow, U-shaped, regularly decoiling, contained 4.08 times in the 



diameter. Aperture subcircular with evenly rounded outer margin, 
inclined about 15 from the shell axis. Height of lectotype 1.28 mm., 
diameter 2.36 mm. 

Lectotype. Fiji: Taveuni Island. Collected by 
Liardet. BMNH 76.9.9.2. 

Range. Taveuni and Kandavu, Fiji Islands. 

Paratypes.BMNH 76.9.9.2. 

Material. Fiji (2 specimens, Brussels): Taveuni (6 
specimens, Zurich); Kandavu (4 specimens, FMNH 
48997, SMF 165749). 

Remarks. Specimens from Kandavu, originating 
from the Rolle collection, are separable (table XXX) 
from the Taveuni specimens only by their narrower 
umbilicus. They are considered to be S. princei, but 
were not included in figuring mean size because I am 
uncertain that the locality is accurate. 

No material of S. princei has been collected in this 
century. 

Sinployea inermis (Mousson, 1870). 

Several populations from islands in the Lau Group 
are considered to be conspecific. They differ from the 
Taveuni and Kandavu S. princei (fig. 65a-c) in their 
higher spire protrusion and much less crowded radial 
ribbing (fig. 66a-e). Lau Archipelago species of similar 
size, S. lauensis (fig. 68a-c) and S. navutuensis (fig. 
68d-f), plus the Viti Levu S. godeffroyana (fig. 69a-c), 
all have much more numerous and crowded radial rib- 
bing plus moderately to greatly different proportions. 
Extralimital species such as the Tongan S. vicaria (fig. 
58a-c) and Ellice Island S. ellicensis (fig. 62a-c) differ 
in details of sculptural character and spacing, but ob- 
viously are closely related. 

Variation among populations on the same island 
and from different islands is relatively extensive (ta- 
bles XXXII- XXX VI) and somewhat difficult to inter- 
pret. Available material was collected in the 1850s, 
1860s, in 1934, 1938, 1973, and 1977. Since the Lau 
Group has unpredictable periods of drought, the possi- 
bility of ecophenotypic dwarfing must be considered in 
accessing the mean differences. Most samples fit rela- 
tively well into one of three geographic variation pat- 
terns that are considered to represent subspecies. 
Specimens from the northern group of Vanua Mbalavu, 
Mango, and Kimbombo Islands are nearest to S. prin- 
cei, having more ribs and intermediate umbilical width 
(fig. 66a-c). They average about nine apical cords. 
Samples from the large, centrally located island of 
Lakemba vary in umbilical width but average a nar- 
row umbilicus, 11.6 apical cords, and have very widely 
spaced and large radial ribs (fig. 66e). Those from the 
more southerly Lau Group, Aiwa, Yangasa Levu, and 
Navutu-i-Loma tend to have a slightly wider um- 
bilicus, average 13.6 apical cords, have a little less 
crowded ribbing than those from the northern area, 
and are easily separable from the Lakemba popula- 
tions (fig. 66d). Frequency distributions of the D/U 
ratio, ribs/mm., and ribs (tables XXXII- XXXIII) illus- 
trate the extent of overlap for any single character. Rib 







a 








FIG. 65. a-c, Sinployea princei (Liardet). Taveuni, Fiji. Probable paratype. Zoologisches Institut der Universitat Zurich; d-f, Sinployea 
irregularis (Garrett). Viti Levu, Fiji. Lectotype. BPBM 7982. Scale lines equal 1 mm. (MM). 



151 



TABLE XXXI. - LOCAL VARIATION IN FIJIAN, MELANESIAN AND MICRONESIAN SINPLOYEA. 



NUMBER OF 
SPECIMENS 



RIBS/MM. 



DIAMETER 



H/D RATIO 



Sta. 3 

BPBM 166642 



Sta. 2 

BPBM 166616 



1.230.025 2.09*0.024 0.58810.0121 

(1.10-1.28) (2.01-2.17) (0.540-0.615) 

1.1810.028 2.1010.031 0.56110.0090 

(1.07-1.28) (2.02-2.25) (0.512-0.591) 



Sta. 27 

BPBM 166967 

lauensis 



12 

4 
5 
3 



Sta. 30 

BPBM 167139 

Sta. 43 

BPBM 167234 

Sta. 28, 29 

BPBM 167020, -55 

navutuensis 

Sta. 29 4 

ex BPBM 167020, 
ex BPBM 167055 

kusaieana 

Sta. 44 11 

BPBM 155874-5, -7, -9 

descendens 

Karlei 24 

ZMB 

Matong 7 

ZMB 



100.412.23 



(95-105) 



119 



142 



11.5910.287 



(10.96-12.62) 



12.51 



11.64 



1.6610.027 2.8110.045 0.58910.0066 

(1.52-1.85) (2.62-3.11) (0.562-0.634) 



1.3810.065 2.6810.112 0.51510.0110 

(1.19-1.49) (2.42-2.95) (0.493-0.544) 

1.4810.076 2.7910.078 



0.53110.0143 



(1.39-1.79) 



(2.65-3.08) 



(0.494-0.581) 



1.6110.145 2.9610.141 0.54210.0242 

(1.32-1.79) (2.68-3.15) (0.494-0.568) 



1.3210.73 2.6610.048 0.49810.0220 

(1.16-1.49) (2.58-2.78) (0.449-0.556) 



90.411.97 11.1910.287 1.3110.027 2.5810.020 0.50610.0074 

(84-103) (9.96-13.17) (1.18-1.48) (2.50-2.73) (0.481-0.542) 

91.711.55 13.0710.232 1.1610.009 2.2510.013 0.51510.0041 

(77-103) (11.16-14.71) (1.05-1.25) (2.14-2.40) (0.470-0.548) 

91.912.62 12.6810.313 1.2210.017 2.3210.037 0.52810.0062 

(80-99) (11.33-13.73) (1.15-1.27) (2.15-2.43) (0.504-0.550) 



novopomme rana 
Karlei 
ZMB 



12 93.813.29 12.7110.499 1.2110.044 2.3310.059 0.52010.0089 

(72-l'04) (9.61-15.10) (1.05-1.61) (2.11-2.85) (0.467-0.567) 



WHORLS 



UMBILICUS 



D/U RATIO 



APICAL CORDS 



Sta. 2 



3 7/8+(3 3/4-4) 
3 7/8-O 3/4-4) 



0.3210.531 

(0.23-0.36) 



6.7110.415 15 
(5.81-8.71) 



0.3810.007 5.5710.099 

(0.35-0.39) (5.21-6.05) 



recursa 
Sta. 27 

lauensis 
Sta. 30 

Sta. 43 
Sta. 28 

navutuensis 
Sta. 29 

kusaieana 
Sta. 44 

descendens 
Karlei 

Ma t ong 



3 7/8-(3 1/2-4 1/8) 0.4210.015 6.7010.211 16.511.50 

(0.33-0.49) (5.78-7.90) (15-18) 



3 5/8+(3 1/2-3 7/8) 0.6110.039 4.4010.120 

(0.53-0.70) (4.15-4.65) 



3 7/8(3 3/4-4 1/8) 0.6810.030 4.1010.085 

(0.63-0.79) (3.88-4.32) 

4 1/8(3 7/8-4 1/4) 0.6910.038 4.2610.150 

(0.63-0.76) (4.00-4.52) 



4 1/8(4-4 3/8) 0.7810.016 3.3810.055 10. Oil. 00 
(0.76-0.82) (3.25-3.52) (9-11) 



4-(3 3/4-4 1/8) 0.7410.009 3.5110.038 9.210.20 

(0.69-0.79) (3.21-3.67) (8-10) 



4 l/8+(4-4 1/4) 0.8310.011 2.7310.029 10.710.30 

(0.76-0.95) (2.43-2.94) (9-14) 

4 3/8-(4 1/8-4 1/2) 0.8510.028 2.7310.046 10.510.29 

(0.74-0.95) (2.55-2.91) (10-11) 



3 3/4(3 1/2-4 1/8) 0.6710.028 3.4910.076 10.710.36 

(0.54-0.92) (3.09-3.96) (10-12) 



152 



SYSTEMATIC REVIEW 153 

TABLE XXXII. - D/U RATIO FREQUENCY IN SINPLOYEA INERMIS SUBSPECIES. TABLE XXXIV. - RIB FREQUENCY IN SINPLOYEA INERMIS SUBSPECIES. 



D/U Ratio 


inermis 


3.00-3 


.39 




3.40-3 


.79 


7 


3.80-4 


.19 


17 


4.20-4 


.59 


17 


4.60-4 


.99 


6 


5.00-5 


.39 




5.40-5 


.79 




5.80-6 


.19 




6.20-6 


.59 




6.60-6 


.99 




7.00-7 


.39 





meridionalis 

7 

17 
2 



S. inermis 
lakembana 



Ribs 

35-41 

42-48 

49-55 

56-62 

63-70 

71-77 

78-84 

85-91 

92-98 

99-105 



S . inermis 
inermis 



meridionalis 



differences are somewhat obscured by the strong ten- 
dency for gerontic crowding and irregularity of rib 
growth in most adult specimens. The differences are 
much clearer on the penultimate and first part of the 
body whorl, but convenient quantification of data from 
this section of the shell was not achieved. Most speci- 
mens are readily assignable to one of the three groups. 
Hence, recognition of three subspecies is proposed: 
Sinployea inermis inermis from Vanua Mbalavu, Kim- 
bombo, and Mango; S. i. lakembana from Lakemba; 
and S. i. meridionalis from Aiwa, Navutu-i-Loma, and 
Yangasa Levu. 

Larger collections from more islands may result in 
altering this classification. The apical cording counts 
and the size differences in ribbing are substantial. Pos- 



TABLE XXXIII. - RIBS/MM. FREQUENCY IN SINPLOYEA INERMIS 
SUBSPECIES. 



S . inermis 



Ribs/mm. 


inermis 


4.50-4.99 




5.00-5.49 




5.50-5.99 




6.00-6.49 




6.50-6.99 




7.00-7.49 




7.50-7.99 


1 


8.00-8.49 


1 


8.50-8.99 


5 


9.00-9.49 


2 


9.50-9.99 


5 


10.00-10.49 


2 


10.50-10.99 


4 


11.00-11.49 


2 


11.50-11.99 


1 



S. inermis 
meridionalis 



lakembana 



sibly specific level differentiation is involved, but the 
presence of samples not clearly referable to a sub- 
species and the amount of overlap (tables XXXII- 
XXXVI) suggest conservative treatment. 

The specimens collected from "Vatou Island" 
(= Vanua Vatu) during the surveys of the Herald in the 
mid-1850s are still preserved in the British Museum 
(Natural History) (BMNH 59.1.14.63). Only one fully 
adult shell is in good condition, but in general appear- 
ance the specimens are intermediate between sub- 
species inermis and subspecies lakembana. They have 
the relatively widely spaced ribbing that is char- 
acteristic of meridionalis and lakembana, although 
somewhat more crowded than typical examples of 
either subspecies. The umbilicus varies widely in 
width from the narrow type of lakembana to the widely 
opened umbilicus of meridionalis. 

Single shells from two islands lying within the 
range of S. i. meridionalis could not be assigned to a 
subspecies with assurance. Their measurements are 
given in Table XXXVI. The Ongea example (BPBM 
161228) has a narrower umbilicus than does the nomi- 
nate race or any specimens of S. i. meridionalis but 
disagrees in sculpture and apical cording with S. i. 
lakembana. An adult shell from Tavu-na-Sithi (BPBM 
78805) could be referred to either typical inermis or 
meridionalis. It has the umbilical size of inermis, but 
the ribbing is closer to that of meridionalis. Collection 
of adequate samples is needed in order to establish the 
position of these populations. 

Derivation of S. adposita (fig. 69d-f) from S. iner- 
mis requires size increase, greater protrusion of the 
spire and its correlated narrowing of the umbilicus, 
plus greater spacing of the ribs. Similarly, increase in 
body whorl descension and development of ribbing ir- 
regularity would be sufficient to derive S. irregularis 
(fig. 65d-f) from either S. inermis or S. princei. 

The penis of Sinployea inermis is slightly more 
than half the size of the penis inS. adposita and differs 
radically in shape and internal proportions. The vergic 



154 



SOLEM: ENDODONTOID LAND SNAILS 



TABLE XXXV. - LOCAL VARIATION IN SINPLOYEA INERMIS, I. 



NUMBER OF 
SPECIMENS 



inermis inermis 



Vanua Mbalavu 
Sta. 70, 72(1938) 

Sta. 69(1938) 
BPBM 179396 

Ex Fulton 
BPBM 167433 

Mango 

Sta. 89(1938) 
BPBM 179941 

Sta. 91(1938) 
BPBM 180025-6 

Kimbombo 

Bryan! (1924) 
BPBM 79103 

Inermis meridionalis 

Yangasa Levu 
Sta. 30(1934) 
BPBM 167138 

Aiwa 

Sta. 40(1934) 
BPBM 167209 



6 
10 



14 



10 



75.35.37 
(65-83) 

76.31.33 
(75-79) 

78.0+8.61 

(65-102) 



78.02.01 
(76-80) 

91.54.72 

(77-104) 

79 



72 



93.011.03 
(82-104) 



HEIGHT 



1.28+0.020 



(1.23-1.36) 



DIAMETER 



2.28+0.013 

(2.25-2.32) 



1.360.032 2.39+0.029 

(1.19-1.56) (2.28-2.55) 

1.48+0.058 2.52+0.067 

(1.36-1.66) (2.38-2.78) 



1.400.065 2.56+0.064 

(1.19-1.62) (2.32-2.78) 

1.67+0.032 2.950.033 

(1.46-1.85) (2.75-3.08) 

1.45+0.067 2.640.140 

(1.32-1.59) (2.28-2.88) 



H/D RATIO 



0.5620.0061 

(0.544-0.586) 

0.5700.0094 

(0.514-0.627) 

0.5850.0134 

(0.547-0.627) 



0.546+0.0127 

(0.506-0.583) 

0.564+0.0075 

(0.530-0.629) 

0.5650.0065 

(0.558-0.580) 



1.81+0.037 3.040.029 0.5950.0149 

(1.59-1.95) (2.95-3.25) (0.505-0.655) 



1.660.027 2.960.109 0.5620.0205 

(1.59-1.72) (2.72-3.21) (0.521-0.610) 



i.iner. 



WHORLS 



Mbalavu 3 5/8+(3 1/2-3 7/8) 
Sta. 70,72 

Sta. 69 3 7/8(3 5/8-4 1/4) 

Ex Fulton 4-(3 7/8-4 1/8) 



Mango 
Sta. 89 



Sta. 91 



Kimbombo 
Bryan! 



i.merid. 

Yangasa 
Sta. 30 



4 l/8+(3 7/8-4 1/2) 
4 l/4-(4-4 5/8) 

3 7/8-(3 1/2-4 1/8) 

4 3/8+(4 1/4-4 5/8) 



Aiwa 

Sta. 40 4 l/4-(4-4 1/4) 



UMBILICUS 



D/U RATIO 



APICAL 
CORDS 



0.53+0.013 4.270.094 8.00+0.71 

(0.49-0.59) (3.89-4.60) (7-10) 

0.5810.016 4.0910.098 9.3810.26 

(0.49-0.66) (3.50-4.60) (8-10) 

0.570.026 4.45+0.112 8.331.20 

(0.53-0.66) (4.17-4.69) (6-10) 



0.65+0.034 3.940.139 9 
(0.53-0.72) (3.55-4.38) 



0.70+0.017 

(0.63-0.82) 



0.63+0.087 

(0.49-0.79) 



4.20+0.081 9.57+0.37 
(3.68-4.65) (8-11) 



4.15+0.260 9 
(3.58-4.60) 



0.8710.025 3.500.092 12.0010.58 
(0.72-0.99) (3.27-4.14) (11-13) 



0.810.044 3.57+0.013 

(0.69-0.89) (3.54-3.60) 



papilla in the former is about a third of the size of the 
latter, and the muscular collar is larger both actually 
and proportionately. Sinployea adposita has the stim- 
ulatory pad smaller and shorter than in S. inermis. The 
latter has a much broader penis that is wider medially 
than apically and then tapers abruptly to atrium. 
Sinployea adposita has the penis enlarged apically and 
medially, then it tapers gradually to atrium. 



Sinployea inermis inermis (Mousson, 1870). Figures 
66a-c, 67e. 

Patula (Patula) inermis Mousson, 1870, J. de. Conchyl., 18, pp. 
118-119, pi. 7, fig. 7 Lomma-Lomma on Vanua-Balavo, Lau 
Group, Fiji Islands. 
Helix inermis (Mousson), PfeifTer, 1876, Monog. helic. viv., 7, p. 

167. 

Patula inermis Mousson, Garrett, 1887, Proc. Zool. Soc. London, 
1887, p. 178. 



SYSTEMATIC REVIEW 

TABLE XXXVI. - LOCAL VARIATION IN SINPLOYEA INERMIS, II. 



155 



NUMBER OF 
SPECIMENS 

inermis meridionalis 

Navutu-I-Loma 12 
Sta. 28-9(1934) 
BPBM 167019, 
-55-6 

inermis lakembana 



Transitional Sets 

Vatou 5 

BMNH 59.1.14. 
63 



Ongea 

Tavua-na- 
Sithi 



RIBS 



68.03.16 
(53-83) 



HEIGHT 



1.480.39 

(1.32-1.69) 



DIAMETER 



2.72+0.047 

(2.52-3.05) 



Lakemba 
Sta. 42(1934) 
BPBM 167222 


4 


54. 


814.61 
(42-62) 


1. 


48+0 
(1 


.083 
.32-1 


.66) 


2, 


.5810 
(2 


.111 
.38-2. 


88) 


Sta. 97(1938) 
BPBM 180101 


10 


49. 


5+1.71 
(38-59) 


1. 


3610 
(1 


.025 
.19-1. 


.49) 


2, 


,2910 
(2 


.028 
.15-2. 


38) 



65.83.40 
(59-70) 



83 
75 



1.3310.093 

(1.13-1.61) 



1.41 
1.58 



2.420.100 

(2.19-2.73) 



2.66 

2.70 



H/D RATIO 



0.544+0.0087 

(0.500-0.595) 



0.5740.0195 

(0.552-0.633) 



0.5940.0085 

(0.545-0.634) 



0.5480.0177 

(0.518-0.616) 



0.531 
0.586 



i.iner. 



WHORLS 



Navutu 

Sta. 28-29 4 l/8+(3 7/8-4 1/2) 



i . lakem. 

Lakemba 

Sta. 42 4 l/8+(3 7/8-4 1/2) 



Sta. 97 



4-(3 3/4-4 1/8) 



Transitional 

Vatou 4(3 7/8-4 1/4) 

Ongea 3 3/4 

Tavua. 4 1/4 - 



UMBILICUS 



D/U RATIO 



APICAL 
RIBS 



0.78+0.021 3.50+0.055 11.430.68 
(0.69-0.92) (3.07-3.81) (9-14) 



0.620.049 4.2110.251 

(0.53-0.76) (3.78-4.93) 

0.3710.015 6.24+0.206 13.57+0.72 
(0.33-0.49) (4.73-7.10) (11-16) 



0.57+0.026 4.30+0.375 

(0.49-0.66) (3.32-5.53) 



0.56 
0.66 



4.77 
4.10 



Helix (Patula) inermis (Mousson), Tryon, 1887, Man. Conchol, (2) 

3, p. 41, pi. 8, figs. 60-61. 

Endodonta (Champa) inermis (Mousson), Pilsbry, 1893, Man. Con- 
chol. (2) 9, p. 35. 

Charopa inermis (Mousson), Gude, 1913, Proc. Malacol. Soc. Lon- 
don, 10 (5), p. 330 Vatou (Herald Expedition!); Germain, 1932, 
Ann. Inst. Ocean., 12 (2), p. 45. 

Diagnosis. Shell relatively small, diameter 2.24-3.06 mm. 
(mean 2.60 mm.), with 3 l /2-4% normally coiled whorls. Apex and 
spire usually moderately and evenly elevated, last whorl descending 
moderately to much more rapidly, spire protrusion about '/fe body 
whorl width, H/D ratio 0.500-0.629 (mean 0.562). Apical whorls with 
6-11 (mean 8.96) prominent spiral cords. Postnuclear whorls with 
high, prominent, narrow, strongly protractively sinuated radial ribs, 
65-104 (mean 80.0) on the body whorl, whose interstices are 3-6 
times their width. Ribs/mm. 8.39-11.75 (mean 9.82). Microsculpture 



of very fine radial riblets, 6-12 between each pair of major ribs, 
crossed by slightly finer and more crowded spiral riblets and a few 
widely spaced secondary spiral cords. Umbilicus open, U-shaped, last 
whorl decoiling much more rapidly, contained 3.47-4.69 times (mean 
4.17) in the diameter, margins rounded. Sutures and whorl contours 
typical, color reddish yellow-horn without separate flammulations. 

Sinployea inermis inermis (fig. 66c) has a narrower 
umbilicus and more crowded major radial ribbing than 
S. i. meridionalis (fig. 66d); S. i. lakembana (fig. 66e) 
has much coarser and more widely spaced major radial 
ribs. Other Fijian species of similar size, S. princei, S. 
lauensis, and S. navutuensis, differ in proportions 
(table XXX) and/or ribbing. The Tongan S. uicaria (fig. 
58a-c) has a higher spire and much more crowded sec- 




a 







FIG. 66. a-c, Sinployea inermis inermis (Mousson). Station 72, Lomaloma, Vanua Mbalavu, Lau, Fiji. BPBM 179460; d, Sinployea inermis 
meridionalis, new subspecies. Station 30, Yangasa Levu, Lau, Fiji. Holotype. BPBM 167138; e, Sinployea inermis lakembana, new subspecies. 
Station 42, Lakemba, Lau, Fiji. Holotype. BPBM 167222. Scale lines equal 1 mm. (a-c, SH: d-e, MM). 



156 



SYSTEMATIC REVIEW 



157 



ondary spiral cording in addition to the much more 
crowded radial ribbing. 

Description. Shell relatively small, with slightly less than 4'/8 
normally coiled whorls. Apex barely emergent, upper spire flat, body 
whorl descending moderately sharply, H/D ratio 0.515. Apical whorls 
1%, sculpture partially eroded with traces of 7 fine, rather widely 
spaced spiral ribs remaining. Postnuclear whorls with prominent, 
rounded, protractively sinuated radial ribs, 69 on the body whorl, 
whose interstices are 2-4 times their width. Microsculpture of very 
fine radial riblets, 4-6 between each pair of major ribs, crossed by 
even finer, more crowded spiral riblets with a secondary sculpture of 
moderately prominent, widely spaced spiral cords developed. Sutures 
deep, whorls strongly rounded above, moderately flattened laterally 
above periphery and on basal margin. Umbilicus open, U-shaped, 
last whorl decoiling more rapidly, contained 3.47 times in the diame- 
ter, rounded margins. Color light reddish brown. Aperture ovate, lip 
slightly deflected above, flattened laterally above periphery and on 
basal margin, inclined about 20 from the shell axis. Height of lec- 
totype 1.38 mm., diameter 2.58 mm. 

Lectotype. Fiji: Lau Group, Vanua Mbalavu. Col- 
lected by Dr. Graeffe. JDCP 3214. 

Range. Vanua Mbalavu, Mango, Kimbombo, 
Tuvutha, Lau Group, Fiji. 

Paratypes. Zurich, FMNH 116998. 

Material. Vanua Mbalavu (6 specimens, BPBM 
167433): Lomaloma (9 specimens, Zurich, J. de Con- 
chyl., FMNH 116998); north end Lomaloma (Station 
69) 10-12 ft. inland at 3 ft. elevation (28 specimens, 
BPBM 179396, BPBM 179398, BPBM 179399); back of 
Lomaloma (Station 67) l /2-% mile inland, 250-700 ft. 
elevation (8 specimens, BPBM 179379); south end 
Lomaloma (Station 72) 15 ft. inland at 3 ft. elevation 
(14 specimens, BPBM 179460, BPBM 179461); south 
end Unione Village (Station 70) 10 ft. inland at 3 ft. 
elevation (2 specimens, BPBM 179426); between Val- 
ika and Mosomo Bay (Station 78) % mile inland at 
200-250 ft. elevation (3 specimens, BPBM 179584, 
BPBM 179585); Station F-8, 300 ft. elevation, P/2 miles 
inland, northwest area (6 specimens, FMNH 168161). 
Mango Island: Marona (Station 92) 100 yd. inland at 10 
ft. elevation (2 specimens, BPBM 180047); south- 
southwest from Marona (Stations 89, 90) about Vz-% 
mile inland at 200-400 ft. elevation (15 specimens, 
BPBM 179941, BPBM 179975, BPBM 179988); %-l 
mile southwest of Marona (Station 91) 1-50 yd. inland, 
at 5-30 ft. elevation (82 specimens, BPBM 180025-9). 
Kimbombo: East Islet, 100 yd. inland at 50-150 ft. ele- 
vation (28 specimens, BPBM 79079, BPBM 79103). 
Tuvutha: west coast (Station F-7) at 100 ft. elevation (5 
specimens, FMNH 193768). 

Remarks. The specimens of Sinployea inermis 
inermis from Vanua Mbalavu, Mango, and Kimbombo 
agree in sculpture and umbilical proportion, although 
exhibiting slight differences in size (table XXXV). 
Shells from Vanua Mbalavu are small, whereas those 
from Kimbombo and Mango are slightly to distinctly 
larger. On Mango, two populations were sampled. 
Shells from Station 89 (Vz-% mile south-southwest of 
Marona at 350-400 ft. elevation) were dead. They are 
noticeably smaller and with fewer radial ribs than 



those from Station 91 (%-l mile southwest of Marona 
at 5-30 ft. elevation), which were collected alive. The 
latter sample contained a higher proportion of gerontic 
individuals, whereas the Station 89 material was 
barely adult. The Mango specimens are larger than 
those from Vanua Mbalavu, but no hesitation is felt in 
referring them to the same subspecies. 

Material from both Mango (BPBM 180025) and 
Vanua Mbalavu (BPBM 179396) was dissected. Only 
differences are noted below. 

Description of soft parts. Pallial region typical, arms of kidney 
subequal, rectal slightly longer, maximum length 1.7 mm., slight 
strip of lung roof visible between ureter arms. 

Genitalia (fig. 67e) typical, free oviduct slightly more slender 
than usual. Penis (P) about 0.8-1.0 mm. long, greatly swollen me- 
dially. Internally with very tiny vergic papilla surrounded by a 
heavy muscular collar. Stimulatory pad very large, walls below pad 
with obscure pilasters. 



Sinployea inermis meridionalis, new subspecies. 
Figure 66d. 

Diagnosis. Shell of average size, diameter 2.50-3.22 mm. 
(mean 2.88 mm.), with 3%-4% rather tightly coiled whorls. Apex and 
spire moderately and evenly elevated, body whorl descending mod- 
erately to much more rapidly, spire protrusion about l k body whorl 
width, H/D ratio 0.505-0.655 (mean 0.566). Apical sculpture of 9-14 
(mean 11.6) prominent, but narrow spiral cords. Postnuclear 
sculpture of narrow, prominent, protractively sinuated, rather 
widely spaced radial ribs, 53-104 (mean 72.2) on the body whorl, 
whose interstices are 3-7 times their width. Ribs/mm. 6.75-11.70 
(mean 8.37). Microsculpture of very fine radial riblets, 8-12 between 
each pair of major ribs, crossed by barely visible spiral riblets, with a 
weak secondary sculpture of fine and widely spaced spiral cords that 
varies in frequency and prominence. Umbilicus broadly U-shaped, 
last whorl decoiling much more rapidly, contained 3.07-3.81 times 
(mean 3.51) in the diameter, margins rounded. Whorls strongly flat- 
tened laterally above periphery, aperture subcircular, inclined about 
20 from shell axis. 

Sinployea inermis meridionalis (fig. 66d) has a 
wider umbilicus and more widely spaced major radial 
ribbing than the nominate race; S. i. lakembana (fig. 
66e) is smaller, has a narrower umbilicus, and much 
more widely spaced radial ribbing. Sinployea vicaria 
paucicosta (fig. 58d) from Tonga is smaller, has the 
spire protruding further, and has a narrower um- 
bilicus. 

Description. Shell slightly smaller than average, with 4'/4 rela- 
tively tightly coiled whorls. Apex barely elevated, lower whorls de- 
scending slowly, body whorl more rapidly, H/D ratio 0.565. Em- 
bryonic whorls l'/2, sculpture of 11 relatively fine spiral ribs with an 
intrusion of faint radial ribbing on the last quarter whorl. Remain- 
ing whorls with rounded, prominent, protractively sinuated radial 
ribs, 67 on the body whorl, whose interstices are about 3-6 times 
their width. Microsculpture of very fine radial riblets, 8-12 between 
each pair of major ribs, with exceedingly fine, inconspicuous spiral 
riblets and a secondary sculpture of low, broadly rounded, relatively 
widely spaced spiral cords. Sutures deeply impressed, whorls 
strongly rounded above, flattened laterally above peripheral margin, 
slightly compressed basally. Umbilicus broadly open, last whorl de- 
coiling more rapidly, contained 3.29 times in the diameter. Color 
light yellow-reddish brown without darker flammulations. Aperture 
circular, strongly flattened laterally above periphery, with sharply 
rounded umbilical basal margin, inclined about 10 from the shell 
axis. Height of holotype 1.71 mm., diameter 3.03 mm. 



158 



SOLEM: ENDODONTOID LAND SNAILS 



DG 




FIG. 67. Anatomy of Fijian, Caroline Island, and New Hebridean Sinployea: a-d, Sinployea irregularis (Garrett). Station 118, Nandarivatu, 
Viti Levu, Fiji. BPBM 178897. a, genitalia. 6, detail of talon-carrefour region, c, partial view of penis interior, d, apical part of penis interior; e, 
genitalia of Sinployea inermis inermis (Mousson). Station 91, Marona, Mango Island, Lau, Fiji. BPBM 180025; f, genitalia of Sinployea inermis 
lakembana, new subspecies. Station 97, Toumba, Lakemba Island, Lau, Fiji. BPBM 180101; g, penis and epiphallus of Sinployea adposita 
(Mousson). Station 97, Toumba, Lakemba Island, Lau, Fiji. BPBM 180099; h, genitalia of Sinployea kusaieana, new species. Station 44, Lele 
Islet, Kusaie, Caroline Islands. BPBM 155874; i-j, Sinployea euryomphala (Solem). Espiritu Santo, New Hebrides. FMNH 109493. (a-c, e-f, h, 
MO; d, MM; g, JC; i-j, PS). 



Holotype. Fiji: Lau Group, Yangasa Levu, Sta- 
tion 30, hillside on south end of island, 150 ft. inland at 
40 ft. elevation. Collected by H. S. Ladd on July 26, 
1934. BPBM 167138. 

Range. Yangasa Levu, Navutu-i-Loma, and 
Aiwa, Lau Group, Fiji. 



Paratypes. Yangasa Levu: hillside at south end 
of island (Station 30), 150 ft. inland at 40 ft. elevation 
(96 specimens, BPBM 167138). Aiwa: Vataniyambia 
on southwest quarter (Station 40) of island, about 50 
yd. inland at 25 ft. elevation (20 specimens, BPBM 
167209). Navutu-i-Loma: ridge on northeast quarter of 



SYSTEMATIC REVIEW 



159 



island (Stations 28, 29), about 150 yd. inland at 100 ft. 
elevation (52 specimens, BPBM 167019, BPBM 
167056, BPBM ex 167055). 

Remarks. The shells of Sinployea inermis mer- 
idionalis from Navutu-i-Loma are distinctly smaller 
and more depressed in H/D ratio than are those from 
Aiwa or Yangasa Levu (tables XXXV-XXXVI). Shells 
from Aiwa have a much higher rib count. Compara- 
tively few of the examples were adult or whole enough 
to measure, so that the significance of this data is un- 
certain. From Navutu-i-Loma there were 23.1% adults; 
from Aiwa 20%; and from Yangasa Levu only 10.3%. I 
would expect that the size difference is real, but more 
complete collections are required. 

Sinployea inermis lakembana, new subspecies. Fig- 
ures 66e, 67f. 

Diagnosis. Shell small, diameter 2.14-2.86 mm. (mean 2.36 
mm.), with 3%-4'/2 rather tightly coiled whorls. Apex and spire mod- 
erately to strongly and evenly elevated, last whorl descending much 
more rapidly, spire protrusion more than l k body whorl width, H/D 
ratio 0.508-0.634 (mean 0.583). Apical sculpture of 11-16 (mean 
13.6) prominent spiral cords. Postnuclear sculpture of high, very 
prominent, sharply defined, protractively sinuated radial ribs, 38-64 
(mean 51.3) on the body whorl, whose interstices are 4-6 times their 
width on the spire and first part of body whorl, becoming crowded 
and irregular on gerontic growth sections. Ribs/mm. 4.67-8.20 
(mean 7.01). Microsculpture of extremely fine radial riblets, more 
than 10 between each pair of major ribs, but usually too fine for 
accurate counting at 96 x magnification, barely visible microspirals, 
and more prominent, rather widely spaced secondary spiral cording. 
Umbilicus variable in size, usually narrow, U-shaped, last whorl 
decoiling slightly to much more rapidly, contained 3.07-7.10 times 
(mean 5.54) in the diameter, margins strongly rounded. Sutures very 
deep, whorls shouldered above, compressed laterally above periphery 
and on basal margin, aperture inclined about 30 from shell axis. 

Sinployea inermis lakembana has much stronger 
and more widely spaced major radial ribbing, plus a 
usually more strongly deflected body whorl and nar- 
rower umbilicus than either of the other races of S. 
inermis (fig. 66a-d). No other Fijian species has such 
strong ribbing that is so widely spaced. 

Description. Shell relatively small, with 4'/2 tightly coiled 
whorls. Apex very slightly elevated, lower spire and body whorl de- 
scending more rapidly, H/D ratio 0.552. Apical whorls l'/2, sculpture 
of 12 prominent spiral cords, surface partially eroded. Postnuclear 
whorls with prominent, relatively narrow, quite widely spaced radial 
ribs, 42 on the body whorl, whose interstices are 3-6 times their 
width. Microsculpture of extremely fine radial and even finer, more 
crowded spiral riblets, crossed by low, rounded, relatively closely 
spaced spiral cords. Sutures deeply impressed, whorls strongly 
rounded, only slightly flattened laterally above periphery and on 
basal margin. Color partially leached from shell. Umbilicus moder- 
ately widely open, U-shaped, last whorl decoiling more rapidly, con- 
tained 3.78 times in the diameter. Aperture circular, slightly flat- 
tened laterally above periphery, lip broken. Height of holotype 1.58 
mm., diameter 2.86 mm. 

Holotype. Fiji: Lau Group, Lakemba, Station 42, 
Tambusue, west side near coast at 10 ft. elevation. Col- 
lected by H. S. Ladd on August 8, 1934. BPBM 167222. 

Range. Lakemba, Lau Group, Fiji. 

Paratypes. Lakemba (2 specimens, AIM, col- 
lected June 30, 1977, by W. Cernohorsky): Tambusue, 
west side near coast (Station 42) at 10 ft. elevation (3 



specimens, BPBM 167222); west end of Toumba 
(=Tubou) (Station 97) to % mile away at 15 ft. eleva- 
tion (10 specimens, BPBM 180101). 

Remarks. The holotype of Sinployea inermis 
lakembana is the largest and most widely umbilicated 
specimen seen, but was chosen because of the clear 
sculpture. Three sets were available, collected in 
different years from different portions of Lakemba. 
Shells taken on the west coast by Harry Ladd in 1934 
(Station 42, BPBM 167222) are larger, less elevated, 
and more widely umbilicated (table XXXVI) than 
those found near Toumba (=Tubou) by Zimmerman 
and Kondo in 1938 (Station 97, BPBM 180101). Cer- 
nohorsky's 1977 specimens (Solem, 1978a) agree with 
the former examples. Apparently Ladd's Station 42 is 
the same as his geological Station L492, shown in Ladd 
& Hoffmeister (1945, p. 99, fig. 19). It was cited as 
"about 3 miles northwest of Tumbou" (Ladd & 
Hoffmeister, 1945, p. 190) and thus is separated by 2.6 
miles from the later collection by Kondo and Zimmer- 
man. Both sets contain only adults showing marked 
gerontic growth, so that the differences are not caused 
by age-biased samples. Further collections are needed 
to determine whether the differences are ecopheno- 
typic or geographic in nature. With the umbilical vari- 
ation showed by the Vanua Vatu material (table 
XXXVI), I suspect that there will be no systematic sig- 
nificance to this difference between the sets. 

Description of soft parts. Pallial region and external body fea- 
tures as in nominate subspecies. Genitalia (fig. 67f) without substan- 
tive differences, more swollen spermathecal base caused by presence 
of sperm packet. Shorter appearance of atrium (Y) caused by drawing 
a retracted specimen with atrium and vagina folded under edge. 

(Based on BPBM 180101, 2 whole and several fragmentary 
examples.) 

Sinployea lauensis, new species. Figure 68a-c. 

Diagnosis. Shell slightly smaller than average, diameter 
2.40-3.13 mm. (mean 2.76 mm.), with 3'/2-4'/4 rather loosely coiled 
whorls. Apex and spire barely to moderately and evenly elevated, 
last whorl descending a little more rapidly, spire protrusion about '/s 
body whorl width, H/D ratio 0.493-0.581 (mean 0.530). Apical 
whorls with 9-11 (mean 9.6) fine spiral cords. Postnuclear sculpture 
of low, rounded, crowded, strongly protractively sinuated radial ribs, 
95-119 (mean 103.9) on the body whorl, whose interstices are 1-2 
times their width. Ribs/mm. 10.93-14.46 (mean 11.53). Micro- 
sculpture of fine radial riblets, 2-5 between each pair of major ribs, 
crossed by slightly finer and more crowded spiral riblets, with fine, 
relatively crowded secondary spiral cording visible on spire and part 
of the body whorl. Umbilicus narrow, V-shaped, last whorl decoiling 
much more rapidly, contained 3.88-4.65 times (mean 4.24) in the 
diameter, margins rounded. Whorls strongly rounded above, moder- 
ately to strongly compressed laterally above periphery, aperture 
subcircular, inclined about 25 from shell axis. Color reddish horn, no 
darker markings. 

Sinployea lauensis differs from S. inermis (fig. 
66a-e) in its much less protruded spire, greater rib 
frequency, and V-shaped, much more rapidly decoiling 
umbilicus. Sinployea princei (fig. 65a-c) is smaller, has 
fewer ribs, and the whorls are more evenly rounded 
and tightly coiled, with the umbilicus decoiling more 
regularly. 








FIG. 68. a-c, Sinployea lauensis, new species. Station 43, Nayau, Lau, Fiji. Holotype. BPBM 167234; d-f, Sinployea navutuensis, new 
species. Station 29, Navutu-i-Loma, Lau, Fiji. Holotype. BPBM 9730. Scale lines equal 1 mm. (MM). 



160 



SYSTEMATIC REVIEW 



161 



Description. Shell of average size, with slightly less than 3% 
loosely coiled whorls. Apex and spire barely protruding, body whorl 
descending moderately, H/D ratio 0.494. Embryonic whorls 1%, 
sculpture of 9 very faint and widely spaced spiral cords. Remaining 
whorls with closely spaced, narrow, rounded, strongly protractively 
sinuated radial ribs, 103 on the body whorl, whose interstices are 1-2 
times their width. Microsculpture of 3-6 fine radial riblets between 
each pair of major ribs, crossed by lower, more crowded spiral riblets, 
with some widely spaced narrow larger spiral cords. Sutures deeply 
impressed, whorls shouldered above, broadly flattened between 
shoulder and periphery, with slightly flattened basal margin. Um- 
bilicus narrow internally, last whorl decoiling very rapidly, con- 
tained 4.05 times in the diameter. Color light reddish brown. Aper- 
ture subcircular, broadly flattened laterally above, inclined about 
25 from the shell axis, with internal basal and columellar callus. 
Columellar margin of aperture slightly reflected. Height of holotype 
1.38 mm., diameter 2.80 mm. 

Holotype. Fiji: Lau Group, Nayau Island, Station 
43, J /2 mile inland from Nauko at 250 ft. elevation. Col- 
lected at base of a limestone cliff by H. S. Ladd on 
August 13, 1934. BPBM 167234. 

Range. Nayau, Yangasa Levu, Namuka, 
Navutu-i-Loma, and Wangava, Lau Group, Fiji. 

Paratypes. Nayau: at base of limestone cliff (Sta- 
tion 43) J /2 mile inland from Nauko at 250 ft. elevation 
(19 specimens, BPBM 167234). Namuka: southeast 
coast (Station F-3) at 50-75 ft. elevation (5 specimens, 
FMNH 168089). Yangasa Levu: hillside at south end of 
island (Station 30) 150 ft. inland at 40 ft. elevation (23 
specimens, BPBM 167139). Navutu-i-Loma: ridge on 
northeast quarter of island (Stations 28, 30) 150 yd. 
inland at 150 ft. elevation (32 specimens, BPBM 
167020, BPBM 167055). Wangava: northeast end of is- 
land (Stations 26, 27) 100 yd. inland at 10-15 ft. eleva- 
tion (7 specimens, BPBM 166955, BPBM 166968, 
BPBM 166997); north end at 100 ft. elevation (38 
specimens, FMNH 168092, FMNH 168101). 

Remarks. Individual specimens of Sinployea 
lauensis approach the measurements of S. inermis 
meridionalis for single characters (table XXX), but the 
different umbilical and whorl shape separate problem 
specimens. Typical examples are immediately separa- 
ble on a number of characters. There is considerable 
similarity to the Samoan Sinployea allecta (fig. 54a-e) 
in general appearance, but that species is much more 
narrowly umbilicated and has less crowded radial rib- 
bing. 

Very little material was adult, and mean dif- 
ferences between the small samples from different is- 
lands are not statistically significant (table XXXI). 

Sinployea navutuensis, new species. Figure 68d-f. 

Diagnosis. Shell smaller than average, diameter 2.58-2.78 
mm. (mean 2.66 mm.), with 4-4% normally coiled whorls. Apex and 
early spire flat to barely protruding, lower spire descending slightly, 
body whorl much more rapidly, spire protrusion V4 body whorl width, 
H/D ratio 0.449-0.556 (mean 0.498). Apical sculpture of 9-11 (mean 
10.3) narrow but prominent spiral cords. Postnuclear whorls with 
narrow, crowded, fine, sharply outlined, protractively sinuated ra- 
dial ribs, 121-147 (mean 136.7) on the body whorl, whose interstices 
are 1-2 times their width. Ribs/mm. 16.94-17.62 (mean 17.3). Mi- 
crosculpture a lattice of barely visible radial riblets, 3-6 between 
each pair of major ribs, crossed by finer spiral riblets. A widely 



spaced, fine sculpture of secondary spiral cording visible under an- 
gled lighting only. Umbilicus broadly open, cup-shaped, regularly 
decoiling, contained 3.25-3.52 times (mean 3.38) in the diameter, 
margins rounded. Whorls strongly rounded, slightly compressed 
laterally above periphery and on basal margin, aperture circular, 
compressed laterally, inclined about 20 from shell axis. 

The nearly flat apex, slight spire protrusion, wide 
umbilicus, and very crowded radial ribbing separate 
Sinployea navutuensis from the other Fijian species. 
Sinployea lauensis (fig. 68a-c) has a much narrower 
umbilicus, fewer and less crowded radial ribs, more 
loosely coiled whorls, and a wider body whorl. Sin- 
ployea godeffroyana (fig. 69a-c) is much smaller, much 
more elevated, has a narrower umbilicus, and even 
more crowded radial ribs. Sinployea kusaieana (fig. 
63a-c) from the Caroline Islands is very similar in size 
and shape, but differs in having only 84-103 ribs on 
the body whorl and a slightly looser coiling pattern. 

Description. Shell smaller than average, with 4'/s moderately 
tightly coiled whorls. Apex and early spire barely elevated, body 
whorl descending moderately, H/D ratio 0.487. Embryonic whorls 
Ite, sculpture of 9 narrow, prominent spiral ribs. Postnuclear whorls 
with low, rounded, narrow, quite crowded, protractively sinuated 
radial ribs, 142 on the body whorl, whose interstices are about 1-2 
times their width. Microsculpture of 3-5 very fine riblets between 
each pair of major ribs, crossed by much finer, lower, spiral riblets. 
Secondary sculpture of widely spaced, rounded spiral cords visible 
under good lighting. Sutures deep, whorls strongly rounded above, 
compressed laterally above periphery and slightly on basal margin. 
Umbilicus broadly cup-shaped, regularly decoiling, contained 3.39 
times in the diameter. Color leached from shell. Aperture ovate, 
compressed laterally above periphery with gently rounded margin, 
inclined about 15 from the shell axis. Height of holotype 1.25 mm., 
diameter 2.57 mm. 

Holotype. Fiji: Lau Group, Navutu-i-Loma, Sta- 
tion 29, on ridge 150 yd. inland at 100 ft. elevation. 
Collected dead among limestone outcrops by H. S. Ladd 
on July 24, 1934. BPBM 9730. 

Range. Navutu-i-Loma and Oneata, Lau Group, 
Fiji Islands. 

Paratypes. Navutu-i-Loma: on ridge 150 yd. in- 
land (Station 29, 1934) at 100 ft. elevation (28 speci- 
mens, BPBM 167103, ex BPBM 167020, ex BPBM 
167055). Oneata: 1 mile north of Dakuiloa, 1 A-1 mile 
inland (Station 99, 1938) at 25-50 ft. elevation (1 
specimen, BPBM 180160). 

Remarks. The shell from Oneata (BPBM 180160) 
is juvenile, height 0.82 mm., diameter 1.81 mm., H/D 
ratio 0.455, whorls 3Vz, umbilicus 0.58 mm., D/U ratio 
3.09, and may not belong to this species. It is closer to 
S. navutuensis than to any other Lau species. 

Only four examples were adult, the remaining 24 
being juvenile or broken. The secondary spiral cording 
is visible only under ideal angular lighting. It is widely 
spaced and distinctly more prominent than the radial 
microribbing. 

Sinployea adposita (Mousson, 1870). Figures 67g, 
69d-f. 

Patula adposita Mousson, 1870, J. de Conchyl., 18, pp. 119-120, pi. 
7, fig. 8 Oneata, Lau Group, Fiji Islands; Garrett, 1887a, Proc. 
Zool. Soc. London, 1887, p. 178. 






a be 





FIG. 69. a-c,Sinployeagodeffroyana, new species. Station 133, Nandarivatu, 2,300-2,500 ft., Viti Levu, Fiji. Holotype. BPBM 179080; d-f, 
Sinployea adposita (Mousson). Station 100, Oneata, Lau, Fiji. BPBM 180177. Scale lines equal 1 mm. (a-c, MM; d-f, SH). 



162 



SYSTEMATIC REVIEW 



163 



Helix adposita (Mousson), Pfeiffer, 1876, Monog. helic. viv., 7, p. 
166. 

Helix iPatula) adposita (Mousson), Tryon, 1887, Man. Conchol., (2) 
3, p. 41, pi. 8, figs. 58-59. 

Endodonta (Champa) adposita (Mousson), Pilsbry, 1894, Man. 
Conchol., (2) 9, p. 35. 

Charopa adposita (Mousson), Gude, 1913, Proc. Malacol. Soc. Lon- 
don, 10 (5), p. 330; Germain, 1932, Ann. Inst. Ocean., 12 (2), p. 
45. 

Diagnosis. Shell large, diameter 2.63-3.75 mm. (mean 3.29 
mm.), with 4-4% rather tightly coiled whorls. Apex and spire moder- 
ately to strongly and evenly elevated, body whorl descending much 
more rapidly, spire protrusion about l k-Yt body whorl width, H/D 
ratio 0.538-0.713 (mean 0.603). Apical sculpture of 12-14 (mean 
13.0) rather fine spiral cords. Postnuclear sculpture of narrow, fine, 
strongly protractively sinuated radial ribs, 36-74 (mean 53.3) on the 
body whorl, whose interstices are 3-6 times their width. Ribs/mm. 
3.35-7.23 (mean 5.28). Microsculpture of extremely fine radial rib- 
lets, usually more than 10 between each pair of major ribs, crossed by 
barely visible spiral riblets. Secondary spiral cords much more prom- 
inent than microradials, interstices about 3-4 times their width. 
Umbilicus very narrow, U-shaped, not or only slightly decoiling, con- 
tained 7.87-17.0 times (mean 11.9) in the diameter, margins 
rounded. Whorls strongly and almost evenly rounded, very slightly 
compressed laterally above periphery and on basal margin, aperture 
ovate, inclined about 30 from shell axis. 

Sinployea adposita differs from all other Fijian 
species in its very narrow umbilicus and presence of 
relatively regularly spaced, major radial ribbing. 
Other Fijian species of similar size have either widely 
open umbilici or if the umbilical range approaches that 
of S. adposita, they have lost the major radial ribbing 
(S. recursa, fig. 64b) or have an angulated periphery (S. 
angularis, fig. 64e). 

Description. Shell relatively large, with slightly less than 4% 
moderately loosely coiled whorls. Apex and spire moderately and 
evenly elevated, last whorl not descending much more sharply, H/D 
ratio 0.622. Apical whorls l'/2, sculpture of about 12 fine spiral ribs 
with a secondary sculpture of much smaller, more closely spaced 
radial riblets. Remaining whorls with thin, protractively sinuated 
radial ribs, 49 on the body whorl, whose interstices are about 3-5 
times their width. Microsculpture a lattice of very fine radial ribs 
crossed by finer, much more crowded spiral riblets with a secondary 
sculpture of much more prominent, moderately closely spaced spiral 
cords. Sutures moderately deep, whorls strongly rounded above, very 
slightly flattened laterally above periphery. Umbilicus very narrow, 
U-shaped, not decoiling, contained 13.60 times in the diameter. Color 
light reddish horn, without darker flammulations. Aperture ovate, 
inclined about 20 from the shell axis. Height of lectotype 1.81 mm., 
diameter 2.91 mm. 

Lectotype. Fiji: Lau Group, Oneata. Collected by 
Dr. Graeffe. JDCP 5246. 

Range. Oneata, Munia, Lakemba, Aiwa, Mothe, 
Tuvutha, Karoni, Nayau, Yangasa Levu, Lau Group, 
Fiji Islands. 

Paratypes. Zurich, FMNH 116979, JDCP. 

Material. Oneata: southwest end of village (Sta- 
tion 100) V* mile inland at 100 ft. elevation (30 speci- 
mens, BPBM 180177-80). Lakemba (1 specimen, 
BPBM 180117; 30 specimens. AIM, FMNH, collected 
June 24-30, 1977, by Walter Cernohorsky): west end 
Toumba (Station 97) 10 ft. to V 4 mile inland at 15 ft. 
elevation (26 specimens, BPBM 180099-102). Munia: 
Hysik's plantation (Station 64), 50 yd. inland, 100-600 



ft. elevation (1 specimen, BPBM 179294); northeast 
middle of island (Station 65), back of Hysik's home, 2 
mile inland at 600-900 ft. elevation (5 specimens, 
BPBM 179322). Nayau: % mile north of Liku Village 
(Station 101) 100 yd. inland at 30-500 ft. elevation (14 
specimens, BPBM 180260); Nauko (Station 43) Vz mile 
inland at 250 ft. elevation (4 specimens, BPBM 
167235). Yangasa Levu: south end Yangasa Levu (Sta- 
tion 30) 150 ft. elevation (67 specimens, BPBM 
167140). Aiwa: Vataniyambia, southwest quarter of is- 
land (Station 40) 50 yd. inland at 25 ft. elevation (7 
specimens, BPBM 167210). Karoni (100 specimens, 
AIM, FMNH, collected July 4, 1977, by Walter Cer- 
nohorsky): slope of peak at 90 ft. elevation (9 speci- 
mens, BPBM 78604-5); Mothe (1 specimen, AIM, col- 
lected July 7, 1977, by Walter Cernohorsky). Tuvutha: 
(Station F-7) west coast at 100 ft. elevation (6 speci- 
mens, FMNH 168132). 

Remarks. Specimens of Sinployea adposita show 
considerable variation in spacing of the major radial 
ribs and some variation in shell height. Mainly dead 
material in small sets was available. 

Shells from Munia and the type set from Oneata 
(table XXXVII) are distinctly Hatter with a lower H/D 
ratio than those from the remaining islands. Possibly 
there is subspeciation, but since most material was 
both dead and juvenile, such separation is premature. 

Shells collected from Oneata Island in 1938 are 
unusual in having the radial ribbing extremely widely 
spaced, with 37-57 (mean 42.7) ribs on the body whorl. 
Those from Munia average 56 ribs on the body whorl; 
one unworn adult from Lakemba had 68; on Yangasa 
Levu they averaged 70.5; the single adult from Karoni 
Islet has 72; and on Nayau shells from two populations 
show the greatest extremes. Rib counts were not made 
on the 1977 collections from Lakemba and Karoni 
(Solem, 1978a). Specimens from % mile north of Liku 
collected in 1938 have an average of 76.3 ribs on the 
body whorl, whereas the one adult collected at Nauko 
in 1934 has only 45 ribs, and the juveniles also have 
very widely spaced ribbing. This cannot be interpreted 
as meaning that the spacing of the ribbing necessarily 
varies seasonally since the two localities on Nayau 
yielded the quite distinct species Vatusila kondoi and 
nayauana. The variation in rib spacing shows no geo- 
graphic pattern, and the populations show practically 
no variation in adult size (table XXXVII). The shells 
from Yangasa Levu, Mothe, and Munia have slightly 
narrower umbilici, but a minor size change produces a 
large numerical shift in this range of measurements. 

Description of soft parts. Fragmentary extracted individuals 
yielded partial genitalia and one pallial region. Latter typical, kid- 
ney 1.45 mm. in greatest length, rectal arm slightly longer than 
pericardia!, arms of ureter widely separated by strip of lung roof. 

Genitalia without unusual external features. Penis (fig. 67g, P) 
about 1.65 mm. long, internally with typical vergic papilla and small 
stimulatory pad. 



TABLE XXXVII. - LOCAL VARIATION IN SINPLOYEA ADPOSITA AND S. IRREGULARIS. 



NUMBER OF 
SPECIMENS 



DIAMETER 



H/D RATIO 



Oneata 
Zurich 


9 45.3*2.17 
(36-54) 


1.87*0.057 
(1.62-2.09) 


3.2310.087 
(2.95-3.64) 


0.57910.0080 
(0.544-0 


.607) 


Sta. 100, 1938 
BPBM 180177-80 


9 42.7*3.04 
(37-57) 


2.03*0.042 
(1.85-2.22) 


3.3010.053 
(3.11-3.64) 


0.61510.0080 
(0.587-0 


.656) 


Munia 
Sta. 64-5, 1938 
BPBM 179322, -294 


4 56.32.33 
(52-60) 


1.80*0.136 
(1.56-2.12) 


3.09+0.192 
(2.72-3.51) 


0.58110.0105 
(0.533-0 


.603) 


Lakemb a 
Sta. 97, 1938 
BPBM 180099 


1 


2.090.099 
(1.89-2.19) 


3.2810.057 
(3.18-3.38) 


0.63610.0329 
(0.575-0 


.687) 


Sta. 98, 1938 
BPBM 180102, -17 


6 68 


2.09*0.045 
(1.95-2.22) 


3.42*0.084 
(3.05-3.64) 


0.613+0.0139 
(0.574-0 


.650) 


Yangasa Levu 
Sta. 30, 1934 
BPBM 167140 


5 70.50.50 
(70-71) 


1.97*0.036 
(1.85-2.05) 


3.21*0.053 
(3.08-3.34) 


0.614+0.0098 
(0.594-0 


.649) 


Nayau 
Sta. 101, 1938 
BPBM 180260 


4 76.3*6.18 
(67-88) 


1.90*0.109 
(1.62-2.12) 


3.2710.092 
(3.01-3.44) 


0.581+0.0205 
(0.538-0 


.616) 


irregularis 
















1 90 + 072 


2oc+ri HAR 


666*0 020 




Zurich 
BPBM 7982 


-] 


(1.71-2.20) 
1. 79*0. 058 


. o_)iU . U4o 
(2.67-3.08) 

2 78+0.041 


(0.613-0 
0. 643+0.014 


.764) 






(1.56-1.99) 


(2.65-2.98) 


(0.588-0 


.705) 


Sta. 133, BPBM 
179076, -9 2300- 
2500' 


18 


1.65*0.026 
(1.51-1.91) 


2.900.022 
(2.70-3.09) 


0.576+0.0091 
(0.528-0, 


.631) 


Sta. 118, BPBM 
178897 3000- 
3600' 


6 


1.71*0.043 
(1.61-1.91) 


2.9710.030 
(2.89-3.09) 


0.575+0.0016 
(0.544-0 


.617) 


Sta. 139, BPBM 
179157-8 2500- 
2900' 


2 


2.040.281 
(1.76-2.32) 


3.23+0.315 
(2.91-3.54) 


0.629+0.0262 
(0.602-0 


.655) 




WHORLS 


UMBILICUS 


D/U RATIO 


APICAL CORDS 




adposita 












Oneata 
Zurich 


4 l/4+(4-4 1/2) 


0.27*0.012 
(0. 23-0.33'" 


12.210.56 
(10.3-14.8) 







Sta. 100 


4 l/2-(4 1/4-4 3/4) 


0.30*0.021 
(0.20-0.36) 


11.410.78 
(8.82-16.7) 


13 




Munia 
Sta. 64 


4 3/8-(4-4 1/2) 


0.24*0.043 
(0.16-0.33) 


13.811.72 
(10.0-17.0) 







Lakemba 
Sta. 97 


4 5/8-K4 1/2-4 7/8) 

L 1/9 ( L I/A L *\ 1 $.\ 


0.30 
36 + 031 


11.010.17 
(10.7-11.3) 


13 




S ta. 98 


4 LI . \Q LI 44 I)/ o ) 


(0.30-0.46) 


9 . 6410 . 66 
(7.85-11.5) 






Yangasa 

c f. a on 


L T/ft ( L 1/R L 1/71 


21 + 080 


15 2*0 63 






b ta * JU 


H Jf Q \^ LI o 4 LI ) 


(0.20-0.23) 


(13.3-16.8) 






Nayau 
Sta. 101 


4 3/8-(4 1/4-4 1/2) 


0.31+0.095 
(0.30-0.33) 


10.410.19 
(10.1-11.0) 







irregularis 












Zurich 


3 7/8-(3 5/8-4) 


0.58*0.019 
(0.49-0.66) 


4.92+0.150 
(4.35-5.36) 







BPBM 7982 


3 5/8+(3 1/2-3 7/8) 


0.58*0.021 
(0.49-0.66) 


4.760.159 
(4.25-5.46) 


18.0+1.16 
(16-20) 




Sta 
Sta. 133 


3 3/4+(3 5/8-4) 


0.6910.013 
(0.59-0.79) 


4.2210.064 
(3.79-4.77) 







Sta. 118 


3 3/4-(3 1/4-4) 


0.680.025 
(0.59-0.76) 


4.3910.143 
(4.13-4.88) 







Sta. 139 


4 1/8(3 7/8-4 3/8) 


0.8110.116 
(0.69-0.92) 


4.0110.184 
(3.82-4.19) 








164 



SYSTEMATIC REVIEW 



165 



Sinployea irregularis (Garrett, 1887). Figures 65d-f, 
67a-d. 

Patula irregularis Garrett, 1887, Proc. Zool. Soc. London, 1887, p. 
179 Vitu Levu, Fiji Islands. 

Patula monstrosa Ancey, 1889, Le Naturaliste, 1889, p. 71- Vitu 
Levu, Fiji Islands. New name for irregularis Garrett, 1887, not 
Endodonta irregularis Semper, 1874; Pilsbry, 1892, Man. Con- 
chol., (2)8, p. 82. 

Endodonta (Charopa) monstrosa (Ancey), Pilsbry, 1893, Man. Con- 
chol., (2) 9, p. 35. Listed irregularis Garrett, 1887, as a synonym, 
assuming it to be preoccupied by Endodonta irregularis Semper, 
1874, now classified in Aaadonta. 

Charopa monstrosa Ancey, Gude, 1913, Proc. Malacol. Soc. Lon- 
don, 10 (5), p. 330, pi. 14, figs. 6a, b, c; Germain, 1932, Ann. Inst. 
Ocean., 12 (2), p. 45. 

Diagnosis. Shell of average size, diameter 2.50-3.52 mm. 
(mean 2.87 mm.), with 3'A-4% rather tightly coiled whorls. Apex and 
early spire flat to slightly elevated, lower spire descending moder- 
ately, body whorl often descending abruptly near aperture, spire 
protrusion variable, from 'Ao-% (usually Vs) body whorl width, H/D 
ratio 0.528-0.764 (mean 0.614). Apical sculpture of 16-20 (mean 
18.5) fine, prominent spiral cords, becoming sinuated on crossing 
radial swellings. Postnuclear sculpture of highly variable, irregu- 
larly spaced, low to high, protractively sinuated radial ribs, rarely 
regular and persistent enough to count. Microsculpture of fine radial 
riblets, finer and more crowded spiral riblets, and rather prominent, 
widely spaced secondary spiral cords. Umbilicus relatively narrow, 
V-shaped, last whorl regularly to more rapidly decoiling, contained 
3.79-6.83 times (mean 4.54) in the diameter, margins rounded. 
Whorls strongly rounded above and on outer margins, flattened to 
weakly channeled laterally above periphery. Aperture circular, in- 
clination varying from 10 to 30 depending on rate of body whorl 
descension. 

The flat spire, irregular ribbing, and often abrupt 
body whorl descension effectively separate typical 
examples of Sinployea irregularis from other Fijian 
species. Sinployea inermis (fig. 66a-e) is separable by 
its regular ribbing, lower apical cord count, and ele- 
vated spire. Sinployea rudis (fig. 47d-f) from 
Rarotonga, Cook Islands, is very similar in sculpture 
and whorl contour, but is much larger (mean diameter 
4.04 mm.), with only 8-12 apical cords, and has the 
apex distinctly elevated. Ba humbugi (fig. 74a-c) has 
the apex strongly elevated, few and very prominent 
apical cords, and the umbilicus a lateral crack or 
closed. 

Description. Shell of average size, with 3% moderately tightly 
coiled whorls. Apex and early spire fiat, last whorl descending 
abruptly, H/D ratio 0.623. Apical whorls l'/2, sculpture of about 16 
wavy, prominent spiral ribs, crossing low, broadly rounded radial 
ribs that are stronger on the last portion of apical whorls. Postapical 
sculpture of irregular, lamellate, protractively sinuated radial ribs, 
greatly reduced on body whorl. Microsculpture of fine, irregularly 
spaced, radial riblets, crossed by finer and more crowded spiral rib- 
lets, plus more prominent, equally spaced spiral cords. Sutures deep, 
whorls strongly rounded above, strongly flattened laterally above 
periphery. Umbilicus narrow, last whorl decoiling more rapidly, con- 
tained 4.25 times in the diameter. Color light yellow-brown, par- 
tially leached from shell. Aperture nearly circular, somewhat de- 
flected and flattened above periphery with protruded umbilical lip. 
Height of lectotype 1.74 mm., diameter 2.80 mm. 

Lectotype. Fiji: Viti Levu, ex Garrett, Museum 
Godeffroy. BPBM 7982. 

Range. Interior of Viti Levu, Fiji. 
Paratypes. Zurich, FMNH 116977, BPBM 7982. 



Material. Viti Levu (22 specimens, BPBM 
167434, FMNH 46376, FMNH 48996, FMNH 46439): 
Saganakoreva area back of Tubarua sawmill (Station 
VL-2, 1965), 5 miles in from Ngaloa, Nuku District, at 
950-1,000 ft. elevation (4 specimens, FMNH 153599, 
FMNH 153611); Mt. Korobamba (Station 60), 2 miles 
in at 1,000-1,300 ft. elevation (2 specimens, BPBM 
178759); Nandarivatu (Station 129), Nukunuku Creek, 
12 miles inland at 2,000 ft. elevation (1 specimen, 
BPBM 179031); Nandarivatu (Station 134), 1 mile east 
of Vatuthere, 12 miles inland at 2,000 ft. elevation (1 
specimen, BPBM 179085); Nandarivatu (Station 133), 
2 miles east of Vatuthere, 14 miles inland at 2,300- 
2,500 ft. elevation (31 specimens, BPBM 179076-9); 
Navai (Station 139), 1 mile west of Nandarivatu, 16 
miles inland at 2,500-2,900 ft. elevation (2 specimens, 
BPBM 179157-8); northwest of Nangaranambulata 
Mt. (Station 122), 10 miles inland at 2,700-3,200 ft. 
elevation (1 specimen, BPBM 178955); Nandarivatu 
(Station 118), 10 miles inland at 3,000-3,600 ft. eleva- 
tion (7 specimens, BPBM 178897-8); 500 ft. ridge (Sta- 
tion F-12) near main highway, Yarawa, southeast re- 
gion (12 specimens, FMNH 168309, FMNH 168311); 
near Tau Village (Station F-14), 2 miles inland at 500 
ft. elevation (1 specimen, FMNH 168336); Nan- 
darivatu (Station F-16), ridge slope at 2,500 ft. eleva- 
tion (2 specimens, FMNH 168363, FMNH 168382). 
Ovalau: upper Totoga Creek (Station F-10), in from 
Levuka (6 specimens, FMNH 168240, FMNH 168250). 
Remarks. We do not know where on Viti Levu 
the original specimens, presumably collected by Dr. 
Graeffe, were obtained. The types have a high H/D 
ratio produced by very sharp descension of the last 
whorl or whorl and one-half. Shells nearest to these, 
but distinctly less elevated and with a wider umbilicus 
(table XXXVII), were obtained by members of the 
Lapham Expedition at 2,500-2,900 ft. elevation near 
Navai lumber camp on the slopes of Mt. Victoria (Sta- 
tion 139). Shells from the vicinity of Nandarivatu (Sta- 
tions 118, 129, 133, 134) and northwest of Mt. Nan- 
garanambulata (Station 122) are much less elevated 
(table XXXVII) and have the radial sculpture greatly 
reduced in size. Shells from Saganakoreva also have 
relatively low H/D ratios and wide umbilici. The im- 
portance of these variations is unknown. The two 
specimens from near the top of Mt. Korobamba (Sta- 
tion 60) are as flat as the Nandarivatu shells, but the 
radial ribs are slightly more prominent and regularly 
spaced with very high periostracal extensions. Much 
more material from the interior of Viti Levu is needed 
in order to access the range of variation in S. ir- 
regularis. 

A single scalariform example, with the last third 
of the body whorl completely detached, was seen. It has 
not been included in the measured material. 

Description of soft parts. Foot and tail retracted in all available 
specimens, general appearance typical. 

Body color light yellow-white, no darker markings. 

Pallial region extending % whorl apically. Lung roof clear, with- 
out granulations. Kidney bilobed, maximum observed length 1.8 



166 



SOLEM: ENDODONTOID LAND SNAILS 



mm., rectal lobe distinctly longer than pericardial. Ureter typical, no 
lung roof visible between arms. Heart % length of pericardial kidney 
arm, lying parallel to hindgut. 

Ovotestis typical, occupying '/a whorl above stomach apex, one 
clump of acini lying parallel to whorl sides. Hermaphroditic duct 
typical. Albumen gland (fig. 67a, GG), talon (GT), and carrefour (X) 
without unusual features (fig. 67b). Prostate (DG) and uterus (UT) 
typical. 

Vas deferens (VD) very slender for entire length. Epiphallus (E) 
with head greatly swollen, tapering rapidly to a narrow neck enter- 
ing penial head lateral to insertion of penial retractor (PR), which is 
very short and arises from diaphragm. Penis (P) with upper section 
enormously swollen, about 0.8-1 mm. long, basal shaft slightly 
shorter and very narrow. Internally (fig. 67c-d) with vergic papilla 
greatly enlarged, stimulatory pad greatly reduced in prominence. 
Atrium (Y) long and narrow. 

Free oviduct (UV) wide, almost equal in diameter to penis. 
Spermatheca (S) with very slender shaft, until just before joining free 
oviduct, head elongately oval, slender. Vagina (V) tapering rapidly 
to atrium. 

(Based on BPBM 178897, 4 fragmentary examples.) 

Sinployea godeffroyana, new species. Figure 69a-c. 

Diagnosis. Shell very small, diameter 1.95-2.32 mm. (mean 
2.12 mm.), with 3%-4 normally coiled whorls. Apex and spire 
slightly and evenly elevated, last whorl descending slightly, spire 
protrusion about l h body whorl width, H/D ratio 0.557-0.619 (mean 
0.595). Apical whorls with 8-11 (mean 9.67) very fine spiral cords. 
Postnuclear sculpture of very fine, narrow, low, protractively 
sinuated radial ribs, 85-152 (mean 117.7) on the body whorl, whose 
interstices are 1-4 times their width. Ribs/mm. 20.99-38.03 (mean 
28.36). Microsculpture of very fine radial riblets, 2-5 between each 
pair of major ribs, crossed by much finer and more crowded spiral 
riblets, with a secondary sculpture of low and rather crowded spiral 
cording. Umbilicus narrow, V-shaped, regularly decoiling, contained 
4.11-6.30 times (mean 5.26) in the diameter, margins rounded. 
Whorl contours typical, strongly rounded above, flattened laterally 
above periphery and weakly flattened on basal margin, aperture 
inclined about 10 from shell axis. 

Small size and very crowded radial ribbing distin- 
guish Sinployea godeffroyana from other Fijian species. 
Sinployea angularis (fig. 64d-f) is similar in size but 
has less than 50 radial ribs, an angulated periphery, 
and markedly elevated spire. 

Description. Shell very small, with 3% moderately loosely 
coiled whorls. Apex and spire only slightly elevated, last whorl 
barely descending, H/D ratio 0.557. Apical whorls 1%, sculpture of 8 
widely spaced, relatively low spiral ribs. Remaining whorls with low, 
rounded, crowded, protractively sinuated radial ribs, 152 on the body 
whorl, whose interstices are 1-2 times their width. Microsculpture of 
2-5 radial riblets between each pair of major ribs, crossed by very 
low and inconspicuous crowded spiral ribs, with a secondary 
sculpture of narrow, widely spaced spiral cords. Sutures relatively 
shallow, whorls strongly rounded above, broadly flattened laterally 
above periphery and on base of shell. Umbilicus narrowly V-shaped, 
last whorl decoiling slightly more rapidly, contained 4.11 times in 
the diameter. Color light yellow-brown without darker flammula- 
tions. Aperture subquadrangular, flattened laterally above 
periphery and on basal margin, inclined about 10 from shell axis. 
Height of holotype 1.28 mm., diameter 2.30 mm. 

Holotype. Fiji: Viti Levu, Station 133, Nan- 
darivatu, 2 miles east of Vatuthere at 2,300-2,500 ft. 
elevation. Collected in damp forest by Yoshio Kondo on 
September 9, 1938. BPBM 179080. 

Range. Viti Levu and satellite islands, Fiji. 

Paratypes. Viti Levu: Mt. Korobamba (Station 
63), in dense forest at 1,000 ft. elevation (1 specimen, 



BPBM 178794); Nangava Island (Station 151), off 
south coast of Viti Levu (1 specimen, BPBM 179256). 

Remarks. The three specimens associated here 
as Sinployea godeffroyana agree in general shape and 
form, but diverge somewhat in sculpture. The shell 
from Nangava Island (BPBM 179256) has the sculp- 
ture much more widely spaced, and the umbilicus is 
much narrower, with the last whorl less expanded. The 
single specimen from Mt. Korobamba (BPBM 178794) 
was injured several times during its growth. The nar- 
rower body whorl and umbilicus may be a result of 
repaired injuries rather than indicating specifically 
important differences. 

Until more material is available I prefer to treat 
these as members of one species, although the Nan- 
gava Island shell may belong to a separate taxonomic 
entity. 

MELANESIAN Sinployea 

Without dissection of the Solomon and Bismarck 
species, their classification in Sinployea rather than in 
the most generalized group of Pilsbrycharopa (see 
Solem, 1970a) will be somewhat arbitrary. In general 
appearance these species have more similarities to 
Sinployea, but either classification could be correct. I 
have taken this opportunity to provide expanded diag- 
noses of the Bismarck Archipelago S. novopommerana 
and S. descendens, plus new descriptions of S. nissani 
and S. solomonensis. 

These species share monochrome coloration, 
rather tight coiling, wider than usual umbilici (except 
S. solomonensis), average ribbing, rather low H/D 
ratios, and are the smallest in average size for any area 
sampled (table XXXVIII). OnlyS. solomonensis andS. 
euryomphala even approach average size for the genus. 
The other four are among the seven smallest Sinployea 
species. Only the Fijian S. angularis and S. godef- 
froyana plus the Ellice Island S. ellicensis nukulae- 
laeana are slightly smaller. 

Dissection of the penial complex in S. euryomphala 
confirmed my impression that Sinployea extends into 
the New Hebrides. Dissection of some Andrefrancia 
from New Caledonia (unpublished data) show that 
they are generically distinct. The only dissected 
Pilsbrycharopa from New Guinea is not one of the 
generalized species, so the closeness of relationship is 
uncertain. 

Less detailed study has been made on this mate- 
rial, but a review is included here rather than making 
a separate report. Obviously collections are fragmen- 
tary and do not indicate actual distributions, but only 
the transitory passages of malacological collectors or 
friends of malacologists. Both Bismarck species came 
from river drift, although they may be sympatric. All 
three known Solomon Island species are allopatric, 
whereas the single New Hebridean species seems to be 
widely distributed in both the New Hebrides and 
Banks Group. Very probably, additional taxa will be 



SYSTEMATIC REVIEW 



167 



TABLE XXXVIII. - RANGE OF VARIATION IN MELANESIAN SINPLOYEA. 



NAME 



NUMBER OF 
SPECIMENS 



RIBS 



kuntzi 

(Solem) 

novopomme rana 
(Rensch) 

descendens 
(Rensch) 

solomonensis 
(Clapp) 

euryomphala 
(Solem) 



38 



90 



10 



87 



102.7(93-117) 
92.7(67-105) 
91.7(77-103) 

89 

74(72-78) 



RIBS/MM. 
12.85 



HEIGHT 
1.00 



15.0(13.64-16.18) 1.17(0.99-1.37) 
13.09(9.61-16.94) 1.18(0.99-1.61) 
12.93(11.13-14.65) 1.18(1.05-1.28) 



10.31 



1.38 



9.43(8.93-9.94) 1.38(1.28-1.48) 



DIAMETER H/D RATIO 

2.15 0.466 

2.18(2.06-2.30) 0.537(0.480-0.593) 

2.27(1.97-2.85) 0.519(0.467-0.567) 

2.27(2.14-2.53) 0.518(0.470-0.550) 



2.75 



0.503 



2.64(2.43-2.93) 0.523(0.494-0.535) 



APICAL 
CORDS 



SPIRE 
ELEVATION 



BODY WHORL 
WIDTH 



SP/BWW 



WHORLS UMBILICUS D/U RATIO 

nissa. 3 1/2 0.67 3.20 10 

kuntzi 3 3/4(3 1/2-4) 0.56(0.51-0.61) 3.91(3.68-4.26) 11.5(11-12) 0.12(0.10-0.15) 0.63(0.56-0.71) 0.191(0.176-0.209) 

novop. 3 3/4-(3 1/2-4 1/8) 0.65(0.53-0.92) 3.49(3.09-3.96) 10.8(10-12) 0.09(0.07-0.13) 0.69(0.61-0.76) 0.124(0.098-0.174) 

desce. 4 l/2-(4-4 1/2) 0.83(0.74-0.95) 2.73(2.43-2.94) 10.6(9-14) 0.04(0.03-0.05) 0.82(0.79-0.86) 0.049(0.038-0.063) 

solom. 4+ 0.64 4.29 12 

0.67(0.59-0.82) 3.97(3.56-4.67) 12.0(11-13) 0.11(0.10-0.13) 0.74(0.63-0.82) 0.149(0.120-0.170) 



euryo. 3 3/4+(3 1/2-4) 



recorded when more adequate collections have been 
made. 



(Solem, 1959). Figures 



Sinployea euryomphala 

67i-j, 70a-c. 

Mocella euryomphala Solem, 1959, Fieldiana: Zoology, 43 (1), pp. 
83-85, pi. 6, figs. 7-8 (radula and penis complex), pi. 31, figs. 
1-5 (shell) Brigstock Point, Espiritu Santo, New Hebrides; 
Solem, 1962, Bull. Brit. Mus. Nat. Hist., Zool., 9 (5), p. 226 
Norovorovo, Maewo, and Gaua, Banks Group, New Hebrides; 
Solem, 1963, Verhandl. Naturf. Ges., Basel, 74 (2), p. 163 Malo 
Island off Espiritu Santo, New Hebrides. 

Range. Espiritu Santo, Maewo, Gaua, Vanua 
Lava, New Hebrides. 

Material. Espiritu Santo: Brigstock Point (4 
specimens, MCZ 186828, FMNH 54906, FMNH 
109493). Maewo (1 specimen, FMNH 109384). Vanua 
Lava (5 specimens, AIM, AMS). 

Remarks. Use of the New Zealand genus Mocella 
as a repository for this species was based on a desire to 
avoid description of a new genus on inadequate data. 
Dissection of Mocella (unpublished data) shows that it 
is not closely related. 

Very few specimens have been seen from any 
single locality. I have not found any significant 
differences and doubt that any separation into local 
races has occurred. A description of the anatomy is 
presented below, but illustrations were prepared only 
for the penial complex (fig. 67i-j). 

Description of soft parts. Foot bluntly rounded anteriorly, 
slightly tapering and bluntly rounded posteriorly. Pedal grooves 
very high on foot, uniting over tail, suprapedal slightly weaker than 
pedal. No caudal foss, caudal horn, or middorsal groove. Foot sole 



undivided and without corrugations. Slime network finely textured, 
rectangular. 

Body color pale yellow in preservative, ommatophores black. 

Mantle collar short, thick, edge tapering. Pneumostome masked 
by a very small anterior left mantle lobe. No other accessory lobes or 
shell laps. Anus opening above and slightly anterior to external 
ureteric pore, which leads into a narrow V-groove, the urinary 
chamber, passing through pneumostome and mantle collar. 

Pallial region typical of subfamily. Kidney bilobed, rectal lobe '/;> 
longer than pericardia! and lapping onto hindgut. Ureter sigmure- 
throus, not expanded at any place, with strip of lung roof between 
arms. Hindgut continuing along parietal-palatal margin well apicad 
of pallial cavity. Heart elongated, slightly more than M> length of 
pericardial kidney lobe. Lung roof clear, without color patches. 

Ovotestis a single clump of finger-like acini buried in digestive 
gland well above stomach-intestine reflexion. Hermaphroditic duct 
an iridescent muscular tube, greenish in tone, narrowing at base of 
albumen gland, then running up to enter duct of talon at midpoint. 
Head of talon a white bulb, duct slender, opening directly into 
prostate-uterus head. Latter area torn in dissection, greatly distorted 
because of contraction, not worked out as to detailed structure. 

Vas deferens a very slender tube passing up along lower half of 
penis, entering expanded tube of epiphallus above midpoint of penis. 
Epiphallus (fig. 67i, E) an expanded tube with clearly defined lon- 
gitudinal pilasters, reflexed, with penial retractor (PR) inserting 
below point of reflexion (? artifact of contraction) and arising on 
diaphragm. Penis (P) showing two bulges externally. Apical portion 
with tiny verge (PV), the walls immediately below expanded to form 
a glandular collar (fig. 67j), lower part with a large glandular 
stimulatory papilla (PP) and a peculiar U-shaped pocket (PC) with 
thin muscular walls. Base of penis narrowing to atrium (Y) with 
weak pilasters present. 

Free oviduct muscular, thick, with terminal glandular collar 
above junction with spermatheca. Latter with grossly expanded, 
thin-walled glandular base, narrowing to a very fine tube about mid- 
point of free oviduct. Position of spermathecal head not observed. 
Vagina very short and thin-walled. 



168 



SOLEM: ENDODONTOID LAND SNAILS 






Digestive and free muscle system not studied. Buccal mass lost 
in dissection. 

The single specimen available for dissection was retracted far 
within the shell and proved quite difficult to dissect. Destruction of 
the shell was mandatory, although the fragments have been saved. 
Diameter of the specimen was 2.27 mm. Radular elements of this 
species were figured previously (Solem, 1959a, pi. 6, fig. 7). 



1923). Figure 



FIG. 70. a-c, Sinployea euryomphala (Solem). Holotype of 
Mocella euryomphala Solem. Under logs at Brigstock Point, Espiritu 
Santo, New Hebrides. UMMZ 186042. Scale line equals 1 mm. 
(Solem, 1959a, pi. 31, fig. 1-3). (EJP). 



Sinployea solomonensis (Clapp, 
71a-e. 

Endodonta (Charopa ) solomonensis Clapp, 1923, Bull. Mus. Comp. 

Zool., Harvard, 65, p. 378, figs. 21-24 Ugi, Solomon Islands. 
Mocella solomonensis (Clapp), Solem, 1959, Fieldiana: Zoology, 43, 

p. 83: Solem, 1960, J. Malacol. Soc. Australia, 4, p. 41, pi. 4, figs. 

1-5. 

Description. Shell of average size, with a little more than 4 
normally coiled whorls. Apex and early spire slightly emergent, body 
whorl descending a little more rapidly, H/D ratio 0.503. Apical 
whorls 1%, sculpture of 12 rather small spiral cords. Postnuclear 
whorls with prominent, rounded, somewhat protractively sinuated 
radial ribs, 89 on the body whorl, whose interstices are 2-4 times 
their width. Ribs/mm. 10.31. Microsculpture mostly eroded, in places 
visible as a lattice of fine radial riblets crossed by even finer and 
more crowded spiral riblets. Sutures deep, whorls somewhat shoul- 
dered above, flattened laterally above periphery with slightly flat- 
tened basal margin. Color light reddish yellow-horn without darker 
markings. Umbilicus V-shaped, regularly decoiling, contained 4.29 
times in the diameter. Aperture subcircular, slightly flattened later- 
ally above periphery, inclined about 15 from the shell axis. No aper- 
tural barriers. Height of holotype 1.38 mm., diameter 2.75 mm. 

Holotype. Solomon Islands: Ugi Island. Collected 
by William Mann. MCZ 36838. 

Range. Ugi, Solomon Islands. 

Remarks. A proportionately thinner body whorl 
and slightly more crowded radial ribbing are the most 
obvious features separating this species from Sinployea 
euryomphala. 

Sinployea kuntzi (Solem, 1960). Figure 72a-c. 

Mocella kuntzi Solem, 1960, J. Malacol. Soc. Australia, 4, pp. 
41-42, pi. 5, A, figs. 1-3 Station FLW 38, peninsula east of 
Halavo, Florida Island, Solomon Islands. 

Range. Florida Island, Solomon Islands. 

Material. Florida Island: peninsula east of 
Halavo (5 specimens, FMNH 54905, MCZ 186827). 

Remarks. Data on variation are presented in 
Solem (1960). Measurements in Table XXXVIII are 
based upon material measured recently and thus differ 
slightly from the data recorded in the original descrip- 
tion. Differences from S. nissani are covered in the re- 
marks under that species. 

Sinployea nissani (Dell, 1955). 

Gyropena nissani Dell, 1955, Pacific Science, 9, p. 328, fig. 1, e- 
g near Tangalan Plantation, Nissan Id., Northern Solomons. 

Mocella nissani (Dell), Solem, 1960, J. Malacol. Soc. Australia, 4, 
pp. 41, 43. 

Description. Shell very small, with 3'/2 moderately loosely 
coiled whorls. Apex barely emergent, spire flat, last whorl not de- 
scending, H/D ratio 0.466. Apical whorls 1%, sculpture partially 
eroded with traces of about 10 relatively widely spaced spiral cords 
remaining. Postnuclear whorls with prominent, rounded, protrac- 
tively sinuated radial ribs, 87 on the body whorl, whose interstices 








FIG. 71. Sinployea solomonensis (Clapp). Holotype of Champa solomonensis Clapp. Ugi Island, Solomon Islands. MCZ 36838: a-c, entire 
shell; d, detail of partly worn apical sculpture, greatly enlarged; e, detail of microsculpture, greatly enlarged, absence from tops of major ribs 
caused by mechanical wear. Scale line equals 1 mm. (Solem, 1960, p. 42, pi. 4). (HJW). 



169 



170 



SOLEM: ENDODONTOID LAND SNAILS 





-- 




FIG. 72. a-c, Sinployea kuntzi (Solem). Holotype of Mocella 
kuntzi Solem. From jungle one mile behind Halavo, Florida Island, 
Solomon Islands. UMMZ 184473. Scale line equals 1 mm. (Solem, 
1960, p. 44, pi. 5, fig. A). (EJP). 



are 2-4 times their width. Ribs/mm. 12.85. Microsculpture of fine 
radial riblets, 4-7 between each pair of major ribs, with much finer, 
more crowded spiral riblets and a relatively prominent secondary 
sculpture of widely spaced spiral cording. Sutures deep, whorls 
evenly rounded above, slightly flattened laterally above periphery 
with gently rounded basal margin. Color uniform yellowish brown, 
slightly faded. Umbilicus widely open, broadly U-shaped, regularly 
decoiling, contained 3.20 times in the diameter. Aperture ovate, lip 
badly broken. Height of holotype 1.01 mm., diameter 2.16 mm. 

Holotype. Solomon Islands: Nissan Island, near 
Tangalan Plantation. Collected in leaf mould by R. K. 
Dell in June, 1944. DMW 2513. 

Range. Nissan Island, Solomon Islands. 

Remarks. As indicated by Solem (1960), 
Sinployea nissani is quite closely related to M. kuntzi 
from Florida Island. The latter is a distinctly higher 
shell (mean height 1.17 mm.), with much narrower 
umbilicus (mean D/U ratio 3.91). Even though the type 
of S. nissani is subadult, the umbilical width is greater 
than inS. kuntzi. Although adult descension of the last 
part of the body whorl would have increased the H/D 
ratio of S. nissani, presumably the more rapid decoil- 
ing of the last umbilical section would have further 
widened the umbilical opening, increasing the 
difference. They are distinct species. 

Sinployea novopommerana (I. Rensch, 1937). Fig- 
ure 73a-c. 

Charopa novopommerana I. Rensch, 1937, Arch. f. Naturgesch., 
n.f., 6 (4), pp. 589-590, figs. 33-34 (shell and radular teeth) 
Karlei, Malkong-Bach, and Nangurup, New Britain, Bismarck 
Archipelago. 

Diagnosis. Shell very small, diameter 1.97-2.85 mm. (mean 
2.27 mm.), with 3'/fe-4Vfe normally coiled whorls. Apex and early spire 
slightly elevated, rounded above, body whorl descending much more 
rapidly, spire protrusion about VH body whorl width, H/D ratio 
0.467-0.567 (mean 0.519). Apical sculpture of 10-12 (mean 10.8) 
very fine spiral cords. Postnuclear whorls with prominent, rounded, 
protractively sinuated radial ribs, 67-105 (mean 92.7) on the body 
whorl, whose interstices are 2-4 times their width. Ribs/mm. 9.61- 
16.94 (mean 13.09). Microsculpture of fine radial riblets, 5-10 be- 
tween each pair of major ribs, slightly finer and more crowded spiral 
riblets, plus rather crowded secondary spiral cording that is slightly 
more prominent than microradials. Umbilicus moderately open, 
V-shaped, last whorl decoiling more rapidly, contained 3.09-3.96 
times (mean 3.49) in the diameter, margins rounded. Sutures very 
deep, whorls strongly rounded above, flattened laterally above 
rounded periphery, with evenly rounded, slightly compressed, basal 
margin. Color light reddish yellow-horn without darker flammula- 
tions. Aperture subcircular, flattened laterally above periphery, in- 
clined about 30 from shell axis. 

Sinployea novopommerana has a much less com- 
pressed body whorl and narrower umbilicus than S. 
descendens (fig. 73d-f). Sinployea kuntzi has a much 
more elevated spire, narrower umbilicus, and less 
marked lateral flattening above the periphery (fig. 
72a-c). 

Range. New Britain, Bismarck Archipelago. 

Material. New Britain: Karlei (34 specimens, 
ZMB, FMNH 146028); Malkong (4 specimens, ZMB). 

Remarks. The slightly elevated apex and rather 
narrow and V-shaped umbilicus immediately separate 






FIG. 73. a-c,Sinployea novopommerana (Rensch). Karlei, Weite Bucht, New Britain, Bismarck Archipelago. Paratype. FMNH 146028; d-f, 
Sinployea descendens (Rensch). Karlei, Weite Bucht, New Britain, Bismarck Archipelago. Paratype. FMNH 146029. Scale line equals 1 mm. 

(MM). 



171 



172 



SOLEM: ENDODONTOID LAND SNAILS 



Sinployea novopommerana (fig. 73a-c) from the sym- 
patric S. descendens (fig. 73d- f). The Solomon Islands. 
solomonensis (Clapp, 1923) appears quite similar but is 
larger, has a slightly narrower umbilicus, the whorls 
are less flattened laterally above the periphery, and 
the whorl count is slightly higher (fig. 71a-e). 
Sinployea nissani (Dell, 1955) and S. kuntzi (Solem, 
1958) are smaller, much more tightly coiled shells. 

Most of the available specimens were not quite 
fully adult, so that the mean diameter recorded in the 
diagnosis may be too small. The range is greatly ex- 
tended in regard to height, diameter, and umbilical 
width by the presence of one very large individual. 
Without this specimen, the maximum height would be 
1.33 mm., maximum diameter 2.47 mm., and 
maximum umbilical width 0.74 mm. If this specimen is 
a freak, there would be a more marked size differential 
between S. novopommerana and S. euryomphala. 

Sinployea descendens (I. Rensch, 1937). Figure 
73d-f. 

Champa descendens I. Rensch, 1937, Arch. f. Naturgesch., n. f., 6 
(4), p. 590, fig. 35 Karlei, Malkong, and Matong, New Britain, 
Bismarck Archipelago. 

Diagnosis. Shell very small, diameter 2.14-2.53 mm. (mean 
2.27 mm.), with 4-4Vi rather tightly coiled whorls. Apex and early 
spire flat or slightly depressed below level of penultimate and first 
half of body whorl, last half of body whorl descending markedly, total 
spire protrusion about l ko of body whorl width, H/D ratio 0.470-0.550 
(mean 0.518). Apical sculpture of 9-14 (mean 10.6) prominent, regu- 
larly spaced spiral cords, whose interstices are less than twice their 
width. Postnuclear sculpture of prominent, high, rounded, rather 
crowded, slightly protractively sinuated radial ribs. 77-103 (mean 
91.7) on the body whorl, whose interstices are 2-4 times their width. 
Ribs/mm. 11.13-14.95 (mean 12.93). Microsculpture of fine radial 
riblets, 5-8 between each pair of major ribs, finer spiral riblets, and 
narrow, quite crowded, secondary spiral cords. Umbilicus broadly 
open, cup-shaped, regularly decoiling, contained 2.43-2.94 times 
(mean 2.73) in the diameter, margins slightly shouldered. Sutures 
deep, whorls shouldered above and on basal margin, evenly rounded 
on columellar margin, strongly compressed laterally above rounded 
periphery. Color reddish horn fading to yellow-white in drift mate- 
rial, no flammulations. Aperture suboval, sharply rounded above 
and at basal margin, evenly rounded laterally, inclined about 10 
from shell axis. 

Sinployea descendens is characterized by its flat 
spire, widely open cup-shaped umbilicus, and thick 
body whorl. The sympatric S. novopommerana (fig. 
73a-c) has a much narrower umbilicus, thinner body 
whorl, distinctly elevated spire, and averages l /z whorl 
less at identical size. Sinployea irregularis (fig. 65d-f) 
from Fiji is very similar in appearance, but is much 
larger, has reduced ribbing, and a narrow umbilicus. 

Range. New Britain, Bismarck Archipelago. 

Material. New Britain: Karlei (76 specimens, 
ZMB, FMNH 146029); Matong (14 specimens, ZMB). 

Remarks. Both Sinployea nissani and S. kuntzi 
are more narrowly umbilicated, have a protruding 
spire, and average one-half whorl less. Size variation 
in the two samples is summarized in Table XXXVIII. 
These were all drift specimens and undoubtedly origi- 
nated from several populations. 



Genus Ba, new genus 

Shell of less than average size with only S'/s-SM; tightly coiled 
whorls. Apex and spire quite elevated, body whorl descending 
rapidly. Apical sculpture of about 12 very prominent spiral cords. 
Postnuclear sculpture reduced to irregularity. Umbilicus closed or 
barely open. Aperture without barriers. Pallial region with mar- 
kedly deflected hindgut and kidney, latter squarish with tightly 
compacted ureter arms. Ovotestis with one acinal clump, hermaph- 
roditic duct short. Prostate-uterine region folded and compacted. 
Penis with unusually complicated penial retractor insertion. Interior 
of penis with large vergic papilla and circular muscle band, pocket 
stimulator greatly reduced in size. 

Type species. Ba humbugi, new species. 

In pattern of sculpture and general appearance, So 
humbugi is extremely similar to the sympatric 
Sinployea irregularis (Garrett). The major conchologi- 
cal differences are the extreme spire elevation, marked 
whorl reduction, and umbilical closure of Ba humbugi. 
Anatomically, the compaction and deflection of the 
kidney and hindgut (fig. 75a) represent a major depar- 
ture from the Charopidae pattern. Similar whorl re- 
duction in the Caroline Island Russatus (fig. 90a) re- 
quired only compaction into a square kidney without 
deflection of either kidney or hindgut from the 
parietal-palatal margin. The New Guinea endemic 
Paryphantopsis (Solem, 1970a, p. 250, fig. 2, a) agrees 
with Ba in showing kidney and hindgut deflection, but 
the pattern of the kidney is quite different in that the 
ureter arms become spread progressively further 
apart. These are different solutions to the same prob- 
lem of visceral hump compaction. Also part of this 
problem is the alteration in penial retractor muscle 
insertion. 

The functioning penial surface structures of Ba 
humbugi show simple character displacement from the 
equivalent structures in Sinployea irregularis, thus 
indicating phyletic closeness. Yet Ba humbugi is the 
only known member of the Pacific Island Charopinae 
to experiment with whorl reduction and visceral hump 
compaction. Because it represents a basic departure 
from the Sinployea pattern, I have chosen to give 
generic level recognition despite its evident relation- 
ships to S. irregularis. 

The only known species, Ba humbugi, has been 
found in Central Viti Levu from Mt. Nangaranam- 
bulata south to a few miles inland from Ngaloa. At the 
latter locality it was found under the same log as 
Sinployea irregularis. Because it occupies a portion of 
Viti Levu that includes the Mba District, I have chosen 
the European spelling of the district for a generic 
name. This was followed by an irresistible impulse to 
use the specific name humbugi. 



Ba humbugi, new species. Figures 74a-c, 75a-h. 

Diagnosis. Shell of slightly less than average size, diameter 
2.30-3.32 mm. (mean 2.72 mm.), with S'/s-S'/i tightly coiled whorls. 
Apex and spire strongly and evenly elevated, body whorl descending 
more rapidly, spire protrusion M)-M> body whorl width, H/D ratio 
0.752-0.842 (mean 0.801). Apical sculpture of 11-13 (mean 12.0) 
high and prominent spiral cords. Postnuclear sculpture of irregular 
growth wrinkles and an occasional identifiable major rib with inter- 



SYSTEMATIC REVIEW 



173 




FIG. 74. a-c,Ba humbugi, new species. Mt. Nangaranambulata, 
2,700-3,200 ft, Viti Levu, Fiji. Holotype. BPBM 178954. Scale line 
equals 1 mm. (MM). 



gradation between these extremes. Microsculpture obscured by sur- 
face irregularities, visible in some areas as consisting of fine radial 
riblets, finer and more crowded spiral riblets, plus low and broadly 
rounded, much larger secondary spiral cords. Umbilicus either im- 
perforate (25%) or with a small lateral crack (75%), possibly latter 
developing in adult stage only, since smallest example had closed 
umbilicus. Sutures deep except near end of body whorl, aperture and 
body whorl flattened laterally above periphery, inclined about 25 
from shell axis. 

Ba humbugi is immediately recognizable by its 
very high spire, low whorl count, reduced radial rib- 
bing, and closed or laterally cracked umbilicus. Other 
Polynesian species with closed or nearly closed um- 
bilici include the Samoan S. clausa and S. clista (fig. 
51c, f) and the Lau Archipelago S. adposita (fig. 69f). 
These species all have prominent radial ribbing and 
higher whorl counts. The Lord Howe Island Mystivagor 
mastersi (Brazier, 1872) is similar in shape and whorl 
count but has vastly different sculpture and anatomy. 

Description. Shell rather small, with 3'/2 relatively tightly 
coiled whorls. Apex and spire markedly and evenly elevated, body 
whorl descending more rapidly, H/D ratio 0.810. Apical whorls I'/z, 
sculpture of 11 very prominent spiral cords. Postnuclear whorls with 
very irregular growth wrinkles and occasional major ribs with high 
periostracal extensions. Microsculpture a lattice of fine radial and 
finer spiral riblets, irregular in spacing, crossed by more prominent 
spiral cords. Sutures deep, whorls flatly rounded above, evenly 
rounded below periphery. Umbilicus closed by expansion of basal lip 
and tight coiling, with only a slight lateral chink visible. Color light 
reddish yellow with dark brown, almost black, periostracal exten- 
sions. Aperture ovate, flattened laterally above periphery, inclined 
about 25 from shell axis. Height of holotype 2.24 mm., diameter 2.76 
mm. 

Holotype. Fiji: Viti Levu, Station 122, Mt. Nan- 
garanambulata at 2,700-3,200 ft. elevation. Collected 
in dense forest by Yoshio Kondo on September 5, 1938. 
BPBM 178954. 

Range. Interior of Viti Levu at 950-3,200 ft. ele- 
vation, Fiji. 

Paratypes. Viti Levu: top of Mt. Korobamba (Sta- 
tion 60) at 1,000-1,300 ft. elevation (2 specimens, 
BPBM 178758); Sanganakoreva area (Station VL-2, 
1965), back of Tubarua saw mill, 950-1,000 ft. eleva- 
tion, 5 miles inland from Ngaloa, Nuku District (1 
specimen, FMNH 153605). 

Remarks. The closed umbilicus, irregular rib- 
bing, and very high spire of Ba humbugi immediately 
separate it from any Pacific Island endodontid. In 
many respects, the shell recalls that of the New 
Caledonian Rhytidopsis minutula (Crosse, 1870) (see 
Solem, 1961, p. 466). Until the latter can be dissected, 
the degree of relationship will remain uncertain. The 
Lord Howe Island species Mystivagor mastersi 
(Brazier, 1872) has gone further toward a succineiform 
shell. It differs in size (6 mm. diameter), strong color 
pattern, smooth shell surface, and apical sculpture of 
widely spaced radial ribs. 

Spire angle varied greatly among the four speci- 
mens, with the one from Saganakoreva (FMNH 
153605) having a much wider angle and lower H/D 
ratio (0.752). It was found under the same log as an 
example of Sinployea irregularis, so that the two 



174 



SOLEM: ENDODONTOID LAND SNAILS 



species are unquestionably sympatric. Lack of any 
black markings on the body probably indicates this is a 
strictly terrestrial species. 

Dissection of two individuals showed major 
changes in the pallial systems that resulted in generic 
recognition. Changes in the genitalia involved folding 
of the prostate-uterus, compaction and broadening of 
the albumen gland, shortening of the hermaphroditic 
duct, altering the simple clump insertion of the penial 
retractor into a complex (fig. 75d-e), enfolding of the 
penis-epiphallus junction, and massive reduction of the 
penial pocket stimulator (fig. 75h). Except for the lat- 
ter change, these are adjustments to the whorl reduc- 
tion and visceral hump compaction. 

Comparison of penial structures in S. irregularis 
(fig. 67c-d) with those of Ba humbugi (fig. 75h) shows 
enlargement of the pocket stimulator in the first and 
reduction in the second. 

A single sperm packet (fig. 75f-g) was taken from 
FNMH 153605, which was collected December 9, 1965, 
the morning after a shower ending a long drought. 

Description of soft parts. Foot and tail equal in length to shell 
diameter. Sole undivided, bluntly rounded and slightly tapering 
posteriorly, truncated anteriorly. Pedal grooves very conspicuous, 
high on foot, uniting above tail, no caudal horn or middorsal groove 
developed. Slime network of large, rectangular sections on tail and 
sides of foot. Head retracted in all individuals examined. Gonopore 
position normal. 

Body color yellow-white, no darker markings. 

Mantle collar (MC) with thickened edge, no glandular extension 
onto pallial roof. Anus (A) opening next to external ureteric pore 
(KX), just inside pneumostome. 

Pallia! region (fig. 75a) extending V 2 whorl apically. Lung roof 
clear, without granulations. Kidney (K) about 1.58 mm. long, 
bilobed, deflected downward from parietal-palatal margin after 0.33 
mm., lobes subequal, pericardial slightly longer than rectal, trun- 
cated anteriorly by ureter. Ureter (KD) compressed between kidney 
lobes, apical part curved downward around tip of kidney, no lung 
surface visible between kidney lobes. Heart (H) % length of kidney, 
lying parallel to deflected plane of hindgut and parallel to lower edge 
of mantle collar. Principal pulmonary vein (HV) very short and in- 
conspicuous, curving around edge of kidney. Hindgut (HG) deflected 
from parietal-palatal margin shortly after kidney apex, intestinal 
loop (dotted line in fig. 75a) partly under heart. 

Ovotestis (fig. 75c, G) occupying V-i whorl above stomach apex, a 
single, tightly compacted clump of palmately clavate acini with short 
collecting tubule. Hermaphroditic duct (GD) kinked initially, nar- 
row, gradually tapering to past middle, grossly expanded, then 
sharply constricting before reflexing up to talon. Albumen gland 
(GG) with highly irregular surface, proportionately large, acini rela- 
tively small. Talon (GT) with globular head, short neck to carrefour, 
both buried in albumen gland. Prostate (DG) with a few long acini 
inserting onto side of uterus, which is bifolded. Uterus (UT) bipar- 
tite, folded upon itself, lower chamber enlarged and with thick, glan- 
dular walls. 

Vas deferens (VD) with thick, glandular walls to penioviducal 
angle, narrowed abruptly and a thin-walled tube to epiphallus, 
loosely bound to penioviducal angle. Epiphallus (E) % length of 
penis, head bulbous, internally with Y valve and longitudinal pilas- 
ters. Folded against head of penis at point of insertion. Penial retrac- 
tor (PR) arising from diaphragm, about Mi length of penis, inserting 
in a broad fan around epiphallus and head of penis (fig. 75d-e), with 
separate strands to penis apex and inner wall of epiphallus. Penis (P) 
club-shaped, about 1.8 mm. long, tapering gradually to atrium, in- 
ternally (fig. 75h) with a bulbous vergic papilla, greatly enlarged 
circular muscle band, and tiny stimulatory pocket pilaster. Lower 



portion with weak glandular ridges into atrium. Atrium (Y) short, 
without unusual features. 

Free oviduct (UV) only slightly longer than vagina, equal in 
diameter to penis at midsection, internally with typical circular 
pilaster at spermathecal junction. Spermatheca (S) with expanded 
head next to albumen gland, slender shaft next to prostate, base 
section abruptly expanding at base of uterus, internally with typical 
glandular pilaster and a larger sperm packet (fig. 75f-g, SP). Vagina 
(V) thin-walled, same diameter as free oviduct, internally with weak 
glandular pilasters. 

Free muscle system typical, somewhat shortened. Right om- 
matophoral retractor passing through penioviducal angle, right 
rhinophoral retractor uniting before tail fan junction. 

Stomach occupying % of a whorl, starting ] Ae of a whorl above 
pallial cavity apex. Intestinal looping compacted into narrow zone 
and protruding partly into pallial cavity area. Hindgut deflected 
from parietal-palatal margin for half of length, reaching margin 
near anterior end of kidney. 

Digestive glands extending past ovotestis to apex, mainly along 
stomach, only one strand near albumen gland. Salivary glands short, 
not uniting above esophagus. 

(Based on FMNH 153605 and BPBM 178954, 2 adult specimens.) 



Genus Maafu, new genus 

Shell relatively large, with 4-4V> normally coiled whorls. Apex 
and spire flatly coiled, depressed below level of postnuclear sub- 
sutural keel. Periphery strongly protruded into a cordlike keel with 
prominent supra- and subperipheral sulci. Umbilicus broadly open, 
cup-shaped, regularly decoiling, contained about 3 times in the 
diameter. Apical sculpture with typical spiral cords. Postnuclear 
sculpture with prominent, widely spaced radial ribs that are en- 
larged on crossing periphery, regular microsculpture reduced, secon- 
dary spiral cording narrow, rather prominent, and quite crowded. No 
apertural barriers. Anatomy unknown. 

Type species. Maafu thaumasius, new species. 

Although the form of Maafu thaumasius is par- 
tially duplicated by such Endodontidae as Austral- 
donta magnasulcata (Solem, 1976b, p. 295, fig. 
127a-b), several Nesodiscus, Endodonta, and many 
juvenile Gambiodonta, no Pacific Island Charopidae 
have a protruded keel. Only Sinployea angularis (fig. 
64e) from Namuka, Lau Group, andHimeroconcha (fig. 
105b, e) from the Marianas have an angulated 
periphery. 

Most of the greatly altered appearance seen in 
Maafu thaumasius relates to the two keels. The sub- 
sutural keel changes what would be a flat-spired shell 
into one that has a depressed apex and early spire. The 
extreme protrusion of the peripheral keel increases the 
diameter and decreases the H/D ratio. Hence, the com- 
paratively large size (mean diameter 3.49 mm.) and 
very low H/D ratio (mean 0.365) are secondary modifi- 
cations resulting from the keels. Efforts to assess the 
relationships of M. thaumasius must discount the im- 
portance of these size and shape shifts. The micro- 
sculpture of Sinployea inermis meridionalis and S. i. 
lakembana is visually very close to that of M. 
thaumasius. Only the greater crowding and emphasis 
of the secondary spiral cording distinguish the latter in 
a gross sense. The pattern of coiling, umbilical shape, 
and major ribbing is very different, but derivation of 
Maafu from the generalized Sinployea inermis-S. prin- 
cei stock would present no insuperable difficulties. 




Fig. 75. Anatomy of the Fijian Ba humbugi, new species. Station VL-2, Tubarua, Viti Levu, Fiji. FMNH 153605: a, pallial region; b, 
genitalia; c, ovotestis and hermaphroditic duct; d-e, insertion details of penial retractor (PR) on epiphallus-penis junction; f-g, spermatophore; 
h, interior of apical penis region. Scale lines equal 1 mm. (SH). 



175 



176 



SOLEM: ENDODONTOID LAND SNAILS 



Besides the keels, the alteration in ribbing pat- 
tern, very broad and regularly decoiling umbilicus, and 
flat spire represent departures from the Sinployea pat- 
tern. 

Thus, generic recognition for Maafu is based on 
several shifts that combine to produce a drastically al- 
tered appearance. Local differentiation from a 
Sinployea-type stock is almost certain, but the 
structural gap is real and large. 

I have chosen to honor the great Tongan leader 
Ma'afu, who unified the Lau Group in the 1850- 1860s, 
by naming this genus after him. The specific name 
thaumasius refers to the most unusual and wonderful 
shape of the shell. 

Maafu thaumasius, new species. Figure 76a-c. 

Diagnosis. Shell relatively large, diameter 3.19-3.98 mm. 
(mean 3.49 mm.), with 3'/2-4'/8 normally coiled whorls. Apex and 
spire depressed below level of subsutural keel, body whorl not de- 
scending or rarely very slightly deflected, H/D ratio 0.343-0.381 
(mean 0.365). Apical sculpture of 10-15 (mean 12.2) narrow but 
prominent spiral cords. Postnuclear sculpture of high, broadly 
rounded, rather widely spaced radial ribs, 36-55 (mean 44.1) on the 
body whorl, whose interstices are about 2-5 times their width, and 
which become higher and more sharply defined on crossing keels. 
Ribs/mm. 3.59-4.74 (mean 4.08). Microsculpture of extremely fine 
radial riblets, barely visible at 96 x magnification and too obscured 
by the secondary spiral cording to be counted accurately, equally fine 
spiral riblets, and much more prominent secondary spiral cording, 
whose interstices are 1-3 times their width. Umbilicus broadly open, 
cup-shaped, regularly decoiling, contained 2.82-3.15 times (mean 
2.99) in the diameter, columellar-basal margin obtusely shouldered. 
Sutures deep, whorls strongly rounded above, after nucleus with a 
gradually increasing subsutural keel followed by a deep supra- 
peripheral sulcus, periphery a sharply protruded cordlike keel, a 
broader and shallower subperipheral sulcus, basal margin rounded 
to an obtusely shouldered columellar-basal angle, walls of umbilicus 
evenly rounded. 

Although the form of Maafu thaumasius recalls 
that of the Navutu-i-Loma endodontid Zyzzyxdonta 
alata Solem (1976b, p. 466, fig. 198a-c), the larger size, 
beaded and numerous apertural barriers, "winglike" 
rib extensions, flat spire, and radial apical sculpture of 
the latter at once separate the two species. No Polyne- 
sian Charopidae have an even slightly similar shape, 
and M. thaumasius cannot be confused with any other 
species. 

Description. Shell relatively large, with slightly less than 4% 
moderately tightly coiled whorls. Apex and early spire depressed 
below plane of subsutural keel on succeeding whorls, last whorl not 
descending, H/D ratio 0.378. Sutures shallow, slightly denticulated 
by radial ribbing. Postapical whorls with a high subsutural ridge, 
becoming slightly less prominent near end of body whorl. Periphery 
of body whorl greatly protruded into a ridgelike keel, denticulated by 
crossing of major ribs, with a very prominent sulcus above and a less 
prominent one below periphery. Embryonic whorls 1%, sculpture of 
about 12 very fine, closely spaced spiral ribs (partially eroded). Post- 
nuclear whorls with widely spaced, low, rounded, protractively 
sinuated radial ribs, 50 on the body whorl, that denticulate the 
periphery and whose interstices are 3-6 times their width. Micro- 
sculpture on early whorls of very fine radial riblets, crossed by 
slightly stronger secondary spiral cords. Microspiral riblets occa- 
sionally visible. On later whorls the radials become obsolete and the 
spirals slightly more prominent. Umbilicus broadly open, cup- 






FIG. 76. a-c, Maafu thaumasius, new species. Station 43, Nayau, 
Lau, Fiji. Holotype. BPBM 167233. (MM). 



SYSTEMATIC REVIEW 



177 



shaped, regularly decoiling, contained 2.98 times in the diameter. 
Color mainly leached from shell. Aperture subquadrangular with 
concave outer margins and rostrate periphery, inclined about 20 
from the shell axis. Height of holotype 1.48 mm., diameter 3.92 mm. 

Holotype. Fiji: Lau Group, Nayau, Station 43, 
hillside '/2 mile inland of Nauko at 250 ft. elevation. 
Collected at base of limestone cliff by H. S. Ladd on 
August 13, 1934. BPBM 167233. 

Range. Nayau, Lau Group, Fiji. 

Paratypes. Nayau, hillside 2 mile inland of 
Nauko (Station 43) at 250 ft. elevation (64 specimens, 
BPBM 167233). 

Remarks. The very striking subsutural ridge 
and peripheral keel immediately separate Maafu 
thaumasius from all other species of Polynesian 
Charopidae. It is most apt to be confused with Zyzzyx- 
donta alata, a species of Endodontidae from Navutu-i- 
Loma that differs in having radial apical sculpture and 
apertural barriers of strong lamellae that are regularly 
beaded above. In addition the major ribs of Z. alata are 
prolonged into cup-shaped winglike structures when 
crossing the periphery (Solem, 1976b, p. 466, fig. 
198a-c). The ribs in M. thaumasius only slightly den- 
ticulate the keel (fig. 76b). 

Only 20% of the specimens were adult. Much of the 
material was fresh, but no specimens were collected 
alive. 

Genus Lauopa, new genus 

Shell extremely large, with about 4% normally coiled whorls. 
Apex and spire flat or slightly depressed, body whorl descending 
rapidly. Apical sculpture of about 22 very prominent spiral cords. 
Postnuclear whorls with high, widely spaced, strongly protractively 
sinuated radial ribs. Microsculpture of fine radials and finer spirals, 
no secondary cording. Umbilicus widely open, cup-shaped, regularly 
decoiling. Whorls flattened laterally above and slightly compressed 
laterally below rounded periphery. Parietal wall with median, high 
barrier, extending about '/i whorl, with abrupt anterior descension, 
not expanded above. No palatal or columellar barriers. Anatomy 
unknown. 

Type species. Lauopa mbalavuana, new species. 

Placement of this species in a distinct genus with- 
out any knowledge of its anatomy is based upon elimi- 
nation of alternative classifications. Key features are 
the very large and numerous apical cords, presence of a 
single parietal barrier, absence of secondary spiral 
cording, very large size, and widely spaced radial rib- 
bing. Species of Graeffedon from Samoa and Tonga are 
much smaller, have many more and shorter barriers, 
more crowded radial ribbing, and much narrower um- 
bilici. They agree in lack of secondary spiral cording 
and the flat or nearly flat apex and spire. Tuimalila 
from Tonga also lacks secondary spiral cording, but 
differs in its narrow umbilicus, distinctly elevated 
spire, prominent and much more crowded radial 
sculpture, greatly reduced apical sculpture, total ab- 
sence of apertural barriers, and smaller size. Sinployea 
complementaria (Mousson) from Upolu, Western 
Samoa, is only 4.92 mm. in mean diameter and lacks 
any apertural barriers (fig. 56b). 



Possibly Lauopa may be related to the Microne- 
sian genus Semperdon, but until live material can be 
obtained and dissected, no affinities can be determined. 
Generic recognition is given because there are major 
differences from any other genus known in this region. 
The name Lauopa refers to its occurrence in the Lau 
Archipelago. For convenience it is associated with the 
generalized Charopinae. 

Lauopa mbalavuana, new species. Figure 77a-c. 

Diagnosis. Shell extremely large, diameter 6.67-8.37 mm. 
(mean 7.52 mm.), with 4>/4-4% normally coiled whorls. Apex and 
spire flat, lower spire descending slightly, body whorl much more 
rapidly, spire protrusion less than 'Ao body whorl width, H/D ratio 
0.422-0.437 (mean 0.430). Apical sculpture of about 22 prominent 
spiral cords whose interstices are about equal to their width. Post- 
nuclear sculpture of high, prominent, protractively sinuated radial 
ribs, 39 on the body whorl in subadult, whose interstices are about 
5-8 times their width, becoming extremely crowded and irregular on 
last third of body whorl in adult (fig. 77a). Ribs/mm, in juvenile 1.86. 
Microsculpture of extremely fine and crowded spiral and radial rib- 
lets, more than 20 radials between each pair of major ribs, spirals 
distinctly finer. No secondary spiral cording. Umbilicus broadly 
open, cup-shaped, regularly decoiling, contained 2.72-2.76 times 
(mean 2.74) in the diameter, margins rounded. Whorls strongly 
rounded above and on periphery, slightly compressed laterally on 
upper and lower palatal margins. Parietal wall with single median, 
high barrier, extending posteriorly less than % whorl, slightly ex- 
panded above, with abrupt descension at anterior end. No columellar 
or palatal barriers. 

Tuimalila pilsbryi and T. infundibulus from 
Tonga (fig. 78a-f) both approach the size of Lauopa 
mbalavuana but lack the parietal barrier, have an ele- 
vated spire, much finer apical cording, and more 
crowded radial ribbing. All other Fijian and Polyne- 
sian species are much smaller in size. 

Description. Shell extremely large, with 4% moderately tightly 
coiled whorls. Apex and early spire slightly depressed below penul- 
timate, last whorl descending rapidly, H/D ratio 0.437. Embryonic 
whorls l'/2, sculpture eroded. Remaining whorls with remnants of 
protractively sinuate, low, rounded, widely spaced radial ribs whose 
interstices are 3-7 times their width, becoming extremely crowded 
on last third of body whorl. Microsculpture of very fine, close-set 
riblets crossed by even more crowded, low spiral ribs. Sutures deeply 
impressed, whorls strongly rounded above, flattened laterally above 
periphery. Color leached from shell with few vague flammulations 
remaining. Aperture circular, flattened laterally above periphery, 
inclined 25 from the shell axis. Parietal wall with one high, 
ridgelike barrier extending a little more than 3 Ae of a whorl. Um- 
bilicus broadly open, cup-shaped, regularly decoiling, contained 2.72 
times in the diameter. Height of holotype 3.66 mm., diameter 8.37 
mm. 

Holotype. Fiji: Lau Group, Vanua Mbalavu, Sta- 
tion 78, a limestone hill between Valika and Mosomo 
Bay, % mile inland at 200-250 ft. elevation. Collected 
under stones and logs in a dense forest by Yoshio 
Kondo and Elwood Zimmerman on August 9, 1938. 
BPBM 179554. 

Paratype. Lau Group: Vanua Mbalavu, a lime- 
stone hill between Valika and Mosomo Bay, % mile 
inland (Station 78) at 200-250 ft. elevation, under 
stones and logs (1 specimen, BPBM 179554). 

Remarks. The adult holotype of Lauopa mbala- 
vuana is a rather badly worn specimen, whereas the 



178 



SOLEM: ENDODONTOID LAND SNAILS 









FIG. 77. a-c, Lauopa mbalavuana, new species. Station 78, 
Vanua Mbalavu. Lati. Fiji. Holotype. BPBM 179554. Scale line 
equals 1 mm. (MMi. 



juvenile subadult retains most of the apical and micro- 
sculpture. It is uncertain whether this is a living or 
extinct species because the better-preserved paratype 
apparently was leached out from coralline rock, frag- 
ments of which are cemented to the shell. 

The only species of similar size, shape, and apical 
sculpture are the Tongan Tuimalila infundibulus and 
T. pilsbryi, neither of which have any apertural bar- 
riers. 

Genus Tuimalila, new genus 

Shell very large, with normally to loosely coiling whorls. Apex 
and spire moderately elevated. Apex with fine and crowded spiral 
cords, at least in one species with a secondary intrusion of radial 
swellings that appear under optical examination as primary radial 
ribs. Postnuclear sculpture of prominent, large, major radials and 
typical microsculpture. Umbilicus narrow, V-shaped, regularly de- 
coiling. Aperture relatively large. Pallial region as in Sinp/oyea, a 
narrow strip of lung roof visible between arms of ureter, kidney lobes 
almost equal in size. Genitalia agreeing with Sinployea except for 
nearly apical pore in the vergic papilla and presence of a well- 
developed retractor muscle attached to the free oviduct head. 

Type species. Tuimalila pilsbryi, new species. 

Although the Upolu Sinployea complementaria 
(Mousson) approaches the size of Tuimalila and several 
Rarotongan species had individuals within its size 
range, the two species grouped here average signifi- 
cantly larger, show a different pattern of whorl incre- 
ment, and have distinctive patterns of apertural open- 
ing. Their apical sculpture has the spiral cords reduced 
in prominence and increased in number over that 
found in Sinployea, whereas at least in T. pilsbryi there 
is developed a secondary intrusion of radial swellings 
(fig. 2c, p. 11). These appear as radial ribs under optical 
microscopic inspection at 96 x and visually mimic the 
characteristic Endodontidae apical sculpture. Whether 
this same feature occurs in T. infundibulus cannot be 
determined until specimens with unworn apical 
sculpture are obtained for study. 

Anatomical features show no characters incompat- 
ible with derivation from Sinp/oyea, whereas the con- 
chological changes are matters of degree rather than 
major shifts. Nevertheless, the change in apical 
sculpture and increase in size are major departures 
from the typical Sinployea pattern. The average mean 
diameter of the two species in Tuimalila, 6.16 mm., 
represents a 547( jump over the median mean diameter 
of Sinployea species and a 25% jump over the mean 
diameter, 4.92 mm., of S. complementaria (Mousson), 
the largest species in that genus. This size increase 
occurred with a slight actual decrease in whorl count 
and a marked increase in relative whorl widths. In 
Sinployea the median mean whorl count is 4V-, 
whereas Tuimalila infundibulus has 3%+ and T. 
pilsbryi 4+ whorls. The result of this size increase 
combined with no increase in whorl count is to alter the 
growth pattern. The much more rapidly increasing 
whorl cross-section combines with the higher pallial 
surface to broaden the kidney (fig. 79a) and to fore- 
shorten the pallial organization. 



SYSTEMATIC REVIEW 



179 



Just as Ba represents a new experiment in altered 
growth pattern by increased spire height, so Tuimallla 
is a new pattern based on great increase in whorl 
width. As such, both merit generic separation from 
Sinployea. 

The name Tuimalila commemorates the deceased 
famous tortoise, Tu'i Malila, which was resident in 
Tonga for many years and supposedly dated from the 
visit of Captain Cook (see Robb & Turbott, 1971). 

Tuimalila pilsbryi, new species, Figures 78a-c, 
79a-e. 

Champa radicalis Boettger, 1916 (not Mousson, 1871), Abhl. 
Senckenb. Naturf. Gesell., 36 (3), p. 290 Eua, Tonga. 

Diagnosis. Shell extremely large, diameter 5.88-6.99 mm. 
(mean 6.64 mm.), with 3%-4% almost normally coiled whorls. Apex 
and spire moderately and evenly elevated, body whorl descending 
slightly more rapidly, spire protrusion about Vs-'/t body whorl width, 
H/D ratio 0.495-0.606 (mean 0.556). Apical sculpture of numerous 
extremely fine and regular spiral cords (fig. 2c, p. 11), with a gradual 
intrusion of low, rounded radial swellings that appear as fine, major 
radial ribs on last portion. Postnuclear sculpture of relatively nar- 
row, prominent, V-shaped, strongly protractively sinuated radial 
ribs, 74-103 (mean 91.2) on the body whorl, whose interstices are 
2-4 times their width. Ribs/mm. 3.55-4.97 (mean 4.37). Micro- 
sculpture of very fine radial riblets, 5-12 between each pair of major 
ribs, crossed by distinctly finer and more crowded spiral riblets (fig. 
2c-d). Umbilicus narrow, V-shaped, regularly decoiling, contained 
4.48-6.50 times (mean 5.62) in the diameter, margins rounded. 
Whorl contours and aperture typical. 

Tuimalila pilsbryi from Eua (fig. 78a-c) is a more 
depressed, larger shell with less descension of the body 
whorl, and has lower, less sharply defined radial ribs 
than does T. infundibulus (fig. 78d-f) from Vavau. 

Description. Shell extremely large, with 4 somewhat loosely 
coiled whorls. Apex and early spire moderately and evenly elevated, 
body whorl not descending more rapidly, H/D ratio 0.529. Embryonic 
whorls I :! /K, first whorl with sculpture of about 33 very fine spiral 
riblets, last % whorl showing a gradual emergence of low, broadly 
rounded, radial swellings, foreshadowing spire sculpture. Radial ribs 
on early spire low, broadly rounded, becoming more prominent and 
higher below. Radial ribs on body whorl protractively sinuated, 
prominent, 94 in number, whose interstices are 2-3 times their 
width. Microsculpture of very fine, closely set radial riblets with 
equally spaced, smaller spiral ribs. Sutures moderately impressed, 
whorls evenly rounded above, body whorl somewhat flattened later- 
ally above periphery and slightly flattened on basal margin. Um- 
bilicus open, V-shaped, last whorl decoiling slightly more rapidly, 
contained 4.86 times in the diameter. Color light yellow-white with 
numerous sinuated irregular reddish brown flammulations. Aper- 
ture ovate with somewhat flattened basal margin, inclined about 20' 
from shell axis. Height of holotype 3.53 mm., diameter 6.67 mm. 

Holotype. Tonga: Eua, Johanssen Plantation, 2 
miles inland at 385 ft. elevation. Collected by H. S. 
Ladd on May 14, 1928. BPBM 87750. 

Range. Eua Island, Tonga. 

Paratypes. Eua (1 specimen, SMF 165341 col- 
lected by E. Wolf on June 5, 1909): Johanssen Planta- 
tion, 2 miles inland at 385-500 ft. elevation (5 speci- 
mens, BPBM 87750, BPBM 87719, BPBM 108565); 
main range on east side (Station T-22) of island at 
1,000 ft. elevation in heavy forest (35 specimens, 
FMNH 152378). 



Remarks. Although there are a few striking 
quantitative differences (table XXXIX) between 
Tuimalila pilsbryi and T. infundibulus, the ribbing 
character and different aperture appearances im- 
mediately separate individuals of the two species. Al- 
though the intrusion of secondary radial ribbing into 
the apex was not seen in any specimens of T. infun- 
dibulus, all specimens of the latter were so worn that 
this may be present on that species also. 

Living examples were common in fallen tree fern 
fronds. 

Great pleasure is taken in dedicating this species 
to the late Henry A. Pilsbry, dean of terrestrial mala- 
cologists and mentor. 

Description of soft parts. Foot and tail equal to shell diameter 
in length, slightly tapering posteriorly, truncated anteriorly. Sole 
undivided longitudinally, transversely corrugated in preservative. 
Pedal grooves typical, suprapedal distinctly weaker than pedal, both 
uniting above tail, no caudal horn or middorsal groove developed. 
Slime network weak, without peculiarities. Head projecting in front 
of foot, ommatophores typical, eyespots large. Gonopore a broad slit, 
opening behind right rhinophore and below right ommatophore. 

Body color yellow-white, a faint trace of grayish color on neck, 
ommatophores distinctly grayish. 

Mantle collar (MO thick, without obvious lobing, a broad black 
zone of color granules along inner margin, no glandular extension 
into pallial roof. Anus opening just inside pneumostome, slightly in 
front of external ureteric pore (KX). 

Pallial region (fig. 79a) extending slightly more than % whorl 
apically. Lung roof clear, no granulations. Kidney (K) about 3.0 mm. 
long, bilobed, with rectal arm slightly longer, reaching across 
hindgut onto parietal wall, very high at end of cavity, dished on 
posterior margin. Distance from anterior end to anus 2.1 mm. Ureter 
(KD) typical, a narrow strip of lung roof visible between arms that 
open at a slight angle. Heart (H) more than M; length of kidney, 
slightly angled to hindgut. Principal pulmonary vein (HV) un- 
branched, relatively conspicuous. Hindgut (HO) without unusual 
features. 

Ovotestis a single clump of palmately clavate alveoli, lying 
parallel to whorl sides and occupying almost '/> whorl above stomach 
apex. Hermaphroditic duct iridescent, slightly kinked in middle 
areas, narrowing abruptly at point where it starts along albumen 
gland, reflexinp up to enter junction of talon and carrefour (fig. 79e). 
Albumen gland with rather large acini, surface deeply indented by 
other organs, general shape squarish. Talon (fig. 79e, GT) globular, 
without a distinct shaft, joining hermaphroditic duct directly at head 
of carrefour (Xl. Latter -<i diameter of talon, tapering into head of 
prostate-uterus. Prostate (DG) with many very prominent and long 
alveoli complexly folded against wall of uterus, opening into groove 
on inner wall of uterine chambers. Prostate-uterus folded back on 
itself for part of length. Uterus (UT) typically bipartite, lower 
chamber with thick, glandular walls. 

Vas deferens (VD) wide in diameter with rather thick, pilas- 
tered, glandular walls until just before penioviducal angle, becoming 
very slender, reflexing upward. Epiphallus (E) about % length of 
penis, inserting into penis to one side of penial retractor, receiving 
vas deferens laterally on very swollen head, lower % more slender. 
Internally with Y valve and plug apically, lower portion with lon- 
gitudinal pilasters running into epiphallic pore (fig. 79c). Penial re- 
tractor (PR) very short, arising on diaphragm, inserting directly onto 
head of penis. Penis (P) about 2.4-2.5 mm. long, club-shaped, swollen 
medially, tapering slightly to atrial junction. Internally with typical 
pocket stimulator producing medial bulge, apically (fig. 79c-d) with 
vergic papilla and circular muscle band on wall below it. Atrium (Y) 
about 'it length of penis, tapering gradually. 




FIG. 78. a-c, Tuimalila pilsbryi, new species. Johanssen Plantation, Eua, Tonga. Holotype. BPBM 87750; d-f, Tuimalila infundibulus 
(Hombron & Jacquinot). Lectotype ofPatula radicalis Mousson. Zoologisches Institut der Universitat Zurich. Scale lines equal 1 mm. (a-c, SH; 
d-f, SG). 



180 



SYSTEMATIC REVIEW 

TABLE XXXIX. - RANGE OF VARIATION IN FIJIAN AND TONGAN ENDEMIC GENERA. 



181 



NUMBER OF 
NAME SPECIMENS RIBS 



RIBS/MM 



HEIGHT 



DIAMETER 



H/D RATIO 




IRREGULAR 



thaumasius 



Lauopa 

mbalavuana 



Tulmallla 

infundibulum 156 



42 



44.1(36-55) 
39 

88.3(75-96) 
91.2(74-103) 



2.17(1.84-2.50) 2.72(2.30-3.32) 

4.08(3.59-4.74) 1.28(1.12-1.51) 3.49(3.19-3.98) 

1.86 3.24(2.81-3.66) 7.52(6.67-8.37) 

4.88(4.17-5.37) 3.49(2.88-4.18) 5.68(4.97-6.86) 

4.37(3.55-4.97) 3.69(3.07-4.12) 6.64(5.88-6.99) 



0.801(0.752-0.842) 
0.365(0.343-0.381) 
0.430(0.422-0.437) 

0.616(0.547-0.710) 
0.556(0.495-0.606) 



humbu. 
thaum. 
mbala. 
inf un. 

pilsb. 



WHORLS 



UMBILICUS 



3 3/8(3 1/8-3 1/2) C L 

4 1/8(4-4 3/8) 1.16(1.05-1.32) 
4 1/2(4 1/4-4 3/4) 2.75(2.42-3.07) 
3 3/4+(3 1/2-4 1/8) 1.04(0.86-1.32) 



4+(3 3/4-4 3/8) 



1.19(1.05-1.24) 



D/U RATIO 

S E D 
2.99(2.82-3.15) 
2.74(2.72-2.76) 
5.47(4.47-7.15) 

5.62(4.48-6.50) 



APICAL 
CORDS 

12.0(11-13) 
12.2(10-15) 

22 

MANY 
REDUCED 

MANY 
REDUCED 



thaum. 



mbala. 



SPIRE 
ELEVATION 

0.37(0.25-0.46) 

DEPRESSED 
0.15(0.07-0.23) 
0.36(0.20-0.53) 
0.47(0.33-0.66) 



BODY WHORL 
WIDTH 

0.97(0.82-1.12) 
0.75(0.66-0.84) 
2.03(1.76-2.28) 
1.94(1.74-2.07) 
2.06(1.91-2.20) 



SP/BWW 
0.379(0.300-0.483) 

0.069(0.037-0.100) 
0.185(0.113-0.258) 
0.223(0.159-0.308) 



Free oviduct (UV) surmounted by a retractor muscle, swollen 
above, tapering rapidly with muscular walls, then suddenly expand- 
ing to form a thin-walled sac gradually tapering to spermathecal 
junction. Spermatheca (S) with ovate head lying next to albumen 
gland, shaft slender to base of uterine fold, lower portion rapidly 
expanding to equal penis in width. Vagina (V) equal in diameter to 
median free oviduct, very short, without prominent internal fea- 
tures. 

Free muscle system unusual only in having a very large muscle 
from the free oviduct apex attached to columellar retractor just above 
tentacular retractors. Right ommatophoral retractor passing 
through penioviducal angle. Tentacular retractors unite with tail fan 
just above junction of latter with buccal retractor. 

Stomach occupying % whorl after reaching expanded size. Intes- 
tinal loops highly compacted, hindgut typical. 

(Based on FMNH 152378, 4 adults 5.9-6.1 mm. in diameter.) 

Tuimalila infundibulus (Hombron & Jacquinot, 
1841). Figure 78d-f. 

Helix infundibulum Hombron & Jacquinot, 1841, Ann. Sci. Nat. 
Zool., (2) 16, p. 64 Vavao, Tonga Islands; Hombron & Jac- 
quinot, 1852, Voy. Pol. Sud, Astrolabe et Zelee, Atlas, pi. 6, figs. 
25-28 Vavao, Tonga; Rousseau, 1854, Voy. Pol. Sud, Astrolabe 
et Zelee, Atlas, 5, p. 20. 

Patula (Patula) radicalis Mousson, 1871, J. de Conchyl., 19, pp. 
12-13, pi. 3, fig. 3 Vavao, Tonga Islands. 



Helix radicalis (Mousson), Pfeiffer, 1876, Monog. helic. viv., 7, p. 

164. 
Helix crebriflammis Pfeiffer, 1876, Monog. helic. viv., 7, p. 148 

lists infundibulum Hombron & Jacquinot, 1841, as a question- 
able synonym of this New Zealand species. 
Charopa radicalis (Mousson), Tryon, 1886, Man. Conchol., (2) 2, p. 

210. 
Endodonta (Flammulina) radicalis (Mousson), Hedley, 1893, 

Nautilus, 7 (3), p. 35. 
Endodonta (Charopa) radicalis (Mousson), Pilsbry, 1893, Man. 

Conchol., (2)9, p. 35. 
Rhytida (Ouagapia) radicalis (Mousson), Mollendorff, 1903, Syst. 

Conchyl. Cab., I, 12, b, p. 83, pi. 13, figs. 12-15. 

Diagnosis. Shell very large, diameter 4.97-6.86 mm. (mean 
5.68 mm.), with 3!^-4'/e rather loosely coiled whorls. Apex and spire 
moderately and evenly elevated, last whorl descending much more 
rapidly, spire protrusion a little less than l h body whorl width, H/D 
ratio 0.547-0.710 (mean 0.616). Apical whorls worn in all examples 
examined, occasionally fine traces of weak and crowded spiral cords 
visible under oblique lighting. Postnuclear sculpture of high, thin, 
sharply defined, strongly protractively sinuated radial ribs, 75-96 
(mean 88.3) on the body whorl, whose interstices are 2-4 times their 
width. Ribs/mm. 4.17-5.37 (mean 4.88). Microsculpture a lattice of 
very fine radial and spiral riblets, 4-10 microradials between each 
pair of major ribs. No secondary spiral cording. Umbilicus narrow, 
V-shaped, regularly decoiling, contained 4.47-7.15 times (mean 
5.47) in the diameter, margins rounded. Whorl contours and aper- 



182 



SOLEM: ENDODONTOID LAND SNAILS 



DG 




PP 



FIG. 79. Anatomy of the Tongan Tuimalila pilsbryi, new species. Station T-22, Eua, Tonga. FMNH 152378: a, pallial region; b, pallial and 
terminal genitalia; c, interior of penis, epiphallus and vas deferens; d, detail of major penial stimulator; e, talon and hermaphroditic duct. Scale 
lines equal 1 mm. (SH). 



tural inclination normal. Color with narrow, wavy reddish flammu- 
lations, a tendency for broad circular supraperipheral and basal red- 
dish zones evident in many individuals. 

Tuimalila infundibulus (fig. 78d-f) is smaller, has 
a less protruded spire, but a greater H/D ratio than 
does T. pilsbryi (fig. 78a-c). The latter has less crowded 
ribbing, and the apical whorls show a clear radial ele- 
ment (fig. 2c, p. 11) that apparently is lacking in T. 
infundibulus. No other Polynesian Charopidae even 
approach the size of this species. 

Description of radicahs. Shell very large, with 4 rather loosely 
coiled whorls. Apex and spire moderately elevated, slightly rounded 
above, body whorl descending more sharply, H/D ratio 0.634. Apical 
whorls 1%, sculpture of extremely fine and crowded spiral ribs that 
are barely visible except under strong lateral lighting. Postnuclear 
whorls with irregular, high, rounded, strongly protractively sinuated 
radial ribs, about 92 on the body whorl, whose interstices are 2-3 
times their width. Microsculpture of very fine, crowded radial riblets, 
crossed by slightly finer and more crowded spiral riblets. Sutures 



deep, whorls shouldered above, strongly flattened laterally above 
periphery, with evenly and gently rounded subperipheral margins. 
Color light yellowish brown with very irregular radial reddish macu- 
lations that coalesce to form a broad and a narrow band above the 
periphery, a wide subperipheral plus a faint umbilical color band. 
Umbilicus narrowly open, V-shaped, regularly decoiling, contained 
5.55 times in the diameter. Aperture subcircular, strongly flattened 
laterally above periphery, inclined almost 25 from the shell axis. 
Height of lectotype 3.82 mm., diameter 6.02 mm. 

Lectotype of radicalis. Tonga: Vavau. Collected 
by Dr. E. Graeffe. Zoologisches Museum der Univer- 
sitat Zurich. 

Range. Vavau Island, Tonga. 
Paratypes. Zurich, FMNH 116982. 

Material. Vavau (20 specimens, RSM, AMS, 
FMNH 46222, SMF 165340, BPBM 54361, BPBM 
167432): Talau Id., 1 mile west (Station T-7) of Neiafu 
at 400 ft. elevation in dense low forest (62 specimens, 



SYSTEMATIC REVIEW 



183 



FMNH 152423); Mo'ungalafa, near Tu'anuku (Station 
T-9), about 14 miles southwest of Neiafu at 700 ft. ele- 
vation in heavy forest (2 specimens, FMNH 152207); 
Muitoulo (Station T-10), about 15 miles west of Neiafu 
at 500 ft. elevation in heavy forest (19 specimens, 
FMNH 152290); Pangai Motu Id. (Station T-ll), 3 
miles south of Neiafu at 200 ft. elevation (4 specimens, 
FMNH 152357); Makave village (Station T-12), 2% 
miles southeast of Neiafu at 200 ft. elevation (1 speci- 
men, FMNH 152348); Longomapu (Station T-14), 16 
miles southwest of Neiafu in heavy forest at 400 ft. 
elevation (14 specimens, FMNH 152380); Leimatu'a 
(Station T-17), 9 miles north of Neiafu at 500 ft. eleva- 
tion (2 specimens, FMNH 152323). "Tongatapu" (error) 
(22 specimens, Zurich, FMNH 116983). Erroneous data 
(3 specimens, FMNH 46243, 1KB). 

Remarks. The name Helix infundibulum gener- 
ally has been overlooked in the literature since Pfeiffer 
(1876) tentatively suggested it might be a synonym of 
the New Zealand Flammulina crebriflammis (Pfeiffer). 
Comparison of the type illustrations of infundibulum 
and radicalis leaves no doubt that they depict the same 
species. I could not locate type material of Helix infun- 
dibulum. 

Mollendorff s reference of this species to Ouagapia 
supposedly was based on work by Semper showing this 
species to have paryphantid dentition. I suspect that 
this was caused by a misreading of Semper (1874, p. 
136), who stated that O. gradata was a paryphantid 
and that probably many of the other "Patula" from the 
same area were also paryphantids. 

Extensive collections on Tongatapu in 1966 by 
Laurie Price failed to reveal any specimens of this 
species, hence I consider the material in the Mousson 
collection to be mislabeled. Unfortunately, all of the 
material taken on Vavau was dead so that Tuimalila 
infundibulus could not be dissected. Collections were 
made during a severe drought, and no living material 
was obtained ofSinployea vicaria paucicosta from some 
of the same stations. 

At Station T-7 one individual had the remains of 
an egg capsule with an embryonic shell dried inside. 
Despite the narrow umbilicus, evidently the eggs are 
deposited within the umbilical opening. Although this 
practice is common in the Endodontidae (Solem, 1976b, 
pp. 26-30), this is the only Polynesian Charopidae that 
showed evidence of this trait. 



Genus Lagivala, new genus 

Generally minute Charopidae with 3%-4% tightly coiled whorls. 
Apex and spire depressed (demani), flat (davidi), to moderately ele- 
vated, body whorl descending much more rapidly. Apical sculpture of 
10-14 fine spiral cords. Postnuclear sculpture of prominent to very 
fine (minusculus) radial ribs, microsculpture typical, no secondary 
spiral cording. Umbilicus very widely open, cup-shaped, narrow only 
in minusculus. Periphery usually strongly compressed laterally (ex- 
cept microglyphis), umbilical margin rounded (microglyphis, minus- 
culus} to strongly shouldered (macroglyphis) . Sutures channeled in 
macroglyphis, microglyphis, and minusculus. Shell unicolored with- 
out darker flammulations. Parietal wall with 2 or 3 large barriers 



(only 1 in minusculus), extending posteriorly from 3 Ae of a whorl to 
line of vision; where 2 parietals only, apparently a 3rd is sometimes 
displaced onto columellar margin by partial detachment of parietal 
wall. Palatal barriers 3 to 6, short, very deeply recessed crescents or 
elongated lamellar blades situated near posterior margin of parie- 
tals, in minusculus only a long single barrier lying partly on col- 
umellar and partly on basal margin. Anatomy unknown. 

Type species. Lagivala vivus, new species. 

Lagivala and Vatusila present generally contrast- 
ing patterns of structure (fig. 80-85). Lagivala has the 
body whorl laterally compressed (except for micro- 
glyphis and some demani); Vatusila has the body whorl 
flattened above the periphery. Lagivala lacks secon- 
dary spiral cording; Vatusila (except vaitupuensis) has 
secondary spiral cording. Lagivala has deeply recessed, 
short, and crescentic palatal barriers (except minus- 
culus and some demani); Vatusila has fewer, much 
longer, and less deeply recessed palatals. Lagivala has 
very long and rather complexly twisted, high parietals; 
Vatusila shows a strong tendency toward fusion and 
reduction of the parietals. Lagivala has a very wide, 
cup-shaped umbilicus (except minusculus); Vatusila a 
U-shaped, narrower umbilicus (cup-shaped only in 
eniwetokensis). Lagivala has a relatively low spire; 
Vatusila, a high spire (except eniwetokensis). They 
agree in major sculpture and rib spacing, but only be- 
cause both taxa exhibit an unusually wide range of 
types. Several Lagivala show a tendency toward chan- 
neling of the sutures, but others show no evidence of 
this change. Despite overlapping distributions and 
lack of anatomical evidence, I have no hesitation in 
considering them to be phyletically separate experi- 
ments in apertural constriction. 

From the above list of comparisons, it is evident 
that two species are atypical. Vatusila eniwetokensis 
has the flat spire and cup-shaped umbilicus of 
Lagivala. Its possession of prominent secondary spiral 
cording (fig. 86f), gradual descension of the upper 
parietal (fig. 85e), rounded body whorl with flattening 
restricted to above the periphery, shorter parietals, 
and relatively narrow umbilicus (table XL) are char- 
acters that I consider to be more significant than the 
probably linked spire and umbilical shape changes. 
Lagivala minusculus may be incorrectly placed here, 
but insufficient data are available to propose generic 
separation. Temporary classification within Lagivala 
is a conservative and reasonable treatment pending 
more information. 

Similarities to other genera are few. Graeffedon 
(figs. 86, 88) is much, much larger with short simple 
parietals and narrower, much less deeply recessed 
palatals that have abrupt anterior descension, a nar- 
rower, V-shaped umbilicus, and prominent, darker 
color flammulations. Other Melanesian and Polyne- 
sian taxa lack any apertural barriers. Of the Microne- 
sian genera, Palikirus (fig. 91d-f) has only a single 
parietal barrier, a much narrower, V-shaped um- 
bilicus, and much more widely spaced ribbing. Jokaj- 
don (fig 92a-f) has very fine ribbing, many complicated 
apertural barriers, and secondary spiral cording. Pal- 



184 



SOLEM: ENDODONTOID LAND SNAILS 



davidi 
(Ladd) 



macroglyphis 
(Rensch) 



microglyphis 
(Rensch) 



TABLE XL. - RANGE OF VARIATION IN LAGIVALA. 



"RHBER OF 
SPECIMENS 



1 
2 

1 

82 
5 



RIBS/:*. 



70 or 87 


15.9 or 19.8 


62.5(62-63) 


11. Ii7(11.39-ll. 55) 


121 


214.9 


62.5(51-71-) 


11.16(9.86-12.1(6) 


95.5(86-101) 


17.9(ll.8-20.8) 



demani 
(Tapperone- 
Canefri ) 



( Ladd ) 
vivus 
minusculus 



macroglyphis 

(Rensch) 



microglyphis 
(Rensch) 



demani 
(Tapperone- 
Canefri ) 



0.8k 
0.63 

0.95(0.89-1.02) 
0.71(0.66-0.714) 

0.85(0.76-0.95) 



79.0(56-106) 

DIAMETER 

1.1. 

1.75(1.72-1.77) 
1.55 

1.73(1.614-1.89) 
1.63(1.55-1.714) 

1.90(1.73-2.07) 



13. 09(9. 85-19. It) 
H/D RATIO 

0.500 

0. It83(0.l477-0. It 90) 

O.l40k 

0.553(0.539-0.580) 
0.14314(0. Ii25-o.lt 57) 

O.ltli8(0.l4l6-0.5l8) 



davldl 
vivus 



davidi 
vivus 
minus, 
mac ro . 
micro. 



WHORLS UMBILICUS 1/U KAIL) WICAL CORDS 

3 1/2 0.51 2.62 UNKNOWN 

4 0.68(0.67-0.69) 2.82(2.61-3.03) UNKNOWN 

3 3/8 0.30 5.22 10 

4 3/8-(4 1/8-4 5/8) 0.85(0.79-0.94) 2.03(2.00-2.08) 11-12 

3 7/8(3 3/4-4) 0.65(0.59-0.72) 2.54(2.41-2.61) Circa 11 

4 1/8+O 3/4-4 1/2) 0.72(0.58-0.82) 2.64(2.48-3.03) 12.1(10-14) 



SPIRE 
ELEVATION 

FLAT 

FLAT 
0.05 

0.07(0.04-0.10) 
0.05 



BODY WHORL 
WIDTH 



SP/BWW 
0.51 
0.59 

0.42 0.118 

0.54(0.53-0.59) 0.131(0.076-0.188) 2 1 
0.54 0.091 2 1 



PR C P 

2 or 1 3 or 4 
30 6 

10 1 

5 



donanl FLAT OR DEPRESSED 0.61(0.58-0.67) FLAT OR DEPRESSED 2-3 0-1 T-l-1 



line (figs. 94-96) has secondary spiral cording, looser 
whorl coiling, and less deeply recessed palatals. 

Known species of Lagivala are from Eastern In- 
donesia and West Irian (L. demani); New Britain, Bis- 
marck Archipelago (L. microglyphis and L. macro- 
glyphis); Viti Levu, Fiji (L. vivus and L. minusculus); 
and Funafuti, Ellice Islands (L. davidi). Except for the 
New Britain collections in shore drift and some sets of 
L. demani, only one or two examples are known of each 
despite intensive collecting efforts. Additional species 
will quite probably be discovered in intermediate areas 
since few attempts have been made at sampling the 
minute litter fauna. 

The similarities of all Lagivala except L. minus- 
culus are covered in the species accounts. Apparently, 
partial detachment of the parietal wall has resulted in 
the 3rd parietal becoming a columellar barrier in the 
Bismarck species and some populations of L. demani. 
Partial palatal barrier reduction in some examples of 



L. microglyphis shows how the reduced number in L. 
davidi could be derived. The channeled suture of L. 
minusculus agrees with the Bismarck species, whereas 
its peculiar columellar-palatal barrier could be derived 
from the 3rd parietal or columellar by downward, 
rather than upward, twisting of the anterior portion. 

In previous publications (Solem, 1958b, 1959a) I 
have used the genus Beilania Preston (1913, p. 433) for 
L. demani and several other species. Despite intensive 
search over 13 years, I have not been able to locate any 
specimens of Beilania demani Preston, 1913, in 
museum or private collections. Although the missing 
types may yet appear, the probabilities are that they 
will not be located in the immediate future. Preston's 
description places the parietals as being subperipheral, 
and the palatal wall is without barriers. In the absence 
of any specimens, I am reluctant to use the name 
Beilania for this group until the structures of the type 
species are known in regard to spire elevation, form 
and length of the parietals, actual rib count, and actual 
absence of the palatal barriers. Possibly Lagivala and 
Beilania are synonymous, but it is equally possible 
that Beilania inopina is more closely related to the 
Philippine-Indonesian species, Beilania philippinensis 
(Semper, 1874) (see Solem, 1957, pp. 7-8, fig. 3). That 
species has a narrower, V-shaped umbilicus, looser 
whorl coiling, a single crescentic parietal with gradual 
anterior descension, and three nodular palatals that 
are near the lip margin and often on a heavy callus 
much as in Palline notera and P. biakensis. The two 
latter species (figs. 94-96) differ in having prominent 
secondary spiral cording, tighter coiling, a much 
thicker body whorl, narrower umbilicus, and very 
differently shaped apertural barriers. Beilania philip- 
pinensis, although not dissected, shows major con- 
chological differences from Lagivala demani and the 
more eastern species of Lagivala. Rather than propose 
an additional generic name for the Philippine- 
Indonesian species, I prefer to use Beilania for that 
species and the genotype, pending availability of addi- 
tional material and soft parts for dissection. 

Lagivala is the Fijian nickname, meaning "the 
long fellow," applied to the famous chief Qaraniqio or 
Dakuwaqa of the Rewa District in Viti Levu, Fiji. 

Lagivala davidi (Ladd, 1968). 

Ptychodon species A, Ladd, 1958, J. Paleont., 32 (1), p. 189, pi. 30, 
figs. 13-15166-170 ft. in deep boring at Funafuti, Ellice Is- 
lands (Pleistocene or Recent). 

Ptychodon davidi Ladd, 1968, J. Paleont., 42 (3), p. 857, fig. 1170 
ft. in deep boring, Funafuti, Ellice Islands. 

Diagnosis. A very small species with less than 3M> tightly 
coiled whorls. Apex and spire flat, body whorl descending slightly, 
H/D ratio about 0.500. Apical whorls IMs, sculpture of fine spiral 
cords. Postnuclear whorls with prominent, rounded, almost verti- 
cally sinuated radial ribs, about 70-87 on the body whorl. Micro- 
sculpture of fine radial and finer spiral riblets. Umbilicus widely 
open, cup-shaped, regularly decoiling, contained about 2.60 times in 
the diameter, margins weakly shouldered. Sutures deep, whorls 
strongly rounded above and on basal margin, compressed laterally, 
with evenly rounded outer margin. Parietal barriers 2, agreeing in 



SYSTEMATIC REVIEW 



185 



structure with the upper pair in L. vivus. Outer wall with 3 deeply 
recessed palatals, short and crescentic, a 4th barrier located at baso- 
columellar margin of adults that may be either a columellar or a 
palatal barrier. 

Lagivala vivus (fig. 80a-c) differs fromL. davidi in 
having three parietals, six palatals, and fewer major 
radial ribs. Lagivala minusculus (fig. 80d-f) has only a 
single parietal and one peculiar palatal. Lagivala mac- 
roglyphis and L. microglyphis (fig. 81a-f) have ele- 
vated spires and more palatal barriers. 

Holotype. Deep boring from Funafuti, Ellice Is- 
lands, at 170 ft. depth. AMS F.52343. 

Range. Fossil at Funafuti, Ellice Islands. 

Material. Only the holotype and lost specimen 
initially studied by Ladd (1958) are known. 

Remarks. The juvenile holotype has three deeply 
recessed palatals in the aperture, whereas the lost 
example had an additional barrier at or near the baso- 
columellar margin. I am uncertain whether this was a 
columellar or palatal barrier. Differences fromL. vivus 
are covered in the diagnosis. 



Lagivala vivus, new species. Figure 80a-c. 

Diagnosis. Shell very small, diameter 1.72-1.77 mm. (mean 
1.75 mm.), with about 4 tightly coiled whorls. Apex and spire flat or 
very slightly depressed below level of penultimate whorl, body whorl 
descending moderately, H/D ratio 0.477-0.490 (mean 0.483). Apical 
sculpture of fine and crowded spiral cords, worn off over most of 
surface. Postnuclear whorls with high, rounded, sharply defined, al- 
most vertically sinuated, rather crowded radial ribs, 62-63 (mean 
62.5) on the body whorl, whose interstices are 2-4 times their width. 
Ribs/mm. 11.3911.55 (mean 11.47). Microsculpture of fine radial 
riblets, 5-8 between each pair of major ribs, crossed by extremely 
fine and crowded spiral riblets. No secondary spiral cording present. 
Umbilicus widely open, cup-shaped, regularly decoiling, contained 
2.61-3.03 times (mean 2.82) in the diameter, margins slightly shoul- 
dered, walls of umbilicus moderately flattened internally. Sutures 
deep, whorls strongly rounded above and on basal margin, com- 
pressed laterally with evenly rounded outer margin. Parietal bar- 
riers 3, extending posteriorly more than Vt whorl: upper a high, thin, 
bladelike lamella, slightly elevated and expanded above on posterior 
eighth, with abrupt descension at anterior end after even elevation 
from posterior portion; 2nd slightly higher and more broadly ex- 
panded on posterior quarter, equal in height to 1st during middle 
section, with sinuated descension over anterior 8th; 3rd located just 
above parietal-columellar margin, much lower than 2nd, weakly ele- 
vated and expanded above on posterior half, with gradual descension 
to point just inside lip edge. Columellar wall without barriers. 
Palatal barriers 6, deeply recessed, short and crescentic: lower at 
baso-columellar margin, relatively low, visible only by tilting of 
aperture; 2nd twice height of 1st, longer, slightly flattened on top, 
with relatively sharp anterior descension; 3rd and 4th slightly 
shorter and % height of 2nd with more gradual anterior descension; 
5th supraperipheral, equal in height to 2nd, with more gradual an- 
terior descension; 6th midway to palatal-parietal margin, a deeply 
recessed, low, and lamellate tubercle. 

Lagivala vivus is readily identified by possessing a 
third parietal and six palatals. Lagivala davidi has 
more ribs, only two parietals, and three palatals, but 
obviously is closely related. The Bismarck L. macro- 
glyphis and L. microglyphis (fig. 81a-f) have much 
more elevated spires, two parietals, a prominent col- 
umellar barrier, and only five palatals. Lagivala de- 
mani from Indonesia and West Irian has only three or 



four palatals, a depressed spire, and usually only two 
parietals. 

Description. Shell very small, with 4 tightly coiled whorls. 
Apex and spire slightly depressed, body whorl descending moder- 
ately, H/D ratio 0.477. Embryonic whorls almost 1%, traces of fine 
spiral cording present, surface mostly eroded. Postnuclear whorls 
with high, prominent, sharply defined, crowded radial ribs, 63 on the 
body whorl, whose interstices are 2-4 times their width. Micro- 
sculpture of fine radial riblets, 5-8 between each pair of major ribs, 
crossed by barely visible and extremely crowded spiral riblets. Su- 
tures deep, whorls strongly rounded above and on basal margin, 
compressed laterally, with evenly rounded outer margin. Umbilicus 
broadly open, cup-shaped, regularly decoiling, contained 2.61 times 
in the diameter, margins shouldered, inside walls flattened. Color 
uniform light reddish yellow, without distinct flammulations. Aper- 
ture ovate, compressed laterally, with evenly rounded outer margin, 
inclined about 5 from shell axis. Apertural barriers as in diagnosis 
above. Height of holotype 0.86 mm., diameter 1.76 mm. 

Holotype. Fiji: Viti Levu, Station 54, hillside at 
Telenaua, 48.3 miles west of Suva by Belt Road at 60 ft. 
elevation, back of Serua village. Collected in leaf 
mould by Yoshio Kondo, Elwood Zimmerman, and C. 
M. Cooke, Jr., on July 26, 1938. BPBM 178598. 

Range. South coast of Viti Levu, Fiji. 

Paratype. Viti Levu: Lami Ridge, west of Suva 
(Station 47) on limestone rock (1 specimen, BPBM 
178488); Nandarivatu (Station F-16), ridge slope at 
2,500 ft. elevation (1 specimen, FMNH 168364). 

Remarks. Only three dead specimens are known. 
They agree in apertural features, which clearly distin- 
guish them from other Lagivala. Attempts in 1962 to 
collect additional specimens were unsuccessful, and in 
1965 L. Price obtained one dead adult. 

The name vivus is to contrast this species with its 
nearest relative, Lagivala davidi (Ladd, 1968) from 
Funafuti Pleistocene or Holocene deposits. 

Lagivala minusculus, new species. Figure 80d-f. 

Diagnosis. A very small species with narrow umbilicus, 
slightly elevated spire, numerous and crowded ribs, 1 parietal, and 1 
sinuately twisted palatal. 

Other Lagivala have much more widely open um- 
bilici, two or three large parietals and three to five 
deeply recessed palatals. The singular palatal of L. 
minusculus immediately separates it from any of the 
Vatusila with reduced barrier number. 

Description. Shell very small, with 3% tightly coiled whorls. 
Apex and spire slightly and evenly elevated, body whorl descending 
slightly more rapidly, spire protrusion about l k> body whorl width, 
H/D ratio 0.404. Apical whorls l'/2, sculpture of 10 fine spiral cords, 
whose interstices are about 3-5 times their width. Postnuclear 
whorls with narrow, prominent, sharply defined, slightly sinuately 
protractive radial ribs, 121 on the body whorl, whose interstices usu- 
ally are less than twice their width. Ribs/mm. 24.9. Microsculpture of 
very fine radial riblets, 2-4 between each pair of major ribs, crossed 
by slightly finer and more crowded spiral riblets. Sutures distinctly 
channeled after embryonic whorls, whorls strongly rounded above, 
moderately compressed laterally and on basal margin. Umbilicus 
relatively narrow, U-shaped, slightly and regularly decoiling, con- 
tained 5.22 times in the diameter, margins rounded. Color light red- 
dish yellow-horn, no flammulations. Aperture ovate, compressed lat- 
erally and slightly on basal margin, inclined about 10 from shell 
axis. Parietal wall with single medial, high, bladelike barrier, 
slightly twisted upward posteriorly, weakly expanded above, extend- 






abc 



h 



def 



FIG. 80. a-c, Lagivala I'iuus, new species. Station 54, Telenaua, west of Suva, Viti Levu, Fiji. Holotype. BPBM 178598; d-f, Lagivala 
minusculus, new species. Nangava Island, Viti Levu, Fiji. Holotype. BPBM 179255. Scale lines equal 1 mm. (MM). 



186 



SYSTEMATIC REVIEW 



187 






ing posteriorly beyond line of vision, without noticeable descension 
until just before anterior end. Outer wall with a single moderately 
elevated, bladelike barrier, posteriorly at line of vision on columellar 
wall, anteriorly twisting down onto basal margin, reaching lip edge 
with sharp anterior descension, buttressed on outer side by a thick, 
tapering callus. Both barriers minutely serrated above. Height of 
holotype 0.63 mm., diameter 1.55 mm. 

Holotype. Fiji: Nangava Island, IVa miles south 
of Viti Levu, near Namuka Island, at 210 ft. elevation. 
Collected under dead hau leaves by Yoshio Kondo on 
September 20, 1938. BPBM 179255. 

Range. Nangava Island, south of Viti Levu, Fiji. 

Remarks. Inclusion of this species in Lagivala is 
based upon the channeled suture (also in L. macro- 
glyphis andL. microglyphis), shape of the parietal bar- 
rier (as inL. viuus andL. davidi), and general similar- 
ity of the columellar-palatal barrier to the third 
parietal in L. vivus (fig. 80b). The absence of any 
palatal barriers, narrow umbilicus, much more 
crowded and narrower ribbing, plus the peculiar ter- 
mination of the columellar-palatal differ from the 
Lagivala pattern. It is quite possible that L. minus- 
culus represents a distinct lineage and should be gen- 
erically separated. Without more specimens and mate- 
rial for dissection I prefer to include it in Lagivala. 

Lagivala macroglyphis (I. Rensch, 1937). Figure 
81d-f. 

Ptychodon macroglyphis I. Rensch, 1937, Arch. f. Naturgesch., n. 
f., 6 (4), p. 592, fig. 36 right Karlei, Malkong, and Matong, New 
Britain, Bismarck Archipelago. 

Diagnosis. Shell minute, diameter 1.64-1.89 mm. (mean 1.73 
mm.), with 4'/8-4 5 /8 rather tightly coiled whorls. Apex and early spire 
flat or slightly depressed to barely elevated, last whorl descending 
much more rapidly, spire protrusion slightly more than Vg body whorl 
width, H/D ratio 0.539-0.580 (mean 0.553). Apical whorls 1%-!%, 
sculpture of 11-12 fine spiral ribs whose interstices are about twice 
their width. Postnuclear sculpture of large, high, vertically sinuated 
radial ribs, 51 and 74 on 2 adult examples, whose interstices are 3-5 
times their width. Ribs/mm. 9.86-12.46 (mean 11.16). Micro- 
sculpture a lattice of very fine, almost coequal spiral and radial rib- 
lets, 6- 10 between each pair of major ribs. Umbilicus broadly open, 
cup-shaped, regularly decoiling, contained 2.00-2.08 times (mean 
2.03) in the diameter, margins distinctly shouldered, inner walls 
flattened, sutures strongly channeled with part of parietal wall de- 
tached. Sutures channeled above, whorls strongly rounded above and 
on basal margin with evenly rounded, strongly compressed laterally 
on outer margin, umbilical margin shouldered. Color partly leached 
from shell, a reddish yellow tint remaining. Aperture compressedly 
ovate, strongly rounded above and below, markedly compressed lat- 
erally, lying parallel to shell axis. Parietal barriers 2, extending 
posteriorly almost to line of vision: upper a high, thin blade with 
abrupt anterior descension, becoming thickened above on posterior 
8th with slight downward twist or weakly bifid; lower parietal with 
anterior 8th a high threadlike ridge, abruptly becoming equal in 
height to upper parietal, greatly thickened, expanded and elevated 
above on posterior quarter. Columellar wall with single bladelike 
barrier situated just below parietal-columellar margin on detached 
parietal wall, identical in form to 2nd parietal, but only half as high 
and extending not as far anteriorly. Palatal wall with 5 thin, deeply 
recessed, very short, crescentic ridges situated opposite posterior end 
of parietal barriers: 1st palatal basal in position, a very small, short 
crescent; 2nd a high crescentic barrier, weakly expanded above, 
whose apex lies opposite columellar; 3rd slightly lower than 2nd, 
situated subperipherally and directly opposite lower parietal; 4th 
equal in size to 3rd, situated opposite upper parietal; 5th smaller 



than 1st, a tiny short, low barrier near upper palatal margin, occa- 
sionally absent. All barriers minutely serrated above. 

The sympatric Lagivala microglyphis (fig. 81a-c) 
differs from L. macroglyphis (fig. 81d-f) in having a 
distinctly narrower umbilicus, lower H/D ratio, finer 
and more crowded radial ribs, plus slightly smaller av- 
erage size. Lagivala vivus from Fiji lacks the partly 
detached parietal wall and thus has three parietals. It 
also has six palatals and a slightly depressed apex and 
spire. 

Holotype. Karlei, Weiten Bucht, New Britain, 
Bismarck Archipelago. Collected by Schneider. ZMB. 

Range. Karlei, Malkong, and Matong, New Brit- 
ain, Bismarck Archipelago. 

Material. New Britain: Karlei (81 specimens, 
ZMB, FMNH 146021); Malkong-Bach, Karlei (25 
specimens, ZMB); Matong (1 specimen, ZMB). 

Remarks. Lagivala macroglyphis has several 
unusual features produced by partial separation of the 
postnuclear whorls during growth. On the spire (fig. 
81d) this is evidenced by deep channeling of the su- 
tures that are partly filled by periostracal material in 
fresh examples. In the umbilicus (fig. 81f) there is the 
same partial detachment of the parietal wall seen in 
the Marquesan Planudonta (Solem, 1976a, figs. 146, 
149). The upper portion of each umbilical volution 
lacks shell sculpture. This results from simple detach- 
ment of the parietal callus. At the same time, the 3rd 
parietal barrier of L. vivus has been lowered to occupy 
the "columellar wall." It is, of course, completely 
homologous with the parietal and is called a columellar 
here only to facilitate identification by nonspecialists. 
At its anterior end there is a pronounced tendency for a 
distinct upward angling of the barrier to the functional 
columellar-parietal margin. 

Lagivala vivus (fig. 80a-c) lacks the whorl de- 
tachment, has a narrower umbilicus, lower H/D ratio, a 
flat spire, and an additional palatal lamella. In size 
and sculpture it is quite similar. 

Lagivala microglyphis (I. Rensch, 1937). Figure 
81a-c. 

Ptychodon microglyphis I. Rensch, 1937, Arch. f. Naturgesch., n. f., 
6 (4), p. 591, fig. 36 left Karlei, Weiten Bucht, New Britain, 
Bismarck Archipelago; Solem, 1957, Fieldiana: Zoology, 42 (1), 
p. 7. 

Diagnosis. Shell minute, diameter 1.55-1.74 mm. (mean 1.63 
mm.), with 3%-4 rather tightly coiled whorls. Apex and early spire 
flat to barely protruding, last Vfe-% of body whorl descending a little 
to much more rapidly, spire protrusion about 'Ai body whorl width, 
H/D ratio 0.425-0.457 (mean 0.434). Apical whorls 1V4, sculpture of 
11 prominent spiral cords in only unworn examples. Postnuclear 
whorls with prominent, rounded, slightly protractively sinuated ra- 
dial ribs, 86-101 (mean 95.5) on the body whorl, whose interstices 
are 1-3 times their width. Microsculpture of very fine radial riblets, 
5-8 between each pair of major ribs, crossed by coequal spiral riblets. 
Umbilicus broadly open, cup-shaped, regularly decoiling, contained 
2.41-2.61 times (mean 2.54) in the diameter, margins rounded. Su- 
tures deep, very weakly channeled, whorls strongly rounded above 
and on basal margin, nearly evenly rounded on outer margin with 
very slight lateral flattening above and below periphery. Color red- 
dish yellow-horn without darker markings. Aperture subovate, 









FIG. 81. a-c, Lagivala microglyphis (Rensch). Karlei, New Britain, Bismarck Archipelago. Paratype. FMNH 146022; d-f, Lagivala mac- 
roglyphis (Rensch). Karlei, New Britain, Bismarck Archipelago. Paratype. FMNH 146021. Scale line equals 1 mm. (SH). 



188 



SYSTEMATIC REVIEW 



189 



strongly rounded above and at basal margin, evenly rounded 
elsewhere with slight lateral flattening, inclined about 20 from shell 
axis. Parietal barriers 2, extending posteriorly to line of vision: upper 
high and bladelike with very sharp anterior descension, slightly 
twisted upward, posterior third to quarter varying from bifid to with 
sharp downward deflection of greatly accentuated expanded portion; 
lower parietal a more rounded ridge with very gradual descension 
over anterior third, becoming higher, slightly bulbous on top, and 
twisted upward on posterior third. Columellar wall with single, 
bluntly rounded, moderately recessed, crescentic barrier extending 
posteriorly about '/ whorl with gradual anterior descension across 
top of columellar callus, slightly twisted upward over anterior third. 
Palatal wall normally with 5 deeply recessed, short, and crescentic 
barriers situated opposite posterior end of parietal barriers: lower 
palatal basal in position, a short crescent, slightly bulbous above, 
less than % height of next barrier; 2nd palatal high and crescentic, 
slightly bulbous above, pointing toward upper margin of lower 
parietal; 3rd palatal slightly smaller than 2nd, subperipheral, point- 
ing between parietal barriers; 4th palatal equal in size to 2nd, 
slightly supraperipheral in position, situated opposite edge of upper 
parietal: 5th palatal equal in size to 1st, situated midway between 
4th palatal and parietal-palatal margin. Occasionally only 3 palatals 
are present, and an accessory trace may be present between various 
lower palatal pairs. Expanded upper portions of each barrier with 
minute crystalline barbs. 

Lagiuala microglyphis has many more radial ribs, 
a much lower H/D ratio, fewer whorls (table XL), and a 
more rounded whorl contour than does the sympatric 
L. macroglyphis (fig. 81d-f). In L. vivus (fig. SOa-c) 
there are three parietals and no columellar, the latter 
in both L. macroglyphis and L. microglyphis being a 
descended third parietal. 

Holotype. Karlei, Weiten Bucht, New Britain, 
Bismarck Archipelago. Collected by Schneider. ZMB. 

Range. Karlei, New Britain, Bismarck Ar- 
chipelago. 

Material. Karlei (5 specimens, ZMB, FMNH 
146022). 

Remarks. The five paratypes showed some bar- 
rier variation. A small accessory crescentic barrier lies 
between the second and third palatals of the largest 
specimen; one adult had the first and fourth palatals 
missing, with the second and third grossly thickened; a 
juvenile lacked the first and fourth palatals, with no 
thickening of the second and third palatals. The ten- 
dency toward loss of two palatals provides a link to the 
condition seen in L. davidi. 

Suture channeling in L. microglyphis is greatly 
reduced, and there is no indication of sculptural change 
in the umbilicus caused by parietal wall descension, 
despite the position of the columellar barrier. The 
sculpture is much finer than in other Lagivala, except 
for the unusual L. minusculus. Whether L. micro- 
glyphis is an advanced or a generalized species cannot 
be determined without additional material for study 
and dissection. 

Lagivala demani (Tapparone-Canefri, 1883). 

Patula demani Tapparone-Canefri, 1883, Ann. Mus. Civ. Stor. 
Nat., Geneva, 19, pp. 95-96, 300, pi. 2, figs. 13-15 Vokan, Aru 
Islands; van Benthem Jutting, 1962, Ann. Mus. Civ. Stor. Nat., 
Genova, 73, p. 5. 



Helix demani (Tapparone-Canefri), Tryon, 1887, Man. Conchol., 

(2) 3, pp. 26-27, pi. 4, figs. 39-41. 
Nesophila demani (Tapparone-Canefri), Boettger, 1922, Abhl. 

Senckenb. Naturf. Ges., 35 (4), pp. 373-374, pi. 21, fig. 5. 
Beilania demani (Tapparone-Canefri), Solem. 1958, Arch. f. Mol- 

lusk., 87 (1-3), p. 22; van Benthem Jutting, 1964, Nova Guinea, 

Zool., 26, pp. 11-13 (partly) localities on Biak. 
Ptychodon demani (Tapparone-Canefri), van Benthem Jutting, 

1958, Nova Guinea, Zool., 9 (2), p. 327 Waima, Misool. 

Diagnosis. Shell relatively large, diameter 1.73-2.07 mm. 
(mean 1.90 mm.), with 3%-4'/4 tightly coiled whorls. Apex and spire 
slightly and evenly elevated to moderately depressed below level of 
penultimate whorl, often flat, body whorl descending much more 
rapidly, spire protrusion at most 'A body whorl width, H/D ratio 
0.416-0.518 (mean 0.448). Apical sculpture of 10-14 (mean 12.1) 
very fine spiral cords, whose interstices are about 3 times their 
width. Postnuclear whorls with prominent to very prominent, high, 
sharply defined, usually slightly protractive radial ribs, 56-106 
(mean 79.0) on the body whorl, whose interstices are 1V4-5 times 
their width. Ribs/mm. 9.85-19.4 (mean 13.1). Microsculpture of fine 
radial riblets, 3-13 between each pair of major ribs, crossed by 
slightly finer and more crowded spiral riblets, no secondary spiral 
cording. Umbilicus very widely open, V-shaped, regularly decoiling, 
contained 2.48-3.03 times (mean 2.64) in the diameter, margins 
rounded. Sutures deep, whorls strongly rounded above and on baso- 
columellar margin, outer margin varying from markedly compressed 
laterally to almost evenly rounded with slight lateral compression 
above periphery. Aperture subcircular to compressedly ovate, in- 
clined about 5-10 from shell axis. Parietal barriers normally 2 
(only one known population with 3). high, extending posteriorly 
3 fiB-Vt whorl: upper very high and slender with abrupt anterior de- 
scension, posterior quarter often slightly more elevated with a 
downward twist to elevated portion; 2nd usually equal in height on 
crescentic posterior 3rd, which may be slightly twisted upward, mid- 
dle 3rd slightly lower and less expanded or descending, with gradual 
descension over anterior 3rd until just before termination slightly in 
front of upper parietal; 3rd, when present, about '/2-% height of 2nd, 
just above parietal-columellar margin. Columellar wall without any 
barrier (Biak and West Irian), with a very small and deeply recessed, 
ridgelike trace (Biak), a medium-size barrier (Ambon and Aru Is- 
lands), or with barrier almost equal in size to 2nd parietal (Timor). 
Palatal barriers 3 or 4, rarely absent, deeply recessed, variable in 
size and position, crescentic and short to elongated lamellar ridges: 
1st through 3rd on basal to outer lip, with 3rd just above periphery 
and opposite upper parietal, sometimes shifted lower on outer wall; 
4th, when present, midway between 3rd palatal and palatal-parietal 
margin. 

Lagivala demani differs from the Fijian L. vivus 
and the two Bismarck species in its reduced number of 
palatal barriers, generally lower H/D ratio, and 
slightly greater size. Lagivala macroglyphis is much 
more widely umbilicated, has a distinctly protruded 
spire, and is very strongly compressed laterally on the 
whorl margin. Lagivala microglyphis has finer radial 
ribbing, is smaller, the body whorl has only slight lat- 
eral compression, and the aperture is inclined about 
20 from the shell axis. Lagivala vivus always has a 3rd 
parietal and 6 palatals. 

Description. Shell minute, with 3% moderately tightly coiled 
whorls. Apex and spire slightly emergent, body whorl descending 
only a little more rapidly, H/D ratio 0.454. Surface of shell heavily 
encrusted with dirt and fungal growth so that sculptural details are 
obscured. Size and position of major ribs as in "Diagnosis" cited 
above. Umbilicus broadly V-shaped, widely open, regularly decoil- 
ing, contained 2.54 times in the diameter. Apertural lip badly bro- 
ken, obscuring shape of aperture, sutures filled with extraneous mat- 
ter. Aperture showing only anterior edges of the 2 parietal barriers, 



190 



SOLEM: ENDODONTOID LAND SNAILS 



otherwise choked with dirt and fungal hyphae. Height of lectotype 
0.74 mm., diameter 1.63 mm. 

Lectotype. Aru Islands: Wokan. Collected by O. 
Beccari. MSNG. 

Range. Ambon, Timor, Wokan in the Aru Is- 
lands, Misool, Biak off West Irian, and coastal areas of 
West Irian. Possibly Celebes. 

Parades. MSNG, SMF 4678. 

Material. Ambon (1 specimen, ZMA collected Oc- 
tober 11-13, 1949, by M. A. Lieftinck). Timor: Araki- 
Vlakte (5 specimens, FMNH 150775, ZMA collected 
April 12, 1950, by B. Polak). Aru Islands: Wokan (5 
specimens, MSNG, SMF 4678). Misool: Waima (6 
specimens, FMNH 150774, ZMA collected September 
13-14, 1948 by M. A. Lieftinck). Biak: Kombai (00 46', 
135 58') (12 specimens, ZMA, RNHL); Korim (00 55', 
136 04') (10 specimens, FMNH 150776, RNHL); Sta- 
tion G-65, Saba (1 specimen, RNHL); Station G-34, 
Owi, South Biak (1 specimen, RNHL); Station G-30, 
Bosnik (1 specimen, RNHL). West Irian: Ugar, Kokas 
District (1 specimen, RNHL collected June 6, 1953); 
Sarmi (01 52', 138 44') (2 specimens, ZMA collected 
December, 1962, by G. J. M. Gerrits). 

Remarks. In the type lot of Lagivala demani 
there were five examples of this species and a single 
Discocharopa aperta (Mollendorff). Tapparone- 
Canefri's original figures show an extremely widely 
umbilicated shell without trace of apertural barriers. 
Although the Lagivala were all juvenile and had the 
aperture choked with dirt so that only the tips of the 
parietals were visible, I suspect that, despite no men- 
tion of the barriers, the description was based on the 
Lagivala and the figures on the Discocharopa. Boettger 
(1922) illustrated one specimen from the type lot with 
the parietals and depressed spire. I prefer to use the 
name in this sense rather than to adopt the Dis- 
cocharopa as type, since its reduced sculpture and ele- 
vated spire agrees more with the type figures. 
Tapparone-Canefri (1883, p. 300) indicated in the ex- 
planation of the plates that the sculpture was omitted. 
The very fine ribs of the Discocharopa could have been 
overlooked much more easily than the coarse ribbing of 
the Lagivala. Recently, van Benthem Jutting (1962, p. 
5) indicated that the lectotype was the "shell figures pi. 
2, fig. 13-15." There is little agreement between the 
lectotype and the figures. 

Unfortunately, most available material was 
juvenile. Only 12 of the Biak examples from Kombai, 
Korim, and Saba plus one shell from Misool showed 
adult shell features. Because of the barrier variation, 
changes in spire elevation and ribbing and variation 
data for several sets of adults and juveniles are sum- 
marized in Table XLI. Size differences should be ig- 
nored, because the sets are not comparable in age com- 
position. Since umbilical decoiling is regular in this 
species and the spire is normally flat or depressed, both 
D/U and H/D ratios would be little affected by the 
change from juvenile to adult growth. The very slightly 
increased H/D ratio for the Timor shells is attributable 



to the slightly elevated spire in that population. Shells 
from Misool are more narrowly umbilicated, whereas 
those from Korim, Biak, have a lower H/D ratio, possi- 
bly as a correlative with the more depressed apex and 
spire of that population. Rib counts vary widely, but 
the more critical factor of rib spacing shows agreement 
between the samples. Timor shells do have more prom- 
inent ribs, but there is only a slight decrease in rib 
spacing that is not statistically significant. Misool 
shells have narrower and thus more numerous and 
slightly more crowded major ribs (mean ribs/mm. 
13.83) than other sets. Measured differences are thus 
minor. 

Changes in spire protrusion and apertural barriers 
are much more significant. In the types from Wokan, 
Aru Islands, the spire is barely elevated, the columel- 
lar barrier is high but deeply recessed, and the three 
palatals are situated in normal position, deeply re- 
cessed, short and high. Juveniles from Timor in addi- 
tion to the enlarged radial ribbing have the spire and 
apex distinctly elevated, the columellar barrier is half 
to two-thirds as high as the second parietal and ex- 
tends to the top of the columellar callus with gradual 
anterior descension, plus four palatals. The lower three 
are high and crescentic, with the first at the baso-lower 
palatal margin, the second pointing toward the lower 
parietal top edge, and the third opposite and nearly 
touching the upper parietal. A much smaller and lower 
fourth palatal lies one-third of the way between the 
third palatal and the palatal-parietal margin. One in- 
dividual has an accessory cordlike trace at the baso- 
columellar margin, more deeply recessed than the first 
palatal and much shorter. All specimens from Biak 
have a flat or noticeably depressed spire, the columel- 
lar reduced to a threadlike or ridged lamellar trace 
visible only by extreme tilting of the aperture, and the 
three palatals longer, much lower, with the second 
situated distinctly above the second parietal level, and 
the third sometimes split into two parts. The Misool 
examples have the spire flat or very slightly depressed, 
no trace of any columellar barrier, and the position of 
the palatals altered so that the first palatal lies at the 
baso-columellar margin, the second at the baso-lower 
palatal margin, and the third pointing between the 
first and second parietals. In size these palatals are 
intermediate between those of the Biak and Timor 
examples. Their downward shift in position probably 
reflects adjustments to the columellar barrier loss and 
serves to restore balanced narrowing of the aperture. 
The broken and worn individual from Ambon has a 
medium-sized columellar barrier, the palatals shifted 
as in the Misool shells, and the spire very slightly ele- 
vated. Two shells from Sarmi, West Irian, have a third 
barrier on the lower edge of the parietal wall and equal 
in height posteriorly to the second parietal. Its anterior 
descension is very gradual and terminates behind the 
end of the first parietal. There is no columellar barrier 
as such. One shell has the typical short and crescentic, 
deeply recessed three palatals, the other lacks any 



SYSTEMATIC REVIEW 

TABLE XLI. - LOCAL VARIATION IN LAGIVALA DEMANI . 



191 



NUMBER OF 
SPECIMENS 



Sarmi, W. Irian 2 
ZMA 

Kombai, Blak 5 

ZMA, RNHL 

Korlm, Biak 7 

RNHL 

Saba, Biak 1 

RNHL 

Waima, Misool 5 
ZMA 

Araki-Vlakte .Timor 4 
ZMA 

Wokan, Aru 5 

SMF 4678.MSNG 

Ambon 1 

ZMA 



RIBS 



RIBS /MM. 



65.510.53 13.01.32 



HEIGHT 



0.76+0.049 



(55-76) 



(11.68-14.31) 



(0.71-0.81) 



DIAMETER 



1.600.099 

(1.50-1.69) 



75.54.01 13.10.66 0.860.032 1.86+0.080 

(56-106) (9.85-19.39) (0.76-0.93) (1.66-2.06) 

70.1+3.37 12.30.47 0.7910.017 1.82+0.042 

(56-85) (9.88-13.87) (0.72-0.86) (1.63-1.96) 



57.7+0.33 



0.95 



2.07 



71.8+6.65 13.8+0.41 0.75+0.044 1.610.101 

(56-87) (12.92-14.85) (0.66-0.90) (1.38-1.97) 

54.52.53 12.70.54 0.660.017 1.37+0.028 

(47-58) (11.33-13.68) (0.63-0.71) (1.32-1.45) 



13.0+0.61 



(57-58) (12.31-14.20) 
50 10.2 



0.68+0.022 1.45+0.056 

(0.64-0.74) (1.30-1.63) 



0.74 



1.56 






H/D RATIO 

Sarmi 0.475+0.0014 

(0.473-0.476) 

Kombai 0.4650.0134 

(0.443-0.518) 

Korim 0.4350.0040 

(0.416-0.448) 



Saba 



0.460 



WHORLS 

4 1/8 

4 1/4(4-4 1/2) 
4(3 1/2-4 1/4) 

4 1/2 

3 l/2-(3 1/8-3 3/4) 



UMBILICUS 



D/U RATIO 



Waima 0.4630.0118 

(0.455-0.476) 

Araki 0.4850.0100 

(0.475-0.488) 



Wokan 0.468+0.0099 3 l/2-(3 1/4-3 7/8) 
(0.438-0.493) 



0.660.033 
(0.63-0.69) 


2.410.030 
(2.39-2.45) 


0.69+0.046 
(0.58-0.81) 


2.71+0.086 
(2.55-3.03) 


0.71+0.020 
(0.64-0.79) 


2.55+0.021 
(2.48-2.63) 



Ambon 



0.473 



0.82 

0.540.040 

(0.41-0.66) 

0.51+0.019 

(0.46-0.54) 

0.5510.031 

(0.48-0.64) 

0.63 



2.52 

2.99+0, 

(2 

2.66+0, 
(2, 

2.670 
(2 

2.50 



,100 
,81-3.36) 

,076 
,49-2.86) 

,054 
,53-2.78) 



trace of palatals. Whorl contours vary with the ribbing, 
the Sarmi and Misool shells having the body whorl 
markedly compressed laterally, the Biak shells show- 
ing only slight lateral compression, and the Timor 
shells having a subcircular body whorl contour with 
some lateral compression above the periphery. Reduc- 
tion of the columellar barrier correlates with increased 
lateral compression of the body whorl. 

Recognizably different populations exist, and the 
pattern of variation in many Wallacean animal groups 
strongly suggests subspeciation has occurred. With 
only single collections of mainly juvenile individuals 
from widely scattered areas I prefer not to give 
nomenclatural recognition to these populations. Varia- 
tional data summarized above and in Table XLI should 
enable interpretation of variation in additional popula- 
tions but are not sufficient for description of subspecific 
units. 

The wide range of this species, extending as far 
west as Timor, Ambon, and Misool, makes it quite 



plausible that Beilania inopina Preston, 1913, from 
Beilan-Beilan Island north of Obi is a member of this 
complex. Reasons for not including it at this time have 
been given above (p. 184). 

A single worn and broken shell from Tjamba Pass, 
near Makassar, South Celebes, was located in a set of 
Discocharopa aperta. It may be an undescribed species 
of Lagivala or represent a population of L. dernani in 
which the palatals are very close to the lip. 

Genus Vatusila, new genus 

Shell relatively small to very small, with 3'/2-5 normally to 
rather tightly coiled whorls. Apex and spire flat (eniwetokensis) to 
rather strongly elevated (nayauana), body whorl usually descending 
much more rapidly. Apical sculpture of 8-12 fine, but prominent 
spiral cords. Postnuclear sculpture of crowded to widely spaced radial 
ribs, strong secondary spiral cording in all but niueana (weakened) 
and vaitupuensis (absent). Umbilicus rather widely open, narrowed 
in kondoi. Periphery rounded, whorls slightly to strongly flattened 
laterally above periphery. Color flammulations, when present, not 
zigzagged, following lines of growth. Parietal wall with 2 barriers, 1 



192 



SOLEM: ENDODONTOID LAND SNAILS 



bifurcated barrier, or greatly reduced single barrier. Columellar wall 
with or without recessed, low ridge. Palatal barriers varying from 
0-3, broadly expanded (tongensis) down to a low, broad remnant 
callus (niueana). Anatomy mostly unknown, penis in tongensis 
differing from Sinployea only in accentuation of semicircular pilas- 
ters forming vergic papilla. 

Type species. Vatusila tongensis, new species. 

Despite the rather widely different appearance of 
the sculpture and large size range, I have no hesitation 
in considering these species congeneric. The two small 
species, V. kondoi (fig. 82a-c) and V. nayauana (fig. 
82d-f) from the Lau Group, and V. eniwetokensis (fig. 
85e-f) from the Miocene of Eniwetok, agree in shell 
microsculpture, although not in parietal barrier size or 
character of major ribbing. The larger V. tongensis (fig. 
83a-c) from Eua has the microsculpture of this group 
and agrees in parietal barrier shape with the fossil. 
Comparison of the figures shows that a callus connect- 
ing the second and third palatals, elaboration of the 
first palatal, and loss of the columellar from V. kondoi 
would duplicate the outer lip barriers in V. tongensis. 
Although no direct linear relationship between these 
species should be implied, all the above species could be 
derived from a common model. Vatusila vaitupuensis 
(fig. 85a-d) and V. niueana (fig. 83d- f) would represent 
successive stages in barrier reduction from a V. 
tongensis barrier stage, although they agree more with 
V. eniwetokensis in major rib spacing. They differ in 
having modified secondary spiral sculpture or none 
(vaitupuensis). 

It is significant that those with the widely spaced 
major sculpture, V. tongensis, V. kondoi, and V. 
nayauana, are from a central area, whereas those with 
more average sculpture, V. niueana, V. vaitupuensis, 
and V. eniwetokensis, have a fringing distribution 
around this core. I suspect that more species of this 
group will be discovered in Fiji and Tonga when addi- 
tional collections become available. 

Comparisons with other genera are unsatisfactory 
because so little anatomical data were available. The 
penis of Vatusila tongensis (fig. 84a-b) represents a 
rather minor change from the pattern seen in Sin- 
ployea. Without more complete material for study, the 
closeness of this relationship will remain uncertain. 
The small species with heavier barriers show consider- 
able sculptural modification from the pattern of 
Sinployea, whereas one of the larger species, V. 
niueana, can be mistaken for that genus very easily. 
Progressive reduction in aperture barriers with in- 
creased size is common in the Endodontidae, but this 
cannot be postulated for Vatusila, because the sizes of 
V. tongensis, V. vaitupuensis, and V. niueana are es- 
sentially identical. Species ofLagivala have much sim- 
pler structure to the parietal barriers; their palatals 
are short, simple, and very deeply recessed. They differ 
in both microsculpture and whorl contours and seem to 
be a group paralleling the structures of Vatusila, but of 
different origin. Microcharopa (fig. 33a-c) is smaller, 
has greatly modified apical shell sculpture, and lacks 
apertural barriers. Graeffedon (figs. 86, 88) is much, 



much larger and has quite different palatal barriers 
(although the parietals have a similar structure to 
those found in V. eniwetokensis) and a very different 
anatomy (fig. 87a-d). 

Vatusila is named after the Fijian tribe (located at 
the headwaters of the Sigatoka River) that killed and 
ate Rev. Thomas Baker, a Wesleyan missionary, on 
July 21, 1867. 

Vatusila kondoi, new species. Figure 82a-c. 

Diagnosis. Shell very small, diameter 1.68-1.84 mm. (mean 
1.75 mm.), with 3%-3% normally coiled whorls. Apex and spire mod- 
erately elevated, slightly rounded above, last whorl descending much 
more rapidly, spire protrusion about Vs body whorl width, H/D ratio 
0.500-0.610 (mean 0.552). Apical sculpture of 9-11 (mean 10.3) 
rather fine spiral cords. Postnuclear whorls with high, prominent, 
very widely spaced, strongly protractively sinuated radial ribs, 
31-48 (mean 39.0) on the body whorl, whose interstices are 5-10 
times their width. Ribs/mm. 5.51-8.06 (mean 7.04). Microsculpture 
of barely visible radial and spiral riblets, almost completely obscured 
by prominent and crowded secondary spiral cording. Umbilicus nar- 
row, U-shaped, slightly and regularly decoiling, contained 4.12-5.40 
times (mean 4.72) in the diameter, margins rounded. Sutures deep, 
whorls strongly rounded above, flattened laterally with a weak su- 
praperipheral sulcus, basal margin evenly rounded and slightly 
compressed. Aperture ovate, inclined about 15 from shell axis. 
Parietal wall with single high barrier, extending Vn- 3 he of a whorl 
posteriorly, crescentic in outline with gradual anterior and posterior 
descension, usually flattened above on middle third, varying from 
slightly thickened medially to broadly bifid, with superior arm of 
barrier twisted upward. Columellar wall with low, broadly rounded, 
cordlike ridge, lying parallel to plane of coiling, reaching top of col- 
umellar callus, occasionally (5.6%) absent. Palatal barriers usually 
3, sometimes (11.1%) only 2, short, moderately to deeply recessed 
within aperture: lower basal in position, crescentic, broadly rounded 
above; 2nd much lower and slightly shorter; 3rd slightly sup- 
raperipheral, longer, narrower, more deeply recessed bladelike bar- 
rier with a very gradual anterior descension. Columellar and lower 
palatal walls with a rather thick callus. 

Vatusila kondoi (fig. 82a-c) is smaller and has 
much smaller apertural barriers than the Eua Island 
V. tongensis (fig. 83a-c). Vatusila nayauana (fig. 
82d-f) has a distinctly higher spire, more crowded ra- 
dial ribbing, a wider umbilicus (table XLII), and re- 
duced apertural barriers. Vatusila eniwetokensis (fig. 
85e-f) has much more numerous and crowded radial 
ribs, two parietals, and a wider umbilicus. 

Description. Shell very small, with 3'/2 moderately loosely 
coiled whorls. Apex and spire moderately elevated, rounded above, 
last whorl descending rapidly, H/D ratio 0.558. Apical whorls 1%, 
sculpture of 11 relatively prominent, narrow spiral ribs, partly 
eroded. Postnuclear whorls with widely spaced, protractively 
sinuated, narrow, rounded radial ribs, 34 on the body whorl, whose 
interstices are 5- 10 times their width. Microsculpture of very fine, 
close-set radial ribs, crossing quite strong, relatively prominent sec- 
ondary spiral cords. Microspiral sculpture barely visible. Major ribs 
beaded on top by crossing of spiral cords. Sutures moderately deep, 
whorls shouldered above, flattened laterally above periphery with 
slight basal flattening, a weak supraperipheral sulcus present. Um- 
bilicus narrowly U-shaped, regularly decoiling, slightly constricted 
by expansion of basal lip, contained 4.72 times in the diameter. Color 
uniform light yellow-brown. Aperture ovate, strongly flattened 
above periphery, slightly so on base, inclined about 15 from the shell 
axis. Parietal wall with a single high barrier, extending about 3 Ae of 
a whorl, gradually descending anteriorly and posteriorly, broadly 
expanded above and bifid in center. Columellar barrier a broad, 
rounded, moderately prominent ridge, lying parallel to plane of coil- 




a 





FIG. 82. a-c, Vatusila kondoi, new species. Station 101, Nayau, Lau, Fiji. Holotype. BPBM 180259; d-f, Vatusila nayauana, new species. 
Station 43, Nayau, Lau, Fiji. Holotype. BPBM 167232. Scale lines equal 1 mm. (MM). 



193 



194 



SOLEM: ENDODONTOID LAND SNAILS 



TABLE XLII. - RANGE OF VARIATION IN VATUSILA. 



NUMBER OF 
NAME SPECIMENS RIBS 



RIBS /MM. 



DIAMETER 



H/D RATIO 



kondoi 


56 


39. 


0(31-48) 


7 


,04(5.51-8.06) 


0. 


97(0.86-1 


,05) 


navauana 


42 


51. 


0(48-54) 


9 


,48(9.11-9.85) 


0. 


95(0.87-1 


.02) 


eniwe tokens is 


1 


89 




15 


,1 


0. 


86 




(Ladd) 


tongensis 


20 


61 


,0(52-72) 


7 


.66(6.89-8.71) 


1, 


36(1.13-1 


.72) 


vaitupuensis 


2 


115 




14 


.27 


1, 


25 




niueana 


1 


89 




11 


.60 


1. 


27 





1.75(1.68-1.84) 0.552(0.500-0.610) 

1.65(1.58-1.74) 0.573(0.552-0.608) 

1.88 0.456 

2.53(2.32-2.75) 0.538(0.472-0.650) 

2.57 0.487 

2.44 0.500 



kondoi 


WHORLS 
3 l/2+(3 3/8-3 7/8) 
4-(3 5/8-4 1/8) 
3 1/2 
4 5/8-(4 1/4-5) 
4 1/2+ 
It 1/4 


nayau. 
eniwe. 


tonge. 
vai tu. 


niuea. 



APICAL 
UMBILICUS D/U RATIO CORDS 

0.37(0.35-0.43) 4.72(4.12-5.40) 10.3(9-11) 

0.46(0.43-0.48) 3.61(3.43-3.69) 10.1(9-11) 

0.53 3.56 12 

0.76(0.66-0.89) 3.31(3.00-4.00) 10.4(8-12) 

0.89 2.89 8 

0.66 3.70 10 







SPIRE 




BODY WHORL 














ELEVATION 




UinTH 




SP/BW 


PR C 


p 


kondoi 





.10 


0. 


56 


0, 


177 


I 1 0-1 


2-3 


navau. 





17(0.13-0.23) 


0. 


53(0.48-0.56) 


0.310(0.260-0.420) 


1 0-1 


0-1 


eniwe. 




FLAT 


0. 


61 




FLAT 


2 ? 


7 


tonge. 





.20(0.16-0.30) 


0. 


76(0.63-0.82) 


0, 


270(0.200-0.375) 


I 1 


2 2 


vaitu. 


0.13 


0, 


79 





,167 


1 


2 


niuea. 





.13 


0. 


76 





.174 


1 


Callu: 



(Reduced) 



1. Bifid 

2. Enlarged laterally 



ing, reaching top of columellar callus. Palatal barriers 3: lower 
reaching edge of moderate apertural callus, short, >Ae whorl in 
length, a high, broadly rounded ridge; middle palatal slightly more 
deeply recessed, lower, narrower, an equally short threadlike ridge; 
upper slightly supraperipheral in position, a broad V-shaped ridge, 
short, with gradual anterior descension about as deeply recessed as 
middle palatal. All barriers minutely serrated above and on sides. 
Height of holotype 0.95 mm., diameter 1.71 mm. 

Holotype. Fiji: Lau Group, Nayau, Station 101, % 
mile north of Liku Village at base of a limestone cliff, 
100 yd. inland at 30-500 ft. elevation. Collected in leaf 
litter by Yoshio Kondo on August 22, 1938. BPBM 
180259. 

Range. Nayau, Lau Group, Fiji. 

Paratypes. Lau Group: Nayau, % mile north of 
Liku Village (Station 101) at base of a limestone cliff, 
100 yd. inland at 30-500 ft. elevation (56 specimens, 
BPBM 180259). 

Remarks. The parietal barrier in Vatusila kon- 
doi varies from a single lamellate ridge to a markedly 
bifid barrier as in the type (fig. 82b). The columellar 
barrier is often reduced in size, occasionally absent, 



and the middle palatal may be absent. These varia- 
tions are relatively minor and do not bridge the gap 
between the barriers of V. kondoi and V. nayauana. 
Differences in proportions (table XLII) are equally 
large, and I have no hesitation in placing them as dis- 
tinct species. 

Great pleasure is taken in naming this species 
after its collector, Dr. Yoshio Kondo. 

Vatusila nayauana, new species. Figure 82d-f. 

Diagnosis. Shell very small, diameter 1.58-1.74 mm. (mean 
1.65 mm.), with 3%-4'/8 rather tightly coiled whorls. Apex and spire 
rather strongly elevated, sometimes rounded above, last whorl de- 
scending distinctly more rapidly, spire protrusion about Va body 
whorl width, H/D ratio 0.552-0.608 (mean 0.573). Apical sculpture of 
9-11 (mean 10.1) fine spiral cords, whose interstices are about 3 
times their width. Postnuclear whorls with narrow, prominent, 
rounded, protractively sinuated radial ribs, 48-54 (mean 51.0) on the 
body whorl, whose interstices are 3-7 times their width. Ribs/mm. 
9.11-9.85 (mean 9.48). Microsculpture of extremely fine radial and 
spiral riblets, obscured by much more prominent, rather crowded 
secondary spiral cords. Umbilicus rather widely open, broadly U- 
shaped, last whorl decoiling more rapidly, contained 3.43-3.69 times 



SYSTEMATIC REVIEW 



195 



(mean 3.61) in the diameter, margins rounded. Sutures deep, whorls 
strongly rounded above, slightly flattened laterally above periphery, 
with evenly rounded lower margins. Aperture ovate, inclined about 
10 from shell axis. Parietal barrier a short, high, crescentic blade, 
extending posteriorly about Vs whorl, expanded and serrated above, 
with gradual anterior and posterior descension, medial in position. 
Columellar wall with (60%) or without (40%) a low, rounded cordlike 
ridge that barely reaches top of columellar callus. Palatal wall with 
(60%) or without (40%) a single, bladelike barrier, relatively low, 
strongly expanded and serrated above, with gradual anterior descen- 
sion, moderately recessed, situated on middle of lower palatal mar- 
gin. 

Vatusila nayauana (fig. 82d-f) differs from V. 
kondoi (fig. 82a-c) in its fewer and smaller apertural 
barriers, much wider umbilicus, more protruded spire, 
more crowded radial ribs, and greater H/D ratio (table 
XLII). Vatusila eniwetokensis (fig. 85e-f) has two 
parietals, much more numerous and crowded ribbing, 
plus a flat spire. Vatusila tongensis (fig. 83a-c) is much 
larger and has grossly developed apertural barriers. 

Description. Shell very small, with slightly over 4 rather 
tightly coiled whorls. Apex and spire moderately and evenly ele- 
vated, last whorl descending more rapidly, H/D ratio 0.608. Apical 
whorls 1%, sculpture of 11 narrow spiral ribs, whose interstices are 
about 3 times their width. Postnuclear whorls with protractively 
sinuated, widely spaced, prominent, rounded radial ribs, 48 on the 
body whorl, whose interstices are 3-5 times their width, becoming 
more crowded on the last half of body whorl. Microsculpture of very 
fine and crowded, inconspicuous radial riblets and lower, much more 
crowded spiral riblets, crossing stronger, rather widely spaced spiral 
cords. Major ribs finely beaded above by crossing of spiral cords. 
Sutures relatively deep, whorls strongly rounded above, slightly flat- 
tened laterally above periphery. Umbilicus open, broadly U-shaped, 
last whorl decoiling more rapidly, contained 3.64 times in the diame- 
ter. Aperture circular, inclined about 10 from the shell axis with a 
moderately prominent basal callus. Parietal wall with a short bar- 
rier extending about Vs whorl, broadly rounded and ridgelike. Col- 
umellar wall with a small, deeply recessed, cordlike tubercle. Palatal 
wall with a subperipheral, broadly rounded ridgelike barrier, mod- 
erately recessed, expanded above, extending about *Ae of a whorl. 
Major palatal and parietal minutely serrated above and on sides. 
Height of holotype 1.02 mm., diameter 1.68 mm. 

Holotype. Fiji: Lau Group, Nayau, northwest 
coast, Station 43, Nauko, hillside Vfe mile inland at 250 
ft. elevation. Collected dead at the base of a limestone 
cliff by H. S. Ladd on August 13, 1934. BPBM 167232. 

Range. Nayau, Lau Group, Fiji. 

Paratypes. Lau Group; Nauko, hillside, Vz mile 
inland (Station 43) at 250 ft. elevation (42 specimens, 
BPBM 167232). 

Remarks. Although Vatusila nayauana can be 
derived from V. kondoi by a combination of barrier 
reduction, greater spire protrusion, rib crowding, and 
umbilical widening, these changes are not correlated. 
Hence specific-level recognition is warranted. 

The exact relationship of the type localities is un- 
certain. Vatusila nayauana was taken on the north- 
west coast of Nayau, either at or on the way to Ladd's 
geological station L502-L504 (Ladd & Hoffmeister, 
1945, pp. 86, 190). Vatusila kondoi was taken "three- 
quarters mile north of Liku Village." The latter is lo- 
cated approximately in the middle of the southwest 
coast of the island. Although this locality is 100 yd. 
inland, Ladd's station was cited as Vfe mile inland. 



Hence, regardless of the relative shore proximity of the 
sites, they are separated at a minimum by about 700- 
800 yd. Thus the snails are isolated colonies. Only five 
examples were adult. Most of the rest had the aperture 
clogged with debris. Hence the proportion of specimens 
that lack the columellar and palatal barriers may be 
altered considerably when new collections are avail- 
able. 

Vatusila eniwetokensis (Ladd, 1958). Figure 85e-f. 

Ptychodon eniwetokensis Ladd, 1958, J. Paleont., 32 (1), p. 190, pi. 
30, figs. 9-12 Drill hole K-1B at 820-831 ft. depth, Engebi 
Islet, Eniwetok Atoll, Marshall Island (Miocene g). 

Description. Shell very small, probably with 4 whorls when 
complete (only 3'/2 remaining). Apex barely emergent, spire flat, last 
whorl descending more rapidly, H/D ratio of remnant 0.456. Apical 
whorls 1%, sculpture of 12 fine spiral ribs. Lower whorls with 
crowded, prominently narrow, protractively sinuated radial ribs, 89 
on last complete whorl, whose interstices are 2-3 times their width. 
Microsculpture with 2 elements: a greatly reduced microlattice visi- 
ble on a few areas under 96 x magnification and a much more promi- 
nent sculpture of rather widely spaced spiral cords (fig. 850. Sutures 
channeled above on postapical spire and body whorl, whorls strongly 
rounded above, slightly flattened laterally above and below rounded 
periphery, with evenly rounded baso-umbilical margin. Very faint 
brownish yellow cast to entire shell. Umbilicus cup-shaped, widely 
opened, last whorl decoiling more rapidly, contained 3.56 times in 
diameter. Last '/> whorl missing, hence status of any columellar or 
palatal dentition unknown. Parietal barriers 2, extending pos- 
teriorly about 3 Ae of a whorl: upper broken off for entire length, only 
remnants of base present; lower a broad, high, humped lamella with 
irregularly gradual anterior descension and even, rather steep, pos- 
terior descension. Aperture of whole specimen probably inclined 
about 15 from shell axis. Height of holotype 0.86 mm., diameter 1.88 
mm. 

Holotype. Marshall Islands: Eniwetok Atoll, En- 
gebi Islet, drill hole K-1B at 820-831 ft. depth 
(Miocene #). USNM 562091. 

Remarks. The spiral apical sculpture, secondary 
microsculpture of strong spiral cords on the lower 
whorls, and shape of the remaining parietal barriers 
immediately place Vatusila eniwetokensis in the 
Charopidae. 

Association with recent species is handicapped by 
the absence of the last half-whorl and resulting loss of 
any palatal barriers. The combination of 12 apical spi- 
ral ribs and a secondary microsculpture of very promi- 
nent and widely spaced spiral cords is relatively 
unusual in the present-day Micronesian charopid 
fauna. Taxa with approximately these features include 
only the Palline notera complex from Palau and the 
Vatusila radiation from Fiji (Nayau, Lau Group), 
Tonga (Eua), Niue, and Vaitupu, Ellice Islands. The 
Palline notera complex has simple parietal barriers 
(probably reduced), whereas Vatusila shows a tendency 
for fusion and reduction of two parietals. The basic 
shape of the parietals in V. tongensis is the same as in 
V. eniwetokensis. The latter could be transformed into 
the former by partial parietal fusion. Vatusila tongen- 
sis, V. kondoi, and V. nayauana share the very strong 
microsculpture of V. eniwetokensis. There are no fea- 
tures allying it to other genera, and I have no hesita- 
tion in placing it with Vatusila. 



196 



SOLEM: ENDODONTOID LAND SNAILS 



Vatusila tongensis, new species. Figures 83a-c, 
84a-b. 

Diagnosis. Shell large, diameter 2.32-2.75 mm. (mean 2.53 
mm.), with 4'A-5 tightly coiled whorls. Apex and spire moderately 
and evenly elevated, body whorl descending much more rapidly, 
spire protrusion more than Vt body whorl width, H/D ratio 0.472- 
0.650 (mean 0.538). Apical sculpture of 8-12 (mean 10.4) narrow but 
prominent spiral cords whose interstices are 3-4 times their width. 
Postnuclear sculpture of high, narrow, rather widely spaced, protrac- 
tively sinuated radial ribs, 52-72 (mean 61) on the body whorl, 
whose interstices are 4-6 times their width. Ribs/mm. 6.89-8.71 
(mean 7.66). Microsculpture of extremely fine radial riblets, more 
than 12 between each pair of major ribs, even finer and more crowded 
spiral riblets, with a secondary microsculpture of prominent spiral 
cords about twice as large as the microradials, whose interstices are 
1-3 times their width. Umbilicus broadly open, U-shaped, regularly 
decoiling, contained 3.00-4.00 times (mean 3.31) in the diameter, 
margins rounded. Sutures deep, whorls strongly rounded above, 
markedly compressed laterally and slightly on basal margin. Aper- 
ture subcircular. inclined about 10 from shell axis. Parietal wall 
with single bifurcated barrier, extending posteriorly more than 3 Ae of 
a whorl: upper blade very high and crescentic, gradual posterior 
descension, rather sharp anterior descension, lower blade shorter 
posteriorly, elevated medially with same anterior descension found 
in Graeffedon. Both blades mounted on a broad callus that tapers 
laterally beyond anterior end of bifurcated blade. Columellar wall 
without barriers. Palatal barriers 2, rather deeply recessed, bulky, 
extending about '/a whorl posteriorly: 1st basal in position, slightly to 
strongly bifurcated, occasionally split into two parts, inner a broad, 
high, crescentic, rounded, knoblike protuberance usually connected 
by a callus to a much lower, cordlike blade; 2nd on outer palatal wall, 
slightly supraperipheral and subperipheral to completely sub- 
peripheral, about 0.25-0.36 mm. wide, crescentic in outline, with 
rather gradual anterior descension, sometimes slightly concave on 
upper surface with threadlike lamellae. 

Vatusila tongensis is immediately recognizable by 
its very large, bifurcated parietal and two very broad 
palatal barriers. All other Vatusila have very slender 
palatals. No other Pacific Island endodontoids match 
the palatal barriers. Vatusila vaitupuensis and V. 
niueana both have a less protruded spire, much more 
crowded radial ribbing, and much less conspicuous 
apertural barriers. 

Description. Shell large, with 4 7 /s tightly coiled whorls. Apex 
and spire very slightly elevated, last whorl descending moderately, 
H/D ratio 0.519. Embryonic whorls l'/2, sculpture mainly eroded with 
only traces of spiral cording remaining in sutures. Postnuclear 
whorls with sculpture of widely spaced, high, protractively sinuated 
radial ribs, rounded above, 72 on the body whorl, whose interstices 
are 3-5 times their width. Microsculpture of very low and faint ra- 
dial and spiral riblets crossing much more prominent, rather widely 
spaced secondary spiral cords. Sutures deep, whorls slightly shoul- 
dered above, laterally compressed on outer margins. Umbilicus 
broadly U-shaped, regularly decoiling, contained 3.00 times in the 
diameter. Color light yellow-white with broad, somewhat vague 
flammulations that follow lines of growth. Aperture ovate, flattened 
laterally, inclined about 5 from shell axis. Parietal wall with a 
large, crescent-shaped bifid barrier, extending about '/s whorl, 
structure as in diagnosis. Columellar wall without barriers. Palatal 
wall with 2 oddly shaped, recessed barriers: lower a moderately high, 
irregularly shaped knoblike protuberance with much lower recessed 
ridgelike outer portion; outer a low, very broad, 0.36 mm. wide cal- 
lus, extending about VLB of a whorl. All barriers are very minutely 
serrated on all portions. Height of holotype 1.38 mm., diameter 2.67 
mm. 

Holotype. Tonga: Eua, hillside on cliffs east of 
Ohunua, 2 miles inland at 390 ft. elevation. Collected 
by H. S. Ladd on May 10, 1928. BPBM 87705. 



Range. Eua Island, Tonga. 

Paratypes. Eua: cliffs east of Ohunua, 2 miles in- 
land at 390 ft. elevation (9 specimens, BPBM 87705, 
BPBM 87660); on cliff, V 2 mile inland from Vaigana at 
760 ft. elevation (9 specimens, BPBM 87632, BPBM 
87693-4); main range on east side (Station T-22) at 
1,000 ft. elevation (1 specimen, FMNH 152376). 

Remarks. The very peculiar barriers of Vatusila 
tongensis immediately identified this species. The bifid 
parietal probably originated through partial fusion of 
two separate barriers, a process that seems to have 
been carried further in the Ellice Island V. uaitupuen- 
sis and the Fijian V. kondoi. The palatal barriers vary 
considerably, the lower one usually being low and 
ridgelike with a lateral fanglike barrier, but often as- 
suming the knoblike form of the type. The outer 
palatal varies from a flattened, even callus as in the 
type, to a slanted, basal callus with three or four low, 
threadlike lamellae on top of it, but generally is with- 
out distinct ribbing above. 

Fragmentary soft parts were available and are de- 
scribed below. 

Description of soft parts. Several fragmentary examples were 
seen, three of which had the penis and epiphallus present. Vas defer- 
ens (fig. 84a, VD) reflexing at penioviducal angle, entering expanded 
head of epiphallus (E). Latter more than % length of penis, entering 
latter through fan of penial retractor muscle (PR). Penis 0.9-1.0 mm. 
long, expanded apically, a strong median bulge, tapering gradually 
to rather long and slender atrium (Y). Interior (fig. 84b) of penis with 
a circularly bilobed verge and superior epiphallic pore, biscuit-like 
stimulator very large and with narrow central crack. 

(Based on BPBM 87660 and BPBM 87705, fragmentary parts.) 

Vatusila vaitupuensis, new species. Figure 85a-d. 

Diagnosis. Vatusila vaitupuensis has 1 parietal and 2 small 
palatal barriers, a wider umbilicus, and much more numerous and 
crowded radial ribs than the Tongan V. tongensis. Vatusila niueana 
has a narrower umbilicus, only traces of apertural barriers, and 
many more radial ribs than V. vaitupuensis. 

Description. Shell large, with over 4'/2 relatively tightly coiled 
whorls. Apex and spire barely protruding above body whorl, last 
whorl descending slightly more rapidly, spire protrusion about Ve 
body whorl width, H/D ratio 0.487. Apical whorls 1%, sculpture of 8 
relatively broad, prominent spiral cords, whose interstices are about 
2*/2 times their width. Postnuclear whorls with low, rounded, 
crowded, protractively sinuated radial ribs, 115 on the body whorl, 
whose interstices are 2-3 times their width. Microsculpture of fine 
radial riblets, 3-5 between each pair of major ribs and very incon- 
spicuous crowded spiral riblets. No secondary spiral cording. Sutures 
relatively shallow, whorls slightly shouldered above, almost evenly 
rounded on outer margins, very slightly compressed laterally above 
and below periphery. Umbilicus broadly U-shaped, regularly decoil- 
ing, contained 2.89 times in the diameter, margins rounded. Color 
yellowish white with relatively regular, broad, reddish flammula- 
tions that parallel lines of growth. Aperture circular, inclined about 
5 from the shell axis. Parietal wall with 1 deeply recessed, low, 
crescentic barrier (fig. 85d), extending less than 'Ae of a whorl. Col- 
umellar wall without barriers. Palatal barriers 2: lower a short, 
broad, cordlike ridge near baso-columellar margin; upper slightly 
supraperipheral, moderately recessed, extending about Vs whorl, a 
low, V-shaped ridge. Height of holotype 1.25 mm., diameter 2.57 mm. 

Holotype. Ellice Islands: Vaitupu. Collected by L. 
Isaacs in May, 1941. BPBM 189681. 
Range. Vaitupu, Ellice Islands. 









FIG. 83. a-c, Vatusila tongensis, new species. East of Ohunua, Eua, Tonga. Holotype. BPBM 87705; d-f, Vatusila niueana, new species. 
Alofi, Niue. Holotype. BPBM 194517. Scale lines equal 1 mm. (a-c, MM; d-f, SH). 



197 



198 



SOLEM: ENDODONTOID LAND SNAILS 



PR 




VD 




VD 



FIG. 84. Partial anatomy of Vatusila tongensis, new species. Eua, 
Tonga. BPBM 87660, BPBM 87705: a, penis and epiphallus; b, vergic 
papilla and stimulatory pad as seen by opening penis at junction 
between collar and pad. Scale line equals 1 mm. (MM). 



Paratype. Vaitupu (1 specimen, BPBM 189681). 

Remarks. Only the adult holotype and a single 
juvenile specimen are known. The barriers in the latter 
are relatively more prominent than in the adult, but 
occupy the same position and have the same shape. The 
parietal (fig. 85d) closely resembles the upper blade of 
V. tongensis in outline (fig. 83b), but is reduced in size. 
Similarly, the palatal barriers in V. vaitupuensis oc- 
cupy the same positions as the palatals of V. tongensis, 
but are greatly reduced in prominence. 



Vatusila niueana, new species. Figure 83d-f. 

Diagnosis. Vatusila niueana has the apertural barriers re- 
duced to barely visible traces. It differs from V. vaitupuensis (fig. 
85a-d) also in having secondary spiral cording, a much narrower 
umbilicus, and fewer, less crowded radial ribs. Vatusila tongensis 
(fig. 83a-c) has very large apertural barriers, much more widely 
spaced radial ribbing, and a distinctly more elevated spire. 
Differences from geographically adjacent Sinployea are discussed 
below. 

Description. Shell large, with 4V4 rather tightly coiled whorls. 
Apex and spire moderately and almost evenly elevated, last whorl 
descending much more rapidly, spire protrusion more than l k body 
whorl width, H/D ratio 0.500. Apical whorls l'/2, sculpture of 10 rela- 
tively prominent spiral cords whose interstices are about 3 times 
their width. Postnuclear whorls with prominent, rounded, rather 
closely spaced, protractively sinuated radial ribs, 89 on the body 
whorl, whose interstices are 2-3 times their width. Ribs/mm. 11.60. 
Microsculpture of extremely fine radial riblets, 5-10 between each 
pair of major ribs, slightly finer spiral riblets, and weak, rather 
widely spaced secondary spiral cording. Sutures moderately im- 
pressed, whorls evenly rounded on outer margins. Umbilicus open, 
broadly U-shaped, last whorl decoiling slightly more rapidly, con- 
tained 3.70 times in the diameter, margins rounded. Color light 
yellow-white, with a few irregular, reddish flammulations that fade 
out below periphery. Aperture circular, inclined about 10 from shell 
axis. Parietal wall with low, broadly angled, slightly submedial 
swelling, visible from normal side view only as an irregularity in 
outline (fig. 83e). Columellar, basal, and lower palatal wall with a 
white callus, narrower and higher at first, becoming broader, lower 
and more deeply recessed up to periphery of palatal wall. Height of 
holotype 1.27 mm., diameter 2.44 mm. 

Holotype. Niue Island: Alofi Village at 20 m. ele- 
vation. Collected on Pilea microphylla Liebm. by T. G. 
Yuncker on February 12, 1940. BPBM 194517. 

Range. Niue Island. 

Remarks. Even with reasonably careful inspec- 
tion, Vatusila niueana (fig. 83d-f) can be misidentified 
as a Sinployea. The parietal swelling and palatal callus 
are visible only by tilting and use of oblique lighting 
into the aperture. I consider it to represent another 
stage in barrier reduction within the genus Vatusila. 
Although the shell measurements fall just within the 
range of Sinployea uicaria and S. ellicensis (table 
XXIX), in form and whorl contour V. niueana is much 
closer to the other Vatusila, the Sinployea showing lat- 
eral whorl flattening and a higher spire protrusion 
(uicaria, fig. 58b) or much narrower umbilicus (ellicen- 
sis, fig. 62c). Unfortunately, only the single example, 
with no soft parts present, was available. 






a be 



FIG. 85. a-d, Vatusila vaitupuensis, new species. Vaitupu, Ellice Islands. Holotype. BPBM 189681: a-c, entire shell, d, parietal barrier; e-f, 
Vatusila eniwetokensis (Ladd). Drillhole K-lb at 820-831 ft. depth, Engebi Islet, Eniwetok, Marshall Islands. Holotype. USNM 562091. e, lateral 
view of parietal barriers;/; diagram of microsculpture. Scale line equals 1 mm.; d-f greatly enlarged. (MM). 



199 



200 



SOLEM: ENDODONTOID LAND SNAILS 



Genus Graeffedon, new genus 

Shell large, with 4V&-5V4 normally coiled whorls. Apex and spire 
flat to slightly elevated, body whorl very slightly to moderately de- 
scending. Apical sculpture of 1 1- 13 narrow, prominent spiral cords. 
Postnuclear sculpture of medium to very prominent, protractively 
sinuated radial ribs, with fine radial and finer microspiral elements, 
but no spiral cording. Periphery rounded, body whorl slightly flat- 
tened laterally above periphery. Umbilicus open, regularly decoiling. 
Color flammulations zigzagged. Parietal wall with 2 barriers, ex- 
tending posteriorly about Vn whorl, upper with rather sharp anterior 
descension, lower with more gradual anterior descension. Columel- 
lar wall with or without a low threadlike ridge parallel to plane of 
coiling, recessed or reaching lip margin. Palatal barriers 4 to 7, 
short, high, almost reaching lip edge, sometimes with weak acces- 
sory traces. Pallial region with rectal kidney lobe much larger and 
longer than pericardia!, a distinct intrusion of mantle gland tissue 
onto pallial roof. Apical genitalia apparently without unusual fea- 
tures. Epiphallus with simple entrance of vas deferens, followed by 2 
high pilasters. Penis with an apical muscle ring, and 3 large pilas- 
ters, 1 probably acting as a vergic papilla. Lower female tract with 
simple pore opening from free oviduct into common chamber of vag- 
ina and spermatheca, which is lined with high, longitudinal pilas- 
ters. 

Type species. Endodonta graeffei Mousson, 1869. 

The most similar shells are those of Semperdon, 
particularly examples of S. heptaptychius (Quadras & 
Mollendorff, 1894) (fig. 99a-c, e) and S. rotanus (fig. 
99d). Their penial anatomy (fig. lOOb, d-g, k) is so 
totally different from that of Graeffedon (fig. 87b c) 
that the shell similarities clearly are convergent. Even 
optical microscopic examination of the apertural bar- 
riers shows that there are differences in microarma- 
ture between the two genera, and the SEM photos of 
Graeffedon (fig. 7a-c) and Semperdon xyleborus (fig. 
8a-e) demonstrate that they are formed quite 
differently (see pp. 19-20). 

Graeffedon stands quite isolated from other Pacific 
Island Charopinae in terms of anatomical structure. 
The simple vas deferens-epiphallus juncture, very 
large penial pilasters, and strong pilasters within the 
vaginal-spermathecal chamber are quite different from 
those found in the typical Charopinae. It is quite possi- 
ble that eventual subfamily separation of Graeffedon 
will be required, but until many more extralimital 
taxa have been studied, retention of Graeffedon within 
a broadly defined Charopinae is preferred. 

The only genus with which I can at present relate 
Graeffedon in terms of anatomy is Helenoconcha 
Pilsbry, 1892, from St. Helena in the South Atlantic. 
The only dissected species, Helenoconcha relicta Solem 
(1977c, figs. 184a-b, 185a-d), agrees in pallial 
structure (except for lacking the mantle gland intru- 
sion) and has exactly the same type of penial structure. 
The two taxa differ only in details of pilaster position 
and size, but the penis complex of Helenoconcha is re- 
markably similar to Graeffedon. At present the inter- 
nal structures of the terminal genitalia in the South 
African endodontoid taxa, Trachycystis Pilsbry, 1893, 
and Afrodonta Melville & Ponsonby, 1908, are un- 
known (see Solem, 1970c), but the external genital fea- 
tures of Trachycystis illustrated by Watson (1934, pis. 
20-21) are compatible with at least the latter genus 
being a member of this complex. 



It is thus premature to attempt formal designation 
of a subfamily unit for Graeffedon. 

Specimens of Graeffedon are rare in collections 
and in the field. The three species differ in size, 
sculpture, and details of the palatal barriers (see Diag- 
noses). Because of their allopatric distribution, no 
structural key is presented. Their known distributions 
are: 

Graeffedon graeffei Upolu, Samoa 

G. sauaiiensis Savai'i, Samoa 

G. pricei Tongatapu, Tonga. 

Graeffedon graeffei (Mousson, 1869). Figures 7a-c, 
86a-c, 87a-d. 

Endodonta graeffei Mousson, 1869, J. de Conchyl. 17, pp. 332-333, 

pi. 14, fig. 3 Upolu, Samoa. 
Pitys graeffei (Mousson), Pease, 1872, Proc. Zool. Soc. London, 

1871, p. 474; Garrett, 1887, Proc. Acad. Nat. Sci. Philadelphia, 

1887, pp. 130-131. 
Helix (Endodonta) graeffei (Mousson), Pfeiffer, 1876, Monog. helic. 

viv., 7, p. 258; Tryon, 1887, Man. Conchol., (2) 3, p. 65, pi. 12, fig. 

31. 
Endodonta (Thaumatodon) graeffei (Mousson), Pilsbry, 1893, Man. 

Conchol., (2)9, p. 27. 

Diagnosis. Shell very large, diameter 4.61-5.92 mm. (mean 
5.23 mm.), with 4%-5'/4 rather tightly coiled whorls. Apex and early 
spire flat or barely protruding, lower spire descending slightly, body 
whorl much more rapidly, spire protrusion about J Ai body whorl 
width, H/D ratio 0.472-0.560 (mean 0.525). Apical sculpture of 
11-13 (mean 12.0) narrow but prominent spiral cords whose in- 
terstices are 4-5 times their width. Postnuclear sculpture of high, 
prominent, V-shaped to U-shaped, crowded, slightly protractively 
sinuated radial ribs, 92-121 (mean 105.0) on the body whorl, whose 
interstices are 2-3 times their width. Ribs/mm. 6.27-6.51 (mean 
6.39). Microsculpture of proportionately fine radial riblets, 5-8 be- 
tween each pair of major ribs, crossed by much finer and more 
crowded spiral riblets. No secondary spiral cording. Umbilicus mod- 
erately open, V-shaped, regularly decoiling, contained 3.53-4.19 
times (mean 3.94) in the diameter, margins rounded. Whorls dis- 
tinctly flattened laterally above periphery and on basal margin, 
aperture inclined about 15 from shell axis. Parietal barriers 2, ex- 
tending posteriorly slightly more than VH whorl: upper very high and 
broadly expanded above posteriorly, this portion deflected down- 
ward, then gradually descending and narrowing to anterior quarter, 
then abruptly descending on anterior 8th; lower broadly expanded, 
serrated, and crescentic on posterior half, narrowing above and with 
even descension anteriorly. Columellar wall with (40% ) or without 
(60% ) a low, broadly rounded, high, threadlike ridge, lying parallel 
to plane of coiling and reaching lip margin. Palatal barriers 5, high 
and prominent, reaching almost to lip margin: lower 4 progressively 
higher, longer, more slender, changing from abrupt to more gradual 
anterior descension, all crescentic, with height about % length, ex- 
tending posteriorly less than Vs whorl, with 1st at basal-columellar 
margin; 4th subperipheral; lower 2 have anterior edge turned inward 
just before reaching lip; upper supraperipheral, greatly reduced in 
height, a bladelike barrier with equally gradual anterior and pos- 
terior descension, longer than 4th, and slightly recessed within aper- 
ture. 

Graeffedon graeffei is much larger than the Savai'i 
Island G. sauaiiensis or the Tongan G. pricei (table 
XLIII). The former has a much wider umbilicus, seven 
palatal barriers, finer sculpture, an elevated apex, and 
more rounded whorls; the latter has fewer and more 
prominent radial ribs, only four palatal barriers, a flat 
apex, and is smaller in size. 



SYSTEMATIC REVIEW 



201 




a 





FIG. 86. a-c, Graeffedon graeffei (Mousson). Upolu, Samoa. Lec- 
totype. Zoologisches Institut der Universitat Zurich. Scale line 
equals 1 mm. (SG). 



Description. Shell very large, with slightly more than 4% mod- 
erately tightly coiled whorls. Apex and early spire flat, later whorls 
descending a little, body whorl more rapidly, H/D ratio 0.527. Apical 
whorls 1%, sculpture of 12 narrow cords. Postnuclear sculpture of 
high, prominent, crowded, protractively sinuated radial ribs, 98 on 
the body whorl, whose interstices are 2-3 times their width. Micro- 
sculpture a lattice of fine, relatively crowded and numerous radial 
riblets, 5-8 between each pair of major ribs, crossed by much finer 
and more crowded spiral riblets. Sutures deeply impressed, whorls 
strongly rounded above, somewhat flattened laterally above periph- 
ery and slightly on basal margin. Umbilicus moderately open, V- 
shaped, regularly decoiling, contained 3.93 times in the diameter. 
Color light yellowish white with very prominent zigzag reddish 
brown flammulations. Aperture subovate, flattened laterally above 
periphery, inclined about 15 from the shell axis. Parietal barriers 2, 
extending a little more than '/ whorl: upper high, bladelike, slightly 
expanded above posteriorly with relatively sharp anterior descen- 
sion; lower more crescentic, more broadly expanded above with com- 
paratively gradual anterior descension. Columellar wall without 
barriers. Palatal barriers 5, reaching lip margin, short and crescen- 
tic: lower at baso-columellar margin, sharply descending to lip edge; 
2nd, 3rd, and 4th crescentic, very high, slightly expanded above, 2nd 
reaching lip margin with very sharp anterior descension, 3rd slightly 
recessed with more gradual descension, 4th a little more recessed 
with much more gradual anterior descension; upper a longer, lower, 
ridgelike lamella, narrow above with very gradual posterior rise and 
anterior descension. All barriers minutely pustulose or serrated 
above. Basal and columellar portion of lip with a low, broad, white 
callus. Height of holotype 2.60 mm., diameter 4.94 mm. 

Holotype. Samoa: Upolu. Collected by E. GraefFe. 
Zoologisches Museum der Universitat, Zurich. 

Range. Probably the uplands of Upolu, Western 
Samoa. 

Material. Upolu (4 specimens, Zurich, Brussels, 
FMNH 116976, FMNH 147544): summit of Mt. Solaua 
(Station 39) under ground debris in heavy forest at 
2,000 ft. elevation (1 specimen, FMNH 153420). 

Remarks. Besides the type in the Mousson col- 
lection, two specimens were found in museums iden- 
tified as Sinployea complementaria (Mousson, 1865). A 
single example was collected under rotting wood in 
1965 together with 18 specimens of S. complementaria. 
In size and general appearance (figs. 56a-f, 86a-c) the 
two species are very similar, with the elevated spire 
and narrower umbilicus of S. complementaria (fig. 
56a-f) not being obvious during field examination. Ex- 
ternally the animal of Graeffedon graeffei differs in 
having the body without any grayish markings on the 
neck and sides of the foot. The absence of apertural 
barriers in S. complementaria and their presence in G. 
graeffei immediately separate the two species. 

In order to dissect the one specimen, partial de- 
struction of the shell was necessary. The soft parts 
broke during extraction, but reasonably full study of 
the anatomy was possible. The significant differences 
have been discussed previously (p. 29). 

Description of soft parts. Foot and tail completely retracted 
into shell. Sole and pedal grooves typical, no caudal horn or middor- 
sal groove developed. Slime network conspicuous. Body color 
yellow-white, no trace of darker markings. 

Mantle collar (MC) partly shredded during extraction because of 
large barriers, a prominent glandular extension (MG) onto pallial 
roof. Anus (A) opening slightly in front of external ureteric pore 
(KX), vague grooves leading through pneumostomal opening. 



202 



SOLEM: ENDODONTOID LAND SNAILS 

TABLE XLIII. - RANGE OF VARIATION IN GRAEFFEDON. 



NUMBER OF 
SPECIMENS 



graeffei 

(Mousson) 

savailensis 
price! 



RIBS 
105(92-121) 

88 
68 



RIBS/MM. 
6.39(6.27-6.51) 



H/D RATIO 

graef. 0.525(0.472-0.560) 
savai. 0.485 
pricei 0.485 



HEIGHT 
2.75(2.37-3.26) 

1.61 
1.61 



DIAMETER 
5.23(4.61-5.92) 

3.32 
3.32 



APICAL 

WHORLS UMBILICUS D/U RATIO CORDS 

5-(4 3/4-5 1/4) 1.36(1.15-1.48) 3.94(3.53-4.19) 12.0(11-13) 

4 1/4 1.02 3.26 11 

4 1/8- 0.92 3.61 WORN 



SPIRE 
ELEVATION 

graef. 0.17(0.10-0.20) 
saval. 0.15 
pricei FLAT 



BODY WHORL 
WIDTH 

1.83(1.68-2.04) 

1.05 

1.13 



SP/BWW PR 

0.092(0.058-0.118) 2 

0.141 2 

2 



P 

5 

7 

4+2 



Pallial region (fig. 87a) extending about 5.5 mm., distance from 
anus to anterior tip of kidney 1.65 mm. Lung roof clear, without 
granulations. Kidney (K) bilobed, pericardia! lobe less than half 
length of rectal lobe, very narrow, rectal lobe crossing hindgut into 
parietal wall, length 3.85 mm. Ureter (KD) with short primary arm 
jammed between lobes of kidney with no lung roof visible, secondary 
ureter widening along hindgut. Heart (H) angled to hindgut, slightly 
more than half length of pericardia! kidney lobe. Principal pulmo- 
nary vein (HV) unbranched and inconspicuous, reaching to edge of 
mantle collar extension. Hindgut (HG) extending well past pallial 
cavity apex before leaving parietal-palatal margin. 

Ovotestis (fig. 87d, G) a single clump of large, long acini, pal- 
mately clavate, lying parallel to whorl sides above stomach apex. 
Hermaphroditic duct (GD) long, expanded evenly for most of length, 
abruptly narrowing before reflexing to enter base of globular talon. 
Albumen gland completely destroyed by parasites in dissected 
example. Talon (not shown) large, globular, with very narrow neck to 
carrefour. Prostate (fig. 87b, DG) of large acini opening into groove in 
inner uterine wall, variously folded. Uterus (UT) bipartite, upper 
chamber thin-walled and narrow, lower grossly expanded and 
thick-walled. 

Vas deferens (VD) tapering and with glandular walls to 
penioviducal angle, narrow and muscular after passing through 
muscle threads, entering epiphallus without special structures. 
Epiphallus (E) larger than penis, curved and twisted, internally (fig. 
87c) with one very high and rounded and a second lower and hemi- 
spherical pilaster that taper into epiphallic pore. Penial retractor 
(PR) a broad muscle band arising from diaphragm and inserting in a 
narrow line onto head of penis along middle zone of apical penial 
pilaster. Penis (P) about 1.48 mm. long, sharply constricted subapi- 
cally, then tapering. Internally (fig. 87c) with a dome-shaped pilaster 
and epiphallic pore (EP) on one side of center occupying area above 
constriction, below with a huge spade-shaped pilaster tapering to 
atrium. Lateral to this a "hoe"-shaped pilaster tapering to a broad 
glandular ridge and a high, longitudinal pilaster tapering into glan- 
dular area. Atrium (Y) short, without marked features. 

Free oviduct (UV) muscular, short, with glandular walls. Sper- 
matheca (S) with expanded head next to albumen gland, slender 
shaft bound to prostate-uterus, expanded basal portion with 2 large 



and several smaller tapered pilasters (fig. 87c). Vagina (V) with 
many narrow pilasters, externally with numerous muscle strands 
holding it to the body wall. 

Esophagus distended, typical in position, stomach occupying % 
of a whorl. Intestinal looping without unusual features. Digestive 
glands mostly replaced by parasites above stomach apex, partly in 
region of esophagus-stomach junction. Salivary glands united above 
esophagus, right gland greatly reduced in size. 

(Based on 1 adult, FMNH 153420.) 

Graeffedon savaiiensis, new species. Figure 88a-c. 

Diagnosis. Graeffedon sauaiiensis differs from the Upolu G. 
graeffei (Mousson, 1869) in its slightly and regularly elevated apex, 
possession of 7 palatal barriers, wider umbilicus, and much smaller 
size. The Tongan G. pricei (fig. 88d-f) has a flat apex and spire, much 
coarser radial ribbing, lower and shorter parietals, and only 4 major 
palatals, although essentially identical in size and proportions. 

Description. Shell slightly larger than average, with 4V* nor- 
mally coiled whorls. Apex and spire slightly and evenly elevated, last 
whorl not descending more rapidly, spire protrusion about l h body 
whorl width, H/D ratio 0.485. Apical whorls 1V4, sculpture of 11 thin, 
relatively high, spiral cords. Postnuclear whorls with thin, lamel- 
late, protractively sinuated radial ribs, 88 on the body whorl, whose 
interstices are 2-4 times their width. Ribs/mm. 8.43. Microsculpture 
of fine radial riblets, 3-6 between each pair of major ribs, with much 
finer and more crowded spiral riblets. Sutures moderately impressed, 
whorls strongly rounded above suture, slightly flattened laterally 
and on basal margin with evenly rounded periphery. Umbilicus 
widely open, V-shaped, last whorl decoiling more rapidly, contained 
3.26 times in the diameter. Color light yellow-white with irregular 
reddish brown flammulations becoming darker on body whorl. Aper- 
ture ovate, slightly flattened laterally above periphery, inclined 
about 10 from shell axis. Parietal barriers 2, extending posteriorly 
% whorl: upper very high and crescentic posteriorly, serrated and 
expanded above, flattening out anteriorly, then descending rather 
sharply on anterior 8th; 2nd a much lower, bladelike lamella, 
broadly expanded above and slightly elevated posteriorly, with 
gradual descension over anterior half. Columellar wall with single, 



H 



VD 



UV 




FIG. 87. Anatomy of the Samoan Graeffedon graeffei (Mousson). Station 39, Mt. Solaua, Upolu, Samoa. FMNH 153420: a, pallial region; b, 
genitalia; c, interior of penis, epiphallus, spermathecal base; d, ovotestis and hermaphroditic duct. Scale lines equal 1 mm. (SH). 



203 








def 




FIG. 88. a-c, Graeffedon savaiiensis, new species. Mataulano Lake, Savai'i, Samoa. Holotype. BPBM 95738; d-f, Graeffedon pricei, new 
species. Manima Id., Tongatapu, Tonga. Holotype. FMNH 152399. Scale lines equal 1 mm. (a-c, MM; d-f, SH). 



204 



SYSTEMATIC REVIEW 



205 



low, threadlike ridge, lying parallel to plane of coiling, reaching top 
of columellar callus. Palatal barriers 7, short, high, reaching lip 
margin for early ones, then later barriers slightly recessed: lower at 
baso-columellar margin, hemispherical, very high, with abrupt an- 
terior descension, curving back inward at lower anterior margin; 
3rd, 4th, and 5th progressively lower, longer, less expanded above, 
changing to more gradual anterior descension with 5th crescentic, all 
subperipheral; 6th equal in height to 5th, slightly supraperipheral, 
with more gradual anterior descension; 7th high, bladelike, much 
lower than 6th, slightly concave above, moderately expanded, with 
gradual anterior descension, injured anteriorly, slightly recessed 
within aperture. All palatals weakly serrated on upper edge and 
sides. Height of holotype 1.61 mm., diameter 3.32 mm. 

Holotype. Samoa: Savai'i, near Mataulano Lake. 
Collected in leaf mould on a hillside by E. Christopher- 
son on October 2-3, 1929. BPBM 95758. 

Range. Interior of Savai'i, Western Samoa. 

Material. Known only from the holotype. 

Remarks. The elevation of Mt. Mataulano is 
2,952 ft., but I have no data as to the elevation of the 
lake at which this species was collected. The single 
shell is probably subadult, but the distinctness from G. 
graeffei is based on the former having smaller nuclear 
whorls, a flat apex and early spire, wider and higher 
major radial ribs, and a noticeably narrower umbilicus. 
Shape of the parietal barriers is identical, but in G. 
savaiiensis they are shorter and proportionately 
higher. Besides the increased number of palatals, in G. 
savaiiensis they are shorter and hemispherical in out- 
line, rather than being crescentic. The columellar in G. 
savaiiensis is lower and more deeply recessed. 

Despite particular effort by Mr. Price, no speci- 
mens were collected on Savai'i in 1965. 

Graeffedon pricei, new species. Figure 88d-f. 

Diagnosis. Graeffedon pricei differs from the Savai'i G. 
savaiiensis in its flat apex and spire, heavier and less crowded rib- 
bing, smaller parietals, and presence of only 4 palatals. Graeffedon 
graeffei from Upolu is much larger, has the body whorl descending 
much more rapidly, and has proportionately much larger apertural 
barriers. 

Description. Shell slightly larger than average, with a little 
less than 4'/s rather tightly coiled whorls. Apex and spire flat, body 
whorl descending slightly, H/D ratio 0.485. Apical whorls 1%, 
sculpture eroded except for traces of narrow spiral cords in sutures 
and umbilicus. Postnuclear whorls with very prominent, wide, pro- 
tractively sinuated radial ribs, 68 on the body whorl, whose in- 
terstices usually are 2-4 times their width. Microsculpture mostly 
obscured, visible on spots as consisting of very fine radial riblets, 
6-12 between each pair of major ribs, crossed by slightly finer and 
more crowded spiral riblets. Sutures impressed, whorls strongly 
rounded above and on periphery, slightly compressed laterally above 
and below periphery. Color leached from shell, with faint traces of 
reddish flammulations visible above periphery. Umbilicus open, 
broadly V-shaped, regularly decoiling, contained 3.61 times in the 
diameter. Aperture circular, slightly compressed laterally above and 
below rounded periphery, inclined about 10 from shell axis. Parietal 
barriers 2, relatively low, extending posteriorly '/* whorl: upper ele- 
vated on posterior quarter, expanded and serrated above, gradually 
descending to anterior threadlike quarter; 2nd equal in height, 
slightly more expanded above on elevated posterior quarter, rather 
sharply descending to midpoint, anterior half a high threadlike 
ridge. Columellar wall without barriers. Palatal barriers 4, high, 
bulbously expanded above, short, reaching lip edge, with two thread- 
like accessory traces: lower at baso-columellar margin, hemispheri- 
cal, anterior descension abrupt, margin deflected inward just before 



lip; 2nd slightly higher, equally expanded above, with sharp anterior 
descension, but no inward deflection; 3rd equal in height, slightly 
subperipheral, with more gradual anterior descension; 4th supra- 
peripheral, reduced in height, a bladelike lamella strongly expanded 
above, slightly recessed, with gradual anterior descension. Accessory 
traces low and threadlike, located midway between 1st and 2nd, 2nd 
and 3rd palatals, slightly recessed, shorter than major palatals. 
Height of holotype 1.61 mm., diameter 3.32 mm. 

Holotype. Tonga: Tongatapu, Manima Island, 4 
miles northeast (Station T-4) of Nuku'alofa in second 
growth on a coral atoll. Collected by Laurie Price on 
January 26, 1966. FMNH 152399. 

Range. Probably Tongatapu and its satellite is- 
lands, Tonga. 

Material. Only the holotype is known. 

Remarks. Graeffedon pricei is identical in height 
and diameter to the single known example of G. 
savaiiensis, but readily separable by the characters 
cited in the diagnosis. Its aperture is more evenly 
rounded, and the ribbing difference is particularly ob- 
vious. The position of the major palatals and two acces- 
sory palatal traces in G. pricei does not correspond to 
the position of the palatals in G. savaiiensis or G. 
graeffei. 

Collections on Tongatapu were made during a se- 
vere drought. No living endodontids were collected and 
comparatively little live material in other groups. 
Great pleasure is taken in naming this species after 
Mr. Laurie Price of Kaitaia, New Zealand, in grateful 
recognition of his collecting activities for Field 
Museum of Natural History. 

Subfamily TRUKCHAROPINAE, new subfamily 

Formal description of this subfamily has been 
given on p. 70. 

Several genera confined to the Palau and Caroline 
Islands show major anatomical modifications. In sharp 
contrast to the Charopinae and Semperdoninae, there 
is no indication of differentiation into a vas deferens 
and an epiphallus. The vas starts as a thin-walled 
transparent tube that rapidly changes into the typi- 
cally muscular shiny state seen in almost all endodon- 
toid species. There may be slight diameter changes at 
the penioviducal angle, but usually the tube retains a 
constant diameter until entering the penis. Except in 
Jokajdon, these genera all agree in having the vas def- 
erens tranverse the penis head, then passing through 
the penial retractor muscle fan before entering the 
penis. In both the Charopinae and Semperdoninae, 
there is a prominent epiphallus developed, with a com- 
plicated valvular entrance present in many 
Charopinae (figs. 29i-j, 57e). 

Although the Palau and Caroline Island genera 
grouped as the Trukcharopinae share this basic pat- 
tern of vas deferens-penis junction, there is much 
greater diversity in internal penial structure and mod- 
erate divergence in other genital features. The degree 
of genital difference between these genera is much 
greater than that which is seen in the Pacific Island 



206 



SOLEM: ENDODONTOID LAND SNAILS 



Charopinae and Semperdoninae, and involves both 
addition and elimination of structures. Shell structure 
is diverse, with an array of large and small, tightly and 
loosely coiled, heavily sculptured and with reduced 
sculpture, with and without barriers, elevated spires to 
deeply depressed apices all represented. Pallial organs 
show variation, but not to an extreme extent. Only 11 
species are involved, but their range of diversity ex- 
ceeds that in the other subfamilies. 

At this time I prefer not to speculate on the mutual 
affinities of these genera. Except for Jokajdon and 
Palikirus, which could be monophyletic, the differences 
are large enough that I am somewhat dubious of local 
derivation. Until enough Australian and New Zealand 
charopids have been dissected to establish the pattern 
of variation within that complex, the possibility that 
the Micronesian genera represent relicts from an early 
faunal migration from Southeast Asia through New 
Guinea to Australia, New Zealand, and New Caledonia 
cannot be excluded. Such an early migration would be 
prior to the spread of the helicarionid taxa presently 
dominant in the Indonesian-New Guinea Archipelago. 
Although the Semperdoninae and generalized Charo- 
pinae appear monophyletic with reasonably clear pat- 
terns of change, the Trukcharopinae share the lack of 
an epiphallus and enfolding of the vas deferens within 
the penial retractor muscle but represent a great vari- 
ety of other experiments. 

Major items of anatomical differences are sum- 
marized in the following paragraphs. Roimontis to- 
lotomensis, Palline biakensis, and Palikirus ponapicus 
have not been dissected and thus are excluded from the 
following discussion. 

Variations in the pallial cavity involve length, de- 
gree of kidney overlap onto the parietal wall, relative 
length of the pericardial and rectal kidney lobes, and 
shape of the kidney. Pallial cavity length is about 
three-quarters of a whorl in Jokajdon (fig. 93a); one- 
half whorl in Palikirus, Palline, Trukcharopa, and 
Kubaryiellus; and one-quarter whorl in Russatus (fig. 
90a). The square shape and equal kidney lobe length in 
Russatus is correlated with the whorl count reduction 
and coiling change. The other genera have elongated 
kidneys with the lobes of unequal length. In Jokajdon 
(fig. 93a), Palline (fig. 93e, g) and Kubaryiellus (fig. 
90e) the rectal lobe is twice the length of the pericar- 
dial; in Palikirus and Trukcharopa the rectal lobe is 
207c-3(y7c longer than the pericardial. There is not a 
direct correlation between lobe length and parietal 
wall overlap, because there is moderate overlap in 
Palline and Palikirus, strong overlap in Jokajdon, 
slight overlap in Trukcharopa, and no overlap in 
Kubaryiellus and Russatus. In Jokajdon and Palline 
the pericardial kidney arm is greatly reduced in width 
as well as length, but in Palikirus, with moderate kid- 
ney overlap, the pericardial arm is not reduced in size. 
I could not correlate the degree of overlap or differences 
in lobe length with shell features such as coiling pat- 
tern or body whorl width. Possibly the size factor is 



critical because the three small genera, Jokajdon, Pal- 
line, and Palikirus, have marked kidney overlap onto 
the parietal wall, whereas the larger genera do not. 
More data are needed. 

Previous extraction of the soft parts in many 
species meant that the apical genitalia could not be 
studied. Often the apical part of the visceral hump 
breaks off when the animal is pulled out. It remains in 
the shell and subsequently dries up. Thus only partial 
data are available on a dichotomy into species having 
one or two alveolar clumps in the ovotestis. No size 
correlation exists, since the very small Jokajdon and 
quite \argeKubaryiellus have a single clump, whereas 
the small Palline and medium-sized Trukcharopa have 
two distinct clumps, one lying completely apicad of the 
other, with both touching the palatal surface of the 
whorl. I could not examine the ovotestis of Palikirus or 
Russatus. In Jokajdon the ovotestis alveoli were fewer 
in number and proportionately much larger and longer 
than in the other genera. Sufficiently accurate mea- 
surements for comparison of midvisceral genitalia 
proportions were not obtained. I saw no qualitative or 
obvious proportional differences in this area. 

Distinctions are present in proportions and rela- 
tive development of the free oviduct, spermatheca, and 
vagina. Normally the basal portion of the sper- 
mathecal shaft is greatly expanded in the Charopidae, 
with the interior having various complicated glandular 
or muscular pilasters and ridges (fig. 87c). Jokajdon is 
radically different in lacking any expansion of the 
lower spermathecal shaft (fig. 93b). It also is the only 
Micronesian genus in which the vagina is distinctly 
longer than the free oviduct. Russatus (fig. 90c) has a 
short vagina and a short free oviduct with the base of 
the spermathecal shaft globosely and suddenly ex- 
panded. These features correlate with the reduced 
whorl count and shortened pallial cavity, because a 
zonal compaction of the midvisceral genitalia typically 
accompanies such an alteration in coiling pattern. The 
other four genera have rather long free oviducts. In 
Palikirus the vagina is quite short; in Kubaryiellus 
(fig. 90f), Trukcharopa (fig. 90i), and Palline (fig. 93f, 
h) the vagina is slightly shorter than the free oviduct. 
Spermathecal shaft expansion is essentially the same 
in Palikirus and Kubaryiellus, with the gradual ta- 
pered expansion occurring over a distance equal to the 
free oviduct length. In Trukcharopa there is a rela- 
tively sharp expansion that occurs over little more 
than half of the free oviduct length, whereas in Palline 
there is very gradual expansion that starts about at the 
mid-uterine level, well above the free oviduct origin. 
The functional significance of these changes is un- 
known. 

Fundamental differences exist in the penial com- 
plex. Most of these are concerned with the functioning 
surfaces of the penis, but even just external shape and 
proportions show major alterations. Palline (fig. 93f, h), 
Trukcharopa (fig. 90i), and Palikirus agree in having 
an ovately to globosely expanded penial head following 



SYSTEMATIC REVIEW 



207 



a much shorter, narrow tube (almost equal to head 
length in Truckcharopa). Russatus is basically the 
same, except that the expanded head is clearly triangu- 
lar in shape (fig. 90b). Kubaryiellus (fig. 90f) has a very 
long, club-shaped penis with the shaft tapering gradu- 
ally to the atrium. Jokajdon (fig. 93b) has the penis 
tapered apically and basally from a median expansion. 
Penial size is best expressed in relation to shell diame- 
ter because Russatus is 214 times the diameter of 
Jokajdon. Three genera, Russatus, Palikirus, and Jo- 
kajdon, have a long penial retractor muscle, and the 
ratio of total penis length to shell diameter is between 
0.15 and 0.30. Palline and Kubaryiellus have short pe- 
nial retractor muscles, and the ratio is between 0.45 
and 0.60. Trukcharopa has a short penial retractor and 
total penis length almost equals the shell diameter. 
Because origin of the penial retractor muscle normally 
lies near the apex of the pallial cavity, increasing rela- 
tive penis length shortens the penial retractor. Unlike 
many Endodontidae, no Pacific Island Charopidae 
have the penial retractor muscle detached from the 
diaphragm and joined to the free retractor system. 

Internally, Palline has a tubular verge that api- 
cally may function as an epiphallic chamber; Kubar- 
yiellus, a trilobed vergic papilla (fig. 90h); Russatus 
(fig. 90d), a simple pore with a lateral, rather long 
caecum on the inner wall of the penis head; Jokajdon, 
Palikirus, and Trukcharopa (fig. 90j), a simple pore 
with no major apical structures in the penis. Some form 
of muscular "pocket" pilaster is found in Jokajdon, 
Palikirus, and Trukcharopa, with the pilaster globular 
in Palikirus, of the Sinployea-type in Jokajdon, and 
greatly changed in Trukcharopa. Weak apical lon- 
gitudinal pilasters are found in Jokajdon and 
Palikirus, strong submedial longitudinal pilasters in 
Kubaryiellus. Russatus has weak, circular glandular 
pilasters, whereas Palline has a lateral pocket with 
concentric glandular pilasters and a large, median 
thick circular or partly circular pilaster. Jokajdon is 
unique in having a separate free retractor muscle at- 
tached to the penioviducal angle. 

Of the above changes, the verge and circular pilas- 
ters in Palline, internal caecum in Russatus, 
penioviducal angle muscle in Jokajdon, and major lon- 
gitudinal pilasters in Kubaryiellus represent entirely 
new structures. Opening of the vas deferens into the 
penis through a simple pore in Russatus, Palikirus, 
Jokajdon, and Trukcharopa represents a loss of 
structure when compared with the Charopinae. Other 
structures could be relatively simple modifications 
from the Sinployea pattern, but the items detailed 
above represent loss or gain of entire structures. 

Factors of shell variation are discussed in greater 
detail elsewhere (pp. 8-23). Here it is necessary to 
note only a few points. It is the smallest shells, Jokaj- 
don and Palline, that have the tightest coiling pattern 
(figs. 92a, d; 94a) and greatest development of aper- 
tural barriers (fig. 92b, e). The largest shells, 
Kubaryiellus and Russatus, have the loosest form of 



coiling (fig. 89a-f) and totally lack apertural barriers. 
Only Palline and Palikirus show the normal charopid 
shell coloration of simple yellow-horn or with traces of 
darker flammulations. The remaining genera have the 
shell reddish (Jokajdon, Trukcharopa), reddish brown 
(Kubaryiellus, Roimontis), or deep reddish purple- 
brown (Russatus). Only Palline (fig. 94b) has a moder- 
ately elevated spire, most of the others have it nor- 
mally flat or slightly elevated, whereas in Kubaryiellus 
(fig. 89b) the apex is distinctly depressed, and in 
Roimontis (fig. 91a-b) it is very depressed. Umbilical 
width shows only moderate variation, with slight nar- 
rowing in Palline (fig. 94c) and great narrowing in 
Russatus (fig. 89f). Sculpture is greatly reduced in 
Russatus, greatly enlarged in Palikirus (fig. 91d-f). In- 
terpretation of these variations is rendered difficult by 
the high concentration of species on Ponape. Palline 
notera is found on Koror, Peleliu, and Babelthuap in 
the Palau Islands; Palline biakensis, on Biak Island, 
West Irian; and Trukcharopa is restricted to the Truk 
Group and Lukunor in the Central Carolines. The re- 
maining eight species are restricted to Ponape. We 
have no exact locality for Palikirus ponapicus (Mollen- 
dorff, 1900), but the other seven species were taken at 
one or more stations by the B. P. Bishop Museum Mic- 
ronesian Expedition in early 1936. Several species live 
together, because three to five species were collected at 
four stations (table XLIV). No data exist concerning 
any ecological separation of the species. The genital 
differences, except between the two species of Jokaj- 
don, cannot be ascribed to a simple character dis- 
placement phenomenon. The size of the penis itself is 
so different between sympatric forms, and both dele- 
tion and addition of structures are involved. 

Ponape offers considerable opportunities for inves- 
tigation of isolating mechanisms in land snails because 
the multitude of sharply differentiated charopids there 
is equivalent to the remarkable radiation of trocho- 
morphid genera and species, Kondoa, Hogolua, and 
Brazieria in the Truk Group (Baker, 1941, pp. 272- 
280), and Trochomorpha itself on Ponape (Baker, 1941, 
pp. 290-296). Rapa shows an even greater radiation of 
Endodontidae (Solem, 1976b, pp. 490-492), but these 
taxa show far less dramatic changes in genital 
structures, a much greater amount of local geographic 
speciation, and probably are extinct. 

Genus Trukcharopa, new genus 

Shell of average size, with about 4% normally coiled whorls. 
Apex and spire flat to slightly elevated, body whorl descending 
slightly. Apical sculpture of about 10 prominent spiral cords. Post- 
nuclear sculpture narrow, crowded, protractively sinuated radial 
ribs, averaging about 18/mm. on body whorl. Microsculpture of fine 
radial riblets, finer spiral, and prominent secondary spiral cords. 
Umbilicus widely open, margins rounded. Whorls only slightly com- 
pressed laterally above and below rounded periphery, aperture in- 
clined about 20" from shell axis. No apertural barriers developed. 
Pallial region extending Vi whorl apically, with rectal kidney arm 
20%-30V< longer than pericardia!. Ovotestis with 2 clumps of alveoli, 
1 above other, hermaphroditic duct partly convoluted. Penial retrac- 
tor muscle relatively short, enfolding vas deferens before inserting 



208 



SOLEM: ENDODONTOID LAND SNAILS 



TABLE XLIV. - SYMPATHY OF CHAROPIDAE 
ON PONAPE, CAROLINE ISLANDS. 



Station 91 
Palikirus cosmetus 



Kubaryiellus kubaryi 
Jokajdon tumidulus 
Jokajdon calllzonus 



Station 92 

Roimontis tolotomensis 
Russatus nigrescens 
Kubaryiellus kubaryi 
Jokaj don tumidulus 
Jokajdon callizonus 

Station 93 

Roimontis tolotomensis 
Russatus nigrescens 
Jokajdon tumidulus 
Jokajdon callizonus 

Station 139 
Palikirus cosmetus 
Kubaryiellus kubaryi 
Jokaj don tumidulus 



on penis head. Penis long, head greatly expanded, internally with a 
huge medial stimulator, no vergic papilla, weak papillose glandular 
walls in lower tube. Spermatheca with basal part of shaft greatly 
expanded, vagina relatively long. 

Type species. Trukcharopa trukana, new species. 

Both Kubaryiellus and Trukcharopa have much 
longer penes (fig. 90f, i) than any of the other Microne- 
sian genera that lack an epiphallus, but the structures 
of the penes are very different. Trukcharopa has the 
head grossly expanded, lacks a vergic papilla, and has 
a huge median muscular stimulatory pad on one wall 
of the expanded chamber (fig. 90i-j). Serial sections of 
the penial region will be required before the exact 
structure of this stimulatory pad can be determined, 
but the essential differences from the other Microne- 
sian genera are obvious. 

Trukcharopa is nearest in shell structure to the 
Polynesian-Melanesian genus Sinployea. Anatomi- 



cally the two differ in Sinployea having a prominent 
epiphallus that is about half the penis in length, the 
penis has a distinct vergic papilla, and the typical 
stimulatory pad occupies the median part of the penis. 
Sinployea kusaieana is the only named Micronesian 
species in that genus. Conchological comparisons with 
Trukcharopa trukana (tables XXVII, XLV) show that 
they are virtually identical in mean diameter, umbili- 
cal width, D/U ratio, and body whorl width. Sinployea 
kusaieana (fig. 63a-c) is a light reddish yellow and has 
an average of 91 major radial ribs, 10.3-13.1 ribs/mm. 
on the body whorl, with five to 10 microradial riblets 
between each pair of major ribs. In contrast, Truk- 
charopa trukana (fig. 63d-f) is dark reddish brown in 
color and has an average of 150 major radial ribs, 
14.1-20.8 ribs/mm, on the body whorl, with only two to 
four microradials between each pair of major ribs. I 
could find no conchological features in Trukcharopa 
that would differentiate it generically from Sinployea. 
Both pallial region and male genital structures (fig. 
90i-j) place T. trukana as being more closely related to 
the Micronesian genera than to Sinployea. 

In choosing Trukcharopa as a name for this genus, 
I am reflecting both its great similarity to the tradi- 
tionally named "Charopa" (=Sinployea) of Polynesia 
and its geographic location. 

Trukcharopa trukana, new species. Figures 63d-f, 



Diagnosis. Shell smaller than average, diameter 2.50-3.08 
mm. (mean 2.66 mm.), with 4'/s-4% rather tightly coiled whorls. 
Apex and early spire normally flat, sometimes slightly protruding, 
rarely moderately elevated, body whorl usually descending slightly 
more rapidly, spire protrusion generally much less than 'Ac body 
whorl width, occasionally moderately elevated, H/D ratio 0.371- 
0.513 (mean 0.450). Apical sculpture of 8-12 (mean 9.68) prominent 
spiral cords, whose interstices are 2-3 times their width. Postnuclear 
sculpture of low, narrow, crowded, slightly protractively sinuated 
radial ribs, 112-197 (mean 150.1) on the body whorl, whose in- 
terstices usually are l'/2-2 times their width. Ribs/mm. 14.1-20.8 
(mean 17.7). Microsculpture of very fine, rather widely spaced radial 
riblets, 2-4 between each pair of major ribs, much finer and more 
crowded spiral riblets, with a secondary sculpture of spiral cords that 
are more prominent than the microradial riblets and whose in- 
terstices are about 3-4 times their width. Umbilicus broadly open, V- 
to U-shaped, regularly decoiling, contained 3.07-4.00 times (mean 
3.43) in the diameter, margins evenly rounded. Sutures deep, whorls 
strongly rounded above, slightly compressed laterally above 
periphery and on basal margin, with evenly rounded outer and col- 
umellar margins. Aperture subovate, compressed laterally above 
periphery and on basal margin, inclined about 20 from shell axis. No 
apertural barriers, but a weak to moderate callus extending from 
columellar region up to periphery in some individuals. 

Trukcharopa trukana is much smaller, has much 
more crowded radial ribbing, more apical cords, and 
lacks the depressed spire of Kubaryiellus kubaryi 
(Mollendorff) (fig. 89a-c). Russatus nigrescens (Moll- 
endorff) has fewer whorls, a looser coiling pattern, and 
is much larger in size (fig. 89d-f). Differences from the 
Kusaie Island Sinployea kusaieana (fig. 63a-c) are 
covered in the generic discussion above. 

Description. Shell rather small, with slightly more than 4'A 
relatively tightly coiled whorls. Apex and spire flat, last whorl barely 



SYSTEMATIC REVIEW 

TABLE XLV. - RANGE OF VARIATION IN MICRONESIAN CHAROPINAE, I. 



209 





NUMBER OF 














NAME 


SPECIMENS 


RIBS 


RIBS/MM 


HEIGHT 


DIAMETER 


Slnployea 
















kusaieana 


61 


91.1(84-103) 


11.25(10.27-13.12) 


1.32(1.19-1.49) 


2.60(2 


.52-2 


.75) 


Trukcharopa 
















trukana 


405 


150.1(112-197) 


17.7(14.07-20.84) 


1.19(0.99-1.42) 


2.66(2 


.50-3, 


.08) 


Kubaryiellus 
















kubaryi 


52 


112.2(99-132) 


9.54(8.39-11.70) 


1.86(1.59-2.25) 


3.74(3 


.36-4 


.60) 


Russatus 
















nigrescens 


30 


R E 


D U C E D 


2.20(1.91-2.66) 


4.81(4 


.34-5. 


53) 


Roimontis 
















tolotomensis 


2 


57.5(57-58) 


6.64(6.27-7.01) 


1.46(1.39-1.52) 


2.78(2 


.65-2, 


.91) 


Palikirus 
















cosmetus 


8 


18.9(17-21) 


2.51(2.29-2.71) 


1.23(1.12-1.38) 


2.40(2 


.17-2. 


58) 


ponapicus 


1 


46 


6.74 


0.99 


2.17 







H/D RATIO WHORLS 

kusel. 0.506(0.462-0.542) 3 7/8+(3 3/4-4 1/8) 

truka. 0.450(0.371-0.513) 4 3/8-(4 1/8-4 3/4) 

kubar. 0.494(0.453-0.540) 

nigres. 0.455(0.418-0.482) 

tolot. 0.524(0.523-0.525) 

cosme . 0.510(0.487-0.542) 

ponap. 0.454 



4 l/8-(3 7/8-4 1/2) 

3 l/8+(3 1/8-3 1/2) 

4 

3 3/4+(3 5/8-4) 

3 3/8 



UMBILICUS D/U RATIO 

0.74(0.69-0.79) 3.51(3.21-3.67) 

0.77(0.63-0.95) 3.43(3.07-4.00) 

1.01(0.83-1.32) 3.70(3.36-4.24) 

0.53(0.30-0.69) 10.6(6.30-16.5) 

0.71(0.69-0.72) 3.91(3.81-4.00) 

0.70(0.63-0.82) 3.44(3.14-3.67) 

0.54 4.00 








APICAL 
CORDS 


SPIRE 
ELEVATION 


BODY WHORL 
WIDTH SP/BWW 


PR 


kusei. 


9.2(8-10) 


0.119(0.099-0.164) 


0, 


72(0, 


79-0 


.86) 0.150(0.125-0.208) 





truka. 
kubar. 


9.68(8-12) 
7.62(6-9) 
REDUCED 
6.50(6-7) 
11.80(10-13) 
9 


0.08(FLAT-0.23) 

DEPRESSED 

SLIGHTLY DEPRESSED 
TO FLAT 

DEEPLY DEPRESSED 
0.17(0.15-0.20) 

NOT 


0, 
1, 
1, 
1. 

A 


80(0, 

41(1. 
29(1 
13(1. 
.76(0 

V A 


72-0 
,32-1 
.18-1 
07-1 
.72-0 
I 


.87) 0.092(FLAT-0.264) 
co\ 





2 
1 



41 1 


nigres. 
tolot. 


-i o \ 


. 18,1 

.79) 0.228(0.205-0.250) 
L A B L E 


cosme . 


ponap . 



descending, H/D ratio 0.430. Embryonic whorls l'/2, sculpture of 8 
prominent, rather closely spaced spiral cords. Postnuclear whorls 
with prominent, narrow, crowded, protractively sinuated radial ribs, 
133 on the body whorl, whose interstices are 2-3 times their width. 
Microsculpture of very fine radial riblets and even finer spiral rib- 
lets, crossed by prominent, widely spaced secondary spiral cords that 
become wider and more numerous on the body whorl. Sutures deep, 
whorls evenly rounded above, markedly flattened laterally above 
periphery and on basal margin. Umbilicus broadly U-shaped, regu- 
larly decoiling, contained 3.59 times in the diameter. Color reddish 
brown. Aperture ovate, flattened basally and laterally above 
periphery, inclined about 5 from the shell axis. Height of holotype 
1.12 mm., diameter 2.60 mm. 

Holotype. Caroline Islands, Truk, Tol Islet, Sta- 
tion A5, on dead leaves, 1,200 m. inland, near top of 
Mt. Urupot. Collected by K. Atoda on February 17, 
1940. BPBM 189318. 

Range. Truk Group and Lukunor, Caroline Is- 
lands. 



Paratypes. Truk: Moen Islet: Nebokos (Station 
16), from shore to 1 A mile inland at 4-5 ft. elevation (4 
specimens, BPBM 153701-2); west side of Mt. Teroken 
(Station 18), 1 mile inland at 900 ft. elevation (2 speci- 
mens, BPBM 153792); Saboku Village (Station A4), 
under dead leaves (105 specimens, BPBM 189263). 
Fefan Islet: Urunna village (Station A3), near shore on 
dead leaves (26 specimens, BPBM 189179). Dublon 
Islet: Mt. Tolowan (Station 9), in native forest at 850 ft. 
elevation (53 specimens, BPBM 153385-92); Mt. Tolo- 
wan (Stations 1, 8, 11) at 300-800 ft. elevation (3 
specimens, BPBM 153072, BPBM 153451); Meseran 
Village (Station 12), northeast part of island, 1-100 ft. 
inland at 1-12 ft. elevation (5 specimens, BPBM 
153506). Param Islet: 200 ft. back of beach (Station 37) 
at 3 ft. elevation (1 specimen, BPBM 155749). Tarik 
Islet (Stations 38, A2; 3 specimens, BPBM 153911, 



210 



SOLEM: ENDODONTOID LAND SNAILS 



BPBM 155762, BPBM 189423). Tol Islet: Mt. Urupot 
(Stations 28, 29, 30, A5, A7), halfway up to summit, 
1,000-1,200 m. inland, 700-1,422 ft. elevation (109 
specimens, BPBM 155550-4, BPBM 153879-82, 
BPBM 155477-8, BPBM 155508, BPBM 189318, 
BPBM 189343); Urifei (Station A6), on dead leaves (9 
specimens, BPBM 189334); Iruku, Tsukuran village 
(Station Al), 100 m. inland on dead leaves (13 speci- 
mens, BPBM 189406); southwest slope of Mt. Urupot, 1 
mile inland above spring, 1,000-2,000 ft. elevation (35 
specimens, BPBM 210864, BPBM 210890-2, BPBM 
210921). Oneap Islet (9 specimens, ANSP 191994). 
Nama Islet (2 specimens, ANSP 191995). Ta Islet (25 
specimens, ANSP 192000). Lukunor (3 specimens, 
AMSC.7561). 

Remarks. Probably Trukcharopa trukana occurs 
on most of the islets in the Truk Group. Numerous 
specimens were collected on Moen, Fefan, Dublon, and 
Tol Islets. Only three living specimens were found on 
Tarik and one dead shell from Param Islet. Several sets 
from the Thaanum collection were obtained on ex- 
change by the ANSP. These sets undoubtedly contain 
selected fragments of larger field samples, and the 
slight size and shape differences shown by these (table 
XLVI) probably reflect sample bias rather than any 
morphologic changes. The cited names of Oneap, 
Nama, and Ta Islets are taken from the ANSP labels. I 
was not able to identify the position of Oneap and Ta. 

Ignoring the differences in the ANSP samples, 
there is only minor shape variation present. Compar- 
ing the specimens from Tol and Moen (table XLVI), the 
differences in height (t = 3.6106 with 21 degrees of 
freedom), H/D ratio (t = 2.8764), and D/U ratio (t = 
2.3826) reach statistical significance. The shells from 
Tol all have a flat spire and hence a lower H/D ratio, 
lower height, and slightly wider umbilicus than those 
from other islets where some individuals have a 
slightly elevated spire. I attach no systematic impor- 
tance to this variation. Specimens from Tol have much 
fewer and more widely spaced major radial ribs than do 
those from Moen or Oneap. Shells from Ta (ANSP 
192000) are intermediate. Possibly there is an ecologi- 
cal correlation involved, but I have no data concerning 
this situation. 

Where large numbers of specimens were taken, 
the percentage of adults is very low. Probably the 
greater number of adults in small samples reflects col- 
lecting bias, at least in part, but the pattern of juvenile 
predominance is clear. Data for six stations are pre- 
sented in Table XLVII. The collecting dates range from 
December to May and from 1935 to 1940. Two different 
collectors were involved. No pattern of seasonality is 
suggested. 

Four specimens from Lukunor (AMS) are referred 
here with some hesitation. They appear to be slightly 
subadult, and if allowance is made for this, they fit 
within the range of variation for Trukcharopa trukana. 
Pending receipt of more adequate material, I classify 
these shells as this species. 



Several penes were opened, but I am not certain of 
the details shown in Figure 90j concerning the exact 
stimulatory pad structure. All specimens had many 
bits and strands of material inside, and determination 
of details was not possible from gross dissections. Time 
did not permit preparation of serial sections. The 
difference between the Sinployea-type stimulator and 
that of Trukcharopa trukana, the former a pocket be- 
hind and above a U-shaped muscle band, with the lat- 
ter a longitudinally oriented structure, combined with 
the presence of an epiphallus in Sinployea and its ab- 
sence in Trukcharopa are sufficient to distinguish the 
two genera without difficulty. 

Description of soft parts. Foot and tail about % shell diameter 
in length, not tapering posteriorly, broadly rounded behind, trun- 
cated anteriorly. Sole undivided, pedal grooves high on foot, pedal 
much deeper, grooves uniting over tail with slight depressions, but 
no caudal horn or middorsal groove developed. Slime network with- 
out unusual features. Gonopore in normal position. Body color light 
yellow-white, without darker markings. 

Mantle collar without glandular extension onto pallial roof. 
Anus and external ureteric pore with same position. Pallial region 
extending V4 whorl apically. Lung roof clear. Kidney about 1.2-1.3 
mm. long, rectal arm longer than pericardial arm, which is about 1 
mm. from anterior end to base. Ureter typical, section between kid- 
ney arms tightly compacted, overlapped by kidney. Heart about half 
length of pericardial kidney arm. Principal pulmonary vein un- 
branched. Hindgut typical. 

Ovotestis (fig. 90i, G) with two clumps of palmately clavate al- 
veoli, located in half whorl above stomach apex, lying parallel to 
whorl margins and occupying most of whorl space, slightly iridescent 
in tone. Hermaphroditic duct partly convoluted, tapering apically, 
constricted just before entering base of talon head. Albumen gland 
(GG) short and compact, acini relatively small. Talon (GT) with 
globular head, very short shaft to carrefour region. Prostate (DG) of 
comparatively few and large acini opening into groove on inner side 
of uterus. Uterus (UT) bipartite, lower chamber much larger and 
with thick, glandular walls. 

Vas deferens (VD) slender for entire length, passing through 
penial retractor muscle while crossing head of penis, then inserting 
laterally on penis head. Penial retractor (PR) arising from dia- 
phragm, slightly shorter than expanded penis head, inserting on 
head of penis after enfolding vas deferens. Penis (P) with expanded 
globular head, about 1.3 mm. long, followed by a narrow tube that is 
equal in length to penis head or slightly shorter, internally (fig. 90j) 
expanded head with an almost hemispherical stimulator (PP) lying 
partly unattached against one wall of penis, no vergic papilla, vas 
entering through a simple pore. Internal structure quite complex and 
should be studied using serial sections. Atrium (Y) narrow and 
rather long. 

Free oviduct (UV) larger than penis head, wider above. Sper- 
matheca (S) with elongately oval head, very slender shaft, strongly 
expanded laterally. Vagina (V) usually longer than illustrated, a 
simple tube. Free muscle and digestive system without unusual fea- 
tures. 

(Based on BPBM 189263, 2 whole individuals 2.43 mm. in 
diameter with 4Vt whorls and 2.57 mm. in diameter with 4V4 + 
whorls, plus several extracted fragments.) 

Genus Kubaryiellus, new genus 

Shell large, with about 4'/s rather loosely coiled whorls. Apex 
and spire depressed, body whorl not descending. Apical sculpture of 
6-9 very prominent spiral cords. Postnuclear whorls with narrow, 
prominent, protractively sinuated radial ribs, averaging less than 
10/mm. on the body whorl. Microsculpture of prominent radial rib- 
lets, finer spirals, and prominent secondary spiral cords. Umbilicus 
moderately open, V-shaped, margins rounded. Whorls flattened lat- 



SYSTEMATIC REVIEW 



211 



NAME 



NUMBER OF 
SPECIMENS 



TABLE XLVI. - LOCAL VARIATION IN TRUKCHAROPA. 



RIBS /MM. 



H/D RATIO 



. 



Oneap 

ANSP 191994 

Nama 

ANSP 191995 

Ta 

ANSP 192000 

Dub Ion Sta. 9 
BPBM 153385, -506 

Tol Sta.A5,A6,A7,30 
BPBM 155550, BPBM 
189318.-34.-43 

Moen Sta.A4, 16 
BPBM 153701, 
BPBM 189263 

Fefan Sta. A3 
BPBM 189179 

1 

Lukunor 
AMS C7561 



11 



11 



Oneap 
Nama 
Ta 

Dub Ion 
Tol 
Moen 
Fefan 
Lukun . 



168.817.46 
(143-197) 


19.0+0.550 
(16.98-20.50) 


1.2710.032 
(1.19-1.39) 


2.8510.055 
(2.70-3.08) 


0.45210.0034 
(0.442-0.460) 


157 


17.98 


1.1310.017 
(1.12-1.15) 


2.7010.083 
(2.62-2.78) 


0.42110.0065 
(0.414-0.427) 


139.313.14 
(124-156) 


17.310.359 
(15.07-19.24) 


1.21+0.019 
(1.08-1.27) 

1 . 16+0 .017 


2.5710.023 
(2.45-2.68) 

2 64+0 029 


0.46910.0078 
(0.420-0.506) 

440*0 0091 






(1.09-1.23) 


(2.52-2.75) 


(0.398-0.468) 


122.5 
(112-133) 


15.2 
(14.09-16.28) 


1.1410.024 
(0.99-1.29) 


2.64+0.036 
(2.52-2.95) 


0.433+0.0090 
(0.371-0.487) 


164.7 
(147-183) 


18.5 
(16.54-20.80) 


1.2610.023 
(1.16-1.42) 


2.7110.041 
(2.55-2.91) 


0.466+0.0069 
(0.432-0.513) 






(1.19-1.32) 


(2.58-2.88) 


(0.437-0.471) 


114.8+2.78 
(107-120) 


16.610.859 
(15.01-19.02) 


1.0610.032 
(0.97-1.12) 


2.20+0.067 
(2.01-2.30) 


0.480+0.0052 
(0.471-0.493) 


WHORLS 


UMBILICUS 


D/U RATIO 


APICAL 
CORDS 




4 l/2-(4 1/4-4 


3/4) 0.86 
(0.76-0, 


3.2910.074 
,95) (3.07-3.59) 


10.67+0.335 
(10-12) 




4 1/4 


0.81+0.050 
(0.76-0. 


3.3610.105 
86) (3.25-3.46) 


10.0 




4 l/4-(4 1/8-4 


3/8) 0.7510.015 
(0.63-0, 


3.4210.066 
82) (3.18-3.97) 


10.36+0.244 
(9-12) 




4 l/4+(4 1/8-4 


1/2) 0.7410.023 
(0.63-0, 


3.5710.087 
79) (3.29-4.00) 


9.8610.261 
(9-11) 




4 3/8-(4 1/4-4 


5/8) 0.7910.019 
(0.72-0, 


3.33+0.052 
95) (3.07-3.63) 


8.7310.305 

(8-11) 




4 3/8-(4 1/8-4 


5/8) 0.7810.017 
(0.69-0. 


3.4810.033 
86) (3.27-3.67) 


9.2010.250 
(8-10) 




4 l/2-(4 1/4-4 


1/2) 0.76+0.028 
(0.69-0. 


3.5710.058 
82) (3.40-3.71) 


9.7510.480 
(9-11) 




4-(3 3/4-4 1/8) 


0.5810.028 
(0.53-0. 


3.79+0.140 
66) (3.40-4.06) 








erally above and below rounded periphery, aperture inclined about 
15 from shell axis. No apertural barriers. Pallial cavity with rectal 
kidney arm about twice length of pericardial, otherwise without un- 
usual features. Ovotestis with one clump of alveoli, hermaphroditic 
duct tightly convoluted. Vas deferens passing through penial retrac- 
tor muscle before entering laterally on penis head. Penis very long, 
club-shaped, tapering, internally with a trilobed vergic papilla fol- 
lowed by 2 long, stimulatory pilasters on a papillose glandular wall. 
A thin muscle sheath surrounds penis. Spermatheca typical, vagina 
somewhat longer than typical. 



TABLE XLVII. - AGE DISTRIBUTION IN TRUKCHAROPA TRUKANA. 



ISLET 
Moen 

Dub Ion 
Tol 



STATION 


DATE OF NUMBER 
COLLECTION OF ADULTS 


NUMBER 

OF SHELLS 
COLLECTED 


% OF 
ADULTS 


A3 

A4 


May 
Mav 


1, 1940 5 
24, 1940 9 


26 
105 


19 

8 


' 
6% 


9 


Dec. 


20. 1935 


53 


13 


.2* 


A5 
A6 
30 


Feb. 
Mav 


17, 1940 5 
9, 1940 2 
4, 1936 3 


24 
9 
47 


20 
22. 
6 


- 
2% 
. 



Type species. Charopa kubaryi Mollendorff, 
1900. 

Roimontis tolotomensis (fig. 91a-c) is most similar 
in shell features, but differs in having spiral grooves 
rather than spiral cords, two parietal barriers, fewer 
and more widely spaced radial ribs, a much more de- 
pressed spire, and it is smaller in size. Russatus nigres- 
cens (fig. 89d-f) has a flat spire, much looser coiling, a 
very narrow umbilicus, reduced radial sculpture, 
greatly altered pallial region, and quite different geni- 
tal structures. Trukcharopa trukana (fig. 63d-f) has a 
flat to slightly elevated spire, much more numerous 
and crowded radial ribs on a much smaller shell, a 
tighter coiling pattern, finer apical sculpture, and the 
penis not only lacks a muscle sheath but has much 
different internal structure. 

Kubaryiellus (fig. 89a-c) has a modified shell. Its 
depressed apex and spire with relatively loose coiling 
are unusual in the Pacific Island Charopidae, but the 



212 



SOLEM: ENDODONTOID LAND SNAILS 



systematically important features are the greatly al- 
tered penial complex. This agrees with most other Mi- 
cronesian genera in having the vas deferens pass 
through the penial retractor muscle, but the distinct 
penial muscle sheath, trilobed vergic papilla, and two 
enlarged longitudinal pilasters (fig. 90g) represent 
major departures from the structural pattern seen in 
the other genera. The penial length of 1.95-2.20 mm. is 
much greater than that seen in the other Micronesian 
genera that lack an epiphallus. The vergic papilla 
could be derived from the type seen in Sinployea ir- 
regularis (fig. 67d) by elongation and size increase, but 
the absence of an epiphallus, development of a distinct 
sheath, and the peculiar pilasters are very different 
from the Sinployea pattern. 

Kubaryiellus is named in honor of I. Kubary, 
pioneer collector of land mollusks in New Guinea and 
the Caroline Islands for Otto von Mollendorff. 

Kubaryiellus kubaryi (Mollendorff, 1900). Figures 
89a-c, 90e-h. 

Charopa kubaryi Mollendorff, 1900, J. Malacol., 7 (5), p. 110 
Ponape, Caroline Islands. 

Diagnosis. Shell large, diameter 3.36-4.60 mm. (mean 3.74 
mm.) with 3 7 /-4'/2 normally coiled whorls. Apex and spire distinctly 
depressed below level of body whorl, last portion of body whorl barely 
descending, H/D ratio 0.453-0.540 (mean 0.494). Apical sculpture of 
6-9 (mean 7.62) prominent spiral cords, whose interstices are about 
twice their width. Postnuclear sculpture of prominent, narrow, 
lamellar, crowded, protractively sinuated radial ribs, 99-132 (mean 
112.2) on the body whorl, whose interstices are 3-6 times their 
width. Ribs/mm. 8.39-11.70 (mean 9.54). Microsculpture of promi- 
nent radial riblets, 3-6 between each pair of major ribs, much finer 
and more crowded spiral riblets, with rather widely spaced secondary 
spiral cords equal in size to or slightly larger than microradial rib- 
lets. Umbilicus broadly V-shaped, regularly and rapidly decoiling, 
contained 3.36-4.24 times (mean 3.70) in the diameter, margins 
evenly and gently rounded. Sutures deep, whorls strongly rounded 
above and on baso-columellar margin, strongly compressed laterally 
above rounded periphery with slightly compressed lower palatal 
margin. Aperture subovate, strongly compressed laterally above 
periphery, inclined about 15 from shell axis. No callus or apertural 
barriers developed. 

In having a distinctly depressed apex and marked 
lateral flattening of the body whorl, Kubaryiellus 
kubaryi is immediately separated from other Microne- 
sian or Polynesian species. Roimontis tolotomensis (fig. 
91a-c) has the apex much more deeply depressed, two 
weak parietals, and is much smaller. 

Description. Shell large, with 4V4 overlapping whorls. Apex 
and early spire distinctly sunken, markedly overlapped by coiling of 
later whorls, last portion of body whorl descending only slightly 
below top of penultimate, H/D ratio 0.458. Apical whorls 1H, 
sculpture of 7 widely spaced, relatively narrow spiral cords. Postnu- 
clear whorls with somewhat irregular, narrow, moderately protrac- 
tively sinuated radial ribs, relatively widely spaced on spire, becom- 
ing crowded on last whorl, about 132 on body whorl, whose 
interstices are 1-3 times their width. Microsculpture of fine, irregu- 
larly spaced radial riblets crossed by finer, slightly more crowded 
spiral riblets with a secondary sculpture of moderately prominent, 
widely spaced spiral cords. Sutures moderately impressed, whorls 
strongly rounded above, strongly flattened laterally above obtusely 
rounded periphery with evenly rounded basal margin. Umbilicus 
broadly V-shaped, regularly decoiling, contained 3.43 times in the 
diameter, with margins slightly shouldered. Color light reddish 



yellow-brown above, apex yellow-horn, and a narrow yellowish white 
spiral band situated Vs of the way between suture and periphery of 
the whorl. Periostracal ends of ribs in sutures and in umbilicus dark 
chocolate brown. Aperture ovate, moderately flattened laterally 
above periphery, inclined about 10 from the shell axis. Height of 
lectotype 1.81 mm., diameter 3.95 mm. 

Lectotype. Caroline Islands: Ponape. Collected by 
Kubary. SMF 165352. 

Range. Ponape, Caroline Islands. 
Paratypes. BPBM 86245, SMF 165353. 

Mater ial Ponape (1 specimen, BPBM 87528): 
back of Naupilo (Station 133) (4 specimens, BPBM 
157564-5); Paishapel Hill, Metalanim (Station 113), at 
650-700 ft. elevation (14 specimens, BPBM 157211-6); 
west side Tolomaim Mt. (Station 91) at 1,000-1,500 ft. 
elevation (3 specimens, BPBM 154002-3); Metalanim 
to Mipit (Station 118), 450 ft. inland at 300-900 ft. 
elevation (6 specimens, BPBM 154157-60); Mt. 
Kuporujo (Station 121) at 500-900 ft. elevation (1 
specimen, BPBM 154338); southwest side of To- 
reairuka Mt. (Station 127) at 250-750 ft. elevation (6 
specimens, BPBM 154451-3); one quarter to halfway 
up Mt. Tolotom (Station 92) at 800 ft. elevation (1 
specimen, BPBM 156677); north slope Mt. Nanalaut 
(Stations 129, 130) at 1,600-2,500 ft. elevation (2 
specimens, BPBM 157443); Mt. Nanalaut (Station 
129A) at 1,900 ft. elevation to top (1 specimen, BPBM 
154523); Palikir, Jokaj (Station 139 = 138), at 600-900 
ft. elevation (3 specimens, BPBM 157797); Palikir, Mt. 
Tamantamansakir (Station 140) at 1,000-1,250 ft. ele- 
vation (6 specimens, BPBM 157864). 

Remarks. Relatively few individuals of Ku- 
baryiellus kubaryi were taken at any single station, 
but it is widely distributed on Ponape. From stations 
where individuals were measured, 39 shells were col- 
lected, of which 20 (51.3%) were adult. This is by far 
the highest percentage of adult specimens for any 
Caroline species of which more than 10 examples were 
collected. 

Spire depression is marked, but much less than in 
Roimontis tolotomensis, and there is no noticeable re- 
duction in umbilical depth. 

Dissection of several penes showed variation in the 
length and exact form of the vergic papilla. The figured 
example (fig. 90g, PV) had the shortest seen, the lower 
lobes are sometimes extended for almost half the penis 
length in a U-shaped form, the upper lobe being rela- 
tively constant in shape. In all examples the opening of 
the vas deferens was beneath the median lobe of the 
vergic papilla, the two lower and lateral lobes serving 
as a channel for sperm transfer. 

No systematic significance should be given to the 
great size variation shown by the individual small sets 
measured (table XLVIII). Because of the depressed 
spire, already crowded ribbing, regular umbilical de- 
coiling, and lack of body whorl descension, separation 
of adult from subadult shells is virtually impossible. 
Probably the specimens from Stations 113 and 127 are 
fairly close to typical adult size. The types are dis- 







def 





a be 



FIG. 89. a-c, Kubaryiellus kubaryi (MbllendorfD. Ponape, Caroline Islands. Paratype. BPBM 87528 ex SMF; d-f, Russatus nigrescens 
(MollendorfT). Station 129A, Mt. Nanalaut, Ponape, Caroline Islands. BPBM 154521. Scale lines equal 1 mm. (MM). 



213 



214 



SOLEM: ENDODONTOID LAND SNAILS 



tinctly smaller and more widely umbilicated than the 
Micronesian Expedition shells. This is not a size- 
correlated change, since almost certainly subadult 
shells from Station 140 agree in umbilical width with 
the other Micronesian Expedition shells. The latter 
were collected at higher elevations than the types 
(Mollendorff, 1900, p. 101). The opposite variation is 
seen in Russatus nigrescens (table XLVIII), where the 
low elevation syntype specimens were much more nar- 
rowly umbilicated than the Micronesian Expedition 
upland shells. Further collections of both species are 
needed. 

Description of soft parts. Foot and tail long and slender, up to 
3.75 mm. long, tapering posteriorly, broadly rounded at end, trun- 
cated in front. Sole undivided. Pedal grooves high on foot, pedal 
much stronger, both uniting above tail, but no caudal horn or mid- 
dorsal groove present. Slime network details not clear in available 
material. Head partly retracted in all specimens examined. Gono- 
pore position not observed. Body color light yellow-white, without 
darker markings. 

Mantle collar without glandular extension onto pallial roof. 
Anus (A) opening at inner edge of pneumostome, external ureteric 
pore (KX) opening well behind anus. Pallial region (fig. 90e) about 
3.6 mm. from inner edge of mantle collar to apex of kidney. Lung roof 
clear, without granulations. Kidney (K) with 2 lobes, pericardial 
barely longer than heart, rectal arm about 2.2 mm. long, more than 
twice length of pericardial arm. Ureter (KD) with arms closely ap- 
proximated, no lung roof visible between, tapering anteriorly. Heart 
(H) lying parallel to hindgut, about equal in length to pericardial 
kidney lobe. Principal pulmonary vein (HV) without obvious branch- 
ing. Intestinal loop just above pallial cavity apex, hindgut (HG) 
reaching parietal-palatal margin just above pallial cavity apex. 

Ovotestis (fig. 90f, G) one clump of 5-8 iridescent, very elon- 
gated acini, lying parallel to sides of whorls, located just above 
stomach apex, extending a fraction of a whorl. Hermaphroditic duct 
(GD) tightly convoluted apically and, in some specimens, just before 
talon, iridescent for most of length, inserting at base of talon head. 
Albumen gland (GG) short, compact. Talon (GT) partly buried in 
surface of albumen gland, head globose, shaft slender to carrefour. 
Prostate (DG) of narrow acini opening into groove on inner surface of 
uterus, acini variable in length. Uterus (UT) bipartite, upper 
chamber much more thin-walled and more slender than thick-walled 
lower chamber (fig. 90h). 

Vas deferens (VD) thin-walled at first, becoming more muscular 
at penioviducal angle, loosely or not bound to side of penis, entering 
penial apex after passing through penial retractor muscle and 
traversing head of penis. Penial retractor (PR) very short, arising 
from diaphragm, inserting on head of penis after enfolding vas defer- 
ens. Penis (P) about 1.95-2.20 mm. long, club-shaped, covered by a 
thin muscle sheath, usually tapered basally (figured example 
bifolded basally), internally (fig. 90g) with a trilobed vergic papilla 
(PV) through which vas deferens enters. Lower part of penis appar- 
ently with two simple to corrugated longitudinal pilasters that form 
one side of the papillate lower penial region. Atrium (Y) long and 
much more slender than penis base. 

Free oviduct (UV) shorter than penis, wider apically and slightly 
expanded before joining spermathecal shaft. Spermatheca (S) with 
ovate head next to albumen gland, shaft slender to level of free 
oviduct, then greatly expanded with thick glandular walls. Vagina 
(V) elongated, slender, with weak internal pilasters. Free muscle 
system without unusual features. Salivary glands fused posteriorly 
above esophagus. 

(Based on BPBM 157211-2, 5 extracted and partial specimens.) 



Genus Russatus, new genus 

Shell very large, with less than 3'/2 very loosely coiled whorls. 
Apex and spire flat or slightly depressed, body whorl descending 
slightly. Apex macroscopically smooth, traces of fine spiral cording 



visible under high magnification. Postnuclear whorls with irregular, 
very crowded, fine radial ribs with high periostracal extensions, on 
lower spire not separable from growth irregularities. Occasional 
microreticulation visible at 96 x magnification. Umbilicus very nar- 
row, V-shaped, regularly decoiling. Whorls compressed laterally 
above periphery and on basal margin, aperture strongly inclined 
from shell axis. No apertural barriers. Pallial cavity extending 1 A 
whorl apically, kidney almost square, very thick in middle, rectal 
and pericardial arms equal in length. Apical genitalia not examined. 
Penial retractor muscle enfolding vas deferens before latter enters 
laterally on penis head through a simple pore. Expanded head of 
penis triangular, internally with a blind caecum lying at apex from 
opening of vas deferens to outer edge of expanded penis head, lower 
parts with circularly elongated glandular papillae on the walls. 
Spermatheca with basal part of stalk globularly expanded, vagina 
relatively long and much thicker than free oviduct. 

Type species. Flammulina (Calymna) nigrescens 
Mollendorff, 1900. 

Except for the Guam Island Himeroconcha fusca 
(fig. 105d-f), Russatus nigrescens is the largest Mi- 
cronesian charopid. Its major distinguishing features 
are the reduced whorl count (table XLV), much looser 
coiling pattern (fig. 89d), great sculptural reduction, 
development of a square kidney (fig. 90a) that remains 
next to the hindgut, and the very peculiar caecum on 
the inner wall of the penis (fig. 90d). No Polynesian or 
Micronesian species is at all similar in shell coiling 
pattern, and the whorl count is lower than all known 
species. Only the minute and very tightly coiled Mi- 
crocharopa mimula (fig. 33a-c) and the very high- 
spired Ba humbugi (fig. 74a-c) from Viti Levu, Fiji, 
come close in terms of whorl count. They differ mark- 
edly in coiling pattern, apical sculpture (Microcharopa, 
fig. 4a-d), and genital anatomy (Ba, fig. 75a-h). 

The New Guinea endemic Paryphantopsis (Solem, 
1970a, fig. la-c, shell, and fig, 2a-e, anatomy) is simi- 
lar in whorl count, coiling pattern, major sculpture re- 
duction, and size, but the similarities are convergent. 
In Paryphantopsis the pallial cavity has the kidney 
partly rotated from the hindgut, the arms of the ureter 
are separated at a 60 angle instead of being in contact, 
the principal pulmonary vein is noticeably branched, 
and the entire pallial cavity only occupies one-sixth 
whorl. Paryphantopsis has quite different terminal 
genitalia, there being an enlarged epiphallus with a 
prominent caecum, the penis with a verge and corru- 
gated longitudinal pilasters, and the vaginal region 
being very short and swollen with thick glandular 
walls. Russatus lacks both an epiphallus and a verge. 
Species of Paryphantopsis have the umbilicus com- 
pletely closed or at most a weak lateral crack, whereas 
the umbilicus of Russatus nigrescens is distinctly open. 

Both genera have started the process of visceral 
hump reduction that can eventually lead to evolution 
of a "slug" genus, but the pallial cavity and genital 
structures indicate that they are not closely related. 

Either Kubaryiellus kubaryi (fig. 90e-h) or Truk- 
charopa trukana (fig. 90i-j) might be compared with 
Russatus nigrescens as a possible ancestral type. The 
first has a large vergic papilla, very prominent 
stimulators, and the pallial cavity with very long rec- 
tal and much shorter pericardial kidney arms. Truk- 



SYSTEMATIC REVIEW 



215 



TABLE XLVIII. - LOCAL VARIATION IN KUBARYIELLUS AND RUSSATUS . 



Kubaryiellus 
kubaryl 

BPBM 86245, 
SMF 165352-3 

Sta.113 

BPBM 157211-2 

Sta. 127 
BPBM 154451-2 

Sta. 140 
BPBM 157864 

Russatus 

nigrescens 
Pre-1900 
SMF 165551-2, 
DMW MF8753 

In 1936 

BPBM 154077, -403, 

-520, BPBM 156678 



117.77.85 

(105-132) 



132.018.27 

(108-168) 



RIBS/MM. 



9.900.431 

(9.15-10.64) 



10.13*0.784 

(9.34-11.70) 



1.78+0.016 

(1.74-1.81) 



2.00+0.064 

(1.88-2.14) 



2.03+0.076 

(1.88-2.24) 



1.71+0.040 

(1.58-1.84) 



2.150.033 

(2.01-2.27) 



2.26+0.177 

(1.92-2.68) 



H/D RATIO 



3.73+0.079 0.479+0.0094 

(3.59-3.95) (0.458-0.495) 



4.09+0.183 0.490+0.0159 

(3.68-4.57) (0.453-0.524) 



3.97+0.091 0.51110.0102 

(3.71-4.14) (0.492-0.540) 



3.47+0.058 0.4930.0068 

(3.36-3.72) (0.471-0.519) 



4.77+0.053 0.450+0.0072 

(4.57-5.00) (0.418-0.475) 



4.82+0.283 0.4640.0125 

(4.37-5.56) (0.430-0.483) 



APICAL 
D/U RATIO CORDS 



4 1/8(4-4 1/4) 






Sta. 113 



Sta. 127 



nigrescens 
Pre-1900 



In 1936 



1.08+0.028 3.46+0.062 

(1.02-1.15) (3.36-3.64) 



4 3/8- (4 1/8-4 1/2) 1.11+0.072 3.70+0.136 7.50.500 

(0.99-1.32) (3.48-4.06) (6-8) 



4 l/4-(4-4 1/2) 
4+(3 7/8-4 1/8) 

3+(3-3 1/8) 



1.05+0.036 3.7910.147 6.8+0.4,79 
(0.95-1.12) (3.53-4.21) (6-8) 

0.9410.041 3.7310.108 8.210.307 
(0.82-1.09) (3.43-4.12) (7-9) 



0.35+0.040 14.22+1.38 

(0.30-0.43) (11.70-16.45) 



3 l/4+(3 1/8-3 1/2) 0.62+0.054 7.92+0.685 REDUCED 
(0.46-0.69) (6.30-9.64) 



charopa has the penis without a vergic papilla, pos- 
sesses a very unusual medial penial stimulator and 
agrees in pallial configurations with Kubaryiellus. Al- 
though their shell form appears intermediate between 
the more typical Trukcharopinae and Russatus, their 
anatomical features indicate no close affinity. 

The name Russatus, meaning clothed in red, refers 
to the very dark color of the shell. 

Russatus nigrescens (Mollendorff, 1900). Figures 
89d-f, 90a-d. 

Flammulina (Calymna) nigrescens Mollendorff, 1900, J. Malacol., 
7 (5), pp. 107-108, figs. 1-3 Naupilo. Ponape, Caroline Islands, 
at 100 m. elevation. 

Diagnosis. Shell very large, diameter 4.34-5.53 mm. (mean 
4.81 mm.), with S'/s-S'/a loosely coiled whorls. Apex and spire flat or 
very slightly depressed, last part of body whorl descending slightly, 
H/D ratio 0.418-0.482 (mean 0.455). Apical whorls with faint traces 
of spiral cords under high magnification, macroscopically smooth. 
Postnuclear whorls with irregular, extremely narrow and crowded, 
protractively sinuated radial ribs with lamellar extensions, becom- 
ing indistinguishable from growth irregularities shortly after apex. 
Occasional traces of microreticulation visible under 96 x magnifica- 
tion. Umbilicus very narrow, broadly V-shaped, regularly decoiling, 



contained 6.30-16.5 times (mean 10.6) in the diameter, margins 
strongly rounded. Sutures deep, whorls strongly rounded above, 
greatly compressed laterally above strongly rounded periphery and 
on basal margin. Aperture elongately ovate, compressed laterally 
above periphery and on basal margin, inclined almost 40 from shell 
axis. No apertural barriers. 

Russatus nigrescens has the lowest whorl count 
and loosest coiling pattern of all Pacific Island En- 
dodontidae and Charopidae. Its large size, tiny um- 
bilicus, reduced apical sculpture, and very crowded, 
lamellar major radial sculpture immediately separate 
it from any Micronesian species. 

Description. Shell very large, with 3!/s very loosely coiled 
whorls. Apex and early spire flat, body whorl descending very 
slightly, H/D ratio 0.466. Apical whorls IVfe, sculpture eroded with 
only faint traces of spiral cording visible in sutures. Postnuclear 
whorls with irregularly spaced radial ribs and growth threads plus a 
fine wavy spiral sculpture of riblets. Sutures moderately impressed, 
whorls flatly rounded above and on basal margin. Color dark 
purplish brown with occasional lighter patches becoming deep purple 
in the sutures and umbilicus. Umbilicus very narrow, greatly con- 
stricted by growth of periostracum, contained 16.45 times in the 
diameter. Aperture elongately ovate, flattened above periphery and 
on basal margin, inclined almost 35 from the shell axis. Height of 
lectotype 2.27 mm., diameter 4.87 mm. 



216 



SOLEM: ENDODONTOID LAND SNAILS 




PP 



FIG. 90. Anatomy of Russatus, Kubaryiellus, and Trukcharopa: a-d, Russatus nigrescens (Mollendorff). Station 129A, Ponape. BPBM 
154522. a, pallia! region, b, genitalia. c, interior of penis with caecum (PC) hiding vas opening, d, interior of penis showing relation of vas 
opening (DP) and pore of caecum; e-h, Kubaryiellus kubaryi (Mollendorff). Station 113, Ponape. BPBM 157212. e, pallial region, f, genitalia. g, 
interior of penis, h, detail of prostate-uterus ending; i-j, Trukcharopa trukana, new species. Station A4, Moen Islet, Truk. BPBM 189263. ;', 
genitalia;j, interior of penis. Scale lines equal 1 mm. (MO). 



Lectotype. Caroline Islands: Ponape, Naupilo, at 
about 100 m. elevation. Collected by Kubary. SMF 
165551. 

Range. Ponape, Caroline Islands. 

Paratypes.SMF 165552. 

Material. Ponape (1 specimen, DMW): Mt. To- 
lotom (Station 92), from V4-% way up, near 800 ft. ele- 



vation (2 specimens, BPBM 156678); Mt. Tolotom (Sta- 
tion 93), % way up, to summit at 1,200 ft. elevation on 
dead leaves (11 specimens, BPBM 154077-80); Mt. 
Nanalaut (Station 129A), from 1,900 ft. to summit, on 
ground (8 specimens, BPBM 154520-2); Mt. Kuporujo, 
near summit (Station 122) at 1,600-1,970 ft. elevation 
(2 specimens, BPBM 154403). 



SYSTEMATIC REVIEW 



217 



Remarks. Only four of 23 specimens collected by 
the Micronesian Expedition were adult, and each came 
from a different station. Although the original collec- 
tion was made at low altitude. 100 m., subsequent finds 
have been at higher elevations from widely scattered 
localities. The types are much more narrowly umbili- 
cated than the more recently collected shells (table 
XLVIII), but I am uncertain as to the significance of 
this difference. Possibly subspecific differentiation is 
involved. 

Suter (in Mollendorff, 1900, p. 108, figs. 1-3) illus- 
trated the tail of a dried, then softened, individual, part 
of the jaw, and a portion of one radular half row. His 
classification of this species with Flammulina was a 
logical guess but is not supported by subsequent dissec- 
tions. Flammulina shows major differences in pallial 
and genital anatomy (fig. 29a-j). The conchological 
similarities clearly are convergent. 

Russatus nigrescens has fewer whorls, mean count 
3Vs + , than any Pacific Island endodontoid. The Fijian 
Ba humbugi has somewhat similar alteration in form, 
but clearly differs in its greatly elevated spire and 
anatomical structure (fig. 75a-h). Ba humbugi does 
have a mean whorl count of 3%, but the pattern of 
coiling (fig. 74a) is very different. 

Description of soft parts. Foot and tail about 5.5 mm. long, 
tapering gradually from visceral stalk to narrowly rounded tip, trun- 
cated anteriorly. Sole undivided. Pedal groove very prominent, su- 
prapedal much weaker, both uniting over tail with marked indenta- 
tions, no caudal horn or middorsal groove developed. Slime network 
weak, more distinct posteriorly, backward slanting grooves on tail. 
Head projecting slightly in front of foot. Ommatophores very long. 
Gonopore located in front of and slightly below right rhinophore. 
Body color yellow-white, without darker markings. 

Mantle collar (MC) rather narrow, thick, no glandular extension 
onto pallial roof. Anus (A) and external ureteric pore (KX) opening 
side by side just inside pneumostome, with anal opening slanted. 
Pallial region (fig. 90a) extending apically about Vt whorl, length 
about 1.7- 1.9 mm. Lung roof clear, without granulations. Kidney (K) 
squarish, about 1.25-1.45 mm. long, compact, much thicker in mid- 
dle with a sharp ridge, narrowing abruptly toward anterior, much 
more gradually posteriorly, rectal and pericardial arms equal in 
length, shape irregular in outline. Ureter (KD) with primary and 
secondary arms compacted and sinuated between kidney lobes, quite 
variable in width, becoming narrow along hindgut. Heart (H) % 
length of kidney, slightly angled from hindgut. Principal pulmonary 
vein (HV) unbranched, inconspicuous. Hindgut (HG) continuing 
along parietal-palatal margin for short distance above pallial cavity 
apex. 

Apical genitalia not seen. Albumen gland (fig. 90b, GG) very 
irregular in shape, with large acini. Talon (GT) small, head globular, 
with a short shaft to slightly more expanded carrefour. Prostate (DG) 
of few long acini entering groove on inner surface of uterus, lower 
rows partly covered by uterine chamber folds. Uterus (UT) bipartite, 
lower chamber very large, completely folded, with thick glandular 
walls. 

Vas deferens (VD) very slender and with thin walls at first, 
becoming thicker and with muscular walls long before reflexion at 
penioviducal angle, not attached to penis during ascent, entering 
head of penis laterally after passing through penial retractor muscle 
(PR), which is fairly long and arises from diaphragm. Penis (P) a 
little less than 1.4 mm. long, apex very broad, upper % roughly 
triangular, lower 3rd a simple tube equal in diameter to free oviduct. 
Interior of penis (fig. 90c-d) with a blind-end muscular tube (PC) 
situated transversely at apex, opening muscular and puckered (fig. 
90d), vas deferens opening (DP) at attached base of caecum. Lower 



portions minutely papillose and glandular, papillae elongate and cir- 
cularly arranged. Atrium (Y) short and rather broad. 

Free oviduct (UV) sharply tapering to union with spermatheca. 
Spermatheca (S) with small, ovate head next to albumen gland, shaft 
slender to near base of uterus, becoming grossly globular with very 
thick muscular and glandular walls. Vagina (V) equal in length to or 
slightly longer than free oviduct, wider in diameter. Free muscle 
system without unusual features, tentacular retractors remain sepa- 
rate and unite with tail fan at columellar muscle. 

(Based on BPBM 154522, several partial specimens.) 

Genus Roimontis, new genus 

Shell of average size, with 4 overlapping whorls. Apex and spire 
deeply sunken beneath level of body whorl, last 3rd of body whorl 
with looser coiling. Apical sculpture of 6-7 very prominent spiral 
cords. Postnuclear sculpture of narrow, protractively sinuated, 
widely spaced radial ribs, about 6.6/mm. on body whorl. Micro- 
sculpture of fine radial riblets, finer spirals, and prominent secon- 
dary spiral grooves. Umbilicus shallow, narrow, V-shaped, margins 
rounded. Whorls very strongly flattened laterally above periphery 
with compressed lower palatal and basal margin, aperture inclined 
about 20 from shell axis. Parietal wall with 2 threadlike ridges 
extending about 3 Ae of a whorl. No columellar or palatal barriers. 
Anatomy unknown. 

Type species. Roimontis tolotomensis, new species. 

Despite the absence of anatomical data, I have no 
hesitation in describing this as a new genus. The 
deeply sunken apex and spire (fig. 91a-b), presence of 
secondary spiral grooves rather than cords, and de- 
velopment of two very low parietal barriers are highly 
distinctive. Kubaryiellus (fig. 89a-c) has a much less 
sunken apex, lacks the parietal barriers, has secondary 
spiral cording, is much larger and more widely umbili- 
cated, but is the most similar species in shell structure. 
Compared with the minor shell differences between 
Trukcharopa trukana (fig. 63d-f) and Sinployea 
kusaieana (fig. 63a-c), these are major changes. With 
the pattern of generic differentiation seen in the other 
Micronesian species, existence of these several major 
conchological changes is acceptable as indicating 
generic level separation. On the basis of the dark shell 
color, similarity in sculpture to the other Caroline Is- 
land species without an epiphallus, and relatively low 
whorl count, I tentatively classify this with the Truk- 
charopinae. I suspect that its anatomy will show 
marked differences from all of the above. 

Roimontis refers to the habitat of this species on 
the slopes of Mt. Tolotom, sometimes known as Roi 
Mountain. 

Roimontis tolotomensis, new species. Figure 91a-c. 

Diagnosis. Shell small, diameter 2.65-2.91 mm. (mean 2.78 
mm.), with 4 very tightly coiled and overlapping whorls. Apex and 
spire deeply sunken, last quarter of body whorl beginning to descend 
slightly, but not reaching level of first part of body whorl, H/D ratio 
0.523-0.525 (mean 0.524). Apical sculpture of 6-7 (mean 6.5) promi- 
nent spiral cords whose interstices are 3-4 times their width. Post- 
nuclear whorls with high, lamellar, very narrow, almost vertically 
sinuated radial ribs, 57-58 (mean 57.5) on the body whorl, whose 
interstices are 4-8 times their width. Ribs/mm. 6.27-7.01 (mean 
6.64). Microsculpture of fine radial riblets, 5-12 between each pair of 
major ribs, crossed by much finer and more crowded spiral riblets, 
with a prominent secondary sculpture of widely spaced spiral grooves 
that become more crowded near sutures and on shell base. Umbilicus 
relatively shallow, V-shaped, regularly decoiling, contained 3.81- 







abc 




FIG. 91. a-c, Roimontis tolotomensis, new species. Station 93, Mt. Tolotom, Caroline Islands. Holotype. BPBM 154081; d-f, Palikirus 
cosmetus, new species. Station 139, Palikir, Ponape, Caroline Islands. Holotype. BPBM 157798. Scale lines equal 1 mm. (a-c, MM; d-f, SH). 



218 



SYSTEMATIC REVIEW 



219 



4.00 times (mean 3.91) in the diameter, margins evenly rounded. 
Sutures deep, whorls shouldered above, strongly flattened laterally 
above rounded periphery, moderately compressed basally, with 
evenly rounded basal margin. Aperture subtriangular, strongly 
compressed and flattened laterally with downwardly moved 
periphery, inclined about 15-20 from shell axis. Parietal barriers 2, 
threadlike ridges extending posteriorly less than 3 Ae of a whorl: 
upper weakly raised medially, located just above periphery; 2nd a 
very low cord about V:i of way to parietal-columellar margin, not 
raised on any portion. 

Other species with a depressed apex include 
Kubaryiellus kubaryi (Mollendorff) (fig. 89a-c), which 
is much larger, has many more radial ribs (table XLV), 
and lacks the parietal barriers; and Russatus nigres- 
cens (fig. 89d-f), which is much, much larger, lacks 
apertural barriers, and has a reduced whorl count. 
Palline micramyla (fig. 95c-e) is much smaller, has 
secondary spiral cording, not grooving, a slightly 
raised apex, and much more crowded and numerous 
radial ribs. 

Description. Shell small, with 4 strongly overlapping whorls. 
Apex deeply sunken, last whorl strongly overlapping former and 
globosely swollen, descending slightly, H/D ratio 0.525. Embryonic 
whorls l'/2, with 7 quite prominent spiral cords. Postnuclear whorls 
with thin, lamellate, slightly protractively sinuated radial ribs, 58 
on the body whorl, whose interstices are 5-8 times their width. Mi- 
crosculpture of fine, widely spaced radial riblets, crossed by very low 
and indistinct spiral riblets, plus secondary sculpture of relatively 
prominent, broadly rounded, spiral grooves. Sutures inset, with 
whorls strongly shouldered above, distinctly flattened laterally 
above periphery with moderately flattened basal margin. Umbilicus 
relatively shallow, cup-shaped, contained 3.81 times in the diameter. 
Color light reddish yellow-brown. Aperture elongately ovate, 
strongly flattened laterally above periphery with gently rounded 
basal margin, inclined about 15 from the shell axis. Parietal wall 
with 2 threadlike barriers extending almost 3 Ae of a whorl, the upper 
slightly raised in middle %. Height of holotype 1.38 mm., diameter 
2.63 mm. 

Holotype. Caroline Islands: Ponape, Station 93, 
three-quarters of the way to summit of Mt. Tolotom at 
about 1,200 ft. elevation. Collected by Yoshio and 
Kiyoko Kondo on February 15, 1936. BPBM 154081. 

Range. Mt. Tolotom, Ponape, Caroline Islands. 

Paratype. (Station 92) one-quarter to three- 
quarters of the way up Mt. Tolotom at 800 ft. elevation 
(1 specimen, BPBM 156681). 

Remarks. At first glance Roimontis tolotomensis 
could be mistaken for a juvenile example of 
Kubaryiellus kubaryi, but the presence of two parietal 
barriers, the much more deeply sunken apex, much 
smaller size at the same whorl count, and much more 
widely spaced radial ribbing easily distinguish the two 
species. 

Both known examples of Roimontis tolotomensis 
were collected on the slopes of Mt. Tolotom between 
800 and 1,200 ft. elevation. The shells are adult, with 
the last third of the body whorl beginning to descend 
from the very overlapping coiling pattern of previous 
whorls, but in neither case does the apertural upper 
edge fall below the level of the penultimate whorl. 

Although the one specimen (BPBM 154081) was 
listed in the BPBM catalogue as having been collected 



alive, no anatomical material of this species could be 
located. 

Genus Palikirus, new genus 

Shell very small, with 3% to about 4 rather tightly coiled whorls. 
Apex and spire barely to slightly elevated, body whorl descending 
more rapidly. Apical sculpture of very fine and widely spaced spiral 
cords. Postnuclear sculpture of very large, prominent, protractively 
sinuated radial ribs, usually quite widely spaced. Microsculpture of 
fine radial riblets, finer spiral riblets, and relatively prominent sec- 
ondary spiral cords. Umbilicus moderately open, margins rounded to 
angulated. Parietal wall with (cosmetus) or without (ponapicus) a 
medial, low, bladelike barrier. Columellar and palatal walls without 
barriers. Pallial region extending '/a whorl apically with rectal kid- 
ney arm crossing hindgut and extending onto parietal wall, about 
30% longer than pericardial kidney arm. Ovotestis not seen. Her- 
maphroditic duct not convoluted. Vas deferens passing through pe- 
nial retractor muscle before entering penis through a simple pore. 
Penis with an expanded head, internally having weak longitudinal 
pilasters to level of an external constriction that marks presence of a 
globose, muscular-walled stimulatory pad, puckered near point of 
attachment to wall. Lower portion of penis with minutely papillose 
glandular walls. Spermathecal shaft markedly expanded on lower 
section, free oviduct relatively long, vagina short. 

Type species. Palikirus cosmetus, new species. 

Discussion of the anatomical differences between 
Palikirus and Jokajdon is given under the latter 
generic name. Conchologically, they are very distinc- 
tive, the few extremely high and large major radial 
ribs of Palikirus cosmetus (fig. 91d-f) contrasting 
greatly with the low and crowded radial ribbing of the 
two Jokajdon (fig. 92a-f). Besides the obvious ribbing 
differences, the larger size, fewer whorls, looser coiling, 
and only single parietal barrier of Palikirus distin- 
guish the two genera. Roimontis tolotomensis (fig. 
91a-c) is the only Micronesian species that might be 
confused with P. cosmetus, and then only on the basis 
of having two weak parietals, comparatively few radial 
ribs, and a moderately wide umbilicus. It has the apex 
and spire deeply depressed, secondary spiral grooving 
rather than cords, the shell is very strongly flattened 
laterally above the periphery, and is distinctly larger 
in size. Semperdon kororensis (fig. 98a-c) has superfi- 
cially similar ribbing, but is much larger in size, has a 
closed umbilicus, and numerous complex apertural 
barriers besides a very different penial structure. 

Inclusion of Charopa ponapica Mollendorff, 1900, 
in this genus is purely for convenience. The holotype 
and only known example of that species may not be 
fully adult. Although it shows more similarities to P. 
cosmetus than to any other Micronesian species, these 
similarities in sculpture and form could be convergent. 
Until living specimens can be collected and dissected, 
classification in Palikirus is a reasonable positioning 
for identification purposes. 

The name Palikirus is taken from the occurrence 
of the type species near Palikir in the Jokaj District of 
Ponape. 

Palikirus cosmetus, new species. Figure 91d-f. 

Diagnosis. Shell very small, diameter 2.17-2.58 mm. (mean 
2.40 mm.), with 3%-4 normally coiled whorls. Apex and spire mod- 



220 



SOLEM: ENDODONTOID LAND SNAILS 



TABLE XLIX. - LOCAL VARIATION IN PALIKIRUS AND JOKAJDON. 



NUMBER OF 
SPECIMENS 



RIBS/MM 



H/D RATIO 



Paliklrus cosmetus 

Sta. 139 

BPBM 157798-9 

Jokaldon callizonus 

TYPES: SMF 165700-1, 
BPBM 86246 

Sta. 93, BPBM 9688 



Sta. 118, BPBM 154161, 7 
BPBM 157282 

.loka-jdon tumidulus 

TYPES: SMF 165702-3, 12 
BPBM 86247 

Sta. 146, Jokaj, Id. 5 
BPBM 157984-5 

Sta. 132 11 

BPBM 157513-7 

Sta. 133 4 

BPBM 157566-7 

Sta. 93 6 

BPBM 154082 

Sta. 95 6 

BPBM 156730-1 



18 


.80.489 
(18-20) 


2.53+0.074 
(2.29-2.71) 


1.19+0 
(1 


.024 
.12-1. 


27) 


2.37+0. 
(2, 


061 
17-2. 


50) 


0.504*0. 
(0. 


0046 
487-0. 


515) 


129 


.3+15.93 
(99-153) 


20.12+1.229 
(17.71-21.73) 


0.92+0 

(0 


.071 
.79-1. 


04) 


2.030, 
(1. 


135 
78-2. 


24) 


0.450+0. 
(0. 


0063 
444-0. 


463) 


127 


.3+9.92 
(109-143) 


18.57*1.181 
(16.22-19.89) 


0.9710 
(0 

0.88+0 
(0 


,041 
.92-1. 

,013 
.84-0, 


05) 

94) 


2.18+0. 

(2, 

1.96+0. 

(1, 


056 
11-2. 

026 
86-2. 


29) 
04) 


0.445*0. 
(0. 

0.450+0. 
(0. 


0087 
430-0. 

0079 
428-0. 


460) 

478) 






99 


.1+4.18 
(79-123) 


13.140.485 
(10.35-15.97) 


1.35+0 
(1 


.031) 

.22-1 


.56) 


2.400 

(2 


.040 
.20-2. 


63) 


0.561*0 
(0 


,0053 
.532-0. 


594) 


114 


.8+7.62 
(92-140) 


14.580.500 
(13.78-16.45) 


1.46+0.094 
(1.12-1. 


64) 


2.50+0, 
(2, 


113 

,09-2. 


71) 


0.58310, 

(0, 


0126 
,535-0. 


606) 


62 


.85.75 
(39-95) 


10.04*1.097 
(5.80-16.18) 


1.20+0.021 
(1.09-1.28) 


2.03+0, 
(1, 


.042 
81-2. 


20) 


0.594+0 
(0, 


.0061 
563-0. 


631) 


97 


.32.96 
(89-105) 


13.71+1.082 
(11.56-16.54) 


1.30+0 
(1 


.082 

.10-1 


48) 


2.2810, 
(2 


.096 
.02-2. 


45) 


0.570+0, 
(0 


0130 
,545-0. 


604) 




REDUCED 


REDUCED 


1.21+0 
(1 


.0130 

.18-1 


.27) 


2.0510 
(1, 


,018 
.97-2. 


11) 


0.59210 
(0 


,0057 
,571-0. 


608) 




REDUCED 


REDUCED 


1.30+0 

(1 


.570 
.25-1 


.48) 


2.120 

(2 


.040 
.01-2. 


30) 


0.615+0 
(0 


.0083 

.585-0. 


642) 



WHORLS 



D/U RATIO 



APICAL RIBS 



cosmetus 



Sta. 139 



callizonus 



Sta. 118 

tumidulus 
TYPES 

Sta. 146 
Sta. 132 
Sta. 133 



Sta. 95 



3 3/4-K3 5/8-3 7/8) 0.69+0.031 3.4610.084 12.310.479 

(0.63-0.79) (3.16-3.67) (11-13) 



4 3/8(4 1/8-4 3/4) 0.66+0.045 3.0610.029 8.310.067 
(0.58-0.72) (3.00-3.09) (7-9) 

4 l/2+(4 1/2-4 3/4) 0.72+0.025 3.0110.044 8.3+0.034 
(0.69-0.77) (2.96-3.10) (8-9) 

4 l/2-(4 1/4-4 1/2) 0.66+0.014 3.0010.077 8.010.258 
(0.61-0.71) (2.84-3.27) (7-9) 



4 3/8+(4 1/8-4 3/4) 0.67+0.023 3.6310.081 10.2+0.250 

(0.54-0.82) (3.19-4.18) (9-11) 

4 l/2+(4 1/8-5) 0.6710.027 3.7710.082 8.810.250 

(0.59-0.71) (3.53-3.90) (8-9) 

4 l/8+(3 7/8-4 1/4) 0.55+0.018 3.70+0.062 9.3+0.289 

(0.48-0.64) (3.28-3.94) (8-11) 

4 1/4(4 1/8-4 1/2) 0.65+0.016 3.5010.166 9.7*0.334 

(0.61-0.67) (3.00-3.70) (9-10) 

4 l/4+(4 1/4-4 3/8) 0.6010.022 3.4610.113 9.310.211 

(0.51-0.67) (3.03-3.87) (9-10) 

4 3/8-(4 1/4-4 1/2) 0.60+0.022 3.55+0.151 9.010.516 

(0.53-0.67) (3.12-4.12) (8-11) 



erately and evenly elevated, body whorl descending more rapidly, 
spire protrusion less than l /i body whorl width, H/D ratio 0.487-0.542 
(mean 0.510). Apical sculpture of 10-13 (mean 11.8) very fine spiral 
cords. Postnuclear whorls with extremely high, prominent, broadly 
U-shaped, strongly protractively sinuated radial ribs, 17-21 (mean 
18.9) on the body whorl, whose interstices are 5-8 times their width. 
Ribs/mm. 2.29-2.71 (mean 2.51). Microsculpture of very fine radial 
riblets, more than 20 between each pair of major ribs, crossed by 
much finer and slightly more crowded spiral riblets, with an inter- 
mixture of crowded secondary spiral cords equal in size to microra- 
dials. Umbilicus broadly V-shaped, regularly decoiling, contained 
3.14-3.67 (mean 3.44) times in the diameter, margins shouldered to 



angled. Sutures impressed, whorls evenly rounded to slight flatten- 
ing laterally above periphery, evenly rounded outer margin, slightly 
compressed lower palatal wall with angled baso-columellar margin. 
Aperture ovate, compressed laterally above periphery, inclined 
about 30 from shell axis. Parietal wall with a single low, bladelike, 
medial barrier extending posteriorly almost 14 whorl, minutely ser- 
rated above, evenly elevated for entire length until last V whorl, 
with sharp to gradual anterior descension. No columellar or palatal 
barriers. 

Palikirus cosmetus differs from P. ponapicus by its 
fewer, larger radial ribs, more crowded secondary spi- 



SYSTEMATIC REVIEW 



221 



ral cording, wider umbilicus, more apical cords, and 
possession of a parietal barrier. All other Caroline and 
Palau species have much finer and more crowded ra- 
dial ribbing or many apertural barriers (Semperdon 
kororensis). 

Description. Shell very small, with 3% normally coiled whorls. 
Apex and spire evenly elevated, last whorl descending moderately, 
H/D ratio 0.487. Embryonic whorls l'/2, sculpture of 13 relatively 
wide spiral cords. Postnuclear whorls with sculpture of very high, 
U-shaped, strongly protractively sinuated, widely spaced radial ribs, 
18 on the body whorl, whose interstices are 4-6 times their width. 
Microsculpture of equally spaced radial and spiral riblets, the spiral 
slightly less prominent, plus obscure secondary spiral cording. Su- 
tures deep, whorls evenly rounded above, somewhat flattened later- 
ally above and below periphery with strongly rounded and slightly 
shouldered basal margin. Umbilicus broadly V-shaped, regularly de- 
coiling, contained 3.45 times in the diameter. Color very light red- 
dish yellow-brown. Aperture ovate, flattened laterally above and 
below periphery and on umbilical margin, inclined about 30 from 
shell axis. Parietal wall with 1 ridgelike barrier, slightly expanded 
above, extending almost Vi whorl. Height of holotype 1.22 mm., 
diameter 2.50 mm. 

Holotype. Caroline Islands: Ponape, Station 139, 
Palikir, Jokaj District, at 600-900 ft. elevation. Col- 
lected on a dead tree by S. Ito and Yoshio Kondo on 
March 14, 1936. BPBM 157798. 

Range. Ponape, Caroline Islands. 

Paratypes. Ponape: Palikir (Station 139), Jokaj 
District, at 600-900 ft. elevation (4 specimens, BPBM 
157798-9); west side of Mt. Tolotom (Station 91) at 
1,000-1,500 ft. elevation (1 specimen, BPBM 154004); 
Palikir (Station 137), more than V4 mile south of school 
at 450 ft. elevation (1 specimen, BPBM 157730); 
Palikir (Station 140), Mt. Tamantamansakir at 
1,000-1,250 ft. elevation (1 specimen, BPBM 157865). 

Remarks. Although Palikirus cosmetus is very 
similar in sculpture to the Hawaiian Cookeconcha 
stellulus (Gould) (see Solem 1976b, p. 218, fig. 93), the 
two species obviously differ in apical sculpture and 
apertural barriers. The Palau Island Semperdon 
kororensis has very similar sculpture but is much 
larger and has numerous large barriers (fig. 98b). 

The name cosmetus, from the Greek meaning 
"adorned," refers to the spectacular nature of the radial 
ribbing. 

Dissection of this species was accomplished during 
a period when I was without an illustrator. It was not 
possible to figure the genitalia or the pallial cavity 
because of the complicated structure and minute size. 

Description of soft parts. Foot and tail much less than shell 
diameter in length, tapering posteriorly. Pedal grooves and slime 
network typical. Gonopore in normal position. Body color yellow- 
white, without darker markings. Mantle collar thin, rather long, 
without glandular extension onto pallial roof. Anus opening very 
slightly anterior of external ureteric pore. 

Pallial region extending '/> whorl apically. Lung roof clear, with- 
out granulations. Kidney typically bilobed, rectal arm about 1.3 mm. 
long, crossing hindgut and extending onto parietal wall, pericardia! 
arm much shorter, less than 1 mm. from base to tip. Ureter tightly 
compacted between arms of kidney, portion along hindgut only 
slightly narrower than hindgut. Heart about % length of pericardia! 
kidney arm, lying parallel to hindgut. Principal pulmonary vein 
large, unbranched. Hindgut without unusual features. 

Ovotestis not seen. Hermaphroditic duct a rather short, thick, 
iridescent tube, slowly tapering apically, strongly constricted before 



reflexing into albumen gland and continuing apical to talon. Albu- 
men gland short and compact, acini quite small. Talon with an ovate 
head and short stalk that leads into a much larger carrefour. Union 
of hermaphroditic duct apparently near junction of talon stalk and 
carrefour, but exact relationship not determined. Prostate of 4 very 
large acini opening into groove on inner uterine wall. Uterus bipar- 
tite, lower chamber expanded and with thick, glandular walls. 

Vas deferens a slender tube, slightly expanded at penioviducal 
angle, becoming slender again during ascension alongside penis, en- 
tering head of penis after passing through penial retractor muscle 
fan. Penial retractor long, arising from diaphragm, attached to head 
of penis after enfolding vas deferens. Penis with expanded head of 0.3 
mm. in length, followed by a much shorter, narrow tube. Head por- 
tion with a supramedial constriction externally. Internally the upper 
section of penis head with numerous narrow longitudinal pilasters 
on the muscular wall, vas deferens entering apex with no special 
vergic papilla formed. Lower part with a huge, globular pilaster with 
smoothly muscled surface, deeply creased near point of attachment 
on one side, with 3 or 4 different creases radiating from a central 
point. Free wall of lower chamber minutely papillose with glandular 
tissue. Atrium short, rather narrow. 

Free oviduct about 0.6 mm. long, distinctly shorter than 
prostate-uterus, about '^i diameter of expanded spermathecal shaft, 
with longitudinal pilasters. Spermatheca with elongately oval head 
lying next to albumen gland, slender shaft lying next to prostatic 
acini, becoming grossly expanded near origin of vas deferens to union 
with free oviduct, expanded portion with thick walls. Vagina short, 
tapering rapidly from glandularized apical section to atrium. 

(Based on BPBM 157799, 2 partial examples.) 

Palikirus ponapicus (Mollendorff, 1900) 

Champa ponapica Mollendorff, 1900, J. Malacol., 7 (5), p. 110 
Ponape, Caroline Islands. 

Description. Shell very small, with 3% normally coiled whorls. 
Apex and spire slightly and evenly elevated, body whorl descending 
a little more rapidly, H/D ratio 0.454. Apical whorls 1%, sculpture 
partially eroded with traces of about 9 narrow, widely spaced, major 
spiral cords. Postnuclear whorls with rounded, prominent, protrac- 
tively sinuated radial ribs, about 46 on the body whorl, whose in- 
terstices are 3-5 times their width. Microsculpture mostly obscured 
by fungal growths, but on portions visible as a fine network of co- 
equal spiral and radial riblets with a secondary sculpture of narrow, 
relatively widely spaced spiral cords. Sutures deep, whorls strongly 
rounded above, slightly flattened laterally above periphery and on 
basal margin. Umbilicus V-shaped, regularly decoiling, contained 
4.00 times in the diameter with evenly rounded margins. Color very 
light reddish yellow-horn. Aperture ovate, slightly flattened later- 
ally above periphery and on basal margin, inclined about 15 from 
the shell axis. Height of holotype 0.99 mm., diameter 2.17 mm. 

Holotype. Caroline Islands: Ponape. Collected by 
Quadras. SMF 165354. 

Range. Ponape, Caroline Islands. 

Remarks. Unfortunately, the microsculpture of 
the holotype and only specimen seen of Palikirus 
ponapicus is partially obscured by fungal growths. De- 
spite this, the appearance is unmistakable, and P. 
ponapicus cannot be confused with any other Caroline 
Island species. 

The relatively few major radial ribs, 46 on the 
body whorl, are lower and more closely spaced, 
ribs/mm. 6.74, than in P. cosmetus, which has only 
17-21 major ribs that are widely spaced, 2.29-2.71 
ribs/mm. The latter has a wider (mean D/U ratio 3.44) 
umbilicus and possesses a single parietal barrier that 
extends posteriorly for one-quarter whorl. 

No specimens of P. ponapicus were obtained by 
members of the Micronesian Expedition. The holotype 



222 



SOLEM: ENDODONTOID LAND SNAILS 



was found in "bags of vegetable mould from the bush." 
These were taken "in the neighbourhood of the Spanish 
settlement on Ponape" by a German trader, Etscheid 
(Mollendorff, 1900, p. 101). 

Genus Jokajdon, new genus 

Shell very small, with 4%-4'/4 very tightly coiled whorls. Apex 
and early spire flat or barely protruding, spire not or sharply de- 
scending. Apical sculpture with variable number of fine spiral cords. 
Postnuclear sculpture of prominent, narrow, slightly protractively 
sinuated radial ribs, usually crowded, sometimes widely spaced or 
greatly reduced. Microsculpture of fine radial riblets, finer spiral 
riblets, and prominent secondary spiral cords. Umbilicus moderately 
to widely open, walls rounded or flattened, margins strongly rounded 
to distinctly shouldered. Periphery rounded, with slight to very 
strong lateral flattening. Parietal barriers 2, with or without acces- 
sory blade on 1st, variable in length. Columellar wall with a low 
knob to high barrier. Palatal barriers 3, variable in height and de- 
gree of twisting, with 3-6 accessory lamellae. Pallial region with 
pericardia! kidney arm greatly reduced, rectal arm enlarged, cover- 
ing hindgut and extending down parietal wall, total length of pallial 
cavity about % whorl. Genitalia with single clump of very few alveoli 
in ovotestis, hermaphroditic duct not convoluted. Penial retractor 
muscle very long, arising on diaphragm, inserting directly onto head 
of penis, vas deferens inserting on side of penis head just below mus- 
cle insertion. Penis variable in length, bitapered from middle, inter- 
nally with weak longitudinal pilasters to middle, plus a Sinployea- 
type medial stimulatory pad just below middle. A separate muscle 
strand from the columellar retractor inserts directly onto 
penioviducal angle. Vagina very long, base of spermathecal shaft not 
expanded. 

Type species. Endodonta (Thaumatodon) cal- 
lizona Mollendorff, 1900. 

Several features in the anatomy of Jokajdon (fig. 
93a-d) are highly unusual in relation to the other Mi- 
cronesian Charopidae. The total absence of broad ex- 
pansion for the spermathecal shaft is unique to these 
two species. In having the vas deferens insert laterally 
on the penis head just below the penial retractor union 
with the penis, Jokajdon differs considerably from 
most Micronesian genera. All those that lack an 
epiphallus, Palikirus, Palline, Russatus, Kubaryiellus, 
and Trukcharopa, have the vas deferens crossing the 
penis head and passing through the penial retractor 
muscle before entering the penis. A comparatively 
simple rotation of the penis head in relation to the 
muscle insertion could change one condition to the 
other, but the division into five genera with the trans- 
verse passage of the vas deferens and one without is a 
remarkable split. Shortening of the pericardial kidney 
arm and extension of the rectal kidney arm onto the 
parietal wall is carried further in Jokajdon than in any 
Pacific Island species. 

Internal penis structure of Jokajdon is most simi- 
lar to that seen in Palikirus cosmetus. Both genera 
have the vas deferens opening through a simple pore 
and with longitudinal pilasters, but in Palikirus the 
penial stimulator is a huge globular pad with a puck- 
ered remnant of a "pocket," whereas in Jokajdon the 
pocketed stimulator is very close to the type seen in 
Sinployea. Probably the differences are exaggerated by 
character displacement, since P. cosmetus and J. 
tumidulus were collected at three of the four stations 



from which P. cosmetus was taken. Gross enlargements 
of the penial pilaster in Jokajdon might account for 
this difference, but the gross spermathecal expansion, 
short vagina, enfolding of the vas deferens by the pe- 
nial retractor, and absence of a special muscle on the 
penioviducal angle are major anatomical differences 
that suggest generic separation. Palikirus cosmetus 
with its single parietal barrier, very few and large ra- 
dial ribs, slanted aperture, and reduced whorl count is 
conchologically very different in appearance. 

Dissection of the two Jokajdon was very difficult, 
the large and sinuated apertural barriers having pro- 
duced numerous distortions in the lower pallial and 
midgenital organs during the many years in preserva- 
tive. The parts were mushy, and some apical organs 
had not been preserved, probably through failure of the 
preservative to penetrate rapidly enough past the pal- 
lial air bubble and barriers during initial field han- 
dling. The only anatomical distinction observed was 
the disparity in penial size, with the penis of J. cal- 
lizonus being less than half the length seen in J. 
tumidulus. Studies of fresh material, probably by use 
of sections, will be required to see if penial size is the 
only distinction between the two species. Both dis- 
sected sets contained fully adult specimens so that no 
question of an age difference arises. 

Conchological differences between the two species 
are covered in the diagnosis of Jokajdon callizonus. 
Besides the shape and size differences enumerated 
there, many barrier details are different (compare fig. 
92b, e). Parietals are longer in J. callizonus, but an 
accessory lateral structure is found in J. tumidulus. 
Jokajdon callizonus has a huge columellar barrier that 
twists up to terminate on the parietal wall, whereas J. 
tumidulus has a low, triangular, knoblike ridge deeply 
recessed on the columellar wall. Less dramatic changes 
distinguish the pattern of palatal barriers in the two 
species, but comparison of the figures and descriptions 
show quite different details of form and proportions. 

Insufficient detail is available for any decision to 
be made, but the possibility that Jokajdon is analogous 
to the Rapan Rhysoconcha should be kept in mind for 
future exploration. The latter clearly represents a 
group derived by size reduction (Solem, 1976b, pp. 
255-258). The contorted palatal barriers of Jokajdon 
(figs. 92b, e) are comparable to the changes seen in 
Rhysoconcha (Solem, 1976b, p. 255, fig. 108a-b). Simi- 
larly, the single clump of very few and long alveoli in 
the ovotestis, lack of expansion in the spermathecal 
base, simplification of the vas-penis-retractor junction, 
and long vagina could be interpreted as resulting from 
differential change during size reduction. The addition 
of a special muscle to the penioviducal angle in Jokaj- 
don may relate to the extreme degree of apertural con- 
striction caused by the very large and complex bar- 
riers. Space between the parietals and palatals is so 
narrow that the animal may have great difficulty in 
effecting withdrawal of the foot and head into the shell. 
Neither time nor material permitted investigation of 



SYSTEMATIC REVIEW 



223 



TABLE L. - RANGE OF VARIATION IN MICRONESIAN CHAROPINAE, II. 



NUMBER OF 
NAME SPECIMENS 


RIBS 


RIBS/MM. 


HEIGHT 


DIAMETER 


Jokajon 
callizonus 


32 
142 


127, 

106 
53, 


,6(99-153) 

,1(74-148) 1 
,6(39-69) 2 


18 

14 
8 


.9(16. 

.61(10 
.13(5. 


2-21.7) 

. 35-22. 9) 1 
80-10. 62) 2 


0.93(0. 
1.30(1, 


79-1 
10-1 


.06) 
.65) 


2.06(1 
2.24(1 


.79-2.40) 
.88-2.73) 


tumidulus 


Palline 


3 


138. 


0(130-146) 


20 


.5(20. 


1-21.0) 


1.02(0. 


97-1 


.06) 


2.15(2 


.07-2.24) 


micramyla 


notera notera 


48 
53 


74, 
78, 


,0(58-95) 
8(63-101) 


11 
11 


.63(9. 
.01(8. 


37-15.66) 
71-12.87) 


1.16(1. 
1.31(1, 


07-1 
15-1 


.48) 
.55) 


2.02(1 
2.26(2 


.89-2.37) 

.04-2.50) 


notera palauana 


notera gianda 


10 


72, 


0(59-83) 


9 


.44(7. 


92-10.57) 


1.33(1. 


25-1 


.41) 


2.43(2 


.34-2.50) 


biakensis 


1 


90 




12 


.90 




1.23 






2.22 





1. Form with normal ribbing 

2. Form with widely spaced ribbing 



H/D RATIO 



WHORLS 



4 1/2(4 1/8-5) 
4 3/8-(4-5) 



calli. 0.448(0.428-0.478) 

tumid. 0.581(0.528-0.642) 

micro. 0.473(0.472-0.474) 4+ 

n.not. 0.572(0.537-0.625) 4+(3 3/4-4 5/8) 

n.pal. 0.581(0.527-0.618) 

n.gia. 0.549(0.526-0.573) 

biake. 0.556 



4 l/8+(4-4 1/2) 



UMBILICUS 



D/U RATIO 



APICAL 
CORDS 



0.69(0.58-0.86) 2.98(2.79-3.27) 8.27(7-9) 

0.63(0.51-0.82) 3.56(2.98-4.18) 9.66(8-13) 

0.58(0.54-0.63) 3.67(3.55-3.79) 9 

0.43(0.38-0.56) 4.75(4.24-5.26) 12.9(11-14) 

0.57(0.49-0.66) 4.00(3.72-4.47) 13.3(11-15) 

4 l/4-(4 1/8-4 3/8) 0.63(0.59-0.69) 3.89(3.57-4.16) 12.4(11-33) 

3 7/8 0.49 4.50 14 



SPIRE 
ELEVATION 

0.06(0.03-0.07) 
0.11(0.05-0.20) 

0-0.75 
0.11(0.07-0.13) 
0.17(0.10-0.28) 
0.13 
0.12 



BODY WHORL 
WIDTH 


SP/BWW 


PR 


C 


P 


calli. 


0.73(0. 
0.83(0. 
0.73(0. 


69-0 

72-0 
72-0 


.76) 
.95) 
.74) 


0. 
0. 
0, 


075(0 
131(0 
068 


.048-0, 
,068-0 


,089) 
,250) 


2 
2 

1 


1 
1 



3+3 
3+5-6 
2 


tumid. 


micro. 


n .not . 


0.75(0. 


71-0 


.81) 


0, 


142(0 


.089-0 


,186) 


3+0-l_ 


-1 


5 


n.pal. 


0.82(0. 


76-0 


.94) 


0, 


212(0 


.107-0, 


298) 


2 


-I 


1-4-5-6. 


n.gia. 


0.87(0. 


84-0 


.92) 


0, 


151(0 


,143-0, 


157) 


3+0-1. 


-I 


1-4-1 


biake. 


0.76 






0, 


152 






1 









this aspect, but it is an important problem for work 
with new and better preserved specimens. 

The name Jokajdon refers to the occurrence of its 
two species in the Jokaj district of Ponape and their 
development of many complicated apertural barriers. 



Jokajdon tumidulus 

92d-f. 



(Mollendorff, 1900). Figure 



Endodonta (Thaumatodon) callizona var. tumidula Mollendorff, 
1900, J. Malacol., 7 (5), p. 110 Mpomp and Naupilo, Ponape, 
Caroline Islands. 

Diagnosis. Shell very small, variable in size, diameter 1.88- 
2.73 mm. (mean 2.24 mm.), with 4-5 very tightly coiled whorls. Apex 
and early spire flat or barely protruding, lower spire descending 
slightly, body whorl much more rapidly, spire protrusion more than 
Ve body whorl width, H/D ratio 0.528-0.642 (mean 0.581). Apical 
sculpture of 8-13 (mean 9.66) fine, low spiral cords, whose interstices 



are 2-3 times their width. Postnuclear sculpture highly variable, 
normally (52%) with high, narrow, crowded, slightly protractively 
sinuated radial ribs, 74148 (mean 106.1) on the body whorl, whose 
interstices are less than twice their width; many examples (29%) 
with the ribs very widely spaced, 39-69 (mean 53.6) on the body 
whorl, whose interstices are 8-12 times their width, and often (19%) 
with major radial ribbing reduced to absent. Ribs/mm. 10.35-22.9 
(mean 14.61) in normal form; 5.80-10.62 (mean 8.13) in widely 
spaced form. Microsculpture of fine radial riblets, 2-5 between each 
pair of major ribs in normally ribbed forms, 10-15 in widely spaced 
ribbed forms, with finer and more crowded spiral riblets, plus secon- 
dary spiral cords that are more prominent than the microradial rib- 
lets. Umbilicus open, broadly U-shaped, regularly decoiling, con- 
tained 2.98-4.18 times (mean 3.56) in the diameter, margins shoul- 
dered, walls of umbilicus distinctly flattened. Sutures deep, whorls 
strongly rounded above, compressed laterally with evenly rounded 
outer margin and strongly rounded basal margin. Aperture ovate, 
compressed laterally, inclined less than 10 from shell axis. Parietal 
barriers 2, extending posteriorly almost Vi whorl: upper a high, 
broadly expanded blade on posterior half, next quarter descending 







FIG. 92. a-c, Jokajdon callizonus (Mollendorff). Station 118, Ponape, Caroline Islands. BPBM 154161; d-f, Jokajdon tumidulus (Mbllen 
dorff). Station 146, Ponape, Caroline Islands. BPBM 157984. Scale line equals 1 mm. (MM). 



224 



SYSTEMATIC REVIEW 



225 



rather sharply with a droplike accessory protrusion on lower side, 
anterior quarter low and broad, tapering to end point; 2nd much 
higher, sinuately twisted and crescentic over posterior %, anterior 
quarter a broad threadlike ridge tapering to a point anterior of upper 
parietal termination. Columellar wall with a single very high, cres- 
centic barrier, very broadly expanded above and slightly sinuated 
posteriorly, anterior end twisted upward along columellar lip edge to 
terminate on lower parietal wall. Palatal barriers 3, very high, ex- 
tending posteriorly more than VH whorl, plus 6, rarely 5, accessory 
traces: lower at baso-columellar margin, very high and broadly ex- 
panded to a T-shape on posterior %, with anterior quarter weakly 
expanded above, descending abruptly to lip margin; 2nd moderately 
recessed, much lower, teardrop-shaped on posterior half, with 
gradual anterior descension of a thin lamellar blade; 3rd equal in 
height to 1st, sinuately twisted transversely upward, a thin blade 
with abrupt anterior descension, moderately recessed. Accessory 
traces, triangular knobs to very short lamellae, normally located 
between columellar and 1st palatal, 1st and 2nd palatals, 2nd and 
3rd palatals, and 3 above 3rd palatal. Rarely one of the lower traces 
absent; upper traces usually short crescents, lower 3 highly variable 
in shape. 

Jokajdon tumidulus (fig. 92d-f, table L) is much 
higher, has a narrower umbilicus, much thicker body 
whorl, and shows many differences in barrier structure 
from J. callizonus (fig. 92a-c). The large number of 
complexly expanded and twisted barriers are unique 
among Pacific Island endodontoid snails and prevent 
confusion with any other species. 

Description. Shell rather small, with slightly less than 4% very 
tightly coiled whorls. Apex and early spire flat, lower spire descend- 
ing a little, body whorl descending sharply, H/D ratio 0.562. Apical 
whorls 1%, sculpture of 10 narrow, rather widely spaced spiral cords. 
Postnuclear whorls with prominent, moderately widely spaced, pro- 
tractively sinuated radial ribs becoming quite crowded on last por- 
tion of body whorl. Microsculpture a lattice of very fine radial riblets, 
crossed by much finer and more crowded spiral riblets with a promi- 
nent secondary sculpture of moderately widely spaced spiral cords. 
Sutures deeply impressed, whorls strongly rounded above, greatly 
compressed laterally, with evenly rounded lateral and basal margin. 
Umbilicus U-shaped, slightly decoiling, contained 3.19 times in the 
diameter, slightly flattened internally with strongly rounded mar- 
gins. Aperture ovate, compressed laterally, inclined less than 5 from 
the shell axis. Parietal barriers 2, extending slightly more than Vi 
whorl: upper moderately high, slightly twisted upward posteriorly 
with a broadly expanded, downwardly pointing, hatchet-shaped ac- 
cessory plate on anterior half; lower quite high, crescentic, thick, 
rounded above, slightly sinuately twisted posteriorly with very 
gradual anterior descension, reaching past lip edge. Columellar bar- 
rier high, broadly rounded, twisting upward anteriorly, reaching 
columellar-parietal margin, sharply descending posteriorly. Major 
palatal barriers 3, with 6 accessory lamellae: lower palatal basal, a 
short, high, sinuately twisted blob greatly thickened in middle, only 
moderately recessed; middle palatal deeply recessed, much smaller 
in size, slightly thickened medially, crescent-shaped; upper palatal 
peripheral in position, hemispheric in shape, roundly expanded 
above, twisting laterally downward posteriorly, moderately recessed 
within aperture. Palatal traces located between columellar and 1st 
palatal, 1st and 2nd palatal, 2nd and 3rd palatal and 3, decreasing in 
size, above upper palatal. All traces short, ridgelike, or knoblike. 
Height of lectotype 1.42 mm., diameter 2.52 mm. 

Lectotype. Caroline Islands, Ponape.SMF 165702. 

Range. Ponape, Caroline Islands. 

Paratypes. SMF 165703, SMF 165756, BPBM 
86247. 

Material Ponape (16 specimens, BPBM 161092, 
BMW, FMNH 159359): one-quarter to three-quarters 
of the way up Mt. Tolotom (Stations 92, 93) at 800- 
1200 ft. elevation (29 specimens, BPBM 154082, BPBM 



156680); Mt. Kuporujo (Station 121) at 500-900 ft. ele- 
vation (1 specimen, BPBM 154339); Wone, south side 
Mt. Tolotom (Stations 95, 98), 1 A mile west at 900- 
1,000 ft. elevation (19 specimens, BPBM 156730-1, 
BPBM 156817-8); north of Sabalop (Station 123), 
south of Alokap-Najijo Mt. range at 60-150 m. eleva- 
tion (4 specimens, BPBM 157318); above Mpomp (Sta- 
tion 125), 2 miles inland, at 75 ft. elevation (8 speci- 
mens, BPBM 157362-4); Toreairuku Mt., southwest 
side (Station 127) 2'/2 miles inland at 250-750 ft. eleva- 
tion (1 specimen, BPBM 157420); Mt. Nanalaut, north 
slope (Stations 130, 132), at 400-1,900 ft. elevation (30 
specimens, BPBM 157469, BPBM 157513-21); Palikir 
(Station 140), Mt. Tamantamansakir at 1,000-1,250 ft. 
elevation (6 specimens, BPBM 157866-7); Palikir (Sta- 
tion 139 = Station 138), Jokaj, 600-900 ft. elevation (1 
specimen, BPBM 157800); Jokaj Island (Station 136), 
Rock, at 300-600 ft. elevation (1 specimen, BPBM 
157704); Mt. Tolomain (Station 91), west side at 
1,000-1,500 ft. elevation (4 specimens, BPBM 
154005-6); back of Naupilo (Station 133) (11 speci- 
mens, BPBM 157566-7); summit of mountain above 
Station 145 (Station 146) at 860 ft. elevation (5 speci- 
mens, BPBM 157984-5). 

Remarks. Although Mollendorff considered that 
Jokajdon tumidulus was a variety of J. callizonus, the 
much thicker body whorl that descends rapidly and the 
more highly developed apertural barriers separate the 
two specifically (fig. 92b, e). At most localities on 
Ponape surveyed by Kondo, they did not occur to- 
gether, but specimens of both species were found at 
Stations 91, 92, 93, and 95 on Roi Peak. 

There is considerable variation in the sculpture of 
the shell in J. tumidulus, some specimens having the 
radial sculpture reduced to vague growth wrinkles. In 
other cases the radial ribs are very widely spaced 
(39-69 on the body whorl), and in most there is very 
strong, relatively crowded radial ribbing (79 to over 
150 on the body whorl). There was some geographic 
basis to this variation, although wherever sufficient 
material was collected, generally two types were found 
together. The shells with greatly reduced radial 
sculpture were all found on the slopes of Mt. Tolotom, 
occurring alone at Station 92, but together with nor- 
mally sculptured shells, although greatly in predomi- 
nance, at Stations 93 and 95. The shells with very 
widely spaced but nevertheless prominent radial ribs 
were concentrated in north central Ponape, occurring 
with normally sculptured shells at Stations 132 and 
140, but alone at Station 91. Specimens with only 
strong radial ribbing were found at Stations 98, 123, 
125, 127, 130, 132, 133, 136, 139, and 146. The number 
of each sculptural type at the stations is summarized in 
Table LI. 

No specimens at all transitional in sculptural type 
were seen. Because the specimens of all three types are 
identical in form and apertural barriers, I consider 
these to be minor genetic variations, probably in the 
process of spreading through the populations. The type 



226 



SOLEM: ENDODONTOID LAND SNAILS 



TABLE LI. - SCULPTURAL VARIATION IN 
JOKAJDON TUMIDULUS 



Station 

91 

92 

93 

95 

98 
121 
123 
125 
127 
130 
132 
133 
136 
139 
140 
146 

TOTALS 



Type of sculpture 
normal reduced widely spaced 
4 



7 

1 
5 
1 
4 
8 
1 
3 

11 
11 
1 
1 
3 
5 

62 



2 

20 
13 



16 



35 



23 



specimens all have normal sculpture. In addition to the 
sculptural variation discussed above, a few individuals 
from Station 146 on Jokaj Island, a single shell from 
Station 132 on Ponape, and specimens remaining in 
the type set are very large in size (table XLIX) and 
have the upper parietal barrier reduced in prominence. 
These probably represent gerontic individuals, and the 
variation is not of systematic importance. 

Most specimens of J. tumidulus were monochrome 
in coloration, a light reddish yellow-brown, but a few 
showed traces of the lighter spiral zones that are char- 
acteristic of J. callizonus. 

Description of soft parts. No fully satisfactory dissections were 
obtained. Two rather squashed penes and poorly preserved apical 
genitalia were observed. Apical genitalia showed no differences from 
the structures seen in Jokajdon callizonus. Penis about 0.8-0.9 mm. 
long, cylindrical, tapering to middle and then to atrium, vas deferens 
and penial retractor as in J. callizonus. Interior of penis with weak 
longitudinal pilasters from apex to midpoint, a Sinployea-type 
stiniulatory pad developed just below midpoint. A muscle band in- 
serts on penioviducal angle. 

(Based on BPBM 156731, 2 whole examples.) 

Jokajdon callizonus (Mollendorff, 1900). Figures 
92a-c, 93a-d. 

Endodonta (Thaumatodon) callizona Mollendorff, 1900, J. 
Malacol, 7 (5), p. 110 Ponape, Caroline Islands. 

Diagnosis. Shell minute, diameter 1.79-2.40 mm. (mean 2.06 
mm.), with 4'/s-5 extremely tightly coiled whorls. Apex and spire 
flat, or barely elevated, body whorl not descending more rapidly, 
spire protrusion about 'A< body whorl width, H/D ratio 0.428-0.478 



(mean 0.448). Apical sculpture of 7-9 (mean 8.27) fine spiral cords, 
whose interstices are 3-5 times their width. Postnuclear sculpture of 
prominent, narrow, rounded, crowded, slightly protractively sin- 
uated radial ribs, 99-153 (mean 127.6) on the body whorl, whose 
interstices are 1-2 times their width. Ribs/mm. 16.2-21.7 (mean 
18.9). Microsculpture of fine radial riblets, 2-5 between each pair of 
major ribs, much finer and more crowded spiral riblets, with 
crowded, somewhat obscure secondary spiral cords equal in size to 
microradial riblets. Umbilicus broadly open, cup-shaped, regularly 
decoiling, contained 2.79-3.27 times (mean 2.98) in the diameter, 
margins strongly rounded to weakly shouldered. Sutures deep, 
whorls strongly rounded to shouldered above, very strongly flattened 
laterally, with evenly rounded periphery and weakly compressed 
basal margin. Aperture subovate, strongly compressed laterally, in- 
clined less than 10 from shell axis. Parietal barriers 2, extending 
posteriorly more than Vi whorl almost to line of vision: upper very 
high and bladelike on posterior half, thin and sinuately twisted, 
posterior 8th twisted downward, abruptly descending about midpoint 
to threadlike anterior half reaching beyond lip margin; 2nd equal in 
height posteriorly to 1st, not sinuately twisted, with sharper descen- 
sion near midpoint to threadlike anterior portion extending beyond 
end of upper. Columellar barrier a low, sometimes triangular knob- 
like ridge, deeply recessed within aperture, visible only by extreme 
tilting of aperture. Palatal barriers 3, moderately recessed, extend- 
ing posteriorly more than Vs whorl, with 3 superior accessory traces: 
lower basal in position, very broadly expanded on posterior third and 
T-shaped, anterior half a thin, high blade with abrupt anterior de- 
scension to point midway across columellar callus; 2nd slightly lower 
than 1st, sinuately twisted outward and upward on posterior %, a 
lateral bifurcation on side of posterior half pointing inward and 
downward, crescentic, with gradual anterior descension, anterior 
end of main blade abruptly descending to point well behind anterior 
end of 1st; 3rd rather deeply recessed, shorter, posterior half grossly 
expanded into teardrop shape, with gradual anterior descension. Ac- 
cessory traces very short, deeply recessed, broadly rounded, low cres- 
centic to cordlike lamellae, evenly distributed between upper palatal 
and palatal-parietal margin, second usually slightly larger. 

Jokajdon callizonus has a much narrower body 
whorl, (mean body whorl width 0.73 mm.), wider um- 
bilicus, lower H/D ratio, and fewer palatal traces than 
does J. tumidulus (table L). In the latter, the body 
whorl descends rapidly and is much wider (mean body 
whorl width 0.83 mm.), the shell height is always 
larger and the umbilicus usually wider (fig. 92a-f). 
Other Micronesian species with numerous apertural 
barriers are much larger (Semperdon kororensis and S. 
xyleborus, fig. 98a-f) or have beaded barriers and usu- 
ally angulated peripheries (Aaadonta, see Solem, 
1976b, pp. 467-487). 

Description. Shell very small, with 4% very tightly coiled 
whorls. Apex and early spire flat, lower whorls descending slightly, 
H/D ratio 0.463. Apical whorls 1%, sculpture of 7 prominent, rather 
widely spaced spiral cords. Postnuclear whorls with slightly protrac- 
tively sinuated, very crowded radial ribs, about 153 on the body 
whorl, whose interstices are less than twice their width. Micro- 
sculpture a lattice of fine microradial and spiral riblets with a secon- 
dary sculpture of much stronger, relatively closely set spiral cords. 
Sutures channeled, whorls evenly rounded above, strongly flattened 
laterally with gently rounded basal margin. Umbilicus broadly V- 
shaped, regularly decoiling, contained 3.09 times in the diameter. 
Color light reddish yellow-brown without darker flammulations or 
obvious spiral banding. Aperture ovate, strongly flattened laterally 
with strongly sinuated margin, inclined less than 10 from shell axis. 
Parietal barriers 2, extending posteriorly more than 1 A whorl, low 
and threadlike anteriorly becoming high and lamellate posteriorly. 
Columellar wall obscured by dried portion of animal. Major palatal 
barriers 3, with 3 accessory traces: lower palatal situated on baso- 
columellar margin, broadly expanded above, sinuately twisting 
downward onto basal lip; middle palatal moderately high, broadly 
expanded above, situated opposite lower parietal; upper palatal 



SYSTEMATIC REVIEW 



227 



peripheral in position, very broadly expanded above, somewhat 
sinuately twisted, moderately deeply recessed within aperture. Ac- 
cessory palatals low, short, broad, deeply and equally recessed within 
aperture, located above upper palatal. A 4th palatal knob, very low 
and broad, located between lower 2 palatals. All major barriers mi- 
nutely pustulose above. Height of lectotype 1.04 mm., diameter 2.24 
mm. 

Lectotype. Caroline Islands, Ponape. Collected by 
Etscheid. SMF 165700. 

Range. Ponape, Caroline Islands. 

Paratypes. BPBM 86246, SMF 165701. 

Material. Ponape: Mt. Tolotom (Stations 92, 93), 
one-quarter to three-quarters of the way up at 800- 
1,200 ft. elevation (10 specimens, BPBM 9688, BPBM 
156679); Metalanim to Nipit (Station 118), 450 ft. in- 
land at 300-900 ft. elevation (14 specimens, BPBM 
154161, BPBM 157282); west side Mt. Tolomain (Sta- 
tion 91) at 1,000-1,500 ft. elevation (1 specimen, 
BPBM 154007); Wone (Station 95), south side of Mt. 
Tolotom at 900 ft. elevation (1 specimen, BPBM 
156732); Paishapal Hill (Station 113), Metalanim, at 
650-700 ft. elevation (1 specimen, BPBM 157210). 

Remarks. Although specimens of Jokajdon cal- 
lizonus and J. tumidulus were taken at Stations 91, 92, 
93, and 95, no intergradation could be detected. The 
differences between them are substantial, and I con- 
sider that they represent valid species. A most unusual 
feature of J. callizonus is the spirally zoned color pat- 
tern seen in most examples. One or two light yellow 
spiral bands will interrupt the basic reddish brown 
color. These zones are highly variable in position and 
width. No attempt was made to analyze this variation. 
Much less size variation was found between local popu- 
lations than in J. tumidulus (table XLIX). 

Description of soft parts. Foot and tail much shorter than shell 
diameter, tapering posteriorly. Pedal grooves and slime network typ- 
ical. Head retracted in all studied material. Body color white with a 
faint yellow tinge, no darker markings. Mantle collar (MC) rather 
wide, edge thickened, no glandular extension onto pallial roof. Anus 
(A) opening slightly behind external ureteric pore (KX). 

Pallial region (fig. 93a) extending apically for more than % 
whorl, quite narrow. Lung roof clear, no granulations. Kidney (K) 
bilobed, rectal arm about 1.51 mm. long, covering hindgut and ex- 
tending halfway down parietal wall. Pericardia! arm very short and 
narrow, scarcely longer than heart. Ureter (KD) without unusual 
features. Heart (H) parallel to hindgut, a little shorter than pericar- 
dial kidney arm. Principal pulmonary vein (HV) narrow, un- 
branched. Hindgut (HG) extending slightly apical of pallial cavity. 

Ovotestis a single clump of 3 or 4 clavate acini above stomach 
apex, lying parallel to whorl sides and weakly iridescent. Hermaph- 
roditic duct very slender, not convoluted. Albumen gland short, com- 
pact, acini small. Talon (fig. 93d, GT) with globular head. Prostate 
(fig. 93b, DG) with a few large acini. Uterus (UT) typically bipartite. 

Vas deferens (VD) highly convoluted, entering laterally on head 
of penis with junction of retractor muscle (fig. 93c). Penial retractor 
(PR) long, inserting directly in head of penis-vas junction. Penis (P) 
short, cylindrically bitapered, about 0.3-0.4 mm. long, upper tapered 
area with a few weak pilasters, internal structure of median bulge 
not worked out in detail, but apparently with a thick muscular pad or 
pilaster, probably as in J. tumidulus. Atrium (Y) short, without un- 
usual features. 

Free oviduct (UV) quite short, tapering. Spermatheca (S) with 
large oval head, slender shaft at most weakly expanded before join- 
ing with free oviduct. Vagina (V) quite long, narrow, with internal 
pilasters. Free muscle system unusual in having a separate muscle 




FIG. 93. Anatomy of Jokajdon and Palline: a-d, Jokajdon cal- 
lizonus (Mollendorff). Sta. 118, Ponape. BPBM 154161. a, pallial re- 
gion, b, genitalia. c, detail of penis-vas deferens junction, d, detail of 
talon-carrefour region; e f, Palline notera notera, new species and 
subspecies. Station 201, Peleliu. BPBM 159946. e, pallial region, f, 
genitalia; g h, Palline micramyla, new species. Station 137, Palikir, 
Ponape. Holotype. BPBM 157731; g, pallial region; h, genitalia. 
Scale lines equal 1 mm. (a-b, e-f, MO; c-d, JC & AS; g-h, PS). 



228 



SOLEM: ENDODONTOID LAND SNAILS 



band inserting on penioviducal angle. Right ommatophoral retractor 
passing through penioviducal angle. 

(Based on BPBM 154161, 4 whole specimens.) 



Genus Palline, new genus 

Shell smaller than average, with 3 7 /s-4i4 normally coiled whorls. 
Apex and spire slightly to moderately elevated, body whorl descend- 
ing more rapidly. Apical sculpture of a variable number of prominent 
spiral cords. Postnuclear sculpture of narrow, moderately spaced to 
crowded, almost vertically sinuated radial ribs. Microsculpture of 
extremely fine radial and spiral riblets with prominent, rather 
widely spaced secondary spiral cording. Umbilicus rather broadly 
V-shaped to narrowly U-shaped, margins rounded. Whorls strongly 
rounded above and compressed laterally to strongly flattened later- 
ally above rounded periphery, aperture inclined less than 10 from 
shell axis. Parietal barriers variable in number, from 3, short and 
high, to 1 or 2. Columellar wall with or without a low threadlike 
ridge. Palatal barriers usually 4-6, reduced to 2 in micramyla. ab- 
sent in bickenxis. Pallia! region extending l /z whorl apically, kidney 
with icctal arm much longer than pericardial and extending over 
hincigut onto parietal wall, pericardial arm about SCX/f longer than 
heart and greatly reduced in size. Ovotestis with two clumps of al- 
veoli. Vas deferens passing through penial retractor muscle before 
latter inserts on head of penis. Penis of medium length, covered by a 
thin muscle sheath, head bulging with external constrictions indicat- 
ing 3 internal zones: a lateral or upper zone with a tubular verge 
leading from vas deferens: an upper part to the side of the verge or 
shifted downward with 2 circular glandular pilasters; and a lower 
zone with thick circular or horseshoe-shaped pilaster tapering into 
narrowed shaft leading to atrium. 

Type species. Palline notera notera, new species 
and subspecies. 

Palline (fig. 93e, g) agrees with Jokajdon (fig. 93a) 
in structure of the pallial region, although the degree 
of rectal kidney arm descension onto the parietal wall 
is greater in the latter genus. Palikirus has the kidney 
arms less disparate in size than in Palline, but the 
degree of crossing onto the parietal wall is about the 
same. The genitalia of Palline (fig. 93f, h) is unique 
among the Pacific Island Charopidae in the possession 
of a distinct tubular verge. Although Kubaryiellus (fig. 
90g) has a modified pillow-like vergic papilla and the 
Sinployea structure is dominant in Polynesia, none of 
the remaining Micronesian genera have other than a 
simple pore from the vas deferens into the penis. De- 
velopment of a true verge is common in New Zealand 
taxa, and the relatives of Palline may be found in the 
Southern Relict endodontoid taxa. The pair of concen- 
tric glandular pilasters and the thick horseshoe pilas- 
ter of Palline also are very different from the structures 
seen in other Micronesian genera. Passage of the vas 
deferens across the penis head and through the penial 
retractor muscle agrees with most of the Micronesian 
taxa. 

Conchologically, Palline shows few unusual fea- 
tures. Palline notera (fig. 94a-f) from the Palau Islands 
has a very strong callus on the columellar and palatal 
walls that at times "buries" the smaller barriers. 
Otherwise it is a small species with quite prominent 
ribbing, a protruded spire, many spiral apical cords, a 
relatively narrow umbilicus, and slight lateral flatten- 
ing of the whorls. Palline micramyla (fig. 95c-e) from 
Ponape has much finer and more crowded radial rib- 



bing, a less protruded spire, fewer spiral apical cords, a 
slightly wider umbilicus, and the whorls are strongly 
flattened laterally above the periphery rather than on 
the outer margin. Both species agree in having a 
somewhat lower whorl count than the other toothed 
Micronesian species and in having the major ribbing 
almost vertically sinuated. Palline biakensis (fig. 
96a-c) from Biak Island, West Irian, has only one 
parietal and no palatal barriers. Otherwise it falls 
within the range of variation for P. notera notera. Shell 
color is light yellow or reddish yellow-horn without 
conspicuous darker markings in all species, although 
some examples of Palline notera gianda show faint 
flammulations. Although the shell form of the Ponape 
species is much more similar to some of the other 
Caroline Island taxa, the anatomy clearly places it 
with Palline notera. 

On the basis of initial conchological study, I had 
assumed that Semperdon uncatus was closely related. 
Discovery that it had such a very different penial 
structure was one of the more surprising aspects of this 
project. Shell differences from Palline notera are dis- 
cussed below. 

The name Palline is manufactured from the first 
three letters of Palau and the last four of Caroline to 
indicate that this genus is found in both areas. 

Palline notera, new species. 

Populations on Babelthuap, Koror, and Peleliu 
agree in basic shell form and structure, but show minor 
differences in barrier size and number plus shell size 
and proportions. No anatomical differences were ob- 
served between the populations, and subspecific recog- 
nition has been given. The exact differences are out- 
lined under the diagnosis of the three races Palline 
notera notera from Peleliu, Palline notera gianda from 
Babelthuap, and Palline notera palauana from Koror. 

Of the Micronesian species, only Semperdon un- 
catus (fig. 97a-c) is apt to be confused with Palline 
notera. At first inspection, shells of S. uncatus could be 
interpreted as gerontic individuals of the latter species. 
Reduction in apertural callus and barrier size in very 
large gerontic individuals is known for Minidonta 
simulata, Solem & Cooke (Solem, 1976b, p. 147, fig. 
70d). Barrier position in S. uncatus agrees withPalline 
notera. The former lacks secondary spiral cording, av- 
erages a half whorl more, has fewer spiral cords on the 
apex, and has a totally different penial structure. The 
similarities are even greater than those between Sin- 
ployea kusaieana and Trukcharopa trukana, but the 
relationship is no closer. 

Conchological differences between Palline notera 
and P. micramyla were outlined above under the 
generic discussion. Anatomical differences are few, 
with the shorter penis and downward shift of the 
thin-walled pocket the most obvious alteration. Palline 
biakensis is almost identical with P. notera notera, but 
is immediately recognizable by having only a single 
parietal and no palatal barriers. 



SYSTEMATIC REVIEW 



229 



Palline notera notera, new species and subspecies. 
Figures 93e-f, 94a-c. 

Diagnosis. Shell very small, diameter 1.89-2.37 mm. (mean 
2.02 mm.), with 3%-4% rather tightly coiled whorls. Apex and spire 
slightly and evenly elevated, body whorl descending more rapidly, 
spire protrusion about l h body whorl width, H/D ratio 0.537-0.625 
(mean 0.572). Apical sculpture of 11-14 (mean 12.9) prominent spi- 
ral cords whose interstices are about 2-3 times their width. Postnu- 
clear sculpture of high, rounded, prominent, almost vertically 
sinuated radial ribs, 58-95 (mean 74.0) on the body whorl, whose 
interstices are 2-5 times their width. Ribs/mm. 9.37-15.66 (mean 
11.63). Microsculpture of barely visible radial riblets, 5-12 between 
each pair of major ribs, equally fine spiral riblets, plus much more 
prominent secondary spiral cords, whose interstices are 3-6 times 
their width. Umbilicus rather narrow, U-shaped, slightly and regu- 
larly decoiling, contained 4.24-5.26 (mean 4.75) times in the diame- 
ter, margins strongly rounded. Sutures deep, whorls strongly 
rounded above and on basal margin, compressed laterally, with 
evenly rounded outer margin. Aperture ovate, compressed laterally, 
inclined about 5 from shell axis. Parietal barriers 3, short, high, 
extending posteriorly about Vs whorl, occasionally with an accessory 
trace: upper with abrupt descension over posterior 8th, very much 
higher and crescentic over next quarter, with gradual anterior de- 
scension becoming rapid over anterior quarter, posterior elevated 
portion slightly twisted downward; 2nd equal in height and posterior 
descension to 1st, crescentic until % point, with gradual anterior 
descension on last 3rd to point beyond termination of upper parietal; 
3rd greatly reduced in height, a raised threadlike ridge with more 
elevated crescentic portion on posterior 3rd, equal in length to 2nd 
parietal. Columellar wall with single threadlike ridge, lying parallel 
to plane of coiling, reaching to top of very thick columellar callus, 
often (2 of 13) covered by callus, which extends on palatal wall al- 
most to periphery. Palatal barriers 5, short, slightly recessed: lower 
basal in position, very high and crescentic, slightly expanded above, 
with abrupt sinuately twisted descension to lip edge, more gradual 
descension posteriorly; 2nd and 3rd less than % height of 1st, cres- 
centic, moderately expanded above, slightly recessed, with much 
more gradual anterior descension; 4th higher than 3rd, lower than 
1st, crescentic, expanded above with rather sharp anterior descen- 
sion, slightly subperipheral; 5th slightly lower than 3rd, but other- 
wise matching it in structure, located above level of upper parietal, 
moderately recessed. 

Palline notera notera (fig. 94a-c) is smaller and 
has a much narrower umbilicus than either P. notera 
gianda (fig. 95a-b), which usually has only four palat- 
als and lacks the columellar, or P. notera palauana (fig. 
94d-f), which has only two parietals, usually no col- 
umellar, and only four palatals. Palline biakensis (fig. 
96a-c) has only one weak parietal and no palatal bar- 
riers, otherwise it is nearly identical to P. notera 
notera. 

Description. Shell very small, with 4Vs moderately tightly 
coiled whorls. Apex and spire almost flat, body whorl descending 
slightly, H/D ratio 0.565. Apical whorls 1%, sculpture of 11 promi- 
nent spiral cords. Postnuclear whorls with prominent, high, rounded, 
vertically sinuated radial ribs, 84 on the body whorl, whose in- 
terstices are 34 times their width. Microsculpture of fine, crowded 
radial riblets, crossing very faint crowded spiral ribs with a secon- 
dary sculpture of prominent, rather widely spaced, spiral cords. Su- 
tures deeply impressed, whorls strong'y rounded above with slightly 
flattened basal margin, compressed laterally. Color light yellow- 
horn, without darker markings. Umbilicus narrowly U-shaped, 
slightly and regularly decoiling, contained 5.24 times in the diame- 
ter. Aperture ovate, inclined less than 5 from the shell axis with 
moderately prominent internal callus. Parietal barriers 3, extending 
posteriorly less than Ve whorl; upper a thin, high barrier with 
gradual anterior descension over anterior half, slightly expanded 
above on posterior 8th and twisted downward; 2nd parietal higher, 
elevated and expanded on posterior half, with more gradual anterior 



descension; 3rd parietal much lower, a threadlike ridge. Columellar 
wall with a low, broad swelling reaching top of heavy callus. Palatal 
barriers 5, extending about Me of a whorl: 1st and 4th relatively 
high, slightly expanded above, crescent-shaped; 2nd, 3rd, and 5th 
much lower, narrower, shorter, more deeply recessed. Height of 
holotype 1.22 mm., diameter 2.16 mm. 

Holotype. Palau Islands: Peleliu, Station 201, 
short distance to north of phosphate mine at 35-200 ft. 
elevation. Collected by Yoshio Kondo and natives on 
April 29, 1936. BPBM 159946. 

Range. Peleliu, Palau Islands. 

Paratypes. Peleliu: short distance to north of 
phosphate mine (Station 201) at 35-200 ft. elevation 
(41 specimens, BPBM 159944-9); Omurbrogol Mt, 
Asias Village (Station 182), Vfe-% mile inland at 300- 
400 ft. elevation (4 specimens, BPBM 159432-3); on 
hill east of Station 176 (Station 184), 100 yd. inland at 
200 ft. elevation (1 specimen, BPBM 159517); 300-400 
yd. north of Station 201 (Station 203) on hillside (1 
specimen, BPBM 159994). 

Remarks. One specimen had an accessory trace 
between the second and third parietals, but otherwise 
there was little variation in the apertural barriers. 
Apparently this species is widely distributed on 
Peleliu. 

It was not possible to illustrate the interior of the 
penis. The verge has only about one-quarter of its 
length as a free tip with central pore and wrinkled 
muscular sides, whereas the upper three-quarters is a 
thinner-walled, slightly expanded chamber that may 
function as an epiphallus in forming a sperm packet. 
Vas deferens enters the apex of this chamber after 
passing through the penial retractor muscle. Lateral to 
the verge is a very thin-walled, bulging pocket with 
two concentric rings of glandular pilasters, with a con- 
stricted opening to the lower chamber with its very 
wide and solid pilaster. 

Description of soft parts. Foot and tail much less than shell 
diameter, shape typical. Sole, pedal grooves, and slime network 
without unusual features. Body color yellow-white, without darker 
markings. Anus (A) opening inside pneumostomal pore, anterior of 
external ureteric pore (KX). 

Pallial region (fig. 93e) extending Vz whorl apically. Lung roof 
without granulations. Kidney (K) short and broad, 1.0-1.1 mm. long, 
rectal lobe much longer than pericardial, partly extending onto 
parietal wall. Ureter (KD) with primary and first part of secondary 
sections compacted between arms of kidney, latter portion of secon- 
dary ureter slightly tapered to anterior. Heart (H) shorter than 
pericardial arm of kidney, lying parallel to hindgut. Principal pul- 
monary vein (HV) unbranched. Hindgut (HG) without unusual fea- 
tures. 

Ovotestis with 2 clumps, 1 above the other, of 3-4, iridescent 
white, long alveoli imbedded in digestive gland above stomach apex. 
Hermaphroditic duct not observed in adult condition. Albumen gland 
(fig. 93f, GG) with relatively large acini. Talon not observed. Pros- 
tate (DG) and uterus (UT) without unusual features. 

Vas deferens (VD) entering penis head after passing through 
penial retractor muscle in transversing penis head. Penial retractor 
(PR) short, arising from diaphragm, inserting onto penis head after 
enfolding vas deferens. Penis (P) with elongated bulging head of 
about 0.7-0.8 mm. length followed by a much shorter, tubular shaft. 
Upper portion covered by a thin muscle sheath, internally very com- 
plex, apparently with a verge occupying one side of upper bulge, 
other half with 2 concentric circular pilasters with exceedingly thin 




a 





a-f 



e 






FIG. 94. a-c,Palline notera notera, new species and subspecies. Station 201, Peleliu, Palau Islands. Holotype. BPBM 159946; A-f,Palline 
notera palauana, new subspecies. Station 220, Ittabagamin, Koror, Palau Islands. Holotype. BPBM 159046. Scale line equals 1 mm. (SG). 



230 



SYSTEMATIC REVIEW 



231 



walls, then a thick pilaster occupying lower bulge of penis head that 
is horseshoe-shaped with ends tapering toward atrium. Atrium (Y) a 
narrow tube. 

Free oviduct (UV) very slender, tapering from uterus. Sper- 
matheca (S) with elongately oval head, lower half of shaft grossly 
expanded. Vagina (V) rather long, equal in diameter to lower part of 
penis. Free muscle and digestive systems without unusual features. 

(Based on BPBM 159946, 2 whole subadults and several frag- 
mentary adults.) 

Palline notera palauana, new subspecies. Figure 
94d-f. 

Diagnosis. Shell small, diameter 2.04-2.50 mm. (mean 2.26 
mm.), with 4-4'/2 tightly coiled whorls. Apex and spire slightly to 
strongly elevated, body whorl descending more rapidly, spire protru- 
sion usually more than Vs body whorl width, H/D ratio 0.527-0.618 
(mean 0.581). Apical sculpture of 11-15 (mean 13.3) prominent spi- 
ral cords, whose interstices are less than twice their width. Postnu- 
clear whorls with high, prominent, rounded, almost vertically 
sinuated radial ribs, 63-101 (mean 78.8) on the body whorl, whose 
interstices are 3-5 times their width. Ribs/mm. 8.71-12.87 (mean 
11.01). Microsculpture of very fine radial riblets, 6-12 between each 
pair of major ribs, crossed by barely visible spiral riblets, with a 
secondary sculpture of much more prominent, rather widely spaced 
spiral cording. Umbilicus rather widely open, U-shaped, slightly and 
regularly decoiling, contained 3.72-4.47 times (mean 4.00) in the 
diameter, margins strongly rounded. Sutures deep, whorls strongly 
rounded above and on basal margin, compressed laterally, outer 
margin evenly rounded. Aperture compressedly ovate, inclined about 
5 from shell axis. Parietal barriers 2, extending posteriorly more 
than '/s whorl, occasionally with a threadlike trace occupying posi- 
tion of 3rd parietal in nominate race: upper high, bladelike, slightly 
more elevated and expanded on posterior quarter, with gradual de- 
scension over anterior half; 2nd usually slightly reduced in height, 
more broadly expanded above on posterior half, with gradual descen- 
sion to point beyond end of upper or no descension until anterior 
quarter. Columellar wall normally without barriers, occasionally 
with a weak threadlike trace. Palatal wall normally (75%) with 4 
rather short, low, crescentic barriers, frequently either 3, 5, or 6: 
lower basal in position, low, broadly rounded, with rather abrupt 
descension across callus to lip margin; 2nd and 3rd slightly lower, a 
little more recessed, with more gradual anterior descension; 4th a 
moderately recessed, supraperipheral, threadlike trace. Callus very 
broad, extending from columellar wall to parietal-palatal margin, 
only slightly decreasing in thickness. 

Palline notera palauana has a relatively pro- 
truding spire, only two parietals, and a wider um- 
bilicus than the nominate race with its three parietals 
and relatively narrow umbilicus. Palline notera gianda 
is larger, has three parietals, and the spire protrusion 
is less (table L). 

Description. Shell small, with 4Va tightly coiled whorls. Apex 
and spire moderately and evenly elevated, last whorl descending 
more rapidly, spire protrusion almost V:> body whorl width, H/D ratio 
0.618. Embryonic whorls 1%, sculpture of 15 prominent, closely 
spaced spiral cords. Postnuclear whorls with prominent, high, verti- 
cally sinuated radial ribs, 101 on the body whorl, whose interstices 
are 2-3 times their width. Microsculpture of fine radial riblets with 
traces of extremely fine spiral riblets and a secondary sculpture of 
relatively prominent, widely spaced spiral cords. Sutures deep, 
whorls strongly rounded above, compressed laterally. Umbilicus 
U-shaped, regularly decoiling, contained 4.00 times in the diameter, 
margins rounded. Color light yellow-horn without darker flammula- 
tions. Aperture ovate, compressed laterally, inclined about 5 from 
shell axis. Parietal barriers 2, extending posteriorly about Vs whorl, 
upper less expanded, slightly reduced in height, with sharper an- 
terior descension. Palatal barriers 4, low and crescentic, extending 
less than Vie whorl: 1st and 3rd palatals prominent, broadly rounded 
short ridges; 2nd and 4th markedly reduced in prominence; 4th a 



threadlike ridge. All palatal barriers situated on a heavy apertural 
callus. Height of holotype 1.55 mm., diameter 2.50 mm. 

Holotype. Palau Islands: Koror, Islet XV, Station 
220, Ittabagamin, at 15-20 ft. elevation. Collected 
under stones and on dead leaves by Yoshio Kondo on 
May 11, 1936. BPBM 159046. 

Range. Koror and satellite islets, Palau Islands. 

Paratypes. Koror: Islet XV (Station 220), It- 
tabagamin, at 15-20 ft. elevation (4 specimens, BPBM 
159046-7); Komakan (Station 219), south below Sta- 
tion 218, 50-75 yd. inland at 75 ft. elevation (1 speci- 
men, BPBM 158783); northeast side, western neck of 
island, Islet XXIX (Station 212), 10 yd. inland at 30- 
100 ft. elevation (2 specimens, BPBM 159107); in forest 
on limestone (1 specimen, BPBM 193146, collected by 
E. V. Hosaka on July 20, 1946); in abandoned quarry, 
Nerubodoru, near lagoon shore, northeast part of is- 
land, 20-25 ft. elevation (44 specimens, USMN, FMNH 
159395, BPBM 212387). 

Remarks. Variation in the degree of spire pro- 
trusion was much more marked than in most species. 
This was caused by two individuals having relatively 
low spires (SP/BWW 0.107 and 0.122) and five having 
high spires (SP/BWW 0.239, 0.250, 0.250, 0.292, 0.298). 
The standard error of the mean for 10 measured exam- 
ples was 0.082, whereas for seven examples of Palline 
notera notera it was 0.013, and for 18 specimens of 
Semperdon xyleborus it was 0.010. Semperdon uncatus, 
with only four examples measured, had a standard 
error of the mean of 0.084 and showed a range of varia- 
tion equal to that seen in P. notera palauana. The grea- 
ter variation in these two taxa is not a function of age 
bias, but a real difference in spire elevation. Further 
study will be required to interpret these changes. 

Specimens were collected both on Koror and on 
two of the satellite islets. Those from Islet XV (Station 
220) are appreciably larger than those from Nerubo- 
doru on Koror (table LII), but the latter are dead 
specimens of uncertain age. Many had been damaged 
during cleaning, and their smaller mean size probably 
has no significance. 

Palline notera gianda, new subspecies. Figure 95a-b. 

Diagnosis. Shell small, diameter 2.34-2.50 mm. (mean 2.43 
mm.), with 4'/s-4% rather tightly coiled whorls. Apex and spire mod- 
erately and evenly elevated, body whorl descending more rapidly, 
spire protrusion more than 'h body whorl width, H/D ratio 0.526- 
0.573 (mean 0.549). Apical sculpture of 11-13 (mean 12.4) prominent 
spiral cords, whose interstices are 2-4 times their width. Postnuclear 
sculpture of prominent, high, rounded, almost vertically sinuated 
radial ribs, 59-83 (mean 72.0) on the body whorl, whose interstices 
are 2-5 times their width. Ribs/mm. 7.92-10.57 (mean 9.44). Micro- 
sculpture of very fine radial riblets, 5-10 between each pair of major 
ribs, barely visible spiral riblets and much more prominent, rela- 
tively crowded secondary spiral cords. Umbilicus rather widely open, 
U-shaped, moderately and regularly decoiling, contained 3.57-4.16 
(mean 3.89) times in the diameter, margins strongly rounded. Su- 
tures deep, whorls strongly rounded above and on basal margins, 
compressed laterally, outer margin evenly rounded. Aperture ovate, 
compressed laterally, inclined about 5 from shell axis. Parietal bar- 
riers 3, extending posteriorly more than '/ whorl, sometimes with an 
accessory trace: upper high and bladelike, abruptly descending on 



232 



SOLEM: ENDODONTOID LAND SNAILS 



TABLE LII. - LOCAL VARIATION IN PALLINE. 



NAME 
notera palauana 



NUMBER OF 
SPECIMENS 



Koror, Sta. 220 3 
BPBM 159046-7 

Nerubodoru, Koror 10 
BPBM 212387 

notera notera 



Sta. 201 9 

BPBM 159945, -7,-9 

notera gianda 

Sta. 15 6 

BPBM 160526-8 



RIBS 



RIBS /MM. 



HEIGHT 



DIAMETER 



92.0i6.25 11.84+0.833 1.41*0.072 2.490.022 

(80-101) (10.19-12.87) (1.32-1.56) (2.45-2.52) 



74.32.19 
(63-84) 



72.13.18 
(58-84) 



72.0t5.17 
(59-83) 



10.740.362 



(8.71-11.93) 



11.38+0.464 



(9.37-13.57) 



7.6110.093 

(7.35-7.85) 



1.280.022 



2.19+0.028 



(1.15-1.38) 



(2.04-2.30) 



1.15+0.018 



2.02+0.025 



(1.07-1.22) 



(1.94-2.15) 



1.3410.022 2.4510.026 

(1.26-1.42) (2.35-2.52) 



H/D RATIO 



n.pal 



WHORLS 



Sta. 220 0.56610.0271 4 1/4(4-4 1/2) 
(0.527-0.618) 

Nerub. 0.586+0.0063 4 1/8(4-4 1/4) 
(0.561-0.618) 



UMBILICUS 



D/U RATIO 



APICAL 
CORDS 



0.6410.011 3.90+0.058 14.3310.664 
(0.63-0.66) (3.80-4.00) (13-15) 

0.5510.011 4.04+0.074 12.83+0.477 
(0.49-0.59) (3.72-4.47) (11-14) 



Sta. 201 0.5700.0063 4+(3 3/4-4 1/4) 
(0.541-0.600) 

n. gia. 



0.4310.006 4.75+0.083 12.56+0.377 

(0.41-0.46) (4.37-5.24) (11-14) 



Sta. 15 0.549+0.0078 4 l/4-(4 1/8-4 3/8) 0.6310.161 3.89+0.082 12.40+0.398 

(0.526-0.573) (0.59-0.69) (3.57-4.16) (11-13) 



posterior 8th, next quarter weakly elevated and moderately ex- 
panded, with gradual anterior descension; 2nd equal in height, ele- 
vated and expanded on posterior 3rd after initial abrupt descension, 
with more gradual anterior descension to point anterior of upper 
parietal; 3rd a very short, deeply recessed, threadlike ridge, some- 
times with posterior half weakly elevated. Columellar wall normally 
without barriers, often (16%) with a weak threadlike ridge on top of 
heavy callus. Palatal barriers normally 4, often 3 (16%) or 5 (16%), 
slightly recessed, extending posteriorly 'Ae-'/s whorl: lower basal in 
position, flatly crescentic, with sharp anterior descension across cal- 
lus almost to lip edge; 2nd slightly lower, a little more recessed, with 
more gradual anterior descension; 3rd slightly subperipheral, point- 
ing between upper 2 parietals, variable in height and width, more 
deeply recessed, equivalent to 4th palatal in nominate race; usually a 
threadlike lump located above 3rd palatal (equivalent to 5th in 
nominate race), often lacking; often a very faint threadlike trace 
present between 2nd and 3rd palatals. 

Palline notera gianda is larger and much more 
widely umbilicated than the nominate race (table L). 
Palline notera palauana has a more strongly protruded 
spire and lacks the 3rd parietal. 

Description. Shell small, with 4V4 tightly coiled whorls. Apex 
flat, spire descending moderately, last whorl descending rapidly, H/D 
ratio 0.547. Apical whorls H4, sculpture of 13 relatively prominent 
spiral cords. Postnuclear whorls with prominent, high, rounded, 
nearly vertically sinuated radial ribs, 78 on the body whorl, whose 
interstices are 2-4 times their width. Microsculpture of fine radial 
riblets, 6-15 between each pair of major ribs, with traces of very fine 
crowded spiral riblets and a secondary sculpture of quite prominent, 
rather widely spaced spiral cords. Sutures deep, whorls strongly 



rounded above, compressed laterally. Umbilicus broadly U-shaped, 
regularly decoiling, contained 4.16 times in the diameter, mar