Rulletin OF THE. Museum of _ Comparative Zoology MOLLUSCAN TAXA AND BIBLIOGRAPHIES OF WILLIAM JAMES CLENCH AND RUTH DIXON TURNER RICHARD |. JOHNSON _ HARVARD UNIVERSITY VOLUME 158, NUMBER 1 CAMBRIDGE, MASSACHUSETTS, U.S.A. 2 September 2003 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Breviora 1952— BULLETIN 1863— Memoirs 1865-1938 Jounsonia, Department of Mollusks, 1941-1974 OcCASIONAL PAPERS ON Mo.uwusks, 1945-— SPECIAL PUBLICATIONS. 1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and Evolution of Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere- dinidea (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution ae Blastozoan funni derms. 284 pp. 4. Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (ede 1983. Advances in Herpetology : and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 7295 pp. 6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the M. C. Z., Vol. 108.) Reprinted 1971. 3 Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter’s Check-list of Birds of the World, vols. 1-16. a ames of the New England Zoological Club 1899-1947. (Complete sets only.) | Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. | This publication has been printed on acid-free permanent paper stock. ff - ‘ 3 ; u % is i . an Pas ae, , * 7 é . sta ¥ wal pau my rr * >! . “ is © The President and Fellows of Harvard College 2003. William J. Clench Taken in 1953. A number of other pictures of Clench appeared in Abbot (1984). Ruth D. Turner Taken in the Department of Mollusks, 1957. A later portrait can be found in Mann (2000). MOLLUSCAN TAXA AND BIBLIOGRAPHIES OF WILLIAM JAMES CLENCH AND RUTH DIXON TURNER RICHARD |. JOHNSON’ CONTENTS INoguaACE wel! Fiat (ee Ree eer ee pe et a ee lintrociichonmpetiees ost “Noh oie stern. Se eT Biographical References and Obituaries Methods and Acknowledgments ____.-_-_ Taxonomic List and Publications __....__. Molluscan Taxa Introduced by William J. Clench and Ruth D. Turner with Their Original References, Type Localities, and Locations of Type Specimens ____- 3 Publications of William J. Clench ___... 25 Publications of Ruth D. Turner _ 38 Publications to which Clench, Turner, or Clench and Turner Made Contributions So RE ES eee KA Oe ee A5 WNW Nee ABSTRACT. Between 1924 and 2002, William J. Clench (1897-1984) and Ruth D. Turner (1914— 2000) introduced over 500 molluscan taxa, almost 70 of them jointly. All of their taxa are included in a single list, giving the original reference, type locality, and location of the type specimens. Their bibliogra- phies are separate. INTRODUCTION Almost immediately after the death of William J. Clench in 1984, Professor Ruth D. Turner began a memorial to him, which was also to include a list of his publications and new taxa. This should have been a simple task since she had available Clench’s own running list of both, which also indicated the catalog number of most of the new species and subspecies he had described. Some 15 years later at the time of her own death in 2000, only the bibli- ography had been made available. She ‘Museum of Comparative Zoology, Harvard Uni- versity, 26 Oxford Street, Cambridge, Massachusetts 02138. had, however, brought all of his papers to- gether, and had prepared an elaborate folder with the title page of each mounted as if she had intended these for publica- tion. The Museum of Comparative Zoology is just over 140 years old, and the mol- lusk collection dates back to the 1830s through the collection of the Boston So- ciety of Natural History. Thus, Clench (1897-1984) and Turner (1914—2000), who were individually or collectively as- sociated with the Museum for nearly 75 years (i.e., 1926-2000), or more than half of its existence, had a remarkable insti- tutional memory of the Museum and the mollusk collection. Because their lives were so intertwined for so long in the history of the department, and because they did much joint work, it seemed ap- propriate to include both in a single study. Complete bibliographies are in- cluded for each, but there is a single list of described taxa. Over 400 were intro- duced by Clench, an occasional few by Clench and others, 68 by Clench and Turner, and only 34 by Turner alone or with authors than Clench. Turners work as a researcher slowed prior to her health problems beginning in about 1995, and she was severely disabled in the last several months of her life. She left an important manuscript on the pho- ladacean bivalve genera, Xylophaga, Xylo- redo, and Xylopholas, that she had been preparing for a number of years through Bull. Mus. Comp. Zool., 158(1): 1-46, September, 2003 I 2 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 active support by the Office of Naval Re- search (ONR). Ms. Helene Ferranti, a long-time coworker and associate of Turn- er, prepared the work for publication (Turner, 2002). The seven new species in- troduced are included here. A Projected Key to the Gastropoda of the Woods Hole, Massachusetts Area, begun by Turner sev- eral decades ago with the collaboration of several others, became so elaborate that it was never made ready for publication. An additional work alluded to by Mann (2000: 8) was also not found in a publishable state. BIOGRAPHICAL REFERENCES AND OBITUARIES ABB Onlin ele ON 3 hee @ lene lav Vallllnerna J(ames), p. 234. [In] American Malacol- ogists. Falls Church, Maryland: Ameri- can Malacologists. . 1984. A farewell to Bill Clench. The Nautilus, 98(2): 55—58. Boss, K. J. 1989. Ruth Dixon Turner. Oc- casional Papers on Mollusks, 4(August): lli—v; portrait. DOWNING, M. B. 1983. Ruth Turner— Benthic biologist. Oceanus, 26(3): 53— 56. MANN, R. 2000. In memoriam: Ruth Dix- on Turner. Journal of Shellfish Re- search, 19(1): 7-12. TURNER, R. D. 1985. William J. Clench October 24, 1897—February 22, 1984. Malacological Review, 18(December): AS — AL, . 2002. On the subfamily Xylopha- gainae (Family Pholadidae, Bivalvia, Mollusca). Bulletin of the Museum of Comparative Zoology, 157(4): 223-308; 38 pls. WEBER, Aho Ee AND Ee Vo Beer: 1991. Ruth D. Turner—investigator of deep sea benthos, pp. 285-286. [In] Marine Biology, 2nd ed. New York: Harper Collins Publishers. METHODS AND ACKNOWLEDGMENTS A few species were described from unique specimens in the possession of oth- ers. The remainder are represented in the Museum of Comparative Zoology by the holotype or paratypes, with the exception of Cepolis garciana, Cerion disforme no- dali, Emoda blanesi, Helicina holguinensis, and Jeanneretia parraiana carinata, all de- scribed by Clench and Aguayo in 1953, paratypes of which are mentioned as from the Museo Poey, Havana, Cuba. They were never received. Present efforts have failed to secure them. If the holotypes are in the Museum of Comparative Zoology, paratypes when pre- sent there or elsewhere are not always list- ed. This is especially true among the Cer- ion, where the paratype lots often consist of staggering numbers. The locality data here are as originally presented. Name changes, especially in Africa and Oceanica, have been so fre- quent that it has been deemed best to let those who might use this list make the changes as required. The following abbreviations have been used in the list of taxa. AMNH American Museum of Natural History, New York, New York ANSP Academy of Natural Sciences of Philadelphia, Pennsylvania Australian Museum Sydney, New South Wales Bishop Museum Bernice P. Bishop Museum, Honolulu, Hawaii Carnegie Museum Pittsburgh, Pennsylvania Florida Museum of Natural History Gainsville, Florida MCZ Museum of Comparative Zo- ology, Cambridge, Massachu- setts MRAC Musée Royal de PAfrique Cen- trale, Tervueren Museo Poey Havana, Cuba UMMZ Museum of Zoology, University of Michigan, Ann Arbor, Mich- igan ZMB Zoologischen Museum, Berlin MOLLUSCA OF CLENCH AND TURNER ° Johnson 3 Thanks are extended to the following for information regarding the type spec- imens of some of the species mentioned in this paper. Mr. John Slapcinsky, Flor- ida Museum of Natural History, Univer- sity of Florida, Gainesville, Florida; Dr. Daniel L. Graf, Museum of Zoology, University of Michigan, Ann Arbor, Michigan; Dr. Didier Vandenspiegal, Musée Royal de l’Afrique Centrale, Ter- vuren, Belgium; Dr. J. L. Van Goethem, Institut royal des Sciences naturelles de Belgique, Bruxelles; Dr. Bernhard Haus- dorf, Zoologisches Museum der Univer- sitaet Hamburg, Germany; Dr. Ole S. Tendal, Zoologisk Museum, Kobenhavns Universitet, Denmark; and Dr. John Stanisic, Queensland South, Brisbane, Australia. Dr. Diarmaid O’Foighil, Mu- seum of Zoology, University of Michigan, kindly supplied a paratype of a Physa de- scribed by Clench not represented in the MCZ. Ms. Mary Sears, Ms. Susan DeSanctis, and Mr. Ronnie Broadfoot of the library of the Museum of Comparative Zoology and Dr. Robert Toll, Dean of the College of Natural Sciences and Mathematics of the University of Central Arkansas, Conway, Arkansas, each helped to locate difficult references. Dr. Alan R. Kabat made numerous helpful suggestions as well as being kind enough to read the manuscript, as did Drs. Kenneth iE Boss and Ernst Mayr. Thanks are also extended to the two anonymous reviewers for carefully cor- recting the manuscript. Mr. Adam J. Bal- dinger was most helpful in many ways, including help with what is new technol- ogy to me, which is being taught to me, most sympathetically, by Ms. Elissa C. Cadillic, Special Library Assistant/Tech- nology, Allston Branch of the Boston Public Library. The cost of preparing the manuscript for publication was made possible by the R. I. Johnson Family Foundation. TAXONOMIC LIST AND PUBLICATIONS Molluscan Taxa Introduced by William J. Clench and Ruth D. Turner with Their Original References, Type Localities, and Locations of Type Specimens Abaconia (subgenus) Clench 1938, Mem. Soc. Cu- bana Hist. Nat., 12: 321 (type species, Leptinaria [Abaconia] naufraga Clench 1938, original desig- nation). [A synonym of Litiopa, teste Clench.] abbotti Clench, Cerion (Strophiops) utowana 1961, Occas. Pap. Mollusks, 2: 251, pl. 43, fig. 4 (Long Island, South Caicos Islands, Bahamas); holotype USNM 610285; paratypes MCZ 189954. abbotti Clench, Conus regius 1942, Johnsonia, 1(6): 6, pl. 4, figs. 2, 3 (Arthurstown, Cat Island, Baha- mas); holotype MCZ 145274. abbotti Clench & Turner, Opalia (Opalia) 1952, Johnsonia, 2: 348, pl. 171 (Atlantis Sta. 3374, 20°45'N, 75°19'W, Puerto Tanamo, Cuba, 300 fth.): holotype MCZ 184511. abbotti Clench & Turner, Taheitia arcasiana 1948, Occas. Pap. Mollusks, 1(13): 187, pl. 22, fig. 3, pl. 23, fig. 2, pl. 24, fig. 1 (first gully from west, north side of Somoloma Hill, 9 mi. S of Lautoka, NW Viti Levu, Fiji Islands); holotype MCZ 179578. achatinus Clench, Liguus fasciatus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 107, fig. 1 (Los Arroyos, Holguin, Oriente, Cuba); holotype MCZ 58803. acklinsensis Clench, Eutrochatella 1963, Bull. Mus. Comp. Zool., 128: 398, pl. 3, fig. 4 (inland from Cornucopia, Acklins, Island, Bahamas); holotype USNM 390633, paratype MCZ 225315. Acrostylus (subgenus) Clench 1935, Nautilus, 48: 126 (type species, Placostylus |Acrostylus| acutus Clench 1935, original designation). acutus Clench, Placostylus (Acrostylus) 1935, Nauti- lus, 48: 126, pl. 7, fig. 6 (Wanderer Bay, Guadal- canal, Solomon Islands); holotype MCZ 93953. aequatorialis Bequaert & Clench, Archachatina (Me- gachatinops) gaboonensis 1936, Rev. Zool. Bot. Afr., 29(1): 90, pl. 1, fig. 4 (Kunungu, near Bolobo, Belgian Congo); holotype MRAC 339. 640; para- type MCZ 103894. africana Clench, Rhinomelania 1929, Bull. Mus. Comp. Zool., 69: 119, text fig. 1 (Kribi, Cameroon); holotype MCZ 15794. Afroditropis (genus) Bequaert & Clench 1936, Rev. Zool. Bot. Afr., 29(1): 102 (type species, Afroditro- pis strongi Bequaert & Clench, 1936, original des- ignation). afromontana Bequaert & Clench Achatina, Rev. Zool. Bot., 24(3): 269, pl. 1, figs. 2, 12 (Western Ankole, Uganda); holotype MCZ 77208. agassizi Clench & Aguayo, Metula 1941, Mem. Soc. Cubana Hist. Nat., 15(2): 179, pl. 14, fig. 4 (Atlan- tis Sta. 3443, 23°22'N, 79°53’W, off Vagua la Grande, Cuba, 325 fth.); holotype MCZ 135248. agglutinans Bequaert & Clench, Potadoma 1941, Bull. Mus. Comp. Zool., 88(1): 10, pl. 1, figs. 1-8, 4 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 10 (estuary of the Congo River, Kala-Kala, near Matadi, Belgian Congo); holotype MCZ 112267. aguayoi Clench, Bathyaurinia 1940, Mem. Soc. Cu- bana Hist. Nat., 14(3): 241, pl. 42, fig. 4, text fig. 1 (Atlantis Sta. 3783, 30°3'N, 78°37'W; 164 mi. off St. Augustine, Florida; 425-330 fth.); holotype MCZ 111858. aguayoi Torre & Clench, Cerion 1932, Nautilus, 45: 89, pl. 6, figs. 6, 7 (road to Caletones about 6 km W Gibana, Oriente, Cuba); holotype MCZ 47902. aguayoi Clench, Cerithium 1934 [in] Johnson, Proc. Boston Soc. Nat. Hist., 40: 110 (new name for Cer- ithium varicosum Sowerby 1834, non Defrance 1817; Valenciennes 1832; Anton 1839; Eudes 1842. aguayoi Clench, Liguus fasciatus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 109, pl. 7, fig. 2 (Punta Roja, Holguin, Oriente, Cuba); holotype MCZ 76696. aguayoi Clench & Pérez Farfante, Murex (Murex) 1945, Johnsonia, 1(17): 15, pl. 8, figs. 1-3 (Atlantis Sta. 3415, 22°51'30"N, 78°55'30"W, off Punta, Ale- gre, Camagiiey, Cuba, 210 fth.); holotype MCZ 147286. alberti Clench & Aguayo, Cerion 1949, Torreia (Univ. Havana), (14): 3, pl. 1, figs. 1-6 (Punta de “El Fuerte,” Entrada de la Bahiaé de Banes, Peninsula de Ramon, Antilla, Cuba); holotype MCZ 166621, paratypes Museo Poey 4159. alberti Clench & Aguayo, Chondropoma 1948, Rev. Soc. Malacél. “Carlos de la Torre,” 6(2): 53, text fig. (0.5 mi. S of Cavo Lucrecia, Banes, Oriente, Cuba); holotype Museo Poey 12974, paratypes MCZali ai a3. albertisi Clench, Westralunio 1957, Breviora, (76): 3, fig. 2 (inland from Daru, Western Division, Papua, New Guinea); holotype MCZ 212908. Alleghenya Clench & Boss, 1967, Nautilus, 80: 101 (new name for Madalia of authors non Haldeman 1840). alleni Clench & Aguayo, Odontosagda 1932, Proc. N. Engl. Zool. Club, 13: 36 (Petionville, Haiti); holo- type MCZ 23425. almiranta Clench, Placostylus (Proaspastas) 1941, Am. Mus. Novit., (1129): 14, fig. 7 (10 mi. inland from Su’u Malaita Island, Solomon Islands, 1,500 ft.); holotype AMNH 79009; paratype MCZ 93001. altenai Turner, Xylopholas 1972, Basteria, 36(2—5): 99, figs. [2127 CerdasStay 1660s oe oun 79°59'W, about 13 mi. SE Fowey Rocks, Florida, 200 fth.); holotype MCZ 279315. amoi Cooke & Clench, Succinea 1945, Occas. Pap. B. P. Bishop Mus., 18(8): 135, fig. 3 (Papenoo Val- ley, Mt. Orofena, Tahiti, Society Islands, 4,000 ft.); holotype Bishop Museum 145634, paratype MCZ 146393. amplior Bequaert & Clench, Archachatima (Mega- chatinops) camerunensis 1936, Rev. Zool. Bot. Afr., 29(1): 89, pl. 1, fig. 12 (Libreville, French Congo); holotype MCZ 59323. andersoni Cooke & Clench, Rapanella 1943, Occas. Pap. B. P. Bishop Mus., 17(20): 254, fig. 3 (Maitua, below Mt. Mangaoa, Rapa Island, Central Pacific); holotype Bishop Museum 189005, paratypes MCZ 140596. angelae Clench & Aguayo, Liguus fasciatus 1934, Oc- cas. Pap. Boston Soc. Nat. Hist., 8: 110, pl. t fig. 3 (Punta Roja, Holguin, Oriente, Cuba); holotype MCZ 76698. aopta Clench & Archer, Helicostyla (Calocochlea) 1933, Pap. Mich. Acad. Sci. Arts Lett., 17: 542, pl. 57, fig. 3 (between Puerto Galera and San Teodoro, Mindoro, Philippines); holotype MCZ 81354. arangoi Clench & Aguayo, Jeanneretia parraiana 1951, Rev. Soc. Malacél. “Carlos de la Torre,” 7: 84, pl. 13, figs. 16, 18 (Sitio del Infierno, Vifales, Cuba); holotype Museo Poey 15079, paratypes MCZ 127393, 105218, 114958. archeri Clench, Helicostyla subcarinata 1936, Nau- tilus, 49: 140, pl. 8, fig. 5 (Tres Reyes Islands, SW coast of Marinduque, Philippine Islands); holotype MCZ 96474. archeri Clench, Liguus fasciatus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 106, pl. 7, fig. 5 (Mogote de Ramon Millo, Vinales, Pinar del Rio, Cuba); ho- lotype MCZ 80901. arenarius Clench & Pérez Farfante, Murex florifer 1945, Johnsonia, 1(17): 34, pl. 19, figs. 1-3 (Sanibel Island, Florida); holotype MCZ 149854. argo Clench & Pérez Farfante, Murex (Chicoreus) 1945, Johnsonia, 1(17): 31, pl. 17, fig. 1 (mew name for imbricatus Higgins & Marrat, 1877; non Broc- chi 1814, Risso 1826, Nardo 1847). ariomus Clench & Pérez Farfante, Murex (Pteryno- tus) 1945, Johnsonia, 1(17): 39, pl. 20, figs. 5-6 (off Hollywood, Florida, 50-60 fth.); holotype MCZ 164734. ariomus Clench, Plysa pomilia 1925, Occas. Pap. Mus. Zool., Univ. Mich., (161): 2, pl. 1, fig. 2 (Gas- tonburg, Wilcox Co., Alabama); holotype UMMZ 32231; paratypes MCZ 55658. ariomus Clench & Aguayo, Potamopyrgus coronatus 1937, Mem. Soc. Cubana Hist. Nat., 11: 68, pl. 7, fig. 10 (Lake Miragoane, Haiti, 2 mi. S of Mira- goane, Haiti); holotype MCZ 108818. armouri Clench, Ceron (Strophiops) 1933, Proc. N. Engl. Zool. Club, 13: 96, pl. 1, fig. 4 (S coast of Mariguana Island, Bahamas); holotype MCZ 101163. armouri Clench, Chondropoma beatensis 1932, Proc. N. Engl. Zool. Club, 12: 106 (Beata Island, Santo Domingo); holotype MCZ 81495. atalanta Clench, Chloritis (Sulcobasis) 1933, Nauti- lus, 47: 23, pl. 3, figs. 9, 10 (New Hanover Island, Bismark Archipelago); holotype MCZ 59844. atalanta Bequaert & Clench, Forbesopomus 1937, Proc. N. Engl. Zool. Club, 17: 54, pl. 2, figs. 1-7 (Lake Lanao, Misamis, Mindoro, Philippine Is- lands); holotype MCZ 74465. atalanta Clench, Papuina williamsi 1936, Nautilus, 50: 54 (Omarakana, North Central Kiriwina Island, Trobriand Islands); holotype MCZ 111151. athearni Clench & Turner, Goniobasis 1956, Bull. MOLLUSCA OF CLENCH AND TURNER * Johnson 5 Fla. State Mus., 1(3): 131, pl. 2, fig. 6 (Chipola River, 2.5 mi. SE of Chason, Calhoun Co., Florida); holotype MCZ 190102. atkinsi Torre & Clench, Urocoptis livida 1930, Nau- tilus, 44: 15, pl. 2, figs. 5, 6 (Vilches Potrero, Cen- tral Soledad, Cienfuegos, Cuba); holotype MCZ 59108. atlantica Clench, Casmaria 1944, Johnsonia, 1(16): 2, pl. 1, figs. 1, 2 (Puerto Sostia, Hispaniola); holotype MCZ 57284. atlanticus Macdonald & Clench, Bigelowia 1934, Oc- cas. Pap. Boston Soc. Nat. Hist., 8: 145, text figs. 1-9 (387°N, 67°12’W, North Atlantic); holotype MCZ 98971. atlanticus Clench & Aguayo, Ficus 1940, Mem. Soc. Cubana Hist. Nat., 14(1): 85, pl. 14, fig. 1 (Hassler Voyage, 11°49’'S, 37°10’W, off Sao Salvador, Bahia Prov., Brasil 1872, 45 fth.); holotype MCZ 104657. atlantis Clench & Aguayo Calliostoma (Calliostoma) 1940, Mem. Soc. Cubana Hist. Nat., 14: 81, pl. 15, fig. 4 (Atlantis Sta. 3306, 23°4'30’N: 82°37'W, off Mariel, Pinar del Rio Prov., Cuba, 330 fth.); holo- type MCZ 135164. atlantis Clench & Aguayo, Columbarium 1938 (At- lantis Sta. 2999, 23°10'N, 81°29'W, off Matanzas, Cuba, 421 fth.); holotype MCZ 135004. atlantis Clench & Pérez Farfante, Murex (Bathymu- rex) 1945, Johnsonia, 1(17): 41, pl. 21, figs. 3-5 (Atlantis Sta. 3333, 22°13’N, 81°11'W, Bahia de Conchinos, Santa Clara, Cuba, 190-200 fth.); ho- lotype MCZ 164684. atlantis Clench & Turner, Nystiella 1952, Johnsonia, 2 OAO DO San ALlantism Stay tOoO 0s 22 290Ne 81°10’W, Bahia de Cochinos, Cuba, 230—265 fth.): holotype MCZ 187988. atlantis Clench, Scaphella (Anrinia) 1946, Johnsonia, 2: 53, pl. 29, fig. 5 (Atlantis Sta. 3415, 22°51’'N, 78°55'30"W, off Punta Alegre, Camagiiez, Cuba, 210 fth.); holotype MCZ 135263. augusti Clench & Aguayo, Opisthosiphon caroli 1949, Rev. Soc. Malacél. “Carlos de la Torre,” 6: 90, pl. 2, fig. 2 (Finca Flores, cerca del Mir, Holguin, Cuba); holotype Museo Poey 12352, paratypes MCZ 174284. aurantius Clench, Liguus crenatus 1929, Nautilus, 43: 19 (Hammock #408[#5] Pincecrest Region, Central Everglades, Florida); holotype MCZ 84624. Auriniopsis (genus) Clench, 1953, Johnsonia, 2: 378 (type species, Scaphella kieneri Clench 1946, orig- inal designation). bahamensis Clench, Byssanodonta 1938, Bull. Mus. Comp. Zool., 80(14): 535, pl. 2, fig. 6 (0.5 mi. due E of Arthurs Town, Cat Island, Bahamas): holotype MCZ 107726. bahamensis Clench Ferrissia (Laevapex) 1938, Mem. Soc. Cubana Hist. Nat., 12: 318, pl. 24, fig. 23 (Eight Mile Road, Grand Bahama Island, Baha- mas); holotype MCZ 116723. bahamensis Clench & Turner Truncatella bilabiata Johnsonia, 2: 155, pl. 67, figs. 1-5 (Northwest Point, Little Inagua Island, Bahamas); holotype MCZ 158794. banesense Clench & Aguayo, Cerion 1949, Torreia (Univ. Havana), (14): 7, pl. 1, figs. 13-15 (Al Este de la Bahia de Sama, Banes, Oreinte, Cuba); ho- lotype MCZ 166623, paratypes Museo Poey 9420. Bankiinae (subfamily) Turner, 1966, A Survey and I- lustrated Catalogue of the Teredinidae, p. 78. Bankiopsis (subgenus) Clench & Turner, 1946, John- sonia, 2(19): 16 (type species, Bankia caribbea Clench & Turner 1946, original designation). barberi Clench, Physa 1925, Occas. Pap. Mus. Zool., Univ. Mich., (164): 2, pl. 1, figs. 1-3 (canal em- bankment, West Palm Beach, Florida); holotype UMMZ 32540, paratypes MZUM 32451; Clench collection 2012 [not located in MCZ]. barbouri Clench, Archegocoptis 1935, Proc. Boston Soc. Nat. Hist., 41: 5, pl. 1, figs. B—F (Tardieu Mt., La Hotte, Haiti, 3,000 ft.); holotype MCZ 108593. barbouri Clench, Brachypodella (Anguliceruix) 1935, Proc. Boston Soc. Nat. Hist., 41: 10, pl. 1, fig. K (La Visite, La Selle Range, Haiti); holotype MCZ 108587. barbouri Clench & Aguayo, Calliostoma (Astele) 1946, Rev. Soc. Malac6él. “Carlos de la Torre,” 4: 89, text fig. (Arenas de la Charrera, Havana [Santa Fé], Cuba, 3-15 fth.); holotype MCZ 178128, para- type Museo Poey 11602. barbouri Clench, Cerion (Strophiops) 1933, Proc. N. Engl. Zool. Club, 13: 95, pl. 1, fig. 5 (S coast of Mariaguana Island, Bahamas); holotype MCZ 101159. barbouri Clench, Eutrochatella calida, Proc. N. Engl. Zool. Club, 13: 82, pl. 1, fig. 13 (SW sides of Mar- iaguana Island, Bahamas); holotype MCZ 101185. barbouri Clench, Liguus crenatus 1929, Nautilus, 43: 18 (Hammock #21, Pinecrest Region, Central Ev- erglades, Florida); holotype MCZ 84527. barbouri Clench & Aguayo, Oocorys 1939, Mem. Soc. Cubana Hist. Nat., 13(3): 193, pl. 29, fig. 2 (Atlantis Sta. 2987, 23°22'N, 79°53’W, off Sagua la Grande, Cuba, 280-300 fth.); holotype MCZ 135055. barbouri Clench, Opisthosiphon bahamense 1933, Proc. N. Engl. Zool. Club, 13: 84, pl. 1, fig. 14 (back of Victoria Hill Settlement, Watling Island, Bahamas); holotype MCZ 101180. barbouri Torre & Clench, Urocoptis livida 1930, Nautilus, 44: 15, pl. 2, figs. 7-9 (La Portuguesa, Central Soledad, Cienfuegos, Cuba); holotype MCZ 59111. bardwelli Clench, Hydridella (Hydridella) 1934, J. Conch., 20(3): 89, pl. 2, figs. 1-5 (lower reaches of Glenelg River, Kimberly Division, Western Austra- lia); holotype MCZ 41999, paratypes MCZ 42000. bardwelli Clench & McLean, Macrocallista (Paradi- one) 1936, J. Conch., 20(7): 201, text fig. (Broome, Western Australia); holotype MCZ 92135. bartletti Clench & Aguayo, Columbarium 1940, Mem. Soc. Cubana Hist. Nat., 14(1): 86, pl. 14, fig. 6 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 3 (Blake Sta. 9, 18°12’N, 78°20’W, off Homer’s Cove, Westmoreland, Jamaica, 254 fth.); holotype MCZ 104729. bartschi Clench & Archer, Helicostyla (Calochlea) 1933, Pap. Mich. Acad. Sci. Arts Lett., 17: 541, pl. 57, fig. 2 (Anduyanan Paltian, Mindoro, Philip- pines); holotype MCZ 81365. Bathyaurinia (genus) Clench & Aguayo, 1940, Mem. Soc. Cubana Hist. Nat., 14(1): 92 (type species Au- rinia torrei Pilsbry 1937, original designation). Bathymurex (subgenus) Clench & Pérez Farfante, 1945, Johnsonia, 1(17): 41 (Murex [Bathymurex] atlantis Clench & Perez 1945, original designa- tion). bayeri Turner, Xylophaga 2002, Bull. Mus. Comp. Zool., 157(4): 260, pls. 30, 31. (26°4’N: 80°4’W, 3.2 mi. off Fort Lauderdale, Florida, 152.4 m); holo- type MCZ 316738, paratype MCZ 316739. beatensis Clench, Chondropoma (Chondropomium) 1932, Proc. N. Engl. Zool. Club, 12: 106 (Beata Island, Santo Domingo); holotype MCZ 81493, paratypes MCZ 81494. beattyi Clench, Brachypodella (Brachypodella) 1951, J. Conchy., 90: 275, fig. 3 (Mona Island, Puerto Rico); holotype MCZ 171015. beattyi Clench, Drymaeus elongatus 1951, J. Conchy., 90: 273, figs. 4-6 (Mona Island, Puerto Rico); ho- lotype MCZ 171032. beattyi Clench, Plagioptycha euclasta 1940, Mem. Soc. Cubana Hist. Nat., 14(3): 244, pl. 42, fig. 3 (Mt. Eagle, St. Croix, Virgin Islands); holotype MCZ 110342. bellonensis Clench, Eustomopsis 1958, Natural His- tory of Rennell Island, British Solomon Islands, Copenhagen, Denmark, 2: 196, pl. 19, figs. 1-3, text fig. 2 (Bellona Island, Solomon Islands); holo- type AMNH 66430, paratype MCZ 92958. bencomoi Clench, Urocoptis (Autocoptis) 1935, Proc. Boston Soc. Nat. Hist., 41: 3, pl. 2, fig. B (vicinity of Port au Prince, Haiti); holotype MCZ 108508. bequaerti Clench & Miller Ashmunella 1966, Bre- viora, (244): 1, pl. 1, text figs. 1, 2 (Goat Cave Can- yon, near Buffalo Trail Boy Scout Camp, NE slope of Black Mt., Davis Mts., Jeff Davis Co., Texas); holotype MCZ 260274. bequaerti Torre & Clench, Cerion aguayoi 1932, Nautilus, 45: 91, pl. 6, fig. 8 (dunes at Lucrecia lighthouse, near Banes, Oriente, Cuba); holotype MCZ 86177, paratypes MCZ 86176. bequaerti Clench & Turner, Megalacron 1964, J. Ma- lacol. Soc. Aust., (8): 48, pl. 9, fig. 10 (new name for Helix trochus Quoy & Gaimard 1832, non Miill- er 1821 and Helix trochoides Deshayes 1838, non Poiret 1789, Gmelin 1790). bequaerti Clench & Pérez Farfante, Murex (Ptero- purpura) 1945, Johnsonia, (17): 40, pl. 21, figs. 1, 2 (off Delray, Florida, 80 fth.); holotype Florida Museum of Natural History (UF) 11986. berevoensis Clench & Archer, Achatuia panthera 1930, Nautilus, 43(3): 85 (forest between Mahabo and Berevo, cotype [lectotype] 31078; Berevo, co- type [paralectotype] MCZ 31077; 10 mi. S of Ber- evo, elevation 850-900 ft., cotype [paralectotype] MCZ 31079, all Madagascar); lectotype MCZ 31078 selected by J. C. Bequaert, 1950, Bull. Mus. Comp. Zool., 105(1): 107, pl. 5, fig. 3. bermudensis Clench, Conus 1942, Johnsonia, 1(6): 34, pl. 13, fig. 4 (Dyer Island, Bermuda); holotype MCZ 141965. bermudezi Clench & Aguayo, Aurinia 1940, Mem. Soc. Cubana Hist. Nat., 14(1): 89, pl. 16, fig. 3 (Atlantis Sta. 2962, 22°7'N, 81°8’W, Bolira Cochi- nos, Santa Clara |Prov.], Cuba, 180—190 fth.); ho- lotype MCZ 135245. bermudezxi Clench & Aguayo, Columbarium 1938, Mem. Soc. Cubana Hist. Nat., 12: 383, pl. 28, fig. 7 (Atlantis Sta. 2989, 23°10'N, 80°4'W, off Sagua la Grande, Cuba, 360 fth.); holotype MCZ 135007. bermudezi Clench, Liguus flammellus 1934, Occas. Pap: Boston) Soe) Nats Hists S: 22) spl: (Gs near (Mogote la Gueca Vinales, Pinar del Rio, Cuba); holotype MCZ 80945. bermudezi Clench & Aguayo, Mecoliotia 1936, Nau- tilus, 49: 92, pl. 5, fig. 3 (0.5 km S of the mouth of Rio Canimar, Maténzas, Cuba); holotype MCZ 110615. bermudezi Clench & Aguayo, Orthaulax 1939, Mem. Soc. Cubana Hist. Nat., 13(5): 357, pls. 47, 48 (Paso Real de San Diego, Pinar del Rio Prov., Cuba, in a well at a depth of 10 ft.); holotype MCZ 27936. bigelowi Clench & Aguayo, Calliostoma (Astele) 1938, Mem. Soc. Cubana Hist. Nat., 12: 378, pl. 28, figs. 4, 5 (Atlantis Sta. 2963-c, Bahia Cochinos, Cuba, 205 fth.); holotype MCZ 135003. Bigelowia (genus) Macdonald & Clench, 1934, Oc- cas. Pap. Boston Soc. Nat. Hist., 8: 145 (type spe- cies, Bigelowia atlanticus, Macdonald and Clench 1934, original designation). blainei Clench & Turner, Epitonium (Boreoscala) 1953, Johnsonia, 2: 361, pl. 180 (about 45 mi. SW of the light house, Boca Grande, Florida); holotype MCZ 189246. blakei Clench & Aguayo, Calliostoma (Calliostoma) 1938, Mem. Soc. Cubana Hist. Nat., 12: 376, pl. 28, fig. 6 (Hassler exped., 41°17'S, off Cape Ber- meja, Argentina, 17 fth.); holotype MCZ 89661. blanesi Clench & Aguayo, Caracolus sagemon 1951, Mem. Soc. Cubana Hist. Nat., 20: 68, pl. 42, figs. 7, 8 (Silla de Gibara, Oriente, Cuba); holotype Mu- seo Poey 17302, paratypes MCZ 76974. blanesi Clench & Agauyo, Cerion 1951, Rev. Soe. Malacél. “Carlos de la Torre,” 8: 70, pl. 11, fig. 1 (Los Cocos, E side of Bahia de Gibara, Cuba); ho- lotype Museo Poey 17266, paratype MCZ 52784. blanesi Clench & Aguayo, Emoda 1953 [in] Aguayo, Mem. Soc. Cubana Hist. Nat., 21: 301, pl. 35, figs. 1, 2 (“Embarradas,” al Norte de Banes, Oriente, Cuba); holotype Museo Poey 17324, paratype MCZ [not received]. bollingi Clench, Oliva reticularis 1934, Nautilus, 47: MOLLUSCA OF CLENCH AND TURNER * Johnson i 142, pl. 7, figs. 3, 4 (off Miami, Florida, crab traps at 200 ft.); holotype MCZ 76656. bottimeri Clench, Physa 1924, Nautilus, 38: 12, text fig. 4 (Comanche Spring, Fort Stockton, Pecos Co., Texas); holotype UMMZ 31617, paratypes MCZ 53866, 86375. boultoni ‘Bequaert and Clench’ Parodiz & Tripp, Vi- viparus (Bellamya): 1988, Ann. Carnegie Mus., 57(5): 130 [nomen nudum].? branchi Clench, Murex (Murex) 1953, Johnsonia, 2: 360, pl. 179 (Gulf of Campeche, Yucatan, Mexico); holotype MCZ 201474. branhamae Clench, Conus jaspideus 1953, Johnsonia, 2: 364, pl. 181, fig. 2 (Green Turtle Cay, Great Abaco, Bahama Islands); holotype AMNH 166926. brasiliensis Clench, Conus 1942, Johnsonia, 1(6): 24, pl. 12, fig. 2 (Thayer exped., Victoria, Brazil); ho- lotype MCZ 146894. brayi Clench, Columbarium 1959, Johnsonia, 3: 330, pl. 173 (Atlantis cruise 240, 11°N, 66°1'W, off Cabo Codera, Venezuela, 150 fth.); holotype MCZ 221601. brazieri Clench, Dublonia 1949, Bull. B. P. Bishop Mus., (196): 8, fig. 2 (Mt. Tolowan, Dublon Island, Truk Islands, Caroline Islands); holotype Bishop Museum 153355, paratypes Bishop Museum 153720, MCZ 140532 from Neborus, Muen Island, Truk Islands. brooksi Bequaert & Clench, Achatina (Pintoa) 1934, RevarZoolesBotawauns 243) sample slp ehigs Sarl (Clauca, Angola); holotype Carnegie Mus., 62.25670 (only specimen). brooksi Gustafson, Turner, Lutz & Vrijenhoek, Bath- ymodiolus 1998, Malacologia, 40(1—2): 78, figs. 11— 15 (Alvin Dive 2211, 26°21.3'N, 94°29.7'W, in the western Gulf of Mexico at a hydrocarbon seep in Alaminos Canyon, 2,222 m); holotype ANSP A1884/18847, paratypes MCZ 316973-6. bullisi Clench & Turner Calliostoma (Elmerlinia) 1960, Johnsonia, 4: 36, pl. 5, fig. 1; pl. 9, fig. 2: pl. 23 (Oregon Sta. 2049, 4°2'N, 50°33’W, about 65 mi. ESE of Cabo Orange, Arnapa, Brasil, 38 fth.); ho- lotype USNM 612702. burryae Clench, Conus floridanus 1942, Johnsonia, 1(6): 29, pl. 14, figs. 3, 4 (off Lower Matecumbe Key, Florida); holotype MCZ 145296. burryi Clench & Pérez Farfante, Murex (Poirieria) 1945, Johnsonia, 1(17): 47, pl. 24, figs. 1-3 (off Fort Walton, Florida, 13-19 fth.); holotype MCZ 164567. burryi Clench & Turner Opalia (Dentiscala) 1950, Johnsonia, 2: 235, pl. 102, figs. 1-3 (4.5 mi. off Carysfort Light, Key Largo, Florida, 92-100 fth.); holotype MCZ 187107. APARODIZ WE SAND |= [io URIPRY 1988) ay pesion Mollusca in the collection of the Carnegie Museum of Natural History Part I. Bivalvia and Gastropoda (Prosobranchia and Opisthobranchia) Annals of the Carnegie Museum, 57(5): 111-154. butleri Clench, Scaphella junonia 1953, Johnsonia, 2: 377, pl. 186, fig. 3 (Bay of Campeche, Yucatan, Mexico); holotype MCZ 193591. cadenasi Clench & Aguayo, Tylotia 1939, Mem. Soc. Cubana Hist. Nat., 13: 195, pl. 29, fig. 5 (Atlantis Sta. 2963, 22°7'N, 81°8’W, Bahia de Cochinos, Cuba, 180 fth.); holotype MCZ 135056. cahabensis Clench, Clappia 1965, Nautilus, 79: 33, text fig. 2 (Cahaba River, 1 mi. N of Centreville, Bibb Co., Alabama); holotype MCZ 251167. caicosense Clench, Cerion (Strophiops) 1937, Proc. N. Engl. Zool. Club, 16: 23, pl. 1, fig. 4 (Cockburn Town, South Caicos Island, Turks Islands); holo- type MCZ 116021. calciphila Cooke & Clench, Fijianella 1943, Occas. Pap. B. P. Bishop Mus., 17(20): 258, fig. 7 (Yaugasa Levu, Lau Islands, Fiji Islands); holotype Bishop Museum 10107, paratypes MCZ 137107. caledoniensis Clench & Jacobson, Emoda 1971, Bull. Mus. Comp. Zool., 141: 112, pl. 2, figs. 11, 12, pl. 5 (Mina Caledonia, Mayari, Oriente, Cuba); holo- type MCZ 261352. candefacta Bequaert & Clench, Archachatina (Me- gachatinops) adelinae 1936, Rev. Zool. Bot. Afr., 29(1): 87 (Johann Albrechtshéhe, Cameroon); ho- lotype MCZ 79972. canetae Clench & Aguayo, Buccinum 1944, Rev. Soc. Malacél. “Carlos de la Torre,” 2(1): 67-68, text fig. (Atlantis Sta. 3008, 23°12'30"N, 82°12’W, off Ha- vana, Cuba, 240-300 fth.); holotype MCZ 145740. carbonarius Clench, Liguus flammellus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 121, pl. 6, fig. 3 (Mogote de Pita, Viriales, Pinar del Rio, Cuba); ho- lotype MCZ 80933. carcellesi Clench & Aguayo, Calliostoma (Calliosto- ma) 1940, Mem. Soc. Cubana Hist. Nat., 14(1): 80, pl. 14, fig. 4 (Hassler voyage, 40°22'S, 60°35’'W, off Punta Rubio, Argentina, 30 fth.); holotype MCZ 104719. caribaea Clench & Aguayo, Poteria 1935, Nautilus, 49: 51, pl. 3, figs. 5, 6 (Spring Mount, St. James, Jamaica); holotype MCZ 109264. caribaeus Clench, Liguus fasciatus 1935, Nautilus, 49: 68, pl. 3, fig. 3 (Moute Dios, Santa Lucia, N coast of Pinar del Rio, Cuba); holotype MCZ 109050. caribbaea Clench & Aguayo, Oocarys sulcata 1939, Sone Soc: Cubana Hist. Nat., 13: 192; pl. 29, fig. 3 (Atlantis Sta. 2953, 21°47'30’N, 84°30'30’W, Ba- hia de Corrientes, Pidelkio, Cuba, 615 fth.); holo- type MCZ 135072. caribbaea Clench, Urocoptis (Urocoptola) 1966, Bre- viora, (245): 9, pl. 1, fig. 3 (1 mi. E of Pointe a Raquette, South Central Gonave Island, Haiti); ho- lotype MCZ 260873. caribbaeum Clench & Turner, Cymatium (Ranularia) 1957, Johnsonia, 3: 204, pl. 111, figs. 3, 4, pl. 113, fig. 3, p. 117, figs. 1, 2 (new name for Triton cy- nocephalum ‘Lam. Kiener 1842, non T. cynoce- phalum Lamarck 1816). 8 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 caribbea Clench & Turner, Bankia (Bankopsis) 1946, Johnsonia, 2(19): 16, pl. 10, figs. 1-4 (Fort Pickens, Pensacola, Florida); holotype MCZ 121065. carinata Clench & Aguayo, Jeanneretia parraina 1951, Rev. Soe. Malacél. “Carlos de la Torre,” 7(3): 86, pl. 14, figs. 8, 9 (Mogote de la Resbalosa, La- guna de Piedras, Viiiales, Cuba); holotype Museo Poey 16001, paratypes MCZ [not received]. carnicolor Clench & Pérez Farfante, Murex (Murex- sul) 1945, Johnsonia, 1(17): 48, pl. 25, figs. 1-4 (Blake Sta. 273, off Barbados, 103 fth.); holotype MCZ 7305. carolae Clench, Ficus 1945, Johnsonia, 18: 3, pl. 2 (5.5 mi. SE of The Elbow, Key Largo, Florida, 92— 100 fth.); holotype MCZ 157501. carribaeus Clench, Conus 1942, Johnsonia, 6: 23, pl. 11, figs. 4, 5 (off Palm Beach, Florida); holotype MCZ 138333. castilloi Clench & Jaume, Helicina (Helicina) 1946, Rev. Soc. Malacol. “Carlos de la Torre,” 4: 7—8, text figs. 1-3 (Rancho Arriba, San José de Ocoa, Prov. de Azua, Républica Dominicana); holotype MCZ 155149. cayemitensis Clench, Urocoptis (Urocoptola) 1966, Breviora, (245): 7, pl. 1, fig. 4 (NE Grande Cay- emite, 6.5 mi. NE of Corail, Dépt. du Sud, Haiti); holotype MCZ 254666. cecilae Wetherbee & Clench, Macroceramus 1984, Caribb. J. Sci., 20(1—2): 11, fig. 1, pl. 1, pt. B, cen- ter (top of Mt. Isabel de Torres, Puerto Plata Prov., Republica Dominicana, 2,500 ft.); holotype MCZ 75450. ceiba Clench, Cerion 1948, Rev. Soc. Malacél. “Car- los de la Torre,” 6(2): 49, figs. 1-3 (N side of Boca del Rio Jibacoa, Santa Cruz del Norte, Havana Prov., Cuba); holotype MCZ 157307. Celetaia (genus) Clench 1966, Nautilus, 79: 137 (type species, Vivipara persculpta P. & F. Sarasin 1898, original designation). cerosa Clench & Aguayo, Hemitrochus 1937, Mem. Soc. Cubana Hist. Nat., 11(2): 76, pl. 7, fig. 5 (La Visite, La Selle Range, Haiti); holotype MCZ 110620. cervus Clench, Liguus flammellus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 120, pl. 6, fig. 4 (Mogote de Vigil, Vifiales, Pinar del Rio, Cuba); holotype MCZ 80935. Chaceia (genus) Turner, 1955, Johnsonia, 3(34): 66 (type species, Pholas ovoidea Gould, 1851, original designation). championi Clench & Turner, Epitonium (Asperiscala) 1952, Johnsonia, 2: 318, pl. 153 (Lewis Bay, Hyan- nis, Cape Cod, Massachusetts); holotype MCZ 182900. chapini Bequaert & Clench, Achatina rugosa 1934, Am. Mus. Novit., (705): 4, text fig. 2 (Lukolela, Bel- gian Congo); holotype AMNHA 6438, paratype MCZ 93211. chapini Bequaert & Clench, Maizania (Maizaniella) lukolelensis 1936, Rev. Zool. Bot. Afr., 29(1): 101 (New Beni [Bungula], Belgian Congo); holotype MCZ 102375. chengi Turner & Santhakumaran, Lignopholas 1989, Ophelia, 30(3): 180, figs. 17-19 (Semariang, Sara- wak about 1.6 km up Semariang River, a tributary of the Santubong River about 8.7 km from the mouth); holotype MCZ 207208, paratypes MCZ 297209. childressi Gustafson, Turner, Lutz & Vrijenhoek, “Bathymodiolus” 1998, Malacologia, 40(1—2): 84, figs. 11-13, 16-20 (Johnson Sea-Link-1, 27°46.9'N, 91°30.4'W, about 210 km SSW of Grand Isle, Lou- isiana, 546 m); holotype ANSP A18848, paratype MCZ 316978. chondrocycloides Bequaert & Clench, Maizania (Maizaniella) 1936, Rev. Zool. Bot. Afr., 29(1): 101 (village of Malela, 5°40’S, 23°45'E, Belgian Congo); holotype MCZ 109548. christophei Clench, Cerion 1937, Proc. N. Engl. Zool. Club, 16: 24, pl. 1, fig. 2 (northeast Point [Chris- tophies Palace], Great Inagua Island, Bahamas); holotype MCZ 116006. ciboney Clench & Pérez Farfante, Murex (Murex) 1945, Johnsonia, 1(17): 20, pl. 10, figs. 1-3 (Atlantis Sta. 3482, 23°9'N, 81°27'30”"W off Matanzas, Cuba, 190 fth.); holotype MCZ 147315. cieba Clench & Turner, Bankia (Plumielella) 1946, Johnsonia, 2(19): 25, pl. 16, figs. 1-4 (Balboa, Ca- nal Zone, Panama); holotype MCZ 168097. cisnerosi Clench & Aguayo, Cerion 1951, Rev. Soc. Malacol. “Carlos de la Torre,” 8: 72, pl. 11, figs. 3, 4 (Playa Morrillo, 11 km W of Bahia Honda, Pinar del Rio, Cuba); holotype Museo Poey 12667, para- types MCZ 187317. clappii Turner, Lignopholas 1955, Johnsonia, 3(34): 99, pl. 60 (Bluefields, Nicaragua); holotype MCZ 200046. clementis Clench & Aguayo, Emoda 1950, Rev. Soc. Malacol. “Carlos de la Torre,” 7: 62, pl. 12, figs. 4, 5 (Cayo del Rey, Mayari, Oriente, Cuba); holotype Museo Poey 12923, paratypes MCZ 185799. clenchi Turner, Hemitrochus 1958, Occas. Pap. Mol- lusks, 2(22): 161-164, pl. 23, figs. 3, 4; pl. 24, figs. 7, 8; pl. 25; pl. 30, fig. 2 (Maricao Forest, Puerto Rico); holotype MCZ 216146. clenchi Turner & Culliney, Xylophaga 1971, Ameri- can Malacological Union Annual Report for 1970, p. 66 (24°54'N, 77°49'W), Tongue of the Ocean, about 4 mi. off NE tip of Andros Island, Bahamas); Turner, 2002, Bull. Mus. Comp. Zool., 157(4): 239 pls. 5-8; holotype MCZ 316743, paratypes MCZ 316744, 316745. cliffordi Clench, Cerion (Strophiops) martensi 1933, Proc. N. Engl. Zool. Club, 13: 91,"pl. 1, fig. 110 (Landrail Pt., Crooked Island, Bahamas); holotype MCZ 101153. coeruleus Clench & Archer, Amphidromus 1932, Oc- cas. Pap. Boston Soc. Nat. Hist., 8: 41, pl. 4, figs. E, F (0.5 mi. above Fort at Long Loba, Tinja River, Sarawak, Borneo); holotype MCZ 44991, paratype MCZ 247891. MOLLUSCA OF CLENCH AND TURNER *¢ Johnson 9 congolensis Bequaert & Clench, Septariellina 1936, Mem. Mus. R. Hist. Nat. Belg., 2nd ser., (3): 163, fig. | (Ango-Ango, 4 km S of Matadi, Congo River, Belgian Congo); holotype MCZ 59807. cookei Clench, Lyropupa 1952, B. P. Bishop Muse- um, Special Publication 42, p. 32 (new name for Lyropupa anceyana Cooke & Pilsbry 1920, non Lyropupa aceyana Pilsbry & Cooke 1920). cookei Clench, Ostodes 1949, Bull. B. P. Bishop Mu- seum 196, p. 10, fig. 3a (Tiavi, Upolu Island, Sa- moan Islands); holotype Bishop Museum 9711, paratype MCZ 140504. cookei Clench, Placostylus (Eumeocostylus) oleryi 1941, Am. Mus. Novit., (1129): 16 (Waiai, San Christoval Island, Solomon Islands); holotype MCZ 65996. cookei Cl & Turner, Taheitia arcasiana 1948, Occas. Pap. Mollusks, 1(13): 186, pl. 23, fig. 3 (Vuni Vatu, central Viti Levu, Fiji Islands); holotype MCZ 181049. cookei Clench, Thaanumella (Jokajia) 1946, Occas. Pap. B. P. Bishop Mus., 18(13): 204, text fig. 4 (base of Peipalop Peak, Ponape, Caroline Islands); holotype Bishop Museum 189004, paratypes MCZ 140590. cookei Clench, Tylotoechus 1949, Bull. B. P. Bishop Mus., 196: 44, fig. 25 (Auki, Malaita Island, Solo- mon Islands); holotype MCZ 141013, paratypes Bishop Museum 188849. cornucopia Cooke & Clench, Fijianella 1943, Occas. Pap. B. P. Bishop Mus., 17(20): 260, fig. 86 (Na- vutu Iloma, Lau Islands, Fiji Islands); holotype Bishop Museum 10109, paratypes MCZ 137108. coultasi Clench, Dendrotrochus 1957, Am. Mus. Nov- it., (1863): 4, fig. 6 (Masahet Island, Lihir Group, Bismarck Archipelago); holotype AMNH 73487, paratypes MCZ 188142. coultasi Clench Pseudocyclotus 1957, Am. Mus. Novy- it., (1863): 1, figs. 1, 2 (Tavi Village, Manus Island, Admiralty Islands); holotype AMNH 73485, para- types MCZ 188141. coultasi Clench, Quirosella 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2: 175, pl. 18, figs. 1, 2, text fig. 2 (Rennell Island, Solomon Islands); holotype AMNH 59011, paratypes MCZ 93018. coutini Clench & Aguayo Caracolus sagemon 1951, Mem. Soc. Cubana Hist. Nat., 20: 66, pl. 42, figs. 1-4 (Nibujén, 4 km NW of Baracoa, Oriente, Cuba); holotype Museo Poey 17300, paratypes MCZ 181892. cramptoni Clench, Partula 1941, Am. Mus. Novit., (1129): 20, fig. 13 (Rennell Island, Solomon Is- lands); holotype AMNH 79014, paratypes MCZ 93201, 100381. Cristovala (subgenus) Clench, 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2: 188 (type species, Crystallop- sis tricolor (Pfeiffer, 1849), original designation). crystallina Clench, Crystallopsis (Crystallopsis) 1958, Natural History of Rennell Island, British Solomon Islands, Copenhagen, Denmark, 2: 182, pl. 16, fig. 5 (Ulawa Island, Solomon Islands); holotype AMNH 66484, paratypes 66441, paratypes MCZ 92945. cuba Clench, Scaphella (Aurinia) 1946, Johnsonia, 2(22): 58, pl. 31, fig. 2 (Atlantis Sta. 2988, 23°15'N, 79°57'W, off Sagua la Grande, Las Villas, Cuba, 380 fth.); holotype MCZ 135229. cubana Clench & Aguayo, Cochlespira (Ancistrosy- rinx) radiata 1940, Mem. Soc. Cubana Hist. Nat., 14(1): 94, pl. 15, fig. 1 (Atlantis Sta. 3332, 22°9'30"N, 81°11’W, Bahia Cochinos, Santa Clara Prov., Cuba, 175-225 fth.); holotype MCZ 135194. cubana Clench & Aguayo, Gaza superba 1940, Mem. Soc. Cubana Hist. Nat., 14(1): 81, pl. 15, fig. 3 (Atlantis Sta. 3448, 23°21'N, 79°56'W, off Sagua la Grande, Santa Clara Prov., Cuba, 380 fth.); holo- type MCZ 135151. cubanum Clench & Aguayo, Calliostoma (Calliosto- ma) 1940, Mem. Soc. Cubana Hist. Nat., 14: 78, pl. 16, fig. 4 (Atlantis Sta. 3474, 23°18'N, 80°46'W. off Cardenas, Matangas Prov., Cuba, 490 fth.); ho- lotype MCZ 135163. cubensis Clench, Liguus flammellus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 123, pl. 6, fig. 1 (Mogote la Gueca, Vifiales, Pinar del Rio, Cuba); holotype MCZ 80951. Culmenella (subgenus) Clench, 1927, Nautilus, 40: 121, fig. 1 (type species, Bulinus [Culmenella] hir- asei Clench 1927, monotypy). Cymatriton (subgenus) Clench & Turner, 1957, John- sonia, 3: 210 (type species [Cymatium|] nicobari- cum Réding 1789, original designation). darlingtoni Clench, Archegocrptis 1935, Proc. Boston Soc. Nat. Hist., 41(1): 6, pl. 1, figs. C, G (Tardieu, Mt. La Hotte, Haiti, 3,000 ft.); holotype MCZ 108592. darlingtoni Clench, Brachypodella (Brachypodella) 1935, Proc. Boston Soc. Nat. Hist., 41: 10, pl. 2, fig. H (Miragoane, Haiti); holotype MCZ 108588. darlingtoni Clench, Hemitrochus 1964, Rev. Mus. Ar- gent. Cienc. Nat. “Bernardino Rivadavia,” 8: 223, pl. 1, fig. 4 (Loma Pelona, near Loma Rucilla [Pico Trujillo] Prov. de Santiago, Républica Dominica, 9,000—10,000 ft.): holotype MCZ 187934. darlingtoni Bequaert & Clench, Mcleania 1939, Mem. Soc. Cubana Hist. Nat., 13: 283, pl. 36, figs. 4—6 (Maricao Forest, western Puerto Rico, 2,500 ft.); holotype MCZ 90736. darlingtoni Clench & Aguayo, Parachondria (Para- chondria) 1937, Mem. Soc. Cubana Hist. Nat., 11(2): 66, pl. 7, fig. 4 (Poste Terre Rouge, Haiti, 2,000 ft.); holotype MCZ 108576. darlingtoni Clench & Archer, Thersites 1938, J. Conch., 21: 20, pl. 1, fig. 2 (MacPherson Range, Queensland Nat. Park, 60 mi. S of Brisbane, Queensland); holotype MCZ 99054, paratype Queensland Museum, South Brisbane, Queensland (UMMO) 1766. deckerti Clench, Liguus fasciatus 1935, Nautilus, 48: 10 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 122, pl. 7, fig. 4 (Hammock 55, E end of Long Pine Key, South Central Everglades, Florida); holotype MCZ 81549. degeneri Clench, Cerion 1948, Rev. Soc. Malacél. “Carlos de la Torre,” 6(2): 50, figs. 4-6 (Fleeming Point, New Providence Island, Bahamas); holotype MCZ 175065. degneri Bequaert & Clench, ee (Magacha- tina) 1936, Rev. Zool. Bot. Afr., 29(1): 78, pl. 1, fig. 1 (Duma, Ubangi Dist., Belgian Congo); holotype Zoological Museum, Hamburg (ZMH) 2893 (only specimen). degneri Bequaert & Clench, Ptychotrema Sneha ma) 1936, Nautilus, 49(3): 96, pl. 6, figs. 3, 4 (Af- rica); holotype MCZ 10939. delli Clench, Palaina (Palaina) 1965, Breviora, (224): 6, pl. 2, fig. 5 (W side of the Lunga River, 0.5 mi. above the Seventh Day Adventist Mission Station on Guadalcanal Island, Solomon Islands); holotype MCZ 258017, paratypes MCZ 258018. demesana Clench & Archer, Helicostyla cincinnifor- mis 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 336, pl. 17, fig. 5 (Lubang Island, Philippines); ho- lotype MCZ 79171, paratypes MCZ 47216. depalmai Turner, Xylophaga 2002, Bull. Mus. Comp. Zool lan (4) 3253: pls. 23, 24 (26°4’N, 80°4’W, about 3.2 mi. off Fort Lauderdale, Florida, 152.4 m); holotype MCZ 316735. depressa Clench & Aguayo Jeanneretia parraiana 1951, Rev. Soc. Malacél. “Carlos de la Torre,” 7: 85, pl. 13, figs. 10-12 (Hoyo del Guama, Vifales, Pina del Rio, Cuba); holotype Museo Poey 17207, paratypes 181710, 181714. destructa Clench & Turner, Bankia (Neobankia) 1946, Johnsonia, 2(19): 20, pl. 13, figs. 1-4 (La Cie- ba, Honduras); holotype MCZ 123303. dewittei Bequaert & Clench, Achatura (Pintoa) 1934, Rev. Zool. Bot. Afr., 24: 270, pl. 1, fig. 428 (Nyon- ga, region of Lahe Upemba, Katanga Dist., Belgian Congo); holotype MRAC 7592, paratypes MRAC 7593-7604, 7605-7625, MCZ 81986. dewulfi Bequaert & Clench, Pila 1933, Rev. Zool. Bot. Afr., 23: 71, pl. 5 (Inkisi River near Kisantu, Belgian Congo); holotype MCZ 92101. diagonalis Clench, Urocoptis (Gongylostoma) 1966, Breviora, (245): 6, pl. 2, fig. 4 (Mina Carlota, Sier- vade San Juan, 8 mi. S of Cumanayagua, Las Villas, Cuba); holotype MCZ 59292. dickinsoni Clench & Turner, Goniobasis 1956, Bull. Fla. State Mus., 1(3): 141, pl. 2, fig. 10 (Holmes Creek, 1 mi. W of Graceville, Blonday: ); holotype MCZ 191771. disforme Clench & Aguayo, Cerion 1946, Rev. Soc. Malacol. “Carlos de la Torre,” 4: 85, text figs. 1-6 (Cercanias de Punta Manolito, Peninsula del Ra- mon, Antilla, Oriente, Cuba); holotype MCZ 166621. dolphin Clench & Cooke, Succinea wallisi 1945, Oc- cas. Pap. B. P. Bishop Museum, 18(8): 134, fig. 2 (Tautira Valley, Taiarupu Peninsula, Tahiti, Society Islands); holotype Bishop Museum, 9715, paratype MCZ 146392. dominicana Clench, Coryda 1964, Rev. Mus. Argent. Cienc. Nat. “Bernardino Rivadavia,” 8: 224, pl. 1, fig. 2 (Yuma, Prov. de El. Seibo, Reptiblica Domi- nicana); holotype MCZ 188239. domincanum Clench & Aguayo, Cerion marielinum 1951, Rev. Soc. Malacoél. “Carlos de la Torre,” 8: 69, pl. 10, figs. 1-3 (E side of Boca del Rio Domini- ca about 10 mi. W of Mariel, Cuba); holotype MCZ 181904, paratype Museo Poey 12670. Dublonia eae Cl., 1949, Bull. B. P. Bishop Mus., (196): 7 (type species, Dublonia brazieri Clench. 1949, peel designation). Echininus Clench & Abbott, 1942, Johnsonia, 1(4): 3 (new name for Nina Gray 1850, non Horsfield 1829). edentula Clench, Coryda 1964, Rev. Mus. Argent. Cienc. Nat. “Bernardino Rivadavia,” 8: 226, pl. 2, fig. 3 (Bahia de San Lorenzo, Bahia de Samana, Proy. de Samana, Républica Dominicana); holotype MCZ 188243. egregiella Bequaert & Clench, Archachatina margin- ata (Megachatina) 1936, Rev. Zool. Bot. Afr., 29 83, pl. 1, fig. 3 (Kiribi, Cameroon); holotype MCZ ola: ekmani Clench, Urocoptis (Urocoptola) 1966, Bre- viora, (245): 8, pl. 1, fig. 5 (La Source, NW Gonave Island, Haiti); holotype MCZ 260871. elegans Clench & Aguayo, Physa (Haitia) 1932, Proc. N. Engl. Zool. Club, 13: 37 (Lake Miragoane, 2 mi. SE of Miragoane, Haiti); holotype MCZ 83736, paratypes MCZ 837382. Elmerlinia (subgenus) Clench & Turner, 1960, John- sonia, 4: 29 (type species, Callistoma jujubinum Gmelin 1791, original designation). eolis Clench & Turner, Opalia (Nodiscala) 1950, Johnsonia, 2: 241, pl. 105, figs. 1-3, pl. 107, fig. 4 (off Looe Key, Lower Florida Keys, 70-90 fth.); holotype MCZ 187110. eriksoni Clench, Cerion (Maynardia) 1959, Bull. Mus. Comp. Zool., 121(2): 48, pl. 1, fig. 1 (South- west Point, Little Inagua, Bahama Islands); holo- type MCZ 189121. escambia Clench & Turner, Fusconaia 1956, Bull. Florida State Museum, 1(3): 152, pl. 7, figs. 3, 4 (Escambia River, 3 mi. SE of Century, Escambia Co., Florida); holotype MCZ 191470. Euaethiops genus Clench & Archer, 1930, Occas. Pap. Boston Soc. Nat. Hist., 5: 295 (type species Euaethiops loveridgei Clench & Archer 1930, orig- inal designation). euglypta Clench & Aguayo, Alcadia 1950, Rev. Soc. Malacél. “Carlos de la Torre,” 7: 64, pl. 12, figs. 6— 8 (Cerro Cariblanco, Holguin, Oriente, Cuba); ho- lotype Museo Poey 12911, paratypes MCZ 185797. evelynae Clench & Aguayo, Terebra 1939, Mem. Soc. Cubana Hist. Nat., 13: 196, pl. 29, fig. 1 (Atlantis Sta. 2981, 22°48’N, 78°48’W, off northern Santa MOLLUSCA OF CLENCH AND TURNER * Johnson ih Clara (Prova) Cubay 225) tth.)) holotype MECZ LS aOTie evelynae Clench, Urocoptis (Autocoptis) 1935, Proc. Boston Soc. Nat. Hist., 41: 4, pl. 1, figs. D-H (north foothills of Mt. La Hotte, Haiti, 3,000 ft.); holotype MCZ 108597. exhuberans Clench & Aguayo, Caracolus sagemon 1951, Mem. Soc. Cubana Hist. Nat., 20: 67, pl. 41, figs. 7, 8 (Cercenias de Baracoa, Oriente, Cuba): holotype Museo Poey 17301, paratypes MCZ 87543 labeled, Boca de Jauco, Gan Tierra, Baracoa, Oriente, Cuba). exquisita Clench, Urocoptis (Gongylostoma) 1966, Breviora, (245): 5, pl. 2, fig. 2 (0.5 mi. E of Gua- bairo, Soledad, Cienfuegos, Cuba); holotype MCZ 09286. eyerdami Clench & Banks, Anguispira kochi 1939, Mem. Soc. Cubana Hist. Nat., 13: 285, pl. 36, fig. 3 (46°30'N, 120°38'W, slope near mountaintop, Horse Haven Hills, Yakima Indian Reservation, Washington. [This locality is] 75 mi. from Satus Creek and 10 mi. S of Alfalfa, Washington); holo- type MCZ 100491. eyerdami Clench & Aguayo, Helisoma 1932, Proc. N. Engl. Zool. Club, 13: 38 (Lake Miragoane, 2 mi. SE of Miragoane, Haiti); holotype MCZ 83887, paratypes MCZ 83883. eyerdami Clench, Nesopoma 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2: 169, pl. 17, fig. 4 (Suu, Ma- laita, Solomon Islands); holotype AMNH 73885, paratypes MCZ 188086. fairchildi Clench, Asolene (Surinamia) 1933, Nauti- lus, 47: 71, pl. 7, figs. 1, 2 (on rocks in the Cataract of the Surinam River below Kedjo, 100 mi. upriver from Paramaribo, Dutch Guiana); holotype MCZ 80515. fairchildi Clench, Cerion (Strophiops) 1933, Proc. N. Engl. Zool. Club, 13: 97, pl. 1, fig. 6 (Conception Island, Bahamas); holotype MCZ 80502, paratypes MCZ 80503. fairchildi Clench, Liguus blainianus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 117, pl. 7, fig. 4 (road between Havana and Santiago de las Vegas, Havana Prov., Cuba); holotype MCZ 91907. fairchildi Bequaert & Clench, Solaropsis gibboni 1938, Nautilus, 51: 115, pl. 9, figs. 6, 7 (Anapolis, Goyas, Brasil); holotype MCZ 65208; paratypes MCZ 65209. farnumi Clench, Liguus crenatus 1929, Nautilus, 43: 19 (Hammock #7, Pinecrest Region, the Ever- glades, Florida); holotype MCZ 84586. fasciata Clench, Sulcobasis (Goldielix) 1957, Am. Mus. Novit., (1863): 6, fig. 4 (Lorengau, Manus Is- land, Admiralty Islands); holotype MCZ 190157. feriai Clench & Aguayo, Cerion vulneratum 1953, Torreia, (18): 3, text fig. 6 (Cayo Largo o de los Muertos, Bahia o Puerto Naranjo, Oriente, Cuba); holotype Museo Poey 17247, paratype MCZ 83152. feriai Clench, Liguus fasciatus 1934, Occas. Pap. Bos- ton Soc. Nat. Hist., 8: 108, pl. 7, fig. 6 (La Sierra, Holguin, Oriente, Cuba); holotype MCZ 72559. fernandina Clench, Cerion 1937, Nautilus, 51: 21, pl. 3, fig. 5 (Millers, 8 mi. SE of Simms, Long Island, Bahamas); holotype MCZ 65131. Fijianella (genus) Cooke & Clench, 1943, Occas. Pap. B. P. Bishop Mus., 17(20): 257 (type species Fiji- anella calciphila Clench & Cooke 1943, original designation). Fijiopoma (genus) Clench, 1949, Bull. B. P. Bishop Mus., (196): 23 (type species Ostodes liberatus Garrett 1887, original designation). finlayi Clench, Murex (Murex) 1955, Breviora, (44): 1-3, text figs. 1-3 (shell trap, Matanzas Bay, Cuba, 100 fth.); holotype MCZ 189939. fisheri Gustafson, Turner, Lutz & Vrijenhoek, Tamu 1998, Malacologia, 40(1—2): 91, figs. 11-13, 21-23 (Johnson Sea-Link-1 Dive 3108, 27°46.91'N, 91°30.36'W, Bush Hill hydrocarbon seep, 210 km SSW of Grand Isle, Louisiana, 548 m); holotype ANSP A18849, paratype MCZ 316979. fitchi Turner, Penitella 1955, Johnsonia, 3(34): 66, pls. 4042 (Bahia San Bartolomé, Baja California, Mex- ico); holotype MCZ 189413. flammellus Clench, Liguus flammellus 1934, Occas. Paps Boston soc: Nat. Hist, 8: 119) pl) 6g: 7 (Mogote de Roja, Vifiales, Pinar de Rio, Cuba); ho- lotype MCZ 80924. flavipellis Clench & Archer, Helicostylus (Chrysallis) mindoroensis 1933, Pap. Mich. Acad. Sci. Arts Lett., 17: 548, pl. 58, fig. 5 (Calawagan, Paltan, Mindoro, Philippines); holotype MCZ 81358, para- type 81357. florida Clench & Aguayo, Aurinia dohrni 1940, Mem. Soc. Cubana Hist. Nat., 14(1): 88, pl. 16, fig. 1, text fig. | (new name for Voluta dubia Dohrn 1879, non Broderip 1828). floridanus Clench, Liguus crenatus 1929, Nautilus, 43: 20 (Hammock #8, Pinecrest Region, Central Everglades, Florida); holotype MCZ 84559. Forbesopomus (genus) Bequaert & Clench 1937, Proc. N. Engl. Zool. Club, 16: 53, pl. 11 (type spe- cies Forbesopomus atalanta Bequaert & Clench 1937, original designation). Forcartia (genus) Clench & Turner 1963, J. Malacol. Soc. Aust., (6): 19 (type species Papuina buehleri Rensch 1933, original designation). fosteri Clench & Turner, Bankia (Plumuletta) 1946, Johnsonia, 2(19): 24, pl. 15, figs. 14 (Santa Marta, Colombia); holotype MCZ 122536. fosteri Clench & Aguayo, Conus 1942, Johnsonia, 1(6): 34, pl. 12, fig. 5 (Atlantis Sta. 3434, 23°10'N, 79°35'W, off Sagua la Grande, Santa Clara Prov., Cuba); holotype MCZ 146003. fosteri Clench, Plagioptycha 1940, Mem. Soc. Cu- bana Hist. Nat., 14(1): 16, pl. 3, fig. 4 (Bret’s Hill, Long Island, Bahama Islands); holotype MCZ MSSOV- foxi Clench, Placostylus (Eumecostylus) 1950, Tor- Te tae Gli) ee 4s ate xct fig. (OmishSenliGile2 7s leu. 12 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 San Cristobal, Solomon Islands); holotype MCZ 187121 Foxidonta (genus) Cl., 1950, Rev. Soc. Malacdl. “Carlos de la Torre,” 7: 59 (type species Foxidonta stevensoni Clench 1930, monotypy). freirei Clench & Aguayo, Emoda (Glyptemoda) 1950, Rev. Soc. Malacél. “Carlos de la Torre,” 7: 61, pl. 12, figs. 1-3 (La Manuela, antes de llegar a la Fur- nia de la Mina los Negros, Baire, Oreinte, Cuba); holotype Museo Poey 12918, paratype MCZ 185798. frisbeyae Clench & Pulley, Conus 1952, Tex. J. Sci. pp. 59-61, pl. A, fig. 1 (Campeche Bank, Yucatan, Mexico, about 23 fth.); holotype MCZ 187708. furva Clench & Archer, Heliostyla (Chrysallis) min- doroensis 1933, Pap. Mich. Acad. Sci. Arts Lett., 17: 546, pl. 58, fig. 3 (Binuangan, Paltian, Mindoro, Philippines); holotype MCZ 81356. fusiformis Clench & Aguayo, Metula 1941, Mem. Soc. Cubana Hist. Nat., 15(2): 179, pl. 14, fig. 1 (Atlantis Sta. 3344, 21°38’N, 80°12’W, off Cienfue- gos, Cuba, 1,440 fth.); holotype MCZ 135290. gaigei Bequaert & Clench, Choanopoma 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 425 (Chichen Itza, Yucatan): holotype UMMZ 49190, paratypes MCZ 79390 and 79783. galathea Clench, Nesopoma 1930, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2: 169, pl. 17, fig. 2 (Suu, Ma- laita, Solomon Islands); holotype AMNH 73884, paratypes MCZ 188085. gallegoi Clench, Placostylus (Proaspastus) 1941, Am. Mus. Novit., (1129): 15, fig. 6 (Mts. of San Chris- toval Island, Solomon Islands); holotype AMNH 79003, paratype MCZ 92901. garciana Clench & Aguayo Cepolis (Hemitrochus) 1953 [in] Aguayo, Mem. Soc. Cubana Hist. Nat., 21: 308, pl. 35, figs. 8-10 (Caletones, Gibara, Or- iente, Cuba); holotype Museo Poey 17303, para- types MCZ (not received). garretti Clench Ostodes 1949, Bull. B. P. Bishop Mus., (196): 18, fig. 7c (Siavao-Auola, Savaii Island, Samoan Islands); holotype Bishop Museum 108334, paratypes MCZ 140524. Gasseisia (genus) Clench, 1949, Bull. B. P. Bishop Mus., (196): 5 (type species Cyclostoma artense Montrouzier 1859, original designation). geophilus Clench & Aguayo, Cerion 1949, Torreia (Univ. Havana), (14): 5, pl. 1, figs. 7-12 (Punta de Piedra, Banes, Cuba); holotype MCZ 184865, para- types Museo Poey 7028. georgiana Clench, Rehderia 1946, Johnsonia, 2: 47, pl 2 fig. 98" (Leliconm Stay WiGeay ole 2onaNy 79°41.5'W, off Darien, Georgia, 45 fth.); holotype USNM 543509, paratype MCZ 179163. gerda Turner, Xylophaga 2002, Bull. Mus. Comp. Zool. 157(4): 237, pls. 2, 3 (Gerda Sta. 499, 26°37'N, 78°56'W, about 3 mi. off Southwest Point, Great Bahama Island, Bahama Islands, 155 fth.): holotype MCZ 328378, paratypes MCZ 316741, 316742. gizoensis Clench, Placostylus (Placostylus) 1941, Am. Mus. Novit., (1129): 5, fig. 10 (Gizo Island, New Georgia Group, Solomon Islands); holotype Bishop Museum 106216, paratype MCZ 106357. Glyptemoda (section) Clench & Aguayo, 1950, Rev. Soc. Malacol. “Carlos de la Torre,” 7: 61 (type spe- cies, Helicina torrei Henderson 1909, original des- ignation). goodrichi Clench, Liguus faciatus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 111, pl. 7, fig. 7 (0.75 mi. below Castillo de la Jagua, entrance of Cien- fuegos Harbor, Santa Clara, Cuba); holotype MCZ DIDOoY goodrichi Clench, Physa 1926, Occas. Pap. Mus. Zool., Univ. Mich., (168): 1, pl. 1, figs. 1, 2 (South Fork, Powell River, Big Stone Gap, Virginia); ho- lotype UMMZ 29520, paratypes MCZ 51118. Goodrichia (genus) Clench, 1929, Bull. Mus. Comp. Zool., 69: 117 (type species, Goodrichia trochifor- mis Clench 1929, original designation). gouldi Clench, Physa 1935, Nautilus, 49: 30; ibid, 48, pl. 7, fig. 5 (Mouse River, 6 mi. N of Towner, Mc- Henry Co., North Dakota); holotype MCZ 103282, paratypes MCZ 103283. greenwayae Clench, Oliva reticularis 1937, Proc. N. Engl. Zool. Club, 16: 21, pl. 1, fig. 3 (Smith Point, Grand Bahama Island, Bahamas); holotype MCZ 115445. greenwayi Clench, Cerion (Strophiops) 1934, Proc. Boston Soc. Nat. Hist., 40(2): 206, fig. H (Black Booby Cay, West Booby, Atwoods on Samana Group, Bahama Islands); holotype MCZ 103001. greenwayi Clench, Drymaeus bahamensis 1933, Proc. N. Engl. Zool. Club, 13: 88, pl. 1, fig. 11 (Landrail Pt., Crooked Island, Bahamas); holotype MCZ 101187. greenwayi Clench, Eutrochatella 1933, Proc. N. Engl. Zool. Club, 13: 81, pl. 1, fig. 16 (Eleuthera Island, Bahamas); holotype MCZ 90181. greenwayt Clench, Microceramus (Spiroceramus) 1938, Mem. Soc. Cubana Hist. Nat., 12: 329, pl. 25, fig. 1 (Foxtown, Little Abaco Island, Bahama Islands); holotype MCZ 116688. Guladentia (subgenus) Clench & Aguayo, 1951, Rev. Soc. Malacol. “Carlos de la Torre,” 7: 87 (type spe- cies, Helix subtussulcata Pfeiffer 1863, original des- ignation). gundlachi Clench & Aguayo, Jeanneretia (Guladen- tia) 1951, Rev. Soc. Malacél. “Carlos de la Torre,” 7: 89, pl. 14, figs. 9-11 (Mogote La Resbalosa, Vi- fales, Cuba); holotype Museo Poey 16003, para- types MCZ 181564. guppyi Clench, Omphalotropis nebulosa 1958, Nat- ural History of Rennell Island, British Solomon Is- lands, Copenhagen, Denmark, 2: 171, pl. 17, fig. 1 (Olimbrui, Malaita, Solomon Islands); holotype AMNH 59010, paratype MCZ 93014. Haita (subgenus) Clench & Aguayo, 1932, Proc. N. MOLLUSCA OF CLENCH AND TURNER ¢ Johnson 13 Engl. Zool. Club, 13: 37 (type species Physa | Hai- ta] elegans Clench & Aguayo 1932, original desig- nation). haitiana Clench & Aguayo, Ferrissia (Laevapex) 1937, Mem. Soc. Cubana Hist. Nat., 11:(2): 70 (Lake Miragoane, 2 mi. S of Miragoane, Haiti); ho- lotype MCZ 83889. haitiana Clench, Urocoptis monilifera 1935, Proc. Boston Soc. Nat. Hist., 41: 2, pl. 2, fig. C (18°43’N, 72°10'W, Mirebalais road, 26 km NE of Port au Prince, Poste Terre Rouge, Haiti, +2,000 ft.); ho- lotype MCZ 108599. haitiensis Clench, Archegocoptis 1966, Breviora, (245): 2, pl. 1, figs. 1, 2 (top of Morne Rochelois, 25 km WSW of Miragoane, Dépt. du Sud, Haiti, 3,000 ft.); holotype MCZ 252056. hassler Clench & Aguayo, Calliostoma (Astele) 1939, Mem. Soc. Cubana Hist. Nat., 13: 191, pl. 28, fig. 3 (Hassler Voyage, off Cabo Frio, Brasil, 75 mi. E of Rio de Janeiro, Brasil, 35 fth.); holotype MCZ 104554. haysae Clench, Thais floridana 1927, Nautilus, 41: 6, pl. 2, fig. 11, 44 (Grand Bayou, Mississippi Delta, Louisiana); holotype MCZ 52203, paratypes MCZ 52204. heckerae Turner, Gustafson, Lutz & Vrijenhoek, Bathymodiolus 1998, Malacologia, 40(1—2): 68, figs. 6113 (Alvin Dive 1343, 26°.3'N, 84°54’W, base of the W Florida Escarpment in the eastern Gulf of Mexico, 2,370 m); holotype ANSP A18846, para- type MCZ 316977 from Alvin Dive 1755, 26°1.5'N, 84°55.3'W, 3,300 m. hedleyi Clench, Isidora newcombi 1926, J. Conch., 18: 12 (new name for Physa inflata Adams and Angus 1864, non Physa inflata Lea 1841, non Phy- sa fontinalis inflata Moquin-Tandon 1855). helenae Clench & Turner, Meliobba 1960, J. Malacol. Soc. Aust., (4): 30-31, pl. 3 (5°S'S, 144°50’E, Asai- Simbai Divide, Schrader Range, a few miles due W of Aiome, Terr. of New Guinea); holotype Aus- tralian Museum C 62378 (only specimen). helianthus Clench, Liguus fasciatus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 112, pl. 7, fig. 9 (Mogote del Palmarito, Vifales, Pinar del Rio, Cuba); ho- lotype MCZ 80911. hendersoni Clench, Physa pomilia 1925, Occas. Pap. Mus. Zool., Univ. Mich., (161): 4, pl. 1, fig. 3 (Ye- massee, Beaufort Co., South Carolina); holotype UMMZ (Walker Collection no. 45640). The Walker number on the UMMZ lot corresponds to para- types from Clench (teste Graf, personal commu- nication). Because the holotype could not be lo- cated, these and other type lots are regarded as syntypes, UMMZ 123646, 129173; MCZ 51151. hessei Clench & Aguayo, Cerion 1949, Torreira (Univ. Habana), (14): 8, pl. 1, figs. 19-22 (Balcén de las Damas, Guarda la Vaca, Banes, Cuba); holotype MCZ 183635, paratypes Museo Poey 6154. hirasei Clench, Bulinus (Culmenella) 1927, Nautilus, 40: 121, text fig. 1 (Prov. of Kawachi, Japan); ho- lotype UMMZ Walker Collection no. 38964, not located in UMMZ (teste Graf, personal communi- cation); syntypes MZUM 86896, MCZ 51425. hispaniola Clench, Brachypodella (Liparotes) 1935, Proc. Boston Soc. Nat. Hist., 41: 9, pl. 2, fig. F (18°22’N, 74°2’W, Tardieu, Mt. La Hotte, Pic de Macaya, 31 km SSE of Jeremie, Haiti, 3,000—4,000 ft.); holotype MCZ 108583. hispaniolae Clench & Aguayo, Chondropoma (Chon- dropomorus ) 1937, Mem. Soc. Cubana Hist. Nat., 11(2): 64, pl. 7, figs. 1, 2 (Milot, Haiti); holotype MCZ 28410. hispaniolana Clench, Cepolis 1964, Rev. Mus. Ar- gent. Cienc. Nat. “Bermardino Rivadavia,” 8: 222, pl. 2, fig. 4 (Grand Cayamite Island, Haiti, Hispan- iola); holotype MCZ 188262. hoeyeri Clench, Charopa 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2: 173, pl. 16, fig. 6 (Lavanggu, Rennell, Solomon Islands); holotype University Zoological Museum, Copenhagen (not numbered, teste O. S. Tendal, personal communication), para- type MCZ 191461. holguinensis Clench & Aguayo, Helicina 1953 [in] Aguayo, Mem. Soc. Cubana Hist. Nat., 21: 300, pl. 33, figs. 6, 7 (La Calera, San German, Holguin, Cuba); holotype Museo Poey 14311, paratypes MCZ (not received). hoodi Clench, Setaepoma 1965, Breviora, (224): 4, pl. 2, fig. 3 (W side of the Tenaru River, about 0.5 mi. above the Catholic Mission, Guadalcanal, Solomon Islands); holotype MCZ 251309. howardi Clench, Hemitrochus 1964, Rev. Mus. Ar- gent. Cienc. Nat. “Bernardino Rivadavia,” 8: 223, pl. 1, fig. 5 (Monteado Nuevo, 20 km SE of Polo, Prov. de Barahona, Républica Dominicana, 4,000 ft.); holotype MCZ 187924. howelli Clench & Aguayo, Ficus 1940 Mem. Soc. Cu- bana Hist. Nat., 14(1): 85, pl. 14, fig. 2 (Atlantis Sta. 3332, 22°9'30’N, 81°11’W, Bahia de Cochinos, Santa Clara Proy., Cuba, 175-225 fth.); holotype MCZ 135140. howelli Clench, Liguus nobilis 1951, Rev. Soc. Ma- lacél. “Carlos de la Torre,” 7: 93, text fig. (Rio Dominica, about 10 mi. W of Mariel, Pinar del Rio, Cuba); holotype MCZ 187133, paratype Museo Poey 12536. Howellia (genus) Clench & Aguayo, 1941, Mem. Soc. Cubana Hist. Nat., 15(2): 177 (type species How- ellia mirabilis Clench & Aguayo 1941, original des- ignation). hubrichti Clench, Lithasia 1965, Nautilus, 79: 30, text fig. 1 (Big Black River, 3 mi. NW of Edwards, Hinds Co., Mississippi); holotype MCZ 250916, paratypes MCZ 233392. hughesi Clench, Cerion 1952, Rev. Soc. Malacdl. “Carlos de la Torre,” 8: 107, pl. 15, figs. 1-3 (Sand Point, Savannah Sound, Eleuthera Island, Baha- mas); holotype MCZ 116026. humberti Clench & Aguayo, Cerion 1949, Torreia, (14): 4, pl. 1, figs. 16-18 (Ensenada de Jiicaro, Ba- 14 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 hia de Banes, Banes, Cuba); holotype MCZ 184618, paratypes Museo Poey 4266. huntingtoni Clench, Cerion 1938, Bull. Mus. Comp. Zool., 80(14): 526, pl. 3, figs. 1-3 (Colombus Point, SE tip of Cat Island, Bahamas); holotype MCZ 106956. huntingtoni Clench, Pupina (Pupina) 1949, Bull. B. P. Bishop Mus., (196): 33, fig. 19 (N coast of Guad- alcanal Island, Solomon Islands); holotype MCZ 161588, paratype Bishop Museum 190021. icterica Bequaert & Clench, Archachatina marginata 1936, Rev. Zool. Bot. Afr., 29(1): 81 ((Gaboon)); holotype MCZ 45526, only specimen. ignava Clench & Archer, Helicina 1931, Occas. Pap. - Boston Soc. Nat. Hist., 5: 338, text fig. | (Lubang Island, Philippines); holotype MCZ 79176, para- types MCZ 47227. ilogana Clench & Archer, Helicostyla (Chrysallis) mindoroensis 1933, Pap. Mich. Acad. Sci. Arts Lett., 17: 545, pl. 58, fig. 4 (Tara Mangyan, Abra de Ilog, Mindoro, Philippines); holotype MCZ 81359, paratypes MCZ 47264. inaguense Clench, Cerion (Diacerion) 1933, Proc. N. Engl. Zool. Club, 13: 98, pl. 1, fig. 9 (Northwest Pt., Gt. Inagua, Bahamas); holotype MCZ 101164. incendium Clench, Pseudocyclotus 1957, Am. Mus. Novit., (1863): 2, fig. 3 (Rambutyo Island, Admi- ralty Islands); holotype AMNH 73486, paratypes MCZ 188140. indianorum Clench, Cerion paucicostatum 1934, Proc. Boston Soc. Nat. Hist., 40(2): 210, pl. 2, fig. F (Wemyso Biglet, Eleuthera Island, Bahama Is- lands); holotype MCZ 103065. ingolfia Turner, Xyloredo 1972, Breviora, (397): 7, pls. 35) (Ingolf Exped. Sta, 672) Oles0 Ne 22230 IW: wood dredged S of Eyrabakki, Iceland, 975 fth. [1,783 m]); holotype MCZ 279636, paratypes MCZ 2.79637. insolutus Clench & Aguayo, Opisthosiphon sainzi 1949, Rev. Soc. Malac6él. “Carlos de la Torre,” 6: 89, fig. 1 (Loma Leal, Los Haticos, Holguin, Cuba); holotype Museo Poey 12975, paratypes MCZ 167951. insula Clench, Brachypodella weinlandi 1935, Proc. Boston Soc. Nat. Hist., 41: 7, pl. 2, fig. E (Gonave Island, Haiti); holotype MCZ 106244. Insulaceramus (subgenus) Clench, 1967, Breviora, (260) 95 (type species, Microceramus | Spirocera- mus | greenwayi Clench 1938, original designation). insulaefelis Clench, Opiothosiphon bahamense 1938, Bull. Mus. Comp. Zool., 80(14): 515, pl. 2, figs. 1, 7 (Port Howe, southern end of Cat Island, Bahama Islands); holotype MCZ 107905. insularis Clench, Charopa 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2: 174, pl. 18, fig. 5 (Te-ava- manggu, Rennell Island, Solomon Islands); holo- type University Zoological Museum, Copenhagen [only specimen] (not numbered, teste O. S. Tendal, personal communication). insularis Clench & Turner, Truncatella thaanumi 1948, Occas. Pap. Mollusks, 1(13): 166 [not fig. ] (near Huma, Tongatabu, Tonga Islands); holotype B. P. Bishop Museum 87796, paratypes MCZ 183889. iselini Macdonald & Clench, Mastigoteuthis 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 150, text figs. 5, 7-9 (Atlantis Sta. 1106, 39°4'N, 71°29’W, N At- lantic); holotype MCZ 98967. jaumei Clench & Aguayo, Calliostoma (Calliostoma) 1946, Rev. Soc. Malacél. “Carlos de la Torre,” 4: 88, text fig. (Arenas de la Chorrera, Habana, Cuba, 3-15 fth.); holotype MCZ 178127, paratypes Mu- seo Poey 2941. jaumei Clench & Aguayo, Cerion peracutum 1953, Torreia, (18): 2, text fig. 3 (La Jijira, between Boca de Jaruco and Santa Cruz del Norte, Habana, Cuba); holotype Museo Poey 12796, paratype MCZ 187287. jaumei Clench & Aguayo Eutrochatella (Troschelvi- ana) 1957 [in] Aguayo, C. G., and M. L. Jaume, Mem. Soc. Cubana Hist. Nat. “Felipe Poey,” 23: 120, pl. 1, fig. 8 (Mogote de la Finca “La Esperan- za’ San Andrés, Consolacién del Norte, Pinar de Rio, Cuba); holotype Museo Poey 13262, paratypes MCZ 212979. jaumei Clench & Aguayo, Jeanneretia 1951, Rev. Soc. Malacol. “Carlos de la Torre,” 7: 87, pl. 13, figs. 14, 17 (Vega del Mamey, Pau de Guajaibon, Pinar del Rio, Cuba); holotype Museo Poey 16002, para- types MCZ 181722. jaumei Clench & Aguayo Liguus blainianus 1932, Nautilus, 45(3): 99, pl. 6, fig. 10 (Mangas, on road between Mangas and Candelaria, Pinar del Rio, Cuba); holotype MCZ 91720. jeanneae Clench & Turner, Calliostoma 1960, John- sonia, 4: 65, pl. 47, figs. 1, 2 (Atlantis, off Habana, Cuba); holotype MCZ 228370. johnsoni Clench, Physa 1926, Occas. Pap. Mus. Zool., Univ. Mich., (168): 2, pl. 1, fig. 3 (Middle Spring, Hot Sulphur Springs, Banff, Alberta, Canada); ho- lotype UMMZ 33066, paratypes MCZ 55308, 53595. johnstoneae Clench, Scaphella junonia 1953, John- sonia, 2: 376, pl. 186, fig. 2 (off Petit Bois Island, Alabama, 10 fth.); holotype MCZ 194190. johnstonei Clench & Turner, Melongena corona 1955, Johnsonia, 3(35): 178, pl. 105, figs. 1-3 (Little La- goon, Gulf Shores, Alabama); holotype MCZ 189687, paratypes MCZ 194022. Jokajia Clench (subgenus) 1946, Occas. Pap. B. P. Bishop Mus., 18(13): 204 (type species Thaanu- mella |Jokajia| cookei Clench 1946, original desig- nation). jonesi Clench, Bulimulus dealbatus 1937, Nautilus, 51(1): 18, pl. 3, fig. 4 (2 mi. N of West Greene, Greene Co., Alabama); holotype Alabama Univ. 89, now in Florida Museum of Natural History (UF) 174304, paratype MCZ 75036. josephi Clench & Aguayo, Cerion 1949, Torreia, (14): MOLLUSCA OF CLENCH AND TURNER * Johnson 15 6, pl. 1, figs. 23-25 (Playa de Uvita, al Oeste de Caletones, Gibara, Oriente, Cuba); holotype MCZ 184620, paratype MCZ 184621, paratypes Museo Poey 4264. josephinae Clench, Cerion (Strophiops) 1935, Nauti- lus, 49: 49, pl. 3, figs. 1, 4 (Tate’s Bay, SE Long Island, Bahama Islands); holotype MCZ 76474. juliae Clench, Cerion (Strophiops) 1936, Nautilus, 49(4): 112, pl. 8, fig. 6 (Gt. Ragged Cay, SE Gt. Bahama Bank, Bahama Islands); holotype MCZ 10369. juliae Clench, Conus 1942, Johnsonia, 1(6): 26, pl. 12, fig. 4 (9 mi. off Ft. Walton, Okaloosa Co., Flor- ida); holotype Florida Museum of Natural History (UF) 13382 [only specimen]. juliae Clench, Helicina 1962, Breviora, (173): 2, pl. 1, fig. 2 (Colonia Ramfis, 20 km W of San Cristobal, Républica Dominicana); holotype MCZ 168267. juliae Clench & Aguayo, Latiaxis 1939, Mem. Soc. Cubana Hist. Nat., 13: 194, pl. 28, fig. 4 (off Sandy Bay, Barbados, 75-100 fth.); holotype MCZ 89969. juliae Clench, Urocoptis (Autocoptis) 1935, Proc. Boston Soc. Nat. Hist., 41: 2, pl. 2, fig. D (Mt. Trou d’Eau, Haiti, 4,500 ft.); holotype MCZ 108601. juliae Clench & Turner Waghia 1959, J. Malac. Soc. Aust., (3): 4, pl. 1, figs. 1-3, text figs.1-3 (12 mi. NE of Mt. Hagen Range, Sepik-Wahgi Divide, New Guinea, 5,000-6,000 ft.); holotype Australian Mus. C62212, paratypes MCZ 191399 and 191400. junaluskana Clench & Banks, Retinella (Glyphogno- mun) 1932, Nautilus, 46: 15, pl. 2, fig. 4 (83°47'W, 35°12'’N, 2.5 mi. E of Andrews, Cherokee Co., North Carolina); holotype MCZ 86429. juttingae Clench & Turner, Megalacron 1964, J. Ma- lac, Soe, Aust, (Se 46, jal. OG me OS (QS, 151°56’E, Kalili Bay, New Ireland, Bismarck Ar- chipelago); holotype Univ. Zool. Mus., Copenhagen (not numbered, teste O. S. Tendal, personal com- munication), paratype MCZ 248834. Kanapa (genus) Clench, 1949, Bull. B. P. Bishop Mus., (196): 41 (type species Registoma torazieri Crosse 1870, original designation). Katastoma (section) Clench, 1935, Proc. Boston Soc. Nat. Hist., 41(1): 8 (type species, Brachypodella lasellensis Clench 1935, monotypy). katherinae Clench & Turner, Bankia (Liliobankia) 1946, Johnsonia, 2(19): 18, pl. 11, figs. 1-6 (Bahia, Brazil); holotype MCZ 168023. kejong Clench & Archer, Leptopoma 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 337, pl. 17, fig. 6 (Lubang Island, Philippines); holotype MCZ 47224, paratypes MCZ 47225. kieneri Clench, Scaphella (Aurinia) 1956, Johnsonia, 2: 58 (new name for Fusus tessellatus Kiener, 1840 non Schubert & Wagner 1829). kiyokoae Clench, Kondorhaphe 1949, Bull. B. P. Bishop Mus., (196): 28, figs. 15b, 16, 28d (Mt. To- lowan, Dublon Island, Truk Islands, Caroline Is- lands, 600-800 ft.); holotype Bishop Museum 9712, paratypes MCZ 140542. klineae Clench, Cerion (Maynardia) 1961, Occas. Pap. Mollusks, 2: 249, pl. 43, fig. 1 (Bonavista Cay, Ragged Islands, Bahama Islands); holotype MCZ 189209. klinei Clench, Eutrochatella 1959, Bull. Mus. Comp. Zool., 121(2): 34, pl. 1, fig. 5 (NE of Southwest Point, Little Inagua, Bahamas); holotype MCZ 189603. knudseni Clench, Quirosella 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2: 177, pl. 18, fig. 4, text figs. 2, 3 (Te-Avamanggu, Rennell Island, Solomon Is- lands); holotype University Zoological Museum, Copenhagen (not numbered, teste O. S. Tendal, personal communication), paratypes MCZ 191452, 191453. kobelti Clench & Archer, Pterocyclos 1932, Occas. Pap. Boston Soc. Nat. Hist., 8: 38, pl. 4, fig. A (Mt. Kina, Balu, North Borneo); holotype MCZ 47890, paratypes MCZ 31650. Kombologion (subgenus) Clench & Turner, 1960, Johnsonia, 4: 37 (type species, Calliostoma bairdii Verrill & Smith 1880, original designation). kondoi Clench, Paramia 1945, Bull. B. P. Bishop Mus., (196): 27, figs. 15a, 28b (Nebokos, Moen Is- land, Truk Islands, Caroline Islands); holotype Bishop Museum 9713, paratypes MCZ 140539. Kondorhaphe (genus) Clench, 1949, Bull. B. P. Bish- op Mus., (196): 28 (type species Kondorhaphe ki- yokoae Clench 1949, original designation). Kubaryia (genus) Clench, 1948, Occas. Pap. B. P. Bishop Mus., 19(8): 191 (type species Kubaryia pi- likia Clench, 1948, original designation). kugleri Clench & Perez Farfante, Murex (Murex) cailleti 1945, Johnsonia, 1(17): 19, pl. 9, figs. 1, 2 (new name for Murex similis Sowerby 1841, non Schroeter 1805). laddi Cooke & Clench, Fijianella 1943, Occas. Pap. B. P. Bishop Mus., 17(20): 259, fig. 8a (Navutu II- oma, Lau Islands, Fiji Islands); holotype Bishop Museum 10108, paratypes MCZ 137109. laddi Clench & Turner, Taheitia turricula 1948, Oc- cas. Pap. Mollusks, 1(13): 189 [not fig.] (S end of Yangasa Levu, Lau Group, Fiji Islands); holotype Bishop Museum 167165, paratypes MCZ 157850. lahottensis Clench, Brachypodella kraussiana 1935, Proc. Boston Soc. Nat. Hist., 41: 7, pl. 2, fig. I (Tardieu, Mt. La Hotte, Haiti, 3-4,000 ft.); holo- type MCZ 108582. langi Clench & Turner, Thais (Thaisella) 1948, Am. Mus. Novit., (1374): 1, figs. 7-9 (Lobito Bay, An- gola, Africa); holotype AMNH 72715, paratypes MCZ 181240. lasellensis Clench, Brachypodella (Brevipodella) 1935, Proc. Boston Soc. Nat. Hist., 41: 8, pl. 2, fig. G (La Visite, La Selle Range, Haiti, 6—7,000 ft.); holotype MCZ 108604. lasellensis Clench & Aguayo, Lucidella (Poeniella) 1937, Mem. Soc. Cubana Hist. Nat., 11(2): 63, pl. 16 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 7, fig. 6 (La Visite, La Selle Range, Haiti); holotype MCZ 110626. lauensis Clench, Gonatorhaphe 1949, Bull. B. P. Bishop Mus., (196): 23, fig. 10 (near Marona, Man- gold, Lau Group, Fiji Islands); holotype Bishop Museum 180060 (and single broken paratype). laureani Clench & Aguayo Cerion 1951, Rev. Soc. Malacol. “Carlos de la Torre,” 8: 74, pl. 11, figs. 7, 8 (Cabo Corrientes, Peninsula de Guanahacabibes South, Pinar del Rio, Cuba); holotype Museo Poey 17232, paratypes MCZ 129443, 188527, 188528. lepidum Clench & Aguyo, Cerion vulneratum 1951, Rev. Soc. Malacél. “Carlos de la Torre,” 8: 76, pl. 11, figs. 10, 11 (Laguna, Punta de Mulas, Banes, Oriente, Cuba); holotype Museo Poey 12665, para- types MCZ 166619, 178328. Leptocalina Bequaert & Clench 1934, Rev. Zool. Bot. Afr., 26(1): 117 (type species, Achatina specularis Morelet 1866, subsequent designation, Bequaert 1950, Bull. Mus. Comp. Zool., 115[1]: 136). lerneri Clench, Cerion eximium 1956, Am. Mus. Nov- it., (1794): 1-3, two text figs. (S tip of East Bimini, Bimini Islands, Bahamas); holotype MCZ 186830. lewisi Clench, Cerion (Multostrophia) 1961, Occas. Pap. Mollusks, 2(26): 255, pl. 43, fig. 5 (Pine Cay, Caicos Islands, Bahamas Islands); holotype USNM 610286, paratypes MCZ 921564. Lignophalas (genus) Turner, 1955, Johnsonia, 3(34): 98 (type species, Lignophalas clappi Turner, 1955, original designation). Liguellus (subgenus) Clench, 1946, Occas. Pap. Mol- lusks, 1(10): 123 (type species Achatina vittata Swainson 1822, original designation). Liliobankia (subgenus) Clench & Turner 1946, John- sonia, 2(19): 17 (type species Bankia katherinae Clench & Turner 1946, original designation). liliorum Clench, Cerion 1938, Bull. Mus. Comp. Zool., 80(14): 527, pl. 2, figs. 2-4 (Next Point, east coast, 1.5 mi. NE of Governors Harbour, Eleuth- era Island, Bahama Islands); holotype MCZ 116086. loveridgei Clench & Archer, Euaethiops 1930, Occas. Pap. Boston Soc. Nat. Hist., 5: 298, pl. 16, fig. A (Bagilo, Uluguru Mts., Tanganyika Territory, Afri- ca); holotype MCZ 58934, paratypes MCZ 258936. loveridgei Bequaert & Clench, Gonaxis (Macrogon- axis) 1936, J. Conch, 20(9): 271, text figs. 1-5 (Bag- ilo, Uluguru Mts., 500 ft., Tanganyika Territory, Af- rica); holotype MCZ 59915. loveridgei Bequaert & Clench, Leptocala (Leptocal- lista) 1934, Rev. Zool. Bot. Afr., 24: 274, pl. 1, figs. 1, 5, 6 (Nyange, 3,000 ft., Uluguru Mts., Tangan- yika Territory, Africa); holotype MCZ 59949. Sec- ondary homonym of Achatina (Euaethiops) lover- idgei Clench & Archer 1930; renamed Achatina ar- thori. Bequaert 1950. louisiadensis Clench, Pupinoa 1949, Bull. B. P. Bish- op Mus., (196): 38, fig. 21C (Louisiade Islands); holotype Bishop Museum 189002, paratypes MCZ 140575. lubanensis Clench & Archer, Helicostyla cincinnifor- mis 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 334, pl. 17, fig. 1 (Lubang Island, Philippines); ho- lotype MCZ 79166, paratypes MCZ 79165. lubanicus Clench & Archer, Cyclophorus reevei 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 337, pl. 17, fig. 7 (Lubang Island, Philippines); holotype MCZ 47221, paratypes MCZ 47221. lucayanorum Clench, Cerion (Strophiops) 1938, Mem. Soc. Cubana Hist. Nat., 12: 326, pl. 25, fig. 2 (NW portion of Mores Island, 32 mi. NW of Southwest Point, Great Abaco Island, Bahamas): holotype MCZ 116017. lucayanorus Clench, Opisthosiphon (Opisthosiphon) 1963, Bull. Mus. Comp. Zool., 128(8): 399, pl. 3, fig. 2 (3.75 mi. NE of Salina Point, Acklins Island, Bahamas); holotype MCZ 221179. lukolelensis Bequaert & Clench, Maizania (Maiza- niella) 1936, Rev. Zool. Bot. Afr., 29(1), p. 100, pl. 2, figs. 10, 11 (Lukolela, Belgian Congo); holotype MCZ 77206. lymani Cl, Conus bermudensis 1942, Johnsonia, 1(6): 30, pl. 13, fig. 3 (off Nellies Pt., South Lake Worth, Palm Beach Co., Florida); holotype Florida Mu- seum of Natural History (UF) 13362 (only speci- men). macdonaldi Gustafson, Turner, Lutz & Vrijenhoek, “Idas” 1998, Malacologia, 40(12): 98, figs. 11-13, 24-2 (Johnson Sea-Link-1 Dive 3149, 27°50'N, 92°10'W, Gulf of Mexico on the Louisiana Conti- nental Slope near Garden Banks block 386, 650 m); holotype ANSP A18850, paratypes ANSP 400783, 400784, MCZ 316980. madaziniana Clench & Archer Achatina 1930, Occas. Pap. Boston Soc. Nat. Hist., 5: 299, pl. 16, fig. B (Madazini, Tazanyika Territory); holotype MCZ 53185, paratypes MCZ 53186. Magachatina Bequaert & Clench 1936, Rev. Zool. Bot. Afr., 29(1): 78, misspelling of Megachatina. magnifica Boss & Turner, Calyptogena 1980, Mala- cologia, 20(1): 161, figs. 1-9, 1OF—G, 11, 12D-—-F, 13 (Alvin Dive 717, 0°47.9'N, 86°8.5’W, Galapagos Rift, 2,459 m); holotype MCZ 288500. Maizaniella (subgenus) Bequaert & Clench, 1936, Rev. Zool. Bot. Afr., 29(1): 99 (type species, Cyclo- phorus leonensis Morelet 1873, original designa- tion). Malaitella Clench, 1941, Am. Mus. Novit., (1129): 11 (new name for Acrotylus Clench 1935, non Coss- mann 1896). malaitensis Clench, Placostylus (Malaitella) 1941, Am. Mus. Novit., (1129): 11, fig. 4 (near Su’u, Ma- laita Island, Solomon Islands); holotype AMNH 79006, paratype MCZ 92905, Aola Mulaita Island, 3,000 ft. malonei Clench, Cerion 1937, Nautilus, 51: 30, pl. 3, fig. 6 (3.5 mi. SE of Simms, Long Island, Baha- mas); holotype MCZ 112706. mamburaoensis Clench & Archer, Obba planulata 1932, Occas. Pap. Boston Soc. Nat. Hist., 8: 40, pl. MOLLUSCA OF CLENCH AND TURNER * Johnson 17 4, fig. D (Calomintao, Mamburao, Mindoro, Phil- ippines); holotype MCZ 92799. manni Clench & Aguayo, Chondropoma (Chondro- pomorus) 1937, Mem. Soc. Cubana Hist. Nat., 11(2): 65, pl. 7, fig. 3 (Furcy, Haiti); holotype MCZ 25415, paratype MCZ 36692. manni Clench, Kanapa 1949, Bull. B. P. Bishop Mus., (196): 42, figs. 23a-24b—d MCZ (Three Sisters Is- lands, Solomon Islands); holotype MCZ 141012, paratypes Bishop Museum 188844. manni Clench & Aguayo, Lucidella 1932, Proc. N. Engl. Zool. Club, 13: 35 (Furcy, Haiti); holotype MCZ 23414. marcanoi Clench, Proserpina 1962, Breviora, (173): 2, pl. 1, fig. 3 (Colonia Ramfis, 20 km W of San Cristobal, Reptiblica Dominicana); holotype MCZ 188911. mariae Clench, Liguus fasciatus 1935, Nautilus, 48: 123, pl. 7, fig. 3 (Ojo de Agua, near Cayo Magueyal, Pinar del Rio, Cuba); holotype MCZ 94224. maricao Clench, Polydontes (Granodormus) 1940, Mem. Soc. Cubana Hist. Nat., 14(3): 243, pl. 42, figs. 2, 5 (Maricao Forest, 2—3,000 ft., W Puerto Rico): holotype MCZ 111636. mariguanense Clench, Cerion (Strophiops) 1933, Proc. N. Engl. Zool. Club, 13: 94, pl. 1, fig. 3 (S coast of Mariguana Island, Bahamas); holotype MCZ 101162. mariguanense Clench, Chondropoma 1937, Proc. N. Engl. Zool. Club, 16: 66, pl. 3, fig. 2 (Abrahams Hill, 2 mi. NE of Abrahams Bay, Mariguana Island, Bahamas); holotype MCZ 57987. matthewsae Clench & Turner, Epitonium multistria- tum 1952, Johnsonia, 2: 295, pl. 135 (Sanibel Is- land, Florida); holotype MCZ 197134. mauritii Bequaert & Clench, Volvatorbis 1936, Mem. Mus. R. Hist. Nat. Belg., 2nd ser., (3): 166, fig. 2 (Ango-Ango, 4 km S of Matadi, Congo River, Bel- gian Congo); holotype MCZ 59803. maya Bequaert & Clench, Spiraxis (Volutaxis) 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 423 (Chi- chenltza, Yucatan); holotype MCZ 85799. mayabigensis Clench & Archer, Obba listeri 1932, Occas. Pap. Boston Soc. Nat. Hist., 8: 39, pl. 4, fig. C (Mayabig, Baco, Mindoro, Philippines); holotype MCZ 92798, paratypes MCZ 93992. mayri Clench, Palaeohelicina (Palaeohelicina) 1958, Natural History of Rennell Island, British Solomon Islands, Copenhagen, Denmark, 2: 163, pl. 1, fig. 7 (Auki, Malaita, Solomon Islands); holotype MCZ 32610. mayri Clench, Placostylus (Aspastus) miltocheilus 1941, Am. Mus. Novit., (1129): 19 (new name for Bulimus (Aspastus) miltocheilus minor Brazier 1894 non Kobelt 1891). mayri Clench, Setaepoma 1958, Natural History of Rennell Islands, British Solomon Islands, Copen- hagen, Denmark, 2: 168, pl. 17, fig. 5, text fig. 1 (Fulakora, Ysebel, Solomon Islands); holotype MCZ 36840. megintyi Clench, Liguus fasciatus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 116, pl. 7, fig. 10 (Cueva del Chivo, Mariel, Pinar del Rio, Cuba); holotype MCZ 47384. mcleani Clench, Alcadia (Alcadia) 1937, Proc. N. Engl. Zool. Club, 16: 74, pl. 3, fig. 6 (Landrail Point, Crooked Island, Bahamas); holotype MCZ 64782. mceleani Clench, Cerion 1937, Nautilus, 51: 22, pl. 3, fig. 7 (1 mi. E of O’Neills, Long Island, Bahamas); holotype MCZ 112701. mcleani Clench, Polydontes (Hispaniolana) 1964, Rev. Mus. Argent. Cienc. Nat. “Bernardino Riva- davia,” 8: 217, pl. 2, fig. 1 (below Valle Nuevo, SE of Constanza, Prov. de la Vega, Républica Dom- inicana); holotype MCZ 188213. mcleani Clench, Trochomorpha 1958, Natural His- tory of Rennell Island, British Solomon Islands, Copenhagen, Denmark, 2: 179, pl. 17, fig. 6 (10 mi. inland from Su’u, Malaita Island, Solomon Is- lands); holotype AMNH 79016, paratype MCZ 93070. Mcleani (genus) Bequaert & Clench, 1939, Mem. Soc. Cubana Hist. Nat., 13: 283 (type species M. darlingtoni Bequaert & Clench 1939, original des- ignation). mcemichaeli Clench & Turner Elliptio 1956, Bull. Fla. State Mus., 1(3): 170, pl. 7, figs. 1, 2 (Chacta- whatchee River, 8 mi. W of Miller Cross Roads, Holmes Co., Florida); holotype MCZ 191922. memichaeli Clench & Turner, Meliobba 1963, J. Ma- lacol. Soc. Aust., (6): 24, pl. 2, fig. 3, text figs. 3(2), 6(4), 7 (Noorweg, Hollandia, Dutch New Guinea); holotype Aust. Museum C. 62377, paratype MCZ 87618. memurrayi Clench & Aguayo, Latirus (Hemipolygo- na) 1941, Mem. Soc. Cubana Hist. Nat., 15: 178, Pilate ten oF WALaniis: Ska OAS25, 230) N, 81°27'30"W, off Matanzas, Cuba, 190 fth.); holo- type MCZ 135285. Megachatina (subgenus) Bequaert & Clench 1926, Rev. Zool. Bot. Afr., 29(1): 76 (type species Acha- tina marginata Swainson 1821, original designa- tion). Megachatinops Bequaert & Clench, 1936, Rev. Zool. Bot. Afr., 29(1): 87, misspelling of Megachatinop- sis. Megachatinopsis (subgenus) Bequaert & Clench, 1936, Rev. Zool. Bot. Afr., 29(1): 76 (type species, Achatina knorrii Jonas 1839, original designation). melanesia Clench & Turner, Megalacron 1964, J. Ma- lac. Soc. Aust., 8: 53, pl. 10, fig. 14, text fig. 5 (wa- terfall, Lorengau, Manus Island, Admiralty Islands, Bismarck Archipelago); holotype University Muse- um, Copenhagen (not numbered, teste O. S. Ten- dal, personal communication), paratypes MCZ 249306. melanostomum Clench, Cerion (Strophiops) 1934, Proc. Boston Soc. Nat. Hist., 40(2): 212, pl. 2, figs. A, C (Mortimer’s, S end of Long Island [Berry Is- land], Bahama Islands); holotype MCZ 103021. Merrilliana (subgenus) Clench & Turner, 1948, Oc- 18 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 cas. Pap. Mollusks, 1(13): 179 (type species, Trun- catela elegans C. B. Adams 1849, original desig- nation). Mesanella (genus) Clench & Turner, 1952, Nautilus, 66: 32 (type species, Helix trailli Pfeiffer 1855, original designation). michiganensis Clench, Physa 1926, Occas. Pap. Mus. Zool., Univ. Mich., (168): 4, pl. 1, fig. 4 (stream 1 mi. W of Geddes, Washtenau Co., Michigan); ho- lotype UMMZ 33068, paratypes MCZ 251177. mirabilis Clench & Aguayo, Howellia 1941, Mem. Soc. Cubana Hist. Nat., 15(2): 177, pl. 14, fig. 2 (Atlantis Sta. 3483, 23°12’N, 81°23’'W, off Matan- zas, Cuba, 285 fth.); holotype MCZ 135291. modica Clench & Aguayo Jeanneretia (Guladentia) 1951, Rev. Soc. Malacél. “Carlos de la Torre,” 7: 89, pl. 14, figs. 7, 8 (Mal Paso, Luis Lazo, Pinar del Rio, Cuba); holotype Museo Poey 17220, paratypes MCZ 181560 from Le Caoba, Luis Lazo, Cuba. monactia Clench & Archer, Helicostyla (Calocochlea) 1933, Pap. Mich. Acad. Sci. Arts Lett., 17: 539, pl. 57, fig. 1 (Binuangan, Paltian, Mindoro, Philip- pines); holotype MCZ 81353. monaense Clench, Cerion 1951, J. Conchy., 90: 274, figs. 7-11 (Mona Island, Puerto Rico); holotype MCZ 171019. montana Clench, Coryda 1964, Rev. Mus. Argent. Cienc. Nat. “Bernardino Rivadavia,” 8: 226, pl. 2, fig. 5 (Loma Vieja, SW of Constanza, Prov. de la Vega, Républica Dominicana); holotype MCZ 187954. montana Clench, Polydontes (Hispaniolana) 1964, Rev. Mus. Argent. Cienc. Nat. “Bernardino Riva- davia,” 8: 216, pl. 2, fig. 2 (Mt. Quita Espuela, 14 km NE of San Francisco de Macoris, Prov. Duarte, Républica Dominicana); holotype MCZ 188209. monticola Clench & Archer, Thersites 1938, J. Conch., 21: 22, pl. 1, fig. 4 (Mt. Spurgeon, 50 mi. NW of Cairns, Queensland); holotype MCZ 99053, paratypes Queensland Museum, South Brisbane, Queensland (QMMO) 1764. moralesi Clench & Aguayo, Cerion torrei 1951, Rev. Soc. Malacél. “Carlos de la Torre,” 8: 77, pl. 11, figs. 13, 14 (Playa de Morales, 11 km SE of Banes, Oriente, Cuba); holotype Museo Poey 17293, para- types MCZ 187476. moreleti Clench & Aguayo, Cerion iostomum 1951, Rev. Soc. Malacél. “Carlos de la Torre,” 8: 73, pl. 11, fig. 6 (Punta del Este, Isle of Pines, Cuba); holotype Museo Poey 17249, paratypes MCZ 129129, 188542. moseleyae Clench, Hojeda 1938, Mem. Soc. Cubana Hist. Nat., 12: 324, pl. 24, figs. 4, 5 (Eight Mile Rock, Grand Bahama Island, Bahamas); holotype MCZ 116738. mossi Clench, Cerion 1952, Rev. Soc. Malacél. “Car- los de la Torre,” 8: 108 (new name for Cerion pau- cicostatum Clench 1934, non Torre 1929). moussoni Clench, Melanoides tuberculata 1927, Nau- tilus, 40: 101 (new name for Melania tuberculata plicifera Mousson 1849, non Melania plicifera Lea 1839). mukongo Bequaert & Clench, Subulina (Nothapalus) paucipira 1941, Bull. Mus. Comp. Zool., 88: 11, pl. 2, figs. 6-8 (Luadi-Soyo, near Neatadi, Belgian Congo); holotype MCZ 112302. muraokai Turner, Xylophaga 2002, Bull. Mus. Comp. Zool. Asm(4) 247 pls. 184 AS) (GO AA NaI20- 45'W, about 81 mi. SW of Port Hueneme, Califor- nia, or about 25 mi. S of San Miguel Island, Santa Barbara Islands, 1,720 m); holotype MCZ 316746, paratypes MCZ 316747—50. Murexiella (subgenus) Clench & Perez Farfante 1945, Johnsonia, (17): 49 (type species, Murex hi- dalgoi Crosse 1896, original designation). murilloi Clench, Musculium 1939, Mem. Soc. Cu- bana Hist. Nat., 13(5): 286, pl. 36, fig. 1 (Soacha, Dept. Cundinamarea, Columbia); holotype MCZ 91374, paratypes MCZ 91373; paratypes MCZ 91375 from Fugene, 82 km N of Bogota, Dept. Cundinamarea, 2,430 m. naceli Turner, Xyloredo 1972, Breviora, (397): 9, pl. 6 (U.S. Naval Civil Eng. Lab STU 1-4, 33°46’N, 120°46'W, about 30 mi. S of San Miguel Island, off Port Hueneme, California, 2,072.6 m); holotype MCZ 279638, paratypes MCZ 279639. nairi Turner & Santhakumaran, Martesia (Particoma) 1989, Ophelia, 30(3): 163, figs. 7-10 (Neendakara, stamudi estuary, 7 km N of Quilon, Keraka, India); holotype MCZ 298201, paratypes MCZ 297202. najazaensis Cl & Jacobson, Troschelviana pfeifferiana 1971, Bull. Mus. Comp. Zool., 141(7): 440, pl. 4, fig. 8 (El Cacaotal, Najaza, Camagiiey, Cuba); ho- lotype MCZ 92091, paratypes MCZ 127517, Ooo Mere najazensis Clench & Aguayo Caracolus 1951, Mem. Soc. Cubana Hist. Nat., 20: 69, pl. 41, figs. 9-11 (El Cacaotal, Najaza, Camagiiey, Cuba); holotype Museo Poey 17304, paratypes MCZ 181501. nantahala Clench & Banks, Polygyra (Triodopsis) 1932, Nautilus, 46: 17, pl. 2, figs. 1, 2, 5 (35°38'N, 83°39'W, Blowing Springs, Cliff Ridge, Nautahala Gorge, Swain Co., North Carolina); holotype MCZ 86429. nanus Clench & Aguayo, Cadulus (Gadila) iota 1939, Mem. Soc. Cubana Hist. Nat., 13: 197 (not fig.) (Gibara, Cuba, 40 fth.); holotype MCZ 89222. naufraga Clench, Leptinaria Abaconia 1938, Mem. Soc. Cubana Hist., 12: 321, pl. 24, figs. 1, 2 (Sand Bank, Crossing Bay, Gt. Abaco Island, Bahamas); holotype MCZ 116705. neptunia Clench & Aguayo, Aurinia 1940, Mem. Soc. Cubana Hist. Nat., 14: 90, pl. 16, fig. 5 (Blake Sta. no. 8, 17°45'N, 77°58'40’W, off Banner Reef, Pe- dro Bank, 75 mi. S of Jamaica, 322 fth.); holotype MCZ 119025. Nesopoma (genus) Clench, 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2: 169 (type species, Nesopoma eyerdami Clench 1958, original designation). MOLLUSCA OF CLENCH AND TURNER ® Johnson 19 neztalia Turner & McKoy, Bankia 1979, J. R. Soc. N. Z., 9(4): 465, figs. 2-9, 17-31 (Wellington Harbor, New Zealand); holotype MCZ 280345. nioba Clench & McLean, Macrocallista (Paradione) 1936, J. Conch., 20: 201 (new name for Venus cos- tata Chemnitz 1795, non Gmelin 1791). nobilis Clench & Aguayo, eee fasciatus 1932, Nautilus, 45(3): 98, pl. 6, fig. 9 (Cayo Juan Tomas, mouth of Cabanas Bay, Pinar del Rio, Cuba); ho- lotype MCZ 47904. nodali Clench & Aguayo, Cerion disforme 1953 [in|] Aguayo, Mem. Soc. Cubana Hist. Nat., 21: 310, pl. 35, fig. 6 (El Cafionde Banes, cerca del Faro, Or- iente, Cuba); holotype Museo Poey 121404, para- types MCZ (not received). nooi Turner Xyloredo 1972, Breviora, (397): 5, pls. 1, 2 (250 54'N, 77°49'W, from test panel, Tongue of the Ocean, off Andros Island, Bahamas, 1,737 m); holotype MCZ 279631, paratypes MCZ 279832-—5. norvegicum Clench & Turner, Epitonium greenlan- dicum 1952, Johnsonia, 2(31): 323, pl. 155, fig. 1 (new name for Scalaria groenlandica ornata Friele & Grieg 1901, non S. ornata Baily 1865). Nystiella (genus) Clench & Turner, 1952, Johnsonia, 2(31): 337 (type species, Epitonium opalinum Dall 1927, original designation). Nystiellinae (subfamily) Clench & Turner, 1952, Johnsonia, 2(31): 337 (type species, Epitonium opalinum Dall 1927, original designation). obesum “Torre’ Clench & Aguayo, Cerion 1951, Rev. Soc. Malacol. “Carlos de la Torre,” 8: 80 ([nomen nudem|] listed as a synonym under Cerion pauci- costatum Torre). occidentale Clench & Turner, Cymatium (Septa) ru- beculum 1957, Johnsonia, 3: 214, pl. 110, fig. 3; pl. 113, fig. 5; pl. 121, figs. 1-3 (St. Thomas, Virgin Islands); holotype ANSP 36874, paratype MCZ 202269 from Carboneras, Matanzas, Cuba. occidentalis Clench & Archer, Cyclophorus fernan- dezi 1932, Occas. Pap. Boston Soc. Nat. Hist., 8: 37, pl. 4, fig. B (Calominatao, Mamburao, Mindoro, Philippines Islands); holotype MCZ 92797, para- types MCZ 93841. Odontocymbiola Clench & Turner, 1964, Johnsonia, 4: 170 (new name for Adelomelon “Dall Pilsbry & Olsson 1954, non Dall 1906). Odontocymbiolinae Clench & Turner, 1964, Johnson- ia, 4: 170 (new name for Adelomeloninae based upon Adelomelon Pilsbry & Olsson 1954, non Dall 1906). ophir Clench, Placostylus (Malaitella) 1941, Am. Mus. Novit., (1129): 11, fig. 9 (20 mi. inland from Swu, Malaita Island, Solomon Islands); holotype AMNH 79005, paratypes MCZ 92902, 92904 from Aurola, Malaita Island. oregon Clench & Turner, Calliostoma (Kombologion) 1960, Johnsonia, 4: 42, pl. 2; pl. 6, fig. 2; pl. 27 (Oregon Sta. 5501, 26°55'N, 96°25’W, about 70 mi. SE of Corpus Christi, Texas, 125 fth.); holotype USNM 612705, paratype MCZ 235480. organensis Clench, Liguus flammellus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 124, pl. 6, fig. 8 (Eusenada del Valle, Vinales, Pinar del Rio, Cuba): holotype MCZ 80958. orientale Clench & Aguayo, Cerion 1951, Rev. Soc. Malacél. “Carlos de la Torre,” 8: 79, pl. 11, fig. 16 (Punta Nigra, 18 km SW of Punta Maisi, Oriente, Cuba); holotype Museo Poey 12534, paratypes MCZ 188582. orites Cooke & Clench, Electrina succinea 1943, Oc- cas. Pap. B. P. Bishop Mus., 17(20): 252, fig. 2 (eastern Ridge of Mt. Perahu, Rapa Island, Central Pacific, 1,300—1,550 ft.); holotype Bishop Museum 10111, paratypes MCZ 144791. orotis Clench & Archer, Helicostyla mindoroensis (Chrysallis) 1933, Pap. Mich. Acad. Sci. Arts Lett., 17: 549, pl. 58, fig. 1 (near Calavite Mt., Binuan- gan, Palian, Mindoro, Philippines): holotype MCZ 81360. orrae Turner, Taheitia 1959, Occas. Pap. Mollusks, 2(23): 186, pls. 31, 32 (sink-hole cave behind air stp. Biak, Dutch New Guinea); holotype ANSP 223656, paratypes MCZ 221170. osmenti Clench, Drymaeus multilineatus 1948, Nau- tilus, 62: 36 (Little Pine Key, Lower Florida Keys): holotype MCZ 160807. osmenti Clench, Liguus fasciatus 1942, Proc. N. Engl. Zool. Club, 19: 69 (Howes Key, Lower F lor- ida Keys); holotype MCZ 137792. ottenwalderi Wetherbee & Clench, Macroceramus 1984, Caribb. J. Sci., 20(1-2): 11, fig. 1, pl. 1, pt. C, right side (Mt. Isabel de Torres, Puerto Plata Prov., Republica Dominicana); holotype MCZ 57490 (rightmost figure), paratypes MCZ 57491. pallidior Bequaert & Clench, Pseudogibbula duponti 1941, Bull. Mus. Comp. Zool., 88(1): 7 [not fig.] (Matadi, Congo River, Belgian Congo); holotype MCZ 112265. palmeri Clench & Rehder, Humboldtiana 1930, Nau- tilus, 44: 12, pl. 2, figs. 1-4 (head of Madera Can- yon, 7,350 ft., Mt. Livermore, Davis Mts., Jeff Da- vis Co., Texas); holotype MCZ 79779. pandion Clench & Turner, Epitonium (Boreoscala) 1952, Johnsonia, 2: 326, pl. 157 (new name for Acirsa gracilis Verrill 1880, non Scalaria gracilis Sowerby, 1844). Papuanella (genus) Clench & Turner, 1959, J. Mala- col. Soc. Aust., (3): 5 (type species Geotrochus ogeramuensis Kobelt 1914, original designation). parallaxis Clench & Archer, Helicostylus mindoroen- sis (Chrysallis) 1933, Pap. Mich. Acad. Sci. Arts Lett., 17: 550, pl. 518, fig. 2 (Paléan, Mindoro, Philippines); holotype MCZ 83275. Paramia (genus) Clench 1949, Bull. B. P. Bishop Mus., (196): 25 (type species, Cyclostoma incisa Hombron & Jacquinot 1854, original designation). Paramiella Clench, 1954, Nautilus, 67: 139 (new name for Paramia Clench 1949, non Bleeker 1863). paravicinii Clench, Partula 1933, Nautilus, 47: 24, pl. 20 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 3, figs. 5, 6 (St. Matthias, Bismarck Archipelago); holotype MCZ 59854. parvulina Clench & Turner, Geomelania (Merrilli- ana) 1948, Occas. Pap. Mollusks, 1(13): 180 (new name for Geomelania parva Chitty 1853, non par- va C. B. Adams 1850). pattersoni Clench, Oliva reticularis 1945, Mollusca, 1(4): 49, figs. 1, 2 on pl. (5 mi. E of Crabbing Point, Grand Bahama Island, Bahamas); holotype MCZ 151166. paucicostatum Clench, Cerion (Strophiops) 1934, Proc. Boston Soc. Nat. Hist., 40(2): 209, pl. 2, fig. E (Millers Hill, southern Eleuthera, Bahama Is- lands); holotype MCZ 103033. paucisculptum Clench & Aguayo, Cerion (Umbonis) 1952, Occas. Pap. Mollusks, 1(17): 425, pl. 54, figs. 1-2 (Punta de Musica, Bahia de Sama, Banes, Cuba); holotype MCZ 192204. peracutum Clench & Aguayo, Cerion sagraianum 1951, Rev. Soc. Malacél. “Carlos de la Torre,” 8: 75, pl. 11, fig. 9 (Boca de Jaruco, Habana Prov., Cuba); holotype Museo Poey 17268, paratype MCZ 187285, 188533. periculosum Clench, Cerion (Strophiops) 1934, Proc. Boston Soc. Nat. Hist., 40(2): 215, pl. 2, fig. B (South Cay, Mira Por Vos Group, Bahama Islands); holotype MCZ 103019. perryae Clench, Conus 1942, Johnsonia, 1(6): 31, pl. 15, fig. 5 (6 mi. off Sanibel Island, Lee Co., Flor- ida); holotype MCZ 206055. pescalia Clench & Turner, Odontocymbiola 1964, Johnsonia, 4: 172, pls. 110, 111 (about 160 mi. ENE of Mar del Plata, Argentina, on a mud bot- tom); holotype MCZ 233795. pihapilia Clench, Ponapella 1946, Occas. Pap. B. P. Bishop Mus., 18: 200, text fig. 1 (Mt. Tolotom, Ponape, Caroline Islands); holotype Bishop Muse- um 189003, paratypes & MCZ 140577. pilikia Clench, Kubaryia 1948, Occas. Pap. B. P. Bishop Mus., 19(8): 191, figs. la-d (Komakau, Ko- ror Island, Palau Islands, Caroline Islands); holo- type Bishop Museum 158887 (only specimen). pilsbryi Clench, Goodrichia trochiformis 1929, Bull. Mus. Comp. Zool., 69: 119, pl. 1, figs. 5, 6 (Sak- bayeme, Cameroon); holotype MCZ 78073, para- types MCZ 78067. pilsbryi Bequaert & Clench, Gubella 1936, Nautilus, 49: 95, pl. 6, figs. 5-7 (Liberia); holotype ANSP 23891, paratype MCZ 106384. pilsbryi Clench, Liguus blainianus 1935, Nautilus, 48: 123, pl. 7, fig. 1 (Campo de Tiro, 7 km N of Fierro, Pinar del Rio, Cuba, near Santa Cruz de los Pinos); holotype MCZ 94212. pilsbryi Clench, Macroceramus 1935, Proc. Boston Soc. Nat. Hist., 41: 11, pl. 1, fig. J (Camp Perrin, Haiti); holotype MCZ 108605. pilsbryi Clench, Spiroceramus (Spiroceramus) 1967, Breviora Mus. Comp. Zool., (26): 4, pl. 1 (Cerro de Tuabagiiey, Sierra de Cubitas, Camagiiey Prov., Cuba); holotype MCZ 188845, paratypes MCZ 188847, 256082. pinarensis Clench, Liguus fasciatus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 115, pl. 6, fig. 9 (Punta del Este, Isle of Pines, Cuba); holotype MCZ 58806. piratica Clench & Aguayo, Bathyaurinia 1940, Mem. Soc. Cubana Hist. Nat., 14: 93, pl. 15, fig. 2 (At- lantis Sta. 2982A, 22°48'N, 78°50’W, off Punta Ale- gre, Camagiiey Prov., Cuba, 210 fth.); holotype MCZM35235: piratica Clench & Turner, Truncatella scalaris 1948, Johnsonia, 2: 161, pl. 72, figs. 14 (St. George's Causeway, Bermuda); holotype MCZ 178985. piraticus Clench, Cerion (Strophiops) 1937, Proc. N. Engl. Zool. Club, 16: 64, pl. 3, fig. 1 (Southeast Point, 12 mi. SE of Abrahams Bay, Mariguana Is- land, Bahamas); holotype MCZ 57926. piraticus Clench, Conus verrucosus 1942, Johnsonia, (6): 14, pl. 11, fig. 1 (off Palm Beach, Florida); ho- lotype McGinty collection, now in Florida Museum of Natural History (UF) 174328, paratype MCZ 145458, from Carysfort Reef, Florida, 10 fth. platei Clench, Cerion (Strophiops) 1933, Proc. N. Engl. Zool. Club, 13: 90, pl. 1, figs. 7, 8 (Bight road, Cat Island, Bahamas); holotype MCZ 95731. plena Bequaert & Clench, Hydrobia 1936, Mem. Mus. R. Hist. Nat. Belg., 2nd ser., (3): 165, pl. 1, fig. 5 (Ango-Ango, 4 km S of Congo River, Belgian Congo); holotype MCZ 59802 (only specimen). plena Clench, Physa 1930, Occas. Pap. Boston Soc. Nat. Hist., 5: 311, text fig. 3 (Reed Spring, Cen- terville, Reynolds Co., Missouri); holotype MCZ 51850, paratypes MCZ 51851. Plesiophysinae (subfamily) Bequaert & Clench, 1939, J. Conch., 21: 175 (type genus Plesiophysa P. Fi- scher 1883). Plumulella (subgenus) Clench & Turner, 1946, John- sonia, 2(19): 22 (type species Teredo fimbriata Jef- freys 1860 [=Bankia fimbriatula Moll and Roch 1931], original designation). polinigi Clench, Eustomopsis (Verdichloritis) 1957, Breviora, (76): 1, fig. 1 (Western New Guinea); ho- lotype MCZ 212319. Ponapella (genus) Clench, 1946, Occas. Pap. B. P. Bishop Mus., 18(13): 199 (type species, Ponapella pihapiha Clench 1946, original designation). ponapensis Clench, Thaanumella cookei 1946, Occas. Pap. B. P. Bishop Mus., 18(13): 205 (Mt. Tolotom, Ponape, Caroline Islands, 800 ft.); holotype Bishop Museum 154096 (only specimen). popondetta Clench & Turner, Meliobba 1963, J. Ma- lacol. Soc. Aust., (6): 26, pl. 2, fig. 1, text figs. 3(3), 6(3), 8 (Mamoo Estate, Near Popondetta, about 12 mi. S of Buna, Papua, New Guinea); holotype Aus- tralian Museum C 62376, paratype MCZ 221417. porphyrotoma Clench & Archer, Strophocheilus (Strophocheilus) 1930, Nautilus, 43(3): 75, text figs. 1-3 (mountain near Jacquaral, Sao Paulo, Brazil); holotype MCZ 79108. pourtalesi Clench & Aguayo, Solariella 1939, Mem. Soc. Cubana Hist. Nat., 13: 190, pl. 28, fig. 2 (At- MOLLUSCA OF CLENCH AND TURNER * Johnson 21 lantis Sta. 2993, 23°24'N, 80°44’W, off Cardenas, N Cuba, 580 fth.); holotype MCZ 135108. prashadi Clench, Camptoceros (Culmenella) 1931, Nautilus, 44(3): 80 (new name for Bulimus [Cul- menella| hirasei Clench 1927, non Camptocerus hirasei Walker 1919). prasinata Jacobson & Clench, Helicina 1971, Nauti- lus, 84(3): 105, text fig. 1 (Monteado, Nuevo 20 km SE of Polo, Barahona Proy., Dominican Republic); holotype MCZ 187927. Proaspastus (subgenus) Clench, 1941, Am. Mus. Novit., (1129): 12 (type species, Bulimus sanchris- tovalensis Cox 1870, original designation). profunda Turner, Xylophaga 2002, Bull. Mus. Comp. Zool., 157(4): 243, pls. 12, 13 (25°54'N, 77°49’'W, Tongue of the Ocean, off NE tip of Andros Island, Bahama Islands, 1,722 m); holotype MCZ 316751. pterinus Clench & Archer, Thersites 1938, J. Conch., 21: 21, pl. 1, fig. 3 (Lake Barrine, 25 mi. SW of Cairns, Queensland); holotype Queensland Muse- um, South Brisbane, Queensland, QMMO 1769, paratype UMMZ from Lake Eacham, close to Lake Barraine. Pupina (genus) Clench, 1949, Bull. B. P. Bishop Mus., (196): 37 (type species, Pupina aurea Hinds 1842, original designation). Pupinesia (genus) Clench, 1949, Bull. B. P. Bishop Mus., (196): 39 (type species, Pupina adamsiana Crosse 1871, original designation). quillingi Turner, Jouanuetia (Pholadopsis) 1955, Johnsonia, 3(34): 139, pl. 84 (N end of Lake Worth Inlet, Lake Worth, Florida); holotype MCZ 189913. quinonesi Clench & Aguayo, Alcadia 1950, Rev. Soc. Malacol. “Carlos de la Torre,” 7: 63, pl. 12, figs. 9— 11 (Los Tambores, entre Playa Puerto Rico y Punta de Mulas, Banes, Oriente, Cuba); holotype Museo Poey 12913, paratypes MCZ 185800. quinonesi Clench & Aguayo, Cepolis (Coryda) alauda 1957 [in] Aguayo, C. G., and M. L. Jaume, Mem. Soc. Cubana Hist. Nat. “Felipe Poey,” 23: 130, pl. 4, figs. 4-6 (coast between Morales and el Cafion, Banes, Oriente, Cuba); holotype Museo Poey 15097, paratypes MCZ 212977. Quirosella (genus) Clench, 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2(27): 175 (type species, Quiro- sella coultasi Clench 1958, original designation). quirosi Clench, Ompahlotropis 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2(27): 171, pl. 17, fig. 3 (Lavang- gu, Rennell, Solomon Islands); holotype University Zoological Museum, Copenhagen (not numbered teste O. S. Tendal, personal communication); para- types MCZ 191443. randi Clench, Zaphysema 1962, Breviora, (173): 4, pl. 1, fig. 1 (Cueva de San Francisco, Cerros de San Francisco, Mun. Pedro Santana, San Rafael, Reptblica Dominicana); holotype MCZ 230503, paratype MCZ 320504. Rapanella (genus) Cooke & Clench, 1943, Occas. Pap. B. P. Bishop Mus., 17(20): 253 (type species, Rapanella andersoni Cooke & Clench, monotypy). rehderi Clench & Aguayo, Cerion (Umbonis) 1943, Occas. Pap. Mollusks, 1(17): 422, pl. 57, fig. 2 (Jackline, 1 mi. W of Conch Shell Point, Gt. Ina- gua, Bahama Islands); holotype MCZ 189032. Rehderia (genus) Clench, 1946, Johnsonia, 2(27): 45 (type species Aurinia schmitti Bartsch 1931, origi- nal designation). remingtoni Clench, Physa 1930, Occas. Pap. Boston Soc. Nat. Hist., 5: 312, text fig. 4 (Round Spring, 12 mi. N of Eminence, Shannon Co., Missouri); holotype UMMZ 32663, paratypes MCZ 50984. rennellensis Clench, Crystallopsis (Cristovala) 1958, Natural History of Rennell Island, British Solomon Islands, Copenhagen, Denmark, 2: 190, pl. 16, fig. 2, text figs. 2, 3 (Te-avamangga, Rennell Island, Solomon Islands); holotype University Zoological Museum, Copenhagen (not numbered, teste O. S. Tendal, personal communication); paratype MCZ 191379, paratype MCZ 191436 from Tenggano, paratype MCZ from Niupani; all Rennell Island. Rennellia (section) Clench, 1941, Am. Mus. Novit., (1129): 20 (type species, Partula cramptoni Clench 1941, original designation). renschi Clench, Eustomopsis 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2(27): 198, pl. 19, figs. 9, 10 (Tu- lagi, Florida Group, Solomon Islands); holotype AMNH 66463, paratype MCZ 92957. rheophila Bequaert & Clench, Hydrobia 1941, Bull. Mus. Comp. Zool., 88: 8, pl. 2, fig. 4 (estuary of Congo River at Kala-Kala, near Matadi, Belgian Congo); holotype MCZ 112310. Rhysema (subgenus) Clench & Turner, 1957, John- sonia, 3: 236 (type species, Triton clathratum La- marck 1816, original designation). ricardi Clench & Aguayo, Cerion 1951, Rev. Soc. Ma- lacol. “Carlos de la Torre,” 8: 71, pl. 11, fig. 2 (Pun- ta de Tararé, Habana Prov., Cuba); holotype Museo Poey 12669, paratypes MCZ 129421, 129402, 188591. richardsi Clench, Hargravesia 1949, Bull. B. P. Bish- op Mus., (196): 47, fig. 27 (N coast of Guadalcanal Island, Solomon Islands); holotype MCZ 161591. riosi Clench & Turner, Adelomelon (Weaveria) 1964, Johnsonia, 4: 162, pls. 83, 101, 102 (about 130 mi. E of Mar del Plata, Argentina, 99 fth.); holotype MCZ 245017. riveroi Clench, Geomelania (Merrilliana) 1962, Bre- viora, (173): 3, pl. 1, fig. 4 (Colonia Ramfis, 20 km W of San Cristobal, Reptiblica Dominicana); ho- lotype MCZ 230505. riveroi Turner, Hemitrochus 1958, Occas. Pap. Mol- lusks, 2(22): 174-177, pl. 24, figs. 5, 6; pl. 27, figs. 1, 2; pl. 29; pl. 30, figs. 3-6 (Cerro Dofia Juana, Toro Negro National Forest, Puerto Rico); holo- type MCZ 216144. robertsoni Clench, Microceramus (Spiroceramus) 1963, Bull. Mus. Comp. Zool., 128: 406, pl. 3, fig. i) bo 1 (Delectable Bay, Settlement, Acklins Island, Ba- hamas); holotype MCZ 225313. rocai Clench & Aguayo, Cerion tridentatum 1953, Torreira, (18): 2, text fig. 2 (Lagunas Salobres de Boca de Guanabo, Habana, Cuba); holotype Museo Poey 13152, paratypes MCZ 78055, 80729, 129418, 129420. roseatus Clench, Drymaeus bahamensis 1933, Proc. N. Engl. Zool. Club, 13: 87, pl. 1, figs. 12, 13 (Gt. Inagua Island, Bahamas); holotype MCZ 90104. rosewateri Clench & Turner, Calliostoma (Kombolo- gion) 1960, Johnsonia, 4: 41, pl. 6, fig. 3, pl. 10, fig. 2, pl. 26 (Oregon Sta. 1985, 09°45'N, 59°47’W, about 85 mi. SE of Galeota Point, Trinidad, 200 fth.); holotype USNM 612704; paratypes MCZ 235483, 235484. russelli Clench, Cerion 1938, Bull. Mus. Comp. Zool., 80(14): 528, pl. 1, figs. 5-8 (near Turtle Cove, 4 mi. NNE of The Bight, Central Cat Island, Bahama Islands); holotype MCZ 107534. russelli Clench, Chondropoma 1937, Proc. N. Engl. Zool. Club, 16: 67, pl. 3, fig. 3 (Gt. Inagua Island, Bahamas); holotype MCZ 90132. russelli Clench, Liguus flammellus 1935, Nautlius, 48(4): 124, pl. 7, fig. 2 (Mogote Mina Constancia, Vinales, Pinar del Rio, Cuba); holotype MCZ 82980. russelli Clench, Microceramus 1937, Proc. N. Engl. Zool. Club, 16: 63, pl. 3, figs. 4, 5 (Blue Hole Hill, 2 mi. NNE of Orange Creek, N Cat Island, Ba- hamas); holotype MCZ 127296. russelli Clench, Polydontes (Salleana) 1962, Rev. Mu. Argent. Cienc. Nat. “Bernardino Rivadavia,” 8: 218, pl. 1, fig. 3 (N of La Rucilla [Pico Trujillo], Cordillera Central, Républica Dominicana, 5,800 ft.): holotype MCZ 118212. salina Clench, Drymaeus bahamensis 1959, Bull. Mus. Comp. Zool., 121(2): 43 (new name for Dry- maeus bahamensis roseatus Clench 1933, non Bu- limus |Drymaeus| roseatus Reeve 1848). salina Clench, Physa 1930, Occas. Pap. Boston Soc. Nat. Hist., 5: 309, text figs. 1, 2 (Brackish Spring, Skeen’s Ranch, Promontory, Box Elder Co., Utah); holotype MCZ 79387, paratypes MCZ 4239. salina Clench, Urocoptis bencomoi 1935, Proc. Bos- ton Soc. Nat. Hist., 41(1): 4, pl. 2, fig. A (Saline Madame Doisy’, Mahautier, SE end of Gonase Is- land, Haiti); holotype MCZ 106243. Salleana (subgenus) Clench, 1962, Rev. Mus. Argent. Cienc Nat. “Bernardino Rivadavia,” 8: 218 (type species, Polydontes (Salleana) russelli Clench 1962, original designation). samana Clench, Brachypodella (Gyraxis) 1966, Bre- viora, (245): 11, pl. 2, fig. 5 (Pefion de Maria Luisa, Santa Bartara de Samana, Républica Dominicana); holotype MCZ 57214. samana Clench, Coryda 1964, Rev. Mus. Argent. Cienc. Nat. “Bernardino Rivadavia,” 8: 225, pl. 1, fig. | (Bahfa de San Lorenzo, Bahia de Samana, Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 Prov. de Samana, Republica Dominicana); holotype MCZ 188240. samba Cl, Strombus 1937, Proc. N. Engl. Zool. Club, 16: 18, pl. 1, fig. 1 (Wood Cay, West End, Grand Bahamas Island, Bahamas); holotype MCZ 116054. sana Clench & Archer, Polygyra (Triodopsis) 1933, Nautilus, 46(3): 88, pl. 7, figs. 4-6 (34°45’/N, 86°31'W, Big Cove, Monte Sano, Huntsville, Mad- ison Co., Alabama); holotype MCZ 95089. sanagaensis Clench, Lauistes (Lauistes) 1929, Bull. Mus. Comp. Zool., 69(6): 120, pl. 1, figs. 7, 8 (San- aga River, near Sakbayeme, Cameroon); holotype MCZ 78064, paratype MCZ 78065. sanchezi Clench & Aguayo, Cerion 1953, Torreia, (18): 3, figs. 4, 5 (Lengua de Pajaro, Bahia de Le- biza, Mayari, Oriente, Cuba); holotype Mus. Poey 3999, paratypes MCZ 189458. santomennoi Wetherbee & Clench, Macroceramus 1984, Caribb. J. Sci., 20(1-2): 11, fig. 1, ph. 1, pt. A, left side (1 mi. S of Copey, Monti Cristi Prov., Reptiblica Dominicana); holotype MCZ 57346 (leftmost figure), paratypes MCZ 57342, 288374. savati Clench, Ostodes 1949, Bull. B. P. Bishop Mus., (196): 14, figs. 5, 6 (Salailua, Savaii Island, Samoan Islands); holotype Bishop Museum 9710, paratypes MCZ 140501. schepmani Turner, Oocorys 1948, Johnsonia, 2(26): 190 (new name for O. elongata Schepman 1909, non Locard 1897). schevilli Clench & Archer, Thersites 1938, J. Conch., 21: 23, pl. 1, fig. 1 (foot of Mt. Walker, S of Hugh- enden, Queensland); holotype MCZ 99055, para- types Queensland Museum, South Brisbane, Queensland (QMMO) 1968. schoutedeni Bequaert & Cl, Lobogenes 1936, Mem. Mus. R. Hist. Nat. Belg., 2nd ser., (3): 164, pl. 1, figs. 1, 2 (Ango-Ango, 4 km S of Matadi, Congo River, Belgian Congo); holotype MCZ 59805. schroederi Clench & Aguayo, Calliostoma (Callios- toma) 1938, Mem. Soc. Cubana Hist. Nat., 12: 377, pl. 2S) fig. 3 (Atlantis Sta. 2981, 22°48'N, 78°48'W, off northern Cuba, 225 fth.); holotype MCZ 135002. schwabi Clench, Egeria 1929, Bull. Mus. Comp. Zool., 69: 122, pl. 1, figs. 1, 2 (Lobetal, Lower San- aga River, Cameroon); holotype MCZ 78072. scotti Clench, Plagioptycha 1963, Bull. Mus. Comp. Zool., 128(8): 411, pl. 3, fig. 3 (2 mi. S of Binnacle Hill Settlement, Acklins Island, Bahamas); holo- type MCZ 225311. Septariellina (genus) Bequaert & Clench, 1936, Mem. Mus. R. Hist. Nat. Belg., 2nd ser., (3): 163, fig. 1 (type species, Septariellina congolensis Be- quaert & Clench 1936, original designation). Setaepoma (genus) Clench, 1955, Nautilus, 68(4): 134 (type species, Japonia hedigeri 1. and B. Rensch 1936, original designation). shattucki Bequaert & Clench, Drymaeus 1931, Oc- cas. Pap. Boston Soc. Nat. Hist., 5: 424 (Chichen Itza, Yucatan); holotype MCZ 79396, paratypes MCZ 79409. MOLLUSCA OF CLENCH AND TURNER * Johnson 23 shrevei Clench & Aguayo, Cerion (Umbonis) 1952, Occas. Pap. Mollusks, 1(17): 436, pl. 57, fig. 4 (near North West Point, Little Inagua Island, Bahama Islands); holotype MCZ 192287. sisal Clench & Aguayo, Cerion (Umbonis) 1952, Oc- cas. Pap. Mollusks, 1(17): 427, pl. 57, fig. 3 (E. side, Boca de Mosquito, Mariel, Pinar del Rio, Cuba): holotype MCZ 181992. soluta Clench & Turner, Taheitia 1948, Occas. Pap. Mollusks, 1(13): 190, pl. 23, fig. 4; pl. 24, figs. 5, 6 (Bavatu, Varna Mbalavu, Lau Group, Fiji Islands); holotype Bishop Museum 179895, paratypes MCZ Sia spelunca Turner & Clench, Physa 1974, Nautilus, 88: 82, figs. 1-19 (cave stream, about 800-900 ft. from the entrance, in Lower Kane Cave, near Kane, about 12 mi. E of Lovell, E side of the Big Horn River, Big Horn Co., Wyoming, 3,000 ft.); holotype MCZ 280016. spinella Clench, Cassis madagascariensis 1944, John- sonia, 1(16): 15, pl. 8 (Tortugas, Florida); holotype MCZ 140761. springeri Clench & Turner, Calliostoma 1960, John- sonia, 4: 71, pl. 52, figs. 1, 2 (Oregon Sta. 1283, 29°6'N, 88°19’W, about 38 mi. E of Southeast Pass, Mississippi Delta, Louisiana, 260 fth.); holotype USNM 612703. stevensoni Clench, Foxidonta 1950, Rev. Soc. Mala- cél. “Carlos de la Torre,” 7(2): 59, text figs. 1-3 (Fiu, Northern Malaita Island, Solomon Islands); holotype MCZ 186268. strongi Bequaert & Clench, Afroditropis 1936, Rev. Zool. Bot. Afr., 29: 103, pl. 2, figs. 8, 9 (Makato, French Congo, on the right bank of the Congo Riv- ers, between Lukolcla and Irebu); holotype MCZ 77196. subflava Clench, Physa fontinalis 1926, J. Conch., 18: 12 (new name for Physfontinalis inflata Moquin- Tandon 1855, non Physa inflata Lea 1841 and Phy- sa inflata Adams & Angus 1864). Surinamia (subgenus) Clench, 1933, Nautilus, 47: 71 (type species, Asolene [Surinamia] fairchildi Clench 1933, original designation). tablasensis Clench & Archer, Helicina ignava 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 339 (Agca- latac, Tablas Island, Philippines); holotype MCZ 47235, paratype MCZ 21536 (only specimens men- tioned). Tamu (genus) Gustafson, Turner, Lutz, Vrijenhoek, 1998, Malacologia, 40(1—2): 90 (type species Tamu fisheri Gustafson, Turner, Lutz & Vrijenhoek, 1998, original designation). Tectininus (subgenus) Clench & Abbott, 1942, John- sonia, 1(4): 4 (type species, Echininus [Tectininus | nodulosus Pfeiffer 1834, original designation). tejedori Clench & Aguayo, Chondropoma (Parachon- drodes) 1946, Rev. Soc. Malacol. “Carlos de la Tor- re,” 4(2): 52-53, text figure (El Ocujal, Mayari, Or- iente, Cuba); holotype Museo Poey 3997, paratype MCZ 178197. Thaanumella (genus) Clench, 1946, Occas. Pap. B. P. Bishop Mus., 18(13): 201 (type species, Diadema carolinarum Méllendorff 1897, original designa- tion). Thaanumellinae (family) Clench, 1946, Occas. Pap. B. P. Bishop Mus., 18(13): 199 (type genus, Thaan- umella Clench 1946). thaanumi Clench & Turner, Truncatella (Truncatella) 1948, Occas. Pap. Mollusks, 1(13): 165, pl. 25, fig. 6 (Ulali Island, Truk Group, Caroline Islands); ho- lotype MCZ 159379. Thaisella (subgenus) Clench, 1947, Johnsonia, 2(23): 69 (type species, Purpura trinitatensis Guppy 1869, original designation). thermalis Bequaert & Clench, Achatina (Pintoa) 1936, Bull. Mus. R. Hist. Nat. Belg., 12(32): 1, figs. 1-10 (Katana, W shore of Lake Kiva, near hot min- eral springs of Kakonds, Belgian Congo); holotype Institut Royal des Sciences naturelles de Belgique, Bruxelles, Type no. 173, inventoried under I. G. 10.010; paratypes MRAC 201309, 201310; para- types MCZ 111711. thielei Bequaert & Clench, Callistoplepa 1934, Rev. Zool. Bot. Afr., 26: 115, pl. 2, figs. 8-10 (Utengule near Langenburg, Tanganyika Territory, Africa); holotype ZMB 53177a, fig. 8, paratypes ZMB 53177b (Kilias, 1992:179) (2), paratypes MCZ 98686. Thomeomaizenia (subgenus) Bequaert & Clench, 1936, Rev. Zool. Bot. Afr., 29(1): 99 (type species, Cyclophorus vandellii Nobre 1886, monotypy). tiburonense Clench & Turner Epitonium 1952, John- sonia, 2: 305, pl. 143 (Aquin Dept. du Sud, Haiti); holotype USNM 440128, paratype MCZ 187959. tiburonica Clench, Archegocoptis 1935, Proc. Boston Soc. Nat. Hist., 41: 6, pl. 1, fig. I (above Tardieu, Mt. La Hotte, Haiti, 3-4,000 ft.); holotype MCZ 108591. tipperi Turner, Xylophaga 2002, Bull. Mus. Comp. Zool. 157(4): 259, pl. 29 (26°04'N, 80°04'W, about 3.2 mi. off Fort Lauderdale, Florida, 152.4 m); ho- lotype MCZ 316736, paratype MCZ 316737. togdensis Bequaert & Clench, Achatina 1934, Rev. Zool. Bot. Afr., 26: 112, pl. 1, figs. 1-4 (Bismarck- burg, Togo, Africa); holotype ZMB 47176a, fig. 2, paratypes ZMB 4717b; (Kilias 1992:179) (2), para- types MCZ 98688. Tomlinella (subgenus) Clench & Turner, 1948, John- sonia, 2: 159 (type species, Truncatella scalaris Mi- chaud 1830, original designation). Tomlinitella Clench & Turner, 1948, Occas. Pap. Mollusks, 1(13): 169 (new name for Tomlinella Cl & Turner 1948, non Tomlinella Viader 1938). tongana Clench & Turner, Taheitia 1948, Occas. Pap. Mollusks, 1(13): 190, pl. 23, fig. 6 (0.5 mi. from Vaigana, Eua Island, Tonga Islands); holotype Bish- op Museum 87703, paratypes MCZ 157757. torrei Clench & Aguayo, Calliostoma (Calliostoma) 1940, Mem. Soc. Cubana Hist. Nat., 14(1): 79, pl. 14, fig. 5 (Atlantis Sta. 3985, 23°13'N, 81°22'W, off 24 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 Matanzas, Matanzas Prov., Cuba, 385 fth.); holo- type MCZ 135165. torrei Clench & Aguayo, Cepolis (Jeanneretia) 1933, Nautilus, 47: 22, pl. 3, figs. 7, 8 (cave in the Sierra Hato Nuevo, near Marti, Matanzas Prov., Cuba): holotype MCZ 59730. torrei Clench, Liguus fasciatus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 114, pl. 6, figs. 5, 6 (Pun- ta del Este, Isle of Pines, Cuba); holotype MCZ 58805. trachydermon Clench & Archer, Thersites 1938, J. Conch., 21: 21, pl. 1, fig. 5 (24 mi. NW of Card- well, Queensland); holotype MCZ 73284. trochiformis Clench, Goodrichia 1929, Bull. Mus. Comp. Zool., 69(6): 118, pl. 1, figs. 3, 4 (Man Riv- er, near Sakbayeme, Cameroon); holotype MCZ 78066, paratypes MCZ 78067, 78068, 78071. trujilloi Clench, Oliva 1938, Nautilus, 51(4): 111, ph. 9, figs. 3-5 (Puerto Plata, Santo Domingo, 5 fth.); holotype MCZ 57240. tuftsensis Turner, Chlamys (Zygochlamys) 1967, N. Z. J. Geol. Geophys., 10(2): 446-455, figs. 1-5 (approx. 77°31'S, 162°E, outwash gravel, Bull Pass, Wright Valley, McMurdo Sound, Antarctica, 1,000 ft. (305 m elev. [from early to middle Pleistocene ]); holotype MCZ 256085, paratype MCZ 256086. tumidula Clench & Aguayo, Eutrochatella (Microvi- ana) 1957 [in] Aguayo, C. G., and M. L. Jaume, Mem. Soc. Cubana Hist. Nat. “Felipe Poey,” 23: 119, pl. 1, fig. 6 (Cerro de los Portales, Camayén, Barrio de Bariay, Gibara, Oriente, Cuba); holotype Museo Poey 13260, paratypes MCZ 212978. turbinella Clench & Archer, Polygyra (Stenotrema) 1933, Nautilus, 46(3): 89, pl. 7, figs. 1-3 (34°39’N, 86°15'W, 2-4 mi. E of Woodville, Jackson Co., Al- abama); holotype MCZ 95140. turgidus Clench & Aguayo, Caracolus sagemon 1951, Mem. Soc. Cubana Hist. Nat., 20: 67, pl. 42, figs. 1—3 (Mina Caledonia, Mayari, Oriente, Cuba); ho- lotype Museo Poey 17205, paratype MCZ 187544. turnerae Clench & Aguayo, Cerion (Umbonis) 1952, Occas. Pap. Mollusks, 1(17): 423, pl. 53, figs. 4-7 (Lydia Point, Gt. Inagua Island, Bahama Islands); holotype MCZ 184623. turnerae Clench, Chondropoma 1951, J. Conchy., 90: 271, fig. 1 (Mona Island, Puerto Rico); holotype MCZ 171039. ultima Clench & Archer, Helicostyla cincinniformis 1931, Occas. Pap. Boston Soc. Nat. Hist., 5: 335, pl. 17, fig. 4 (Lubang Island, Philippines); holotype MCZ 79169, paratypes MCZ 79170. ulugurensis Bequaert & Clench, Edentulina 1936, J. Conch., 20: 213, text figs. 11, 12 (Vituri, Ulugura Mts., Tanganyika Territory, Africa, 2,500 ft.); holo- type MCZ 59926. unidentatus Clench, Nesonanina 1957, Am. Mus. Novit., (1863): 4, figs. 5, 7 (Rambutyo Island, Ad- miralty Islands); holotype AMNH 73488, paratypes MCZ 188138. Urocoptola (section) Clench, 1935, Proc. Boston Soc. Nat. Hist., 41(1): 3 (type species, Cylindrella gru- neri Dunker 1844, original designation). usambarensis Bequaert & Clench, Edentulina 1936, J. Conch., 20: 115, text fig. 13 (Lutindi, Usambara Mts., Tanganyika Territory, Africa, 4,000 ft.); holo- type MCZ 59919. utahensis Clench, Physa lordi 1925, Occas. Pap. Mus. Zool., Univ. Mich., (161): 8, pl. 1, fig. 5 (Utah Lake, 2 mi. S of Lehi, Utah); holotype UMMZ Walker collection 4257. The Walker number on the UMNZ lot does not correspond to this species. It is a Clausilia from France (teste Graf, personal communication). Since the holotype could not be located, the other type lots are regarded as syn- types, UMMZ 229549, MCZ 36541, 30982. utowana Clench, Cerion (Strophiops) 1933, Proc. N. Engl. Zool. Club, 13: 92, pl. 1, figs. 1, 2 (East Plana Key, Bahamas); holotype MCZ 101157. utowana Clench, Plagioptycha macrodon 1952, Rev. Soc. Malacél. “Carlos de la Torre,” 8: 114, pl. 15, figs. 4-6 (Bannermantown, Eleuthera Island, Ba- hamas); holotype MCZ 103111, paratypes MCZ 103111 (P duclusiana utowana Clench on labels). utowana Clench, Succinea 1933, Proc. N. Engl. Zool. Club, 13: 85, pl. 1, figs. 18, 19 (near Victoria Hill Settlement, Watling Island, Bahamas); holotype MCZ 101171. utowanae Clench, Brachypodella 1932, Proc. N. Engl. Zool. Club, 12: 104 (Beata Island, Santo Do- mingo); holotype MCZ 81492. Valvatorbis (genus) Bequaert & Clench, 1936, Mem. Mus. R. Hist. Nat. Belg., 2nd ser., (3): 166 (type species, Valvatorbis mauritii Bequaert & Clench 1936, original designation). vanattai Clench & Aguayo, Cerion 1951, Rev. Soc. Malacol “Carlos de la Torre,” 8: 78, pl. 11, fig. 12 (Playa Larga, Boca de Janco, Baracoa, Oriente, Cuba); holotype Museo Poey 12671, paratypes MCZ 66713, 128950, 188592. vanattai Clench, Spiroceramus (Spiroceramus ) 1967, Breviora, Mus. Comp. Zool., (260): 5, pl. 1, fig. 4 (Los Cangilones, Sierra de Cubitas, Camagiiey Prov., Cuba); holotype MCZ 256083, paratypes MCZ 131387, 131391, 188844. vanderrieti Clench, Kalendyma 1966, Nautilus, 80: 52, pl. 3, figs. 1, 2 (Ata’a, Malaita, Solomon Is- lands); holotype MCZ 260885. vanderrieti Clench, Trochomorpha 1965, Breviora, (224): 3, pl. 2, figs. 1, 2 (Ata’a, Malaita Island, Sol- omon Islands); holotype MCZ 251176, paratypes MCZ 251177, 247959. vanhyningi Clench & Smith, Dinocardium robustum 1944, Johnsonia, 1(13): 11, pl. 7 (Sanibel Island, Florida); holotype MCZ 151146. vegai Clench & Turner, Lyria (Lyria) 1967, Nautilus, 80(3): 83-84, fig. 1 (fish trap, Cabo Rojo, Prov. Pedernales, Républica Dominicana, Hispaniola); holotype MCZ 256494. verae ‘Clench’ Branham, Natica canrena 1954, Shells Along the Florida Shore, St. Petersburg, p. 26, fig. MOLLUSCA OF CLENCH AND TURNER * Johnson 251 (Marco Island, Florida [error for Naticarius verae Rehder 1947]}). Verdichloritis (subgenus) Clench, 1957, Breviora, (76): 1 (type species, Eastomopsis [Verdichloritis | polingi Clench 1957, original designation). vigoensis Clench & Archer, Chloraea thersites 1933, Pap. Mich Acad. Sci. Arts Lett., 17: 537, pl. 57, figs. 4, 5 (Monte Vigo, Lubang, Philippines); ho- lotype MCZ 81384. viridis Clench, Liguus fasciatus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 105, pl. 6, fig. 11 (La Caoba, near Dolores, Central Soledad, Cienfuegos, Santa Clara, Cuba); holotype MCZ 59507. voluminosa Clench & Banks, Polygyra (Strenotrema) 1932, Nautilus, 46: 16, pl. 2, figs. 6, 7 (35°38’N, 83°39'W, Blowing Springs, Cliff Ridge, Nantahala Gorge, Swain Co., North Carolina); holotype MCZ $2530. Wahgia (genus) Clench & Turner, 1959, J. Malacol. Soc. Aust., (3): 4 (type species Wahgia juliae Clench & Turner 1959, original designation). walkeri Clench, Liguus fasciatus 1933, Nautilus, 46: 91, pl. 7, figs. 7-9 (Hammock 9, Pinecrest Region, Cen- tral Everglades, Florida); holotype MCZ 79299. wallisi Clench & Cooke, Succinea 1945, Occas. Pap. B. P. Bishop Mus., 18(8): 133, fig. 1 (Papenoo Val- ley, Tahiti, Niu (Greater Tahiti), Society Islands, at 50-500 ft.); holotype Bishop Museum 87027, para- type MCZ 146394. wansoni Bequaert & Clench, Potadoma 1941, Bull. Mus. Comp. Zool., 88: 9, pl. 1, fig. 9, 11, 12 (es- tuary of the Congo River, Matadi, Belgian Congo): holotype MCZ 112268. warrenae Clench & Turner, Megalacron tabarensis 1964, J. Malacol. Soc. Aust., 8: 43, pl. 9, figs. 3, 4 (Boang Island, Tanga Group, off E coast of New Ireland, Bismarck Archipelago); holotype AMNH 111300, paratypes MCZ 181333, 181334. Weaveria (subgenus) Clench & Turner, 1964, John- sonia, 4: 162 (type species Adelomelon |Weaveria] riosi Clench & Turner 1946, original designation). whitneyi Clench, Taheitia 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2: 166, pl. 16, fig. 1 (Bambatana, Choiseul, Solomon Islands); holotype AMNH 73883, paratype MCZ 188089. whittumi Clench, Urocoptis (Gongylostoma) 1966, Breviora, (245): 4, pl. 2, fig. 1 (Guabairo, near So- ledad, Cienfuegos, Las Villas, Cuba); holotype MCZ 59580, paratypes MCZ 59284, 86289. whoi Turner, Xylophaga 2002, Bull. Mus. Comp. Zool., 157(4): 242, pls. 10, 11 (Atlantis Sta. 3471 23°21'N, 80°56’W., off Cardenas, Mantanzas Prov., Cuba, 500 fth.) holotype MCZ 275015. williamsi Clench & Archer, Papuina 1936, Nautilus, 49(3): 88, pl. 5, fig. 4 (Omarakana, Central Kiri- wina, Trobriand Islands); holotype MCZ 110396. williamsi Clench, Plagioptycha 1937, Proc. N. Engl. Zool. Club, 16: 61, pl. 3, fig. 7 (Abraham’s Hill, 2 bo Ul mi. NE of Abraham’s Bay, Mariguana Island, Ba- hamas); holotype MCZ 75042. wolffi Clench, Quirosella 1958, Natural History of Rennell Island, British Solomon Islands, Copen- hagen, Denmark, 2: 176, pl. 18, fig. 3 (Te-Ava- manggu, Rennell Island, Solomon Islands); holo- type University Zoological Museum, Copenhagen (not numbered, teste O. S. Tendal, personal com- munication), paratypes MCZ 191447, 191448. woodringi Clench & Pérez Farfante Murex (Murex) 1945, Johnsonia, (17): 9, pl. 4, figs. 1-3 (Jamaica): holotype MCZ 156087. Wrayanna (genus) Clench, 1948, Occas. Pap. B. P. Bishop Mus., 19(8): 192 (type species Diadema so- luta Méllendorff 1897, monotypy). xanthus Clench, Liguus fasciatus 1934, Occas. Pap. Boston Soc. Nat. Hist., 8: 113, pl. 6, fig. 12 (road between San Nicolas and Guines, Havana Prov., Cuba): holotype MCZ 47380. Xylopholas (genus) Turner, 1972, Basteria, 36(2—5): 99 (type species, Xylopholas altenia Turner, 1972. original designation). Xyloredo (genus) Turner, 1972, Breviora, (397): 3 (type species, Xyloredo nooi Turner, 1972, original designation). zairensis Bequaert & Clench, Lobogenes 1936, Mem. Mus. R. Hist. Nat. Belg., 2nd ser., (3): 165, pl. 1, fig. 4 (Kalawanga Island, 2 km N of Matadi, Congo River, Belgian Congo): holotype MCZ 93258. zanzibarica Bequaert & Clench, Maizania 1936, Rev. Zool. Bot. Afr., 29(1): 100, pl. 1, figs. 3-7 (Chuaca, E side of Zanzibar Island); holotype MCZ 72326. zenkeri Bequaert & Clench, Leptocala mollicella 1934, Rev. Zool. Bot. Afr., 26: 118, pl. 1, figs. 5-7, pl. 2, figs. 13 (Yaunde, Cameroon); holotype ZMB 47196a, figs. 6, 13; paratypes ZMB 47196b (Kilias 1992:174, 180) (2): paratype MCZ 98687. (2) Kilias, R. 1992. Die Typen und Typoide der Mol- lusken-Sammlung des Zoologischen Museums in Berlin (11). Il. Euthyneura, Stylommatophora, Sig- murethra: Achatinoidea, 2, Achatinidae. Mitteilun- gen aus dem Zoologischen Museum in Berlin, 68 Gi}: Ae7=t3o: Publications of William J. Clench Publication dates of The Nautilus, al- most always to the exact day of issue, were compiled by: Coan, E. V., and M. G. Har- asewych. 1993. The Nautilus, 106(4): 174— 180. 1923. Observations on the ovipositing of Tetraopes tetrophthalmus Frost. Papers Michigan Academy of Sciences, Arts and Letters, 3: 367. 1923. The use of sodium silicate as a mounting me- dium. Transactions American Microscopical Soci- ety, 42(1): 69-71 (C. W. Creaser, Ist author) [Jan- uary]. 26 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 1923. The marine shells of Sanibel, Florida. Nautilus, 37(2): 52-56 [11 October]. 1924. A new species of Physa from Texas. Nautilus, 38(1): 12-13, text fig. 4 [14 July]. 1924. Radular technique for Physa. Nautilus, 38(1): 13-14 [14 July]. 1924. The Physidae of the Au Sable River, Michigan. Papers Michigan Academy of Science, Arts and Letters, 5: 399-403. 1925. Additions to the list of marine shells from San- ibel, Florida. Nautilus, 38(3): 93-95 [19 January]. 1925. Vagabonding for shells. Nautilus, 38(4): 127— 143 (P. S. Remington, 2nd author) [2 May]. 1925. Notes on the genus Physa with descriptions of three new subspecies. Occasional Papers Michigan Museum of Zoology, University of Michigan, (161): 1-10, pl. 1 [27 May]. 1925. A possible manner of snake distribution. Cop- eia, (142): 40 [20 May]. 1925. A short list of land and freshwater mollusks from southeastern Texas. Nautilus, 39(1): 11-12 [8 July]. 1925. Snails eaten by shrews. Nautilus, 39(1): 28 [8 July]. 1925. Nahant beach shells. Nautilus, 39(1): 29 [8 July]. 1925. [Review] C. W. M. Poynter and Alan Moritz, The effects of ultraviolet light on pond snails. Nau- tilus, 39(1): 34-35 [8 July]. 1925. Description of a new species of Physa from the Pleistocene of Florida. Occasional Papers of the Museum of Zoology, University of Michigan, (164): 1-4, pl. 1, figs. 1-3 [3 August]. 1925. Collecting in Kentucky. Nautilus, 39(2): 71-72 [31 October]. 1926. Three new species of Physa. Occasional Papers of the Museum of Zoology, University of Michigan, (168): 1-8 pl. 1, figs. 1-4 [18 February]. 1926. A change of names of Physa and Isidora. Jour- nal of Conchology, 18(1): 12 [March]. 1926. Some notes and a list of shells from Rio, Ken- tucky. Nautilus, 40(1): 7-12 [12 July]. 1926. [Review] H. A. Pilsbry, A freshwater snail, Phy- sa zionis, living under unusual conditions. Nautilus, 40(1): 34 [12 July]. 1926. Some notes and a list of shells from Rio, Ken- tucky [con't]. Nautilus, 40(2): 65-67 [22 October]. 1927. Malanoides |sic| tuberculata moussoni, new name. Nautilus, 40(3): 101 [1 February]. 1927. [Review] H. A. Pilsbry, The land mollusks of the Republic of Panama and the Canal Zone. Nau- tilus, 40(3): 105 [1 February]. 1927. [Review] H. A. Pilsbry, Costa Rican land shells collected by A. A. Olsson. Nautilus, 40(3): 106 [1 February]. 1927. [Review] F. G. Cawston 1926, Environmental influences which favor the development and spread of the Bilharzia parasite in South Africa. Nautilus, 40(3): 106 [1 February]. 1927. A new subgenus and species of Bulinus from Japan. Nautilus, 40(4): April]. 1927. A new subspecies of Thais from Louisiana. Nautilus, 41(1): 6-8 [23 July]; figured 1930, Nau- tilus, 44(1): pl. 2, figs. 10-11 [17 July]. 1927. Collecting [letter to Dr. Pilsbry]. Nautilus, 41(2): 70 [27 October]. 1928. Venus mercenaria var. notata Say. Nautilus, ANA) al 2O— 22m April]. 1928. Ix fluvialis turrita Anthony. Nautilus, 42(1): 36 [14 July]. 1929. Manly D. Barber. Nautilus, 42(3): 101-102 [15 January]. 1929. Arion ater ater (Linné) in Maine. Nautilus, 42(3): 104 [15 January]. 1929. [Review] H. A. Pilsbry and J. C. Bequaert 1927, The aquatic mollusks of the Belgian Congo, with a geographical account of the Congo malacology. Nautilus, 42(3): 107 [15 January]. 1929. Some records and descriptions of new fresh- water mollusks from Cameroon. Bulletin of the Museum of Comparative Zoology, 69(6): 117-123, pl. 1, text fig. 1 [March]. 1929. [Review] A. W. B. Powell, The Recent and ter- tiary cassids of New Zealand and a study of hy- bridization. Nautilus, 42(4): 144 [11 April]. 1929. Freshwater shells of New England. Bulletin Boston Society of Natural History, (52): 3-8, 2 pls. [11 July]. 1929. Some new Liguus from the Florida Everglades. Nautilus, 43(1): 18-21 [11 July]. 1929. A field method of preserving. Nautilus, 43(1): 33-34 [11 July]. 1929. Some land and marine shells from the Missis- sippi Delta region. Nautilus, 43(1): 34-35 [11 July]. 1929. Concerning a policy. Nautilus, 43(2): 69-70 [17 October]. 1930. A new species of Strophocheilus from Brazil. Nautilus, 43(3): 75-77, text figs. 1-3 [15 January]. 1930. A new variety of Achatina panthera from Mad- agascar. Nautilus, 43(3): 85-86 (A. F. Archer, 2nd author) [15 January]. 1930. Physa and Bulinus of Mauritius. Nautilus, 43(3): 92-93 [15 January]. 1930. Littorina littorea Linn. Nautilus, 43(3): 105 [15 121-122, text fig. 1 [29 January]. 1930. The Harvard expedition to Navassa Island. Harvard Alumni Bulletin, 32(24): 684—687 | March]. 1930. [Review] E. Step 1927, Shell life. Nautilus, 43(4): 143 [24 April]. 1930. On the status of Penion Fischer. Journal of Conchology, 19(1): 21 [April]. 1930. West Indian Mollusks no. I—two new varieties of Urocoptis livida Torre. Nautilus, 44(1): 15-16, pl. 2 (C. de la Torre, Ist author) [17 July]. 1930. [Review] “The Venus.” Nautilus, 44(1): 34-35 [17 July]. 1930. New land snails from Tanganyika Territory. Oc- casional Papers Boston Society of Natural History, MOLLUSCA OF CLENCH AND TURNER * Johnson 27 5: 295-300, pl. 16, text figs. 1-3 (A. F. Archer, 2nd author) [July]. 1930. A new Humboldtiana from Texas. Nautilus, 44(1): 10-13, pl. D. figs. 1-4 (H. A. Rehder, 2nd author) [17 July]. 1930. Additional notes on the colony of Helix ner- moralis at Marion, Mass. Nautilus, 44(1): 13-14 [17 July]. 1930. Notes on Physidae with descriptions of new species. Occasional Papers Boston Society of Nat- ural History, 5: 301-315, text figs. 1-4 [8 August]. 1930. A note on the habit of Viviparus subpupu- reus(Say). Nautilus, 44(2): 70 [21 October]. 1931. Land shells from Lubang Island, Philippines. Occasional Papers Boston Society of Natural His- tory, 5: 333-359, pl. 17 (A. F. Archer, 2nd author) [9 January]. 1931. Camptoceros (Culmenella) prashadi, nom. nov. Nautilus, 44(3): 80 [27 January]. 1931. Studies of African land and freshwater mol- lusks. I. On some African species of Bulinus. Oc- casional Papers Boston Society of Natural History, 5: 357-367 (J. C. Bequaert, Ist author) [31 Janu- ary]. 1931. Cephalopods. Bulletin Boston Society of Nat- ural History, (60): 10-14, 4 text figs. [July]. 1931. Exploring the lower Everglades in Florida. Harvard Alumni Bulletin, 33(38): 1211-1213, 4 text figs. [2 July]. 1931. “Ligging” in the Everglades of Florida. Nauti- lus, 45(1): 10-15 [13 July]. 1931. A preventative for the scaling of the periostra- cum. Nautilus, 45(1): 30-31 [13 July]. 1931. Three new terrestrial snails from Yucatan. Oc- casional Papers Boston Society of Natural History, 5: 423-426 (J. C. Bequaert, 1st author) [10 Octo- ber]. 1931. The genus Goodrichia. Nautilus, 45(2): 71-72 [14 October]. 1931. Entovalva (Devonia) perrieri (Malard) in the western Atlantic. Occasional Papers Boston Society of Natural History, 8: 5-8 (C. G. Aguayo, 2nd au- thor) [7 November]. 1932. West Indian mollusks no. II. New Cuban cer- ions. Nautilus, 45(3): 89-91, pl. 6, figs. 6, 7 (C. de la Torre, Ist author) [9 January]. 1932. West Indian mollusks no. III. Two new sub- species of Cuban Liguus. Nautilus, 45(3): 98-100, pl. 6, fig. 9 (C. G. Aguayo, 2nd author) [9 January]. 1932. Achatina murrea Reeve. Nautilus, 45(3): 106 [9 January]. 1932. A new Liguus from Florida. Nautilus, 46(3): 91-92, pl. 7, figs. 7-10 [25 January]. 1932. West Indian mollusks no. IV. Some land mol- lusks from Beata Island, Santo Domingo. Proceed- ings New England Zoological Club, 12: 103-107 [8 February]. 1932. Thomas Say—his first paper on American mol- lusks. Nautilus, 45(4): 112 [9 April]. 1932. Viviparus japonicus v. Mart. Nautilus, 45(4): 136 [9 April]. 1932. [Review] A. W. B. Powell 1930, The Paryphan- tidae of New Zealand, their hypothetical ancestry, with descriptions of a new species and a new genus. Nautilus, 45(4): 142-143 [9 April]. 1932. West Indian mollusks no. V. New Haitian mol- lusks. Proceedings New England Zoological Club, 13: 35-38 [5 July]. 1932. Descriptions of some land snails of southwest- ern North Carolina. Nautilus, 46(1): 14-18, pl. 2 figs. 1-7 (G. S. Banks, 2nd author) [20 July]. 1932. Land shells collected in southwestern North Carolina. Nautilus, 46(2): 58-59 (G. S. Banks, 2nd author) [22 October]. 1932. Diplomorpha coxi (Pease). Nautilus, 46(2): 68— OO} ple 25 fie 8122" October: 1932. Some new land mollusks from Borneo and the Philippines. Occasional Papers Boston Society of Natural History, 8: 37-42, pl. 4 (A. F. Archer, 2nd author) [10 November]. 1933. Two new land shells from the southern Appa- lachians. Nautilus, 46(3): 86-91, pl. 7, figs. 1-6 (A. F. Archer, 2nd author) [25 January]. 1933. Land mollusks from the islands of Mindoro and Lubang, Philippines. Papers Michigan Academy of Science, Arts and Letters, 17: 535-552, pls. 57, 58 (A. F. Archer, 2nd author) [March]. 1933. Non-marine mollusks of Yucatan, pp. 525-545. [In] G. C. Shattuck (ed.), The Peninsula of Yuca- tan: Medical, Biological and Sociological Studies. Part 4. Carnegie Institution of Washington, Publi- cation 431, pl. 68 (J. C. Bequaert, Ist author) [29 June]. 1933. Studies on African land and freshwater mol- lusks: Il. A new ampullariid snail from the lower Belgian Congo. Revue de Zoologie et de Botanique Africaines, 23(2): 71-73, pl. 5 (J. C. Bequaert, 1st author) [June]. 1933. A new fossil Cepolis from Cuba. West Indian mollusks no. VI. Nautilus, 47(1): 21-22, pl. 3, figs. 7, 8 (C. G. Aguayo, 2nd author) [16 June]. 1933. Two new land snails from the Bismark Archi- pelago. Nautilus, 47(1): 23-24, pl. 3 [16 June]. 1933. Notes and descriptions of land mollusks from the Bahama Islands, based mainly upon collections obtained during the Utowana Expeditions of 1932 and 1933. West Indian mollusks no. VII. Proceed- ings New England Zoological Club, 13: 77—100, pl. 1 [15 August]. 1933. Surinamia, a new ampullariid from Dutch Gui- ana. Nautilus, 47(2): 71-72, pl. 7 figs. 1, 2 [1 No- vember]. 1933. The Thomas Bland collection of American land shells. Nautilus, 47(2): 79-80 [1 November]. 1934. New mollusks in the genus Liguus from Cuba and the Isle of Pines. West Indian mollusks no. VIII. Occasional Papers Boston Society of Natural History, 8: 101-124, pls. 5-7 [21 February]. 1934. Studies of African land and fresh-water mol- lusks III, West African Achatinae related to Acha- tina balteata Reeve. American Museum Novitates, 28 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 (705): 1-16, figs. 1-7 (J. C. Bequaert, 1st author) [15 March]. 1934. A new subspecies of Oliva reticularis from southern Florida. Nautilus, 47(4): 142-143, pl. 7, figs. 3-4 [4 May]. 1934. Paludina humerosa Anthony. Nautilus, 47(4): 151 [4 May]. 1934. Studies of African land and fresh-water mol- lusks IV. Descriptions of four presumably new Achatinidae; with notes on the genus Leptocala. Revue Zoologique et Botanique Africaines, 24(3): 269-276, pl. 1, figs. 1-12 (J. C. Bequaert, Ist au- thor) [May]. 1934. [Editorial work on C. W. Johnson’s unfinished manuscript.| List of marine Mollusca of the Atlan- tic coast from Labrador to Texas. Proceedings Bos- ton Society of Natural History, 40(1): 1-204 [July]. 1934. Physa acuta Draparnaud in Massachusetts. Nautilus, 48(1): 33 [10 July]. 1934. Descriptions of a new genus and two new spe- cies of squids from the North Atlantic. Occasional Papers Boston Society of Natural History, 8: 145— 152, 9 text figs. (R. Macdonald, Ist author) [13 July]. 1934. Descriptions of new cerions from Hispaniola and the Bahama Islands, based mainly upon col- lections obtained during the Utowana Expedition of 1934. Proceedings Boston Society of Natural History, 40(2): 205-218, pls. 1, 2 [August]. 1934. A new Hyridella (Mutelidae) from Australia. Journal of Conchology, 20(3): 89-90, pl. 2, figs. 1— 5 [December]. 1934. Studies of African land and freshwater mollusks V. Three new Achatinidae in the collections of the Berlin Zoological Museum. Revue de Zoologie et de Botanique Africaines, 26(1): 112-119, pls. 1, 2 (J. C. Bequaert, Ist author) [29 December]. 1934. |[Molluscan. In Websters New International Dictionary of the English Language, 2nd ed. Springfield, Massachusetts: A. C. Mirriam Com- any. 1935. New races of Liguus from Florida and Cuba. Nautilus, 48(4): 121-125, pl. 7 [24 April]. 1935. A new subgenus and new species of Placostylus from the Solomon Islands. Nautilus, 48(4): 126, pl. 7, fig. 6 [24 April]. 1935. Collecting mollusks, pp. 315-316. [In] G. C. Shattuck (ed.), Handbook of Travel. Chapter 20. Natural History Collecting. Cambridge, Massachu- setts: Harvard University Press [April]. 1935. Physa ampullacea ‘Gould’ Binney. Nautilus, 49(1): 30-32, [22 July]; (illustrated) Nautilus, 48(4): pl. 7, fig. 5 [24 April]. 1935. Some new Urocoptidae from Hispaniola. Pro- ceedings Boston Society of Natural History, 41(1): 1-12, pls. 1, 2 [May]. 1935. A new species of Cerion from Long Island, Ba- hamas, and a note on Cerion milleri (Pfeiffer). Nautilus, 49(2): 49-50, pl. 3, figs. 1, 4 [8 Novem- ber]. 1935. A new Jamaican Poteria. Nautilus, 49(2): 51, pl. 3, figs. 5, 6 (C. G. Aguayo, 2nd author) [8 No- vember]. 1935. Liguus fasciatus caribaeus. Nautilus, 49(2): 68, pl. 3, fig. 3 [8 November]. 1936. A new species of Papuina with notes on P. tay- loriana. Nautilus, 49(3): 88-91, pl. 5, fig. 4 (A. F. Archer, 2nd author) [30 January]. 1936. A new Pleistocene Mecoliotia from Cuba. Nau- tilus, 49(3): 91-93, pl. 5, fig. 3 (C. G. Aguayo, 2nd author) [30 January]. 1936. Studies of African land and freshwater mollusks X. Gulella pumilio (Gould) and two species con- fused with it. Nautilus, 49(3): 93-97, pl. 6, figs. 1- 7 (J. C. Bequaert, 1st author) [30 January]. 1936. A second contribution to the molluscan fauna of Yucatan. Carnegie Institution of Washington, Publication 457: 61-75, pls. 1, 2 (J. C. Bequaert, 1st author) [5 February]. 1936. Description of a new species of Macrocallista. Journal of Conchology, 20(7): 201-204, text fig. (R. A. McLean, 2nd author) [March]. 1936. Studies of African land and freshwater mollusks VI. Notes of Edentulina Pfeiffer, with descriptions of new species. Journal of Conchology, 20(7): 204— 216, pl. 8, figs. 1-10 (J. Bequaert, Ist author) | March]. 1936. Studies on African land and freshwater mol- lusks IX. Rheophilous mollusks of the estuary of the Congo River. Memoires Musée _ Royal d’ Histoire Naturelle de Belgique, 2nd ser (3): 161— 168, pl. 1, figs. 1-6, text figs. 1, 2 (J. C. Bequaert, 1st author) [April]. 1936. A new Cerion from the Bahamas. Nautilus, 49(4): 112-113, pl. 8, fig. 6 [1 May]. 1936. [Obituary] Leopoldo A. Faustino. Nautilus, 49(4): 139 [1 May]. 1936. Helicostyla subcarinata archeri, new subspe- cies. Nautilus, 49(4): 140-141, pl. 8, fig. 5 [1 May]. 1936. [Review] P. W. Reinhart, Classification of the pelecypod family Arcidae. Nautilus, 49(4): 142 [1 May]. 1936. The collection of mollusks, pp. 69-72. [In] Notes Concerning the History and Contents of the Museum of Comparative Zoology, Cambridge, Massachusetts [June]. 1936. Marine bivalves collected by the Harvard-Ba- hama Expedition of 1935. Memorias de la Sociedad Cubana de Historia Natural, 10(3): 157-168, pl. 7 (R. A. McLean, 2nd author) [12 July]. 1936. “Basteria,” A new journal in malacology. Nau- tilus, 50(1): 33 [14 July]. 1936. A new subspecies of Papuina with records of land mollusks from eastern Papua and associated islands. Nautilus, 50(2): 53-54 [29 October]. 1936. Studies in African land and freshwater mollusks XII. An unusually thick-shelled Achatina from the Kivu Region, Belgian Congo. Bulletin Musée Royal d Histoire Naturelle de Belgique, 12(32): 1-4, text figs. 1-10 (J. C. Bequaert, 1st author) [October]. 1936. Studies on African land and freshwater mol- lusks XI. Notes on Gonaxis Taylor, with description MOLLUSCA OF CLENCH AND TURNER * Johnson 29 of a new species. Journal of Conchology, 20(9): 263-273, figs. 1-5 (J. C. Bequaert, Ist author) [| No- vember]. 1936. Studies of African land and freshwater mollusks no. VII. A revision of the genus Archachatina Al- bers. Revue de Zoologie et de Botanique Africai- nes, 29(1): 73-96, pls. 1, 2 (J. C. Bequaert, Ist author) [20 November]. 1936. Studies of African land and freshwater mollusks no. VIII. New species of land operculates, with de- scriptions of a new genus and two new subgenera. Revue de Zoologie et de Botanique Africaines, 29(1): 97-104, pls. 1, 2 (J. C. Bequaert, 1st author) [20 November]. 1936. The Physidae of the West Indies. Memorias de la Sociedad Cubana de Historia Naturelle, 10(5): 335-342, pl. 24 [December]. 1937. Onchidium (Onchidella) floridanum Dall. Nau- tilus, 50(3): 85-86 [29 January]. 1937. Lampsilis cariosa (Say). Nautilus, 50(3): 105 (H. Vander Schalie, 2nd author) [29 January]. 1937. Descriptions of new land and marine shells from the Bahama Islands. Proceedings New Eng- land Zoological Club, 16: 17—26, pl. 1, figs. 1-4 [5 February]. 1937. Marine bivalves from Little and Great Abaco, Grand Bahama and Eleuthera, Bahama Islands. Memorias de la Sociedad Cubana de Historia Nat- ural, 11(1): 31-42, pls. 5, 6 (R. A. McLean, 2nd author) [6 March]. 1937. Physa canadensis Whiteaves. Nautilus, 50(4): 143-144 [4 May]. 1937. Notes and descriptions of some new land and freshwater mollusks from Hispaniola. Memorias de la Sociedad Cubana de Historia Natural, 11(2): 61— 76, pl. 7 (C. G. Aguayo, 2nd author) [8 May]. 1937. Notes on three rare American Polygyra. Nau- tilus, 51(1): 17-18 [3 July]. 1937. A new variety of Bulimulus dealbatus from Al- abama. Nautilus, 51(1): 18-19, pl. 3, fig. 4 [3 July]. 1937. Three new species of Cerion from Long Island, Bahamas. Nautilus, 51(1): 19-23, pl. 3, figs. 5-10 [3 July]. 1937. The supposed introduction of an African Ar chachatina into the West Indies. Nautilus, 51(1): 33-34 (J. C. Bequaert, Ist author) [3 July]. 1937. Forbesopomus, a new genus in the family Pili- dae (Ampullariidae) from the Philippine Islands. Proceedings New England Zoological Club, 16: 53-56, pl. 2, figs. 1-7 (J. C. Bequaert, Ist author) [17 July]. 1937. Shells of Mariguana Island, with a review of the Bahama Helcinidae and descriptions of new Bahama species. Proceedings New England Zoo- logical Club, 16: 57-59, pl. 3, figs. 1-7 [31 July]. 1938. Some new Australian Thersites. Journal of Con- chology, 21(1): 20-24, pl. 1, figs. 1-6 (A. F. Archer, 2nd author) [February]. 1938. A new species of Oliva from Santo Domingo, with notes on other marine forms. Nautilus, 51(4): 109-114, pl. 9, figs. 1-5 [21 April]. 1938. A new subspecies of Solaropsis gibboni from Brazil. Nautilus, 51(4): 115, pl. 9, figs. 6, 7 (J. C. Bequaert, Ist author) [21 April]. 1938. Robert Swift [a note on the obituary noted by T. Bland]. Nautilus, 51(4): 142-143 [21 April]. 1938. Origin of the land and freshwater mollusk fau- na of the Bahamas, with a list of the species oc- curring on Cat and Little San Salvador Islands. Bulletin of the Museum of Comparative Zoology, 80(14): 481-541, pls. 1-3, text figs. 1, 2 [June]. 1938. A third contribution to the molluscan fauna of Yucatan. Carnegie Institution of Washington, Pub- lication 491: 257-260 (J. C. Bequaert, 1st author) [June]. 1938. Further notes on Archachatina. Nautilus, 52(1): 26-27 (J. C. Bequaert, Ist author) [22 July]. 1938. Land and freshwater mollusks of Grand Ba- hama and the Abaco Islands, Bahama Islands. Me- morias de la Sociedad Cubana de Historia Natural, 12(4): 303-333, pls. 24, 25, 1 map [30 September]. 1938. Freshwater shells of New Hampshire, pp. 201— 206. [In] E. E. Hoover (ed.), Biological survey of the Merrimack Watershed. New Hampshire Fish and Game Commission. Survey Report 3, pls. A, B, fig. 1 (H. D. Russell, 2nd author) [December]. 1938. Notes and descriptions of new species of Cal- liostoma, Gaza and Columbarium (Mollusca); ob- tained by the Harvard-Habana Expedition off the coast of Cuba. Memorias de la Sociedad Cubana de Historia Natural, 15(5): 375-384, pl. 28, figs.1— 7 (C. G. Aguayo, 2nd author) [17 December]. 1939. The Oliver P. Emerson collection of Achati- nellidae. Nautilus, 52(3): 107-108 [23 January]. 1939. Helix nemoralis L. at Warm Springs, Va. Nau- tilus, 52(3): 108 [23 January]. 1939. Discus rotundata (Mull.) in Massachusetts. Nautilus, 52(3): 108 (G. Banks, 2nd author) [23 January]. 1939. Notas sobre las Onchidellas de Cuba. Memo- rias de la Sociedad Cubana de Historia Natural, 13(1): 5-7 (C. G. Aguayo, 2nd author) [15 Febru- ary]. 1836. Notes and descriptions of new deepwater Mol- lusca obtained by the Harvard-Havana Expedition off the coast of Cuba II. Memorias de la Sociedad Cubana de Historia Natural, 13(3): 189-197, pls. 28, 29 (C. G. Aguayo, 2nd author) [17 April]. 1939. New records for West Indian Physidae (Mol- lusca). Memorias de la Sociedad Cubana de His- toria Natural, 13(2): 77-78 [April]. 1939. The classification of Florida Liguus. Proceed- ings New England Zoological Club, 17: 77—86 (G. B. Fairchild, 2nd author) [25 May]. 1939. Philippine Lymnaeidae and Planorbidae. The Philippine Journal of Science, 69(1): 7-21, pls. 1, 2 (J. C. Bequaert, Ist author) [May]. 1939. Mollusks that “muscle in.” New England Nat- uralist, (3): 12-13, 4 figs. [June]. 1939. Additional notes on Terebra flammea Lamarck. Nautilus, 53(1): 7-8 [21 July]. 1939. The genus Plesiophysa P. Fischer. Journal of 30 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 Conchology, 21: 175-178, fig. 1 (J. C. Bequaert, lst author) [September]. 1939. Mcleania, a new genus of land mollusks from Puerto Rico. Memorias de la Sociedad Cubana de Historia Natural, 13(5): 283-284, pl. 36, figs. 4-6 (J. C. Bequaert, Ist author) [December]. 1939. A new subspecies of Anguispira kochi from Washington. Memorias de la Sociedad Cubana de Historia Natural, 13(5): 285, pl. 36, fig. 3 (G. Banks, 2nd author) [December]. 1939. A new Musculium from Colombia. Memorias de la Sociedad Cubana de Historia Natural, 13(5): 286, pl. 36, fig. 1 [December]. 1939. Land shells of Guana Island, Virgin Islands, West Indies. Memorias de la Sociedad Cubana His- toria Natural, 13(5): 287-288, pl. 36, fig. 2 [De- cember]. 1939. A new species of Orthaulax from western Cuba. Memorias de la Sociedad Cubana de His- toria Natural, 13(5): 357-358, pls. 47, 48 (C. G. Aguayo, 2nd author) [December]. 1940. Land and freshwater mollusks of Long Island, Bahama Islands. Memorias de la Sociedad Cubana de Historia Natural, 14(1): 3-17, pls. 2, 3 [30 March]. 1940. Notes and descriptions of new deepwater Mol- lusca obtained by the Harvard-Havana Expedition off the coast of Cuba, III. Memorias de la Sociedad Cubana de Historia Natural, 14(1): 77-94, pls. 14— 16, text figs. 1, 2 (C. G. Aguayo, 2nd author) [30 March]. 1940. Freshwater shells of New Hampshire. Biolog- ical Survey of the Connecticut Watershed. New Hampshire Fish and Game Department Survey Report 4: 222-297, pls. 3, 4, text figs. 83, 84 (H. D. Russell, 2nd author) [March]. 1940. Oxystyla of Sanibel Island, Florida. Nautilus, 53(4): 122-123 [29 April]. 1940. Pyrgulopsis nevadensis Stearns in Oregon. Nautilus, 53(4): 137 [29 April]. 1940. Freshwater shells of New Hampshire. Nautilus, 54(2): 52-53 (H. D. Russell, 2nd author) [2 No- vember]. 1940. Another record for Viviparus malleatus in Mas- sachusetts. Nautilus, 54(2): 69-70 [2 November]. 1940. A new Bathyaurinia from off northern Florida. Memorias de la Sociedad Cubana de Historia Nat- ural, 14(3): 241-242, pl. 42, fig. 4, text fig. 1 [19 October]. 1940. Two new land shells from Puerto Rico and St. Croix. Memorias de la Sociedad Cubana de His- toria Natural, 14(3): 243-244, pl. 42, figs. 1-3, 5, 6 [19 October]. 1941. Additions to the rheophilous mollusk fauna of the Congo Estuary. Bulletin Museum of Compar- ative Zoology, 88(1): 3-13, pls. 1, 2 (J. C. Bequaert, Ist author) [April]. 1941. Notes and descriptions of new deepwater Mol- lusca obtained by the Harvard-Havana Expedition off the coast of Cuba, IV. Memorias de la Sociedad Cubana de Historia Natural, 15(2): 177-180, pl. 14, figs. 1-4 (C. G. Aguayo, 2nd author) [10 July]. 1941. The land Mollusca of the Solomon Islands (Succineidae, Bilimulidae and Partulidae). Ameri- can Museum Novitates, (1129): 1-21, figs. 1-13 [31 July]. 1941. The genus Strombus in the western Atlantic. Johnsonia, 1(1): 1-16, pls. 1-10 (R. T. Abbott, 2nd author) [25 October]. 1941. Concerning gastropods adhering to foreign ob- jects. Science, 94(2448): 514 (J. C. Bequaert, Ist author) [28 November]. 1942. A remarkable development of pseudosculpture on a bivalve. Nautilus, 55(3): 73-74, pl. 6, figs. 1- 4 [12 January]. 1942. The genus Ficus in the western Atlantic. John- sonia, 1(2): 1-2, 1 pl. [15 February] 1942. The genera Dosinia, Macrocallista and Amian- tis in the western Atlantic. Johnsonia, 1(3): 1-8, pls. 1-6 [28 February], and Johnsonia, 1(5): 11 [25 May]. 1942. Land shell collecting, pp. 15-16. In Report, 11th annual meeting. Buffalo, New York: American Malacological Union. | March]. 1942. Land shells of the Bimini Islands. Proceedings New England Zoological Club, 19: 53-67 [28 April]. 1942. The mollusks, 45-48. [In] The Boylston Street Fishweir. Papers of the Robert S. Peabody Foun- dation for Archaeology, vol. 2 [April]. 1942. The genera Tectarius and Echininus in the western Atlantic. Johnsonia, 1(4): 1-4, pls. 1-3 (R. T. Abbott, 2nd author) [1 May]. 1942. A new race of Liguus from the Lower Keys of Florida. Proceedings, New England Zoological Club, 19: 69-71 [9 May]. 1942. The genus Conus in the western Atlantic. John- sonia, 1(6): 1-40, pls. 1-15 [5 December]. 1943. The poison cone shell. American Journal of Tropical Medicine, 23(1): 105-120, text figs. 1-7, pl. 1, figs. 1-6 (Y. Kondo, 2nd author) [January]. 1943. Cienfuegos, Cuba. Johnsonia, 1(7): 28 [10 March]. 1943. The genera Xenophora and Tugurium in the western Atlantic. Johnsonia, 1(8): 1—6, pl. 1, figs. 1-6 [24 June]. 1943. The voyage of the H.M.S. “Challenger.” John- sonia, 1(8): 7-8, 1 text fig. [24 June]. 1943. The genera Cypraecassis, Morum, Sconsia and Dalium in the western Atlantic. Johnsonia, 1(9): 1— 8, pls. 14 (R. T. Abbott, 2nd author) [7 July]. 1943. Helicostyla from the Talaud Islands, Molucca Islands, East Indies. Nautilus, 57(1): 17-19 [23 July]. 1943. Papuina gartneriana Pfeiffer. Nautilus, 57(2): 64—66 [30 October]. 1943. Land shells (Synceridae) from the southern and western Pacific. Occasional Papers B. P. Bishop Museum, 17(20): 249-262, text figs. 1-9 (C. M. Cooke, Jr., Ist author) [8 December]. 1943. The genera Gaza and Livona in the western MOLLUSCA OF CLENCH AND TURNER * Johnson 31 Atlantic. Johnsonia, 1(12): 1-9, pls. 14 (R. T. Ab- bott, 2nd author) [31 December]. 1943. [Review] J. F. Whiteaves 1901, Catalogue of the marine invertebrata of eastern Canada. John- sonia, 1(12): 9 [31 December]. 1943. [Review] Nicholas Gualtieri 1742, Index Tes- tarum Conchyliorum 1742. Johnsonia, 1(12): 11 [31 December]. 1943. The voyage of the “Argo.” Johnsonia, 1(12): 12 [31 December]. 1943. Voyage of the “Chazalie.” Johnsonia, 1(12): 12 [31 December]. 1944. Hemphill’s catalogue of the land and freshwater shells of Utah. Nautilus, 57(3): 108 [9 February]. 1944. Shells collected by Fairchild Garden Expedi- tion. American Malacological Union, Annual Re- port for 1943: 16 [16 April]. 1944. Notes on naiades from the Green, Salt and Tradewater rivers in Kentucky. Papers Michigan Academy of Science, Arts and Letters, 29: 223-228 (H. van der Schalie, 2nd author). 1944. The family Cardiidae in the western Atlantic. Johnsonia, 1(13): 1-32, pls. 1-13 (L. C. Smith, 2nd author) [5 June]. 1944. [Review] Austin C. Apgar 1891, Mollusks of the Atlantic coast of the United States south of Cape Hatteras. Johnsonia, 1(13): 32 [5 June]. 1944. A large specimen of Ensis directus Conrad. Nautilus, 58(1): 31 [17 August]. 1944. Anoma flexuosa (Pfeiffer). Nautilus, 58(1): 31— 32 [17 August]. 1944. Giant snail of Malaya. Minutes of the Concho- logical Club of Southern California, (39, p. 2 [Sep- tember]. 1944. [Review] W. H. Dall and C. T. Simpson 1901, The Mollusca of Puerto Rico. Johnsonia, 1(14): 6 [16 October]. 1944. Harvard-Bahama Expedition, 1904. Johnsonia, 1(14): 8 [16 October]. 1944. Harvard-Grand Bahama Expedition, 1936. Johnsonia, 1(14): 8 [16 October]. 1944. The genus Columbarium in the western Atlan- tic. Johnsonia, 1(15): 1-4, pl. 1, figs. 1-5 [16 Oc- tober]. 1944. The genera Casmaria, Galeodea, Phalium, and Cassis in the western Atlantic. Johnsonia, 1(16): 1— 16, pls. 1-8 [28 October]. 1944. Otala lactea (Muller) in Texas. Nautilus, 58(2): 67 [24 November]. 1944. A new tropical Buccinum from Cuba. Revista de la Sociedad Malacologica “Carlos de la Torre,” 2(2): 67-68, fig. 1 [December]. 1945. A new subspecies of Oliva reticularis Lamarck, from the Bahamas. Mollusca, 1(4): 49, 1 plate [10 April]. 1945. The genus Murex in the western Atlantic. John- sonia, 1(17): 1-58, pls. 1-29 [29 May]. 1945. Review of the genera completed for the west- ern Atlantic. Johnsonia, 1(17): 29 [29 May]. 1945. New species of Succinea from Tahiti, with re- marks on other Polynesian species. Occasional Pa- pers, B. P. Bishop Museum, 18(8): 133-138 [31 May]. 1945. Harvard Navassa Expedition. Mollusca, 1(5): 64-65 [10 June]. 1945. Some notes on the life and explorations of Hugh Cuming. Occasional Papers on Mollusks, 1(3): 17-28, pl. 7 [30 July]. 1945. The West Indian fauna in southern Florida. Nautilus, 59(1): 33-34 [6 September]. 1945. [Supplements to] The genus Strombus, the ge- nus Ficus, and the genus Conus in the western At- lantic. Johnsonia, 1(18): 1-4, pls. 1, 2 [23 Novem- ber]. 1945. I knew Lermond. American Malacological Union, Annual Report for 1944-1945: 1-3 |[No- vember]. 1946. The poison cone shell. Occasional Papers on Mollusks, 1(7): 49-80, pls. 11-14 (republication of the 1943 paper [pp. 52-77. Y. Kondo, 2nd author] with five pages of additional text and 1 new plate) [15 March]. 1946. The genus Bankia in the western Atlantic. Johnsonia, 2(19): 1-28, pls. 1-16 (R. D. Turmer, 2nd author) [27 April]. 1946. A new Helicina from Hispaniola. Revista de la Sociedad Malacologica “Carlos de la Torre,” 4(1): 7-8, text figs. 1-3 (M. L. Jaume, 2nd author) [May]. 1946. New genera and species of Synceridae from Ponape, Caroline Islands. Occasional Papers B. P. Bishop Museum, 18(13): 199-206, 5 text figs. [12 June]. 1946. Praticolella griseola Pfeiffer in Hispaniola. Nautilus, 59(4): 144 [27 June]. 1946. A catalogue of the genus Liguus with a descrip- tion of a new subgenus. Occasional Papers on Mol- lusks, 1(10): 117-128 [10 August]. 1946. Nuevo operculado de la region oriental de Cuba. Revista de la Sociedad Malacologica “Carlos de la Torre,” 4(2): 51-53, text fig. [August]. 1946. Republication of descriptions of North Ameri- can mollusks by Verkruzen, Kurtz and DeTar and Beecher. Nautilus, 60(2): 69—71 [18 December]. 1946. The genera Bathyaurinia, Rehderia and Sca- phella in the western Atlantic. Johnsonia, 2(22): 41-60, pls. 24-31 [21 December]. 1946. Una interesante es pecie nueva del genero Cer- ion. Revista de la Sociedad Malacologica “Carlos de la Torre,” 4(3): 85-87, text figs. 1-6 (C. G. Aguayo, 2nd author) [27 December]. 1946. Notes and descriptions of two new species of Calliostoma from Cuba. Revista de la Sociedad Malacologica “Carlos de la Torre,” 4(3): 88-90, text figs. 1, 2 (C. G. Aguayo, 2nd author) [27 Decem- ber]. 1947. The genera Purpura and Thais in the western Atlantic. Johnsonia, 2(23): 61-91, pls. 32-40 [10 March]. 1947. [Review] J. B. Henderson, A monograph of the east American scaphopod mollusks. Johnsonia, 2(23): 92 [10 March]. 32 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 1947. [Republication] Henry Krebs, The West Indian marine shells. Revista de la Sociedad Malacologica “Carlos de la Torre,” 5(1): 23-40 (C. G. Aguayo, 2nd; R. D. Turner, 3rd authors) [April]. 1947. [Republication] Henry Krebs, The West Indian marine shells. Revista de la Sociedad Malacologica “Carlos de la Torre,” 5(2): 59-80 (C. G. Aguayo, 2nd: R. D. Turner, 3rd authors) [October]. 1947. [Republication] Henry Krebs, The West Indian marine shells. Revista de la Sociedad Malacologica “Carlos de la Torre,” 5(3): 91-116 (C. G. Aguayo, 2nd author; R. D. Turner, 3rd author) [31 Decem- ber]. 1947. A new record for Drymaeus multilineatus Say. Nautilus, 61(2): 71 [18 December]. 1948. The genus Truncatella in the western Atlantic. Johnsonia, 2(25): 149-164, pls. 65-73 (R. D. Turn- er, 2nd author) [30 January]. 1948. Torinia canalifera ‘C. B. Adams’ Dall. Nautilus, 61(3): 104-105 [2 March]. 1948. The genotype of Potamolithus Pilsbry. Nautilus, 61(3): 105 [2 March]. 1948. Two new genera and a new species of Syncer- idae from the Caroline Islands. Occasional Papers B. P. Bishop Museum, 19(8): 191-194, text figs. 1, 2 [14 May]. 1948. A new Thais from Angola and notes on Thais haemastoma Linné. American Museum Novitates, (1374): 1-4, 1 pl. (R. D. Turner, 2nd author) [27 May]. 1948. A remarkable malformed specimen of Venus campechiensis Gmelin. Revista de la Sociedad Mal- acological “Carlos de la Torre,” 6(1): 10, text figs 1-3 [10 June]. 1948. [Republication] Henry Krebs, The West Indian marine shells. Revista de la Sociedad Malacologica “Carlos de la Torre,” 6(1): 11-43 (C. G. Aguayo, 2nd; R. D. Turner, 3rd authors) [10 June]. 1948. A catalogue of the family Truncatellidae with notes and descriptions of new species. Occasional Papers on Mollusks, 1(13): 157-212, pls. 22-25 (R. D. Turner, 2nd author) [22 June]. 1948. [Collation of] Paetel’s Catalogue (Catalog der Conchylien-Sammlung, 1887-1891). Occasional Papers on Mollusks, 1(13): 212 [22 June]. 1948. The Hirase Collections of mollusks. Nautilus, 62(1): 34-35 [22 July]. 1948. Drymaeus multilineatus osmenti, new form. Nautilus, 62(1): 36 [22 July]. 1948. [Review] Conchologia Asiatica. Johnsonia, 2(26): 192 [30 October]. 1948. [Review] W. G. Mazyck 1913, Catalogue of Mollusca of South Carolina. Johnsonia, 2(26): 192 [October]. 1948. [Republication] Henry Krebs, The West Indian marine shells. Revista de la Sociedad Malacologica “Carlos de la Torre,” 6(2): 45-48 (C. G Aguayo, 2nd; R. D. Turner, 3rd authors) [November]. 1948. Two new species of Cerion from Cuba and New Providence, Bahamas. Revista de la Sociedad Malacologica “Carlos de la Torre,” 6(2) 49-51, text figs. 1-6 [November]. 1948. Un nuevo Chondropoma de la Provincia de Or- iente, Cuba. Revista de la Sociedad Malacologica “Carlos de la Torre,” 6(2): 53-54, text fig. (C. G. Aguayo, 2nd author) [November]. 1948. Mr. J. R. le B. Tomlin of St. Leonards-on-Sea [note on the editorship of the Journal of Conchol- ogy]. Nautilus, 62(2): 71 [8 December]. 1948. [Review] John Oughton, A zoogeographical study of the land snails of Ontario. Nautilus, 62(2): 72 [8 December]. 1949. Sea shells. Life Magazine, 27(7): 72-75 (R. D. Turner, 2nd author) [14 February]. 1949. The collection of the Marquis de Monterosato. Nautilus, 62(3): 102 [18 March]. 1949. Cyclophoridae and Pupinidae of the Caroline, Fijian and Samoan Islands. Bulletin B. P. Bishop Museum (196): 1-52, text figs. 1-28 [18 May]. 1949. C. Montague Cooke, Jr. Revista de la Sociedad Malacologica “Carlos de la Torre,” 6(3): 83-84 [June]. 1949. Dos nuevos moluscos Cubanos del genero Op- isthosiphon. Revista de la Sociedad Malacologica “Carlos de la Torre,” 6(3): 89-90, 1 plate (C. G. Aguayo, 2nd author) [June]. 1949. [Obituary] Charles Montague Cooke, Jr. 1874— 1948. Bulletin B. P. Bishop Museum (197): 32-33, with photograph [June]. 1949. [Obituary] C. Montague Cooke, Jr. 1874-1948. Proceedings Malacological Society of London, 28(1): 8-9 [24 June]. 1949. Algunas especies de Cerionidae de la costa nor- te de Oriente, Cuba. Torreia (14): 1-10, pl. 1. figs. 1-25 (C. G. Aguayo, 2nd author) [10 December]. 1950. The western Atlantic marine mollusks de- scribed by C. B. Adams. Occasional Papers on Mol- lusks, 1(15): 233-403, pls. 28-49 (R. D. Turner, 2nd author) [26 June]. 1950. [Review] Kikutaro Baba 1949, Opisthobranchia of Sagami Bay collected by His Majesty the Em- peror of Japan. Occasional Papers on Mollusks, 1(15): 404 [26 June]. 1950. A new genus and species of Endodontidae from the Solomon Islands. Revista de la Sociedad Ma- lacologica “Carlos de la Torre,” 7(2): 59-60, text figs. 1-3 [15 July]. 1950. Nuevos Helicinidos de la Provincia de Oriente, Cuba. Revista de la Sociedad Malacologica “Carlos de la Torre,” 7(2): 61-66, pl. 12, figs. 1-11 (C. G, Aguayo, 2nd author) [15 July]. 1950. Edward Chitty with a bibliography and cata- logue of his species of Jamaica land mollusks. Oc- casional Papers Museum of the Institute of Jamaica (1): 1-12, 1 pl. (R. D. Turner, 2nd author) [1 Au- gust]. 1950. The genera Sthenorytis, Cirsotrema, Acirsa, Opalia, and Amaea in the western Atlantic. John- sonia, 2(29): 221-248, pls. 96-107 (R. D. Turner, 2nd author) [30 September]. 1950. [Review] Gunnar Thorson, Reproduction and MOLLUSCA OF CLENCH AND TURNER * Johnson 33 larval development of Danish marine bottom in- vertebrates. Johnsonia, 2(29): 247 [30 September]. 1950. [Review] T. Kuroda and T. Habe, Illustrated Catalogue of Japanese Shells, Kyoto, Japan. John- sonia, 2(29): 247 [30 September]. 1950. A new species of Placostylus from San Cristo- bal, Solomon Islands. Torreia (15): 1-4, text figs. 1— 2 [October]. 1951. The Cuban genus Jeanneretia. Revista de la Sociedad Malacologica “Carlos de la Torre,” 7(3): 81-92, 2 pls. (C. G. Aguayo, 2nd author) [15 Jan- uary]. 1951. Land shells of Mona Island, Puerto Rico. Jour- nal de Conchyliologie, 90(4): 269-276, 1 plate [25 January]. 1951. A new color form of Liguus from Pinar del Rio, Cuba. Revista de la Sociedad Malacologica “Carlos de la Torre,” 7(2): 93-94, text figs. 14 [15 June]. 1951. Novedades en el genero Caracolus en Cuba. Memorias de la Sociedad Cubana de Historia Nat- ural, 20(2): 65-69, pls. 41-42 (C. G. Aguayo, 2nd author) [25 June]. 1951. Busycon coarctatum Sowerby. Occasional Pa- pers on Mollusks, 1(16): 405-409, pl. 50 [11 July]. 1951. The genus Epitonium in the western Atlantic (Part I), Johnsonia, 2(30): 249-288, pls. 108—130 (R. D. Turner, 2nd author) [28 September]. 1951. Some new cerionids from Cuba. Revista de la Sociedad Malacologica “Carlos de la Torre,” 8(2): 69-82, pls. 10, 11 (C. G. Aguayo, 2nd author) [12 November]. 1951. Bulimulus diaphanus Pfeiffer. Nautilus, 65(2): 69 [9 November]. 1951. Drymaeus multilineatus, form osmenti Clench. Nautilus, 65(2): 69 [9 November]. 1951. Oxychilus draparnaldi (Beck) in Cambridge, Massachusetts. Nautilus, 65(2): 70 [9 November]. 1952. Trends in Malacology. Bulletin American Mal- acological Union for 1951: 1-2 [January]. 1952. The scalarinum species complex (Umbonis) in the genus Cerion. Occasional Papers on Mollusks, 1(17): 413-440, pls. 51-57 (C. G. Aguayo, 2nd au- thor) [27 March]. 1952. Notes on some marine shells from the Gulf of Mexico with a description of a new species of Co- nus. Texas Journal of Science, 4(1): 59-61, pl. A, figs. 1, 2 (T. E. Pulley, 2nd author) [30 March]. 1952. Land and freshwater mollusks of Eleuthera Is- land, Bahama Islands. Revista de la Sociedad Ma- lacologica “Carlos de la Torre,” 8(3): 97-119, 3 pls. [25 April]. 1952. The genera Epitonium (Part II), Depressiscala, Cylindriscala, Nystiella and Solutiscala in the west- ern Atlantic. Johnsonia, 2(31): 289-356, pls. 131- 177 (R. D. Turner, 2nd author) [23 July]. 1952. [Obituary] William F. Clapp. Nautilus, 66(1): 31 [25 July]. 1952. Mesanella, a new genus in the Camaenidae. Nautilus, 66(1): 32 (R. D. Turner, 2nd author) [25 July]. 1952. Dates of publication of Johannes Thiele: Hand- buch der Systematischen Weichtierkunde, Jena, Germany. Nautilus, 66(1): 33 [25 July]. 1952. Charles Montague Cooke, Jr., A biobibliogra- phy. B. P. Bishop Museum, Special Publication no. 42, pp. 1-56 [October] (Biography, pp. 1—29, by Y. Kondo; Bibliography and list of taxa by W. J. Clench pp. 31—56). 1952. Cantharis mollis (Gould). Revista de la Socie- dad Malacologica “Carlos de la Torre,” 9(1): 5-6, text fig. [December]. 1953. Nuevos molluscos Cubanos de genero Cerion. Torreia (18): 1-5, figs. 1-6 (C. G. Aguayo, 2nd au- thor) [February]. 1953. [Introduction] William F. Clapp. Johnsonia, 2: 1953. [Supplement to] The genus Murex in the west- ern Atlantic. Johnsonia, 2(32): 359-361, pl. 179 [13 April]. 1953. [Supplement to] The genera Epitonium, Opa- lia, and Cylindriscala in the western Atlantic. John- sonia, 2(32): 361-363, pl. 180 (R. D. Turner, 2nd author) [13 April]. 1953. [Supplement to] The genus Conus in the west- ern Atlantic. Johnsonia, 2(32): 363-376, pls. 181— 185 [13 April]. 1953. [Supplement to] The genera Scaphella and Au- riniopsis in the western Atlantic. Johnsonia, 2: 376— 380, pls. 186, 187 [13 April]. 1953. [in] Aguayo, C. G. Algunos nuevos Moluscos terrestres de Cuba oriental. Memorias de la Socie- dad Cubana de Historia Natural, 21(3): 290-310, pls. 33-35 [May] 1953. A note on Thomas Martyn. Minutes of the Conchological Club of Southern California (133): 2 [ November]. 1954. Galeodes, Busycon, and Melongena. Nautilus, 67(4): 139 [17 May]. 1954. Paramiella, new name for Paramia. Nautilus, 67(4): 139 [17 May]. 1954. [Review] R. Tucker Abbott 1954, American Sea Shells. Johnsonia, 3(33): 63 [17 May]. 1954. [Review] A LaRocque 1953, Catalogue of the Recent Mollusca of Canada. Johnsonia, 3(33): 64 [17 May]. 1954. Mesodon thyroidus (Say) in Florida. Nautilus, 68(1): 23-24 [24 July]. 1954. Introduction to Occasional Papers on Mollusks, Vol. 1, with Preface and Dedication to Merrill E. Champion. Occasional Papers on Mollusks, 1: i—xii [14 August]. 1954. [Supplement to] The Catalogue of the genus Liguus. Occasional Papers on Mollusks, 1(18): 449-444 [14 August]. 1954. [Supplement to] The Catalogue of the family Truncatellidae. Occasional Papers on Mollusks, 1(18): 445 (R. D. Turner, 2nd author) [14 August]. 1954. [Supplement to] The western Atlantic Marine Mollusks described by C. B. Adams. Occasional Pa- pers on Mollusks, 1(18): 447 (R. D. Turner, 2nd author) [14 August]. 34 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 1954. [Review] R. Tucker Abbott 1954, American Sea Shells. Natural History, 63(7): 293 [September]. 1954. The occurrence of clines in molluscan popu- lations. Systematic Zoology, 3(3): 122-125 [Sep- tember]. 1955. [Abstract] Chattahoochee Survey. American Malacological Union, Annual Report for 1954: 3 [January]. 1955. A freshwater mollusk survey of north Florida rivers. Nautilus, 68(3): 95-98 [11 February]. 1955. The Unionidae of B. H. and S. H. Wright. Nau- tilus, 68(3): 104 [11 February]. 1955. Melania cancellata Say. Nautilus, 68(3): 107 [11 Febraury]. 1955. A new Murex from Matanzas, Cuba. Breviora (44): 1-3, pl. 1 [8 April]. 1955. Setaepoma, a new genus in the Synceridae from the Solomon Islands. Nautilus, 68(4): 134 [28 April]. 1955. Land shell collecting, pp. 59-60. [In] How to Collect Shells. American Malacological Union [Special publication]. 1955. The North American genus Lioplax in the fam- ily Viviparidae. Occasional Papers on Mollusks, 2(19): 1-20, pls. 1-4 (R. D. Turner, 2nd author) [13 August]. 1955. Our disappearing fauna. American Malacolog- ical Union, Annual Report for 1955: 11 [Decem- ber]. 1956. The family Melongenidae in the western Atlan- tic. Johnsonia, 3(35): 161-188, pls. 94-109 (R. D. Turner, 2nd author) [30 January]. 1956. A new Cerion from Bimini, Bahamas. Ameri- can Museum Novitates (1794): 1-3, text figs. 1, 2 [3 October]. 1956. Freshwater mollusks of Alabama, Georgia and Florida from the Escambia to the Suwannee River. Bulletin Florida State Museum (Biological Series), 1(3): 99-239, 9 pls. (R. D. Turner, 2nd author) [3 October]. 1956. Land shells of Barbuda Island, Lesser Antilles. Nautilus, 70(2): 69-70 [12 November]. 1957. [in] C. G. Aguayo and M. L. Jaume. Adiciones a la fauna malacologica Cubana—1. Memorias de la Sociedad Cubana de Historia Natural “Felipe Poey,” 23(2): 119-121, 130-131 [March]. 1957. A catalogue of the Cerionidae (Mollusca: Pul- monata). Bulletin Museum of Comparative Zoolo- gy, 116(2): 121-169 [April]. 1957. [Review] Kathleen Johnstone, Sea Treasure. Natural History, 66(5): 228-229 | May]. 1957. Two new land and freshwater mollusks from New Guinea. Breviora (76): 1-4, figs. 1, 2 [18 June]. 1957. New land Mollusca from the Admiralty and Bismarck Islands. American Museum Novitates (1863): 1-4, figs. 1-7 [6 December]. 1957. The family Cymatiidae in the western Atlantic. Johnsonia, 3(36): 189-244, pls. 110-135 (R. D. Turner, 2nd author) [20 December]. 1958. The “Galathea’s” great Discovery. American Malacological Union, Annual Report for 1957: 3-4 [1 January]. 1958. The importance of the amateur. American Mal- acological Union, Annual Report for 1957: 9-10 [1 January]. 1958. New records of West Indian Streptaxidae. Nau- tilus, 72(1): 19-20 [21 July]. 1958. Physa compacta Pease. Nautilus, 72(2): 68 [1 October]. 1958. Quickella vagans (Pilsbry). Nautilus, 72(2): 69— 70 [1 October]. 1958. Fulgoraria kaneko Hirase. Nautilus, 72(2): 69— 70 [1 October]. 1958. The land and freshwater Mollusca of Rennell Island, Solomon Islands. Natural History of Ren- nell Island, British Solomon Islands, Copenhagen, Denmark, 2(27): 155-202, pls. 16-19 [15 Decem- ber]. 1959. John T. Gulick’s Hawaiian Land Shells. Nauti- lus, 72(3): 95-98 [15 January]. 1959. Methods used by C. B. Adams in describing and measuring shells. Nautilus, 72(3): 105-106 [15 January]. 1959. Land and freshwater mollusks of Great and Lit- tle Inagua, Bahama Islands. Bulletin of the Muse- um of Comparative Zoology, 121(2): 29-54, pl. 1 [May]. 1959. [Supplement to] The genus Conus in the west- ern Atlantic. Johnsonia, 3(39): 329 [26 June]. 1959. [Supplement to] The genus Sconsia in the western Atlantic. Johnsonia, 3(39): 329-330, pl. 172 [26 June]. 1959. [Supplement to] The genus Columbarium in the western Atlantic. Johnsonia, 3(39): 330-331, pl. 173) 26 June}: 1959. [Supplement to] The genus Murex in the west- ern Atlantic. Johnsonia, 3(39): 331-334, pls. 174, 175 [26 June]. 1959. Mollusca, pp. 1117-1160, 119 text figs. [In] T. Edmondson (ed.), Freshwater Biology, 2nd ed. New York: John Wiley & Sons, Inc. [August]. 1959. Zachrysia auricoma (Ferussac) in Miami, Flor- ida. Nautilus, 73(2): 76 [3 October]. 1959. Two new genera of land mollusks (Papuininae) from the central highlands of New Guinea. Journal Malacological Society of Australia (3): 4-9, pl. 1, text figs. 1-3 (R. D. Turner, 2nd author) [13 No- vember]. 1960. The green land mollusk from New Guinea. Nautilus, 73(3): 114 [25 January]. 1960. Origin of the land and freshwater mollusks of the Bahamas. American Malacological Union, An- nual Report for 1959: 4-5 [1 January]. 1960. Zachrysia provisoria (Pfeiffer) in Homestead, Florida. Nautilus, 73(4): 161 [4 April]. 1960. Cepaea nemoralis (Linné) from Newport, Rhode Island. Nautilus, 74(2); 82 [5 October]. 1960. Cypraea leucodon Broderip 1820. Journal Mal- acological Society of Australia (4): 14—15, pl. 2 [13 November]. 1960. A new Meliobba from the Scrader Range, New MOLLUSCA OF CLENCH AND TURNER * Johnson 35 Guinea. Journal Malacological Society of Australia (4): 30-31, 1 pl. (R. D. Tumer, 2nd author) [13 November]. 1960. The genus Calliostoma in the western Atlantic. Johnsonia, 4(40): 1-80, pls. 1-56, text fig. 1 (R. D. Turner, 2nd author) [25 November]. 1960. Bivalves, p. 27; Gastropoda, pp. 333-335; Mol- lusca, pp. 327-329, 329a. In Encyclopedia Ameri- cana. Montreal: Americana Inc. of Canada [The 1999 edition of the Encyclopedia had at least three molluscan entries: Gastropod, Mollusk, and marine snails. They were rewritten but still attributed to Clench. | 1961. A record size for Mya arenaria. Nautilus, 74(3): 122 [11 January]. 1961. The bivalve gastropod and the opisthobranchs. American Malacological Union, Annual Report for 1960: 17-18 [March]. 1961. Land shell collecting, pp. 66-67. [In] How to Collect Shells, 2nd ed. American Malacological Union [Special publication]. 1961. Land and freshwater mollusks of Caicos, Turks, Ragged Islands and islands on the Cay Sal Bank, Bahamas. Occasional Papers on Mollusks, 2(26): 229-259, pls. 40-43 [3 May]. 1962. Bivalve gastropod, p. 247. [In] McGraw Hill Yearbook of Science and Technology. 1962. A catalogue of the Viviparidae of North Amer- ica with notes on the distribution of Viviparous georgianus Lea. Occasional Papers on Mollusks, 2(27): 261-287, pls. 44-46 [26 February]. 1962. New records for the genus Lioplax. Occasional Papers on Mollusks, 2(27): 288 [26 February]. 1962. New names introduced by H. A. Pilsbry in the Mollusca and Crustacea. Academy Natural Scienc- es, Philadelphia, 4: 1-218 [Special publication] (R. D. Turner, 2nd author) [June]. 1962. Collecting freshwater mollusks in south central Georgia. Shells and their neighbors. 12: 1, 7, figs. 1, 2 [August]. 1962. New land mollusks in the families Camaenidae and Fruticicolidae from Hispaniola. Revista del Museo Argentino de Ciencias Naturales “Bernar- dino Rivadavia,” 8(17): 213-227, pls. 1, 2 [Decem- ber]. 1962. New species of land mollusks from the Repub- lica Dominicana. Breviora (173): 1-5, 1 pl. [24 De- cember]. 1962. Mollusca, p. 274. [In] Robert J. Rodden (ed.), Excavations at the Early Neolithic Site at Nea Ni- komedeia, Greek Macedonia. Proceedings Prehis- toric Society for 1962, vol. 28. Gloucester, UK: John Bellows, Ltd. 1963. Monographs of the genera Papustyla, Forcar- tia, and Meliobba (Papuininae: Camenidae). Jour- nal Malacological Society of Australia (6): 3-33, pls. 1—3, text figs. 1-9 (R. D. Turmer, 2nd author) [Jan- uary]. 1963. Land and freshwater mollusks of the Crooked Island Group, Bahamas. Bulletin Museum of Com- parative Zoology, 128: 393-416, pls. 1-3 [18 Feb- ruary]. 1963. Some shell malformations. Shells and their neighbors (16): 1-2 (A. S. Merrill, 2nd author) [April]. 1964. The genera Pedipes and Laemodonta in the western Atlantic. Johnsonia, 4(42): 117-127, pls. 76-79 [13 February]. 1964. The Portland Catalogue. Johnsonia, 4(42): 127— 128 [13 February]. 1964. The subfamilies Volutinae, Zidoninae, Odon- tocymbiolinae and Calliotectinae in the western At- lantic. Johnsonia, 4(43): 129-180, pls. 80-114 (R. D. Turner, 2nd author) [13 February]. 1964. Gulella, (Huttonella) bicolor (Hutton). Nauti- lus, 77(4): 149-143 [14 April]. 1964. Dr. Clench qualifies Solander. Hawaiian Shells News, 12(6): 6 [April]. 1964. [Supplement to] The poison cone shell. Occa- sional Papers on Mollusks, 2(30): 344 [7 August]. 1964. Land and freshwater Mollusca of the Cayman Islands, West Indies. Occasional Papers on Mol- lusks, 2(31): 345-380, pls. 61-63 [25 September]. 1964. Who's who in the Malacological Society of Aus- tralia—Donald F. McMichael. Australian News Letter, 12(47): 7 [31 October]. 1964. Monographs of the genera Megalacron and Rhytidoconcha (Papuininae: Camaenidae). Journal Malacological Society of Australia, (8): 36-71, pls. S—11, text figs. 1-9 (R. D. Turner, 2nd author) [15 December]. 1965. Notes on Gastropota tappaniana (C. B. Ad- ams). Nautilus, 78(3): 106-107 [25 January]. 1965. Correction—new land mollusks in the families Camaenidae and Fruticicolidae from Hispaniola. Nautilus, 78(3): 108 [25 January]. 1965. [Review] A. A. Gould 1841, Report on the In- vertebrata of Massachusetts. Johnsonia, 4(44): 216 [12 February]. 1965. [Review] A. A. Gould 1870, edited by W. J. Binney, Report on the Invertebrata of Massachu- setts. Johnsonia, 4(44): 216 [12 February]. 1965. The genus Viviparus (Viviparidae) in North America. Occasional Papers on Mollusks, 2(32): 385-412, pls. 64-68 (S. L. H. Fuller, 2nd author) [9 July]. 1965. A new species of Lithasia from Mississippi. Nautilus, 79(1): 30-33, text fig. 1 [9 July]. 1965. A new species of Clappia from Alabama. Nau- tilus, 79(1): 33-34, text fig. 2 [9 July]. 1965. New species of land mollusks with notes on other species from the Solomon Islands. Breviora (224): 1-8, pls. 1, 2 [15 July]. 1965. Amblema Rafinesque 1820 (Lamellibranchiata): proposed addition to the Official List and proposed suppression of Amblema Rafinesque 1819. Bulletin Zoological Nomenclature, 22(3): 196-197 (A. H. Clarke, Ist author) [August]. 1965. Introduction. Occasional Papers on Mollusks, 2: i-xvi (R. D. Turner, 2nd author) [8 November]. 36 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 (Includes dedication to and notes on the life of J. C. Bequaert, pp. iii—ix.) 1965. [Supplement to] The Catalogue of the genus Liguus. Occasional Papers on Mollusks, 2(34): 425 [S November]. 1965. [Supplement to] The Catalogue of the Family Truncatellidae. Occasional Papers on Mollusks, 2(34): 425 [8 November]. 1965. [Supplement to] The North American genus Lioplax. Occasional Papers on Mollusks, 2(34): 426 [S November]. 1966. Pomacea bridgesi (Reeve) in Florida. Nautilus, 79(3): 105 [25 January]. 1966. Pomacea bridgesi in Puerto Rico. Nautilus, 79(3): 106 [25 January]. 1966. Monograph of the genus Rhynchotrochus (Pa- puininae: Camaenidae). Journal Malacological So- ciety of Australia, (9): 59-65, pls. 15—22, figs. 1-6 (R. D. Turner, 2nd author) [28 January]. 1966. Neopilina and Berthelinia. Program 19th An- nual Shell Show, Treasure Island, St. Petersburg, Florida, pp. 12-13 [March]. 1966. Celetaia, new genus of Viviparidae from the island of Celebes, Indonesia. Nautilus, 79(4): 137 [25 April]. 1966. A new species of Ashmunella from west Texas (Mollusca: Pulmonata). Breviora (244): 1-6, pl. 1, text figs. 1, 2 (W. B. Miller, 2nd author) [29 April]. 1966. Notes and descriptions of new Urocoptidae from Cuba and Hispaniola (Mollusca: Pulmonata). Breviora (245): 1-14, pls. 1, 2 [29 April]. 1966. Panopea bitruncata (Conrad). Nautilus, 80(1): 36 [6 July]. 1966. A new species of Kalendyma from Malaita, Sol- omon Islands. Nautilus, 80(2): 52-53, pl. 3, figs. 1, 2 [11 October]. 1966. Extension in range for Dosinia discus. Nautilus, 80(2): 70 [11 October]. 1966. On Ampullina Guppy. Nautilus, 80(2): 71 (M. K. Jacobsen, 2nd author) [11 October]. 1966. Land shell collecting, pp. 67-68. [In] How to Collect Shells, 3rd ed. American Malacological Union [Special Publication]. 1967. A new species of Lyria (Volutidae) from His- paniola. Nautilus, 80(3): 83-84, figs. 1-3 (R. D. Turner, 2nd author) [29 January]. 1967. Freshwater Mollusca from James River, Virgin- ia, and a new name for Mudalia of authors. Nau- tilus, 80(3): 99-102 (Kenneth J. Boss, 2nd author) [29 January]. 1967. Monograph of the genus Spiroceramus (Mol- lusca: Pulmonata: Urocoptidae). Breviora (260): 1— 10, pls. 1, 2 [3 February]. 1967. The Henry G. Frampton collection of Florida Liguus. Occasional Papers on Mollusks, 3(35): 36 (Kenneth J. Boss, 2nd author) [8 February]. 1967. William Gaillard Mazyck, (1846-1942) with a bibliography and catalogue of his species. Occa- sional Papers on Mollusks, 3(36): 37-43, pl. 14 [15 February]. 1967. [Review] S. Peter Dance, Shell Collecting, An Illustrated History. Occasional Papers on Mollusks, 3(36): 44 [15 February]. 1967. Notes on the Pleurobema collina (Conrad) from the James River, Virginia. Occasional Papers on Mollusks, 3(37): 45-52, pl. 15 (Kenneth J. Boss, Ist author) [24 February]. 1967. Henry G. Frampton 1902-1966. Nautilus, 81(1): 31-32 [6 July]. 1967. Heilprin, Angelo. Explorations on the west coast of Florida and the Okeechobee wilderness. A note on the republication by the Paleontological Research Institution, Ithaca, New York in 1964. Nautilus, 81(2): 68 [16 October]. 1968. Fiscus carolae and F. floridensis. Nautilus, 81(3): 107 [25 January]. 1968. Revision of proposals concerning Ablema Raf- inesque, 1820. Bulletin Zoological Nomenclature, 22(5/6): 341 (A. H. Clarke, 1st author) [January]. 1968. Tree snails (Liguus) of Cuba, Hispaniola and Florida. American Malacological Union, Annual Report for 1967: 48—49 [20 March]. 1968. The economic importance of the Mollusca. Pittsburgh Shell Club Bulletin (3): 6 [March]. 1968. Monograph of the genus Letitia (Papuininae: Camaenidae). Journal Malacological Society of Australia, (11): 32-49, pls. 3-7, text figs. 1, 2 (R. D. Turner, 2nd author) [22 March]. 1968. Notes on the species of Urocoptis described by George C. Spence. Proceedings Malacological So- ciety London, 38(1): 101-102 [April]. 1968. Monograph of the Cuban genus Viana (Mol- lusca: Archeogastropoda: Helicinidae). Breviora, (298): 1-25, pls. 1-4, maps 1—5 (M. K. Jacobsen, 2nd author) [11 October]. 1968. A progress report on a revision of the Cuban Helicinidae (Mollusca: Prosobranchia: Archaeogas- tropoda). American Malacological Union, Annual Report for 1968: 40-41 (M. K. Jacobsen, 2nd au- thor) [December]. 1968. Additions to the land and freshwater Mollusca of Rennell Island, Solomon Islands. The Natural History of Rennell Island, British Solomon Islands, Copenhagen, 5: 173-179 [30 December]. 1969. [Review] R. T. Abbott 1968, Seashells of North America: A Guide to Field Identification. Johnson- ia, 4(47): 368 [7 February]. 1969. Land shells of Jost Van Dyke, Virgin Islands. Nautilus, 82(4): 144-145 [28 April]. 1969. Corbicula manilensis Philippi in the Noli- chucky River, Tennessee. Nautilus, 82(4): 146 (D. H. Stansbery, 2nd author) [28 April]. 1969. Melanoides tuberculata (Muller) in Florida. Nautilus, 83(2): 72 [31 October]. 1969. The Physa striata complex. Nautilus, 83(2): 73 [31 October]. 1969. [Review] C. T. Menke, Synopsis Methodica Molluscorum. Sterkiana (36): 19-20 [December]. 1969. Early workers on the North American naiades. American Malacological Union, Annual Report for 1969: 24-25 [December]. 1970. Land Mollusca of Saba Island, Lesser Antilles. MOLLUSCA OF CLENCH AND TURNER * Johnson 37 Occasional Papers on Mollusks, 3(38): 53-60, pl. 16 [6 February]. 1970. The genus Priotrochatella (Mollusca: Helicini- dae) of the Isle of Pines and Jamaica, West Indies. Occasional Papers on Mollusks, 3(39): 61-80, pls. 17-21 (M. K. Jacobsen, 2nd author) [6 February]. 1970. [Supplement to] The family Volutidae in the western Atlantic. Johnsonia, 4(48): 369-372, pls. 172-174 (R. D. Turner, 2nd author) [29 June]. 1970. [Supplement to] The genus Conus in the west- ern Atlantic. Johnsonia, 4(48): 372-379, pls. 175— 178 (R. C. Bullock, 2nd author) [29 June]. 1970. Corbicula manilensis (Philippi) in lower Flori- da. Nautilus, 84(1): 36 [16 July]. 1970. Rare and endangered mollusks, no. 5, eastern land snails. Malacologia, 10(1): 35 [20 September]. 1970. New records of land and freshwater Mollusca of Rennell Island, Solomon Islands. The Natural History of Rennell Island, British Solomon Islands, Copenhagen, 6: 169-170 [30 December]. 1971. On some Helicina from the Dominican Repub- lic. Nautilus, 84(3): 101-107, text figs. 1, 2 (M. K. Jacobsen, Ist author) [25 January]. 1971. Monograph of the Cuban genera Emoda and Glyptemoda (Mollusca: Archaeogastropoda: Hel- cinidae). Bulletin Museum of Comparative Zoolo- gy, 141(3): 99-130, pls. 1-6 (M. K. Jacobsen, 2nd author) [4 February]. 1971. A monograph of the genera Calidviana, Us- tronia, Trochelviana and Semitrochetella (Mollusca: Archaeogastropoda: Helcinidae). Bulletin Museum of Comparative Zoology, 141(7): 403-463, pls. 1-8 (M. K. Jacobsen, 2nd author) [6 August]. 1971. Additions to the Cayman Islands land mollusks. Nautilus, 85(2): 69-70 [4 October]. 1972. The value of localized data. The shell case. Na- ples Shell Club, 2(1): 6 [February]. 1972. Vagabonding for shells—in retrospect. Ameri- can Malacological Union, Bulletin for 1971: 42 [February]. 1972. Corbicula manilensis (Philippi) in Oklahoma. Nautilus, 85(4): 145 [April]. 1972. [Review] R. T. Abbott 1972, Kingdom of the Seashell. Nautilus, 86(1): 50 [27 July]. 1972. Land and freshwater snails of Savo Island, with anatomical descriptions (Mollusca: Gastropoda). Steenstrupia, 2(15): 207-232 (R. D. Turner, 1st au- thor) [20 August]. 1973. Bibliography of African papers. Achatina (4): 66-67 [September]. 1973. [Scientific editor] Perey A. Morris, A field guide to the shells of the Atlantic and Gulf coasts and the West Indies, 3rd ed. Boston: Houghton Mifflin Company, 330 pp., 76 pls. [September]. 1974. [Review] R. T. Abbott, American Malacologists. Nautilus, 88(1): 27 [29 January]. 1974. The Pleuroceridae and Unionidae of the North Fork of the Holston River above Saltville, Virginia. American Malacological Union. Bulletin for 1973: 33-36 (D. H. Stansbery, 1st author) [22 July]. 1974. A blind Physa from Wyoming with notes on its adaptation to the cave environment. Nautilus, 88(3): 80-85, figs. 1-19 (R. D. Turner, Ist author) [22 July]. 1974. Land shell collecting, pp. 67-68. [In] How to Study and Collect Shells. American Malacological Union. 1975. Mollusca from Russell Cave, pp. 87-90. [In] Griffin, J. W. Investigation in Russell Cave [Ala- bama]. National Park Service, U.S. Department of Interior. Publications in Archaeology, no. 13, Wash- ington, D.C. [1974]. 1975. The Pleuroceridae and Unionidae of the Mid- dle Fork of the Holston River in Virginia. American Malacological Union, Bulletin for 1974: 51-54 (D. H. Stansbery, Ist author) [May]. 1975. Part II, Catalogue of species and bibliography of William Harper Pease. [In] E. A. Kay and W. J. Clench (eds.), A Biobibliography of William Harp- er Pease, Malacologist of Polynesia. Nemouria, Oc- casional Papers of the Delaware Museum of Nat- ural History (16): 22-50 [30 December]. 1976. Land Shell Collecting in Cuba. Pittsburgh, Pennsylvania: Pittsburgh Shell Club Bulletin. [March]. 1976. Ligging in the Everglades of Florida. Tide-ings, 6(1): 10-11 [March]. 1976. Shell rivers of Georgia. Jacksonville Shell Club, Bicentennial Shell Show Booklet, p. 11 [31 July]. 1977. River collecting for freshwater shells. The Shell Case, Naples Shell Club, 5(1): 16-17 [April]. 1977. Forward p. xiii [in] Andrews, J. Shells and Shores of Texas. University of Texas Press/Austin and London. 1978. The Pleuroceridae and Unionidae of the Upper South Fork, Holston River in Virginia. American Malacological Union, Bulletin for 1977: 75-78, fig. 1 (D. H. Stansbery, 2nd author). 1978. Freshwater mollusks of North America. Aqua- sphere, Journal New England Aquarium, Boston, 12(2): 16-21, text figs. [October]. 1979. A biography of Andrew Garrett, early naturalist of Polynesia, Part Il Catalogue of Mollusks. Nau- tilus, 93(2/3): 96-102 [23 April]. 1980. Introduction to “Shells on postage stamps’ by Karl Emmerich. Fullerton, California: Topical Phil- atelic Publishers, p. 15. 1981. Catalogo de la fauna malacologica terreste y fluviatile de la Espanol. (Catalogue de faune ma- lacologique terrestre et fluviatile de l'Isle de Saint Domingue.) (Catalog of the terrestrial and fluviatile mollusk fauna of Hispaniola.) Privately issued [by] (David K. Wetherbee, 2nd author), pp. 1-70 [24 October]. This mimeographed listing was “pre- pared by the junior author largely from the files [in the Museum of Comparative Zoology] (MCZ) and presented to Dr. Clench on the occasion of his en- tering his eighty-fifth year.” This is an uncritical list. It was unauthorized and surreptitiously purloined from various sources without the knowledge of Clench or anyone else in the Department of Mol- lusks. 38 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 1982. [Obituary] Joseph Charles Bequaert, 1886— 1982. Nautilus, 96(2): 35 [21 April]. 1984. Three new species of Macroceramus (Mollusca: Urocoptidae) from the Dominican Republic. Ca- ribbean Journal Science, 20(1—2): 9-12, 1 pl., 1 ta- ble (David K. Wetherbee, Ist author). Although Wetherbee acknowledged that Dr. Ruth D. Turner had allowed him to explore the MCZ collection, neither Turner nor Kenneth J. Boss authorized Wetherbee to describe new species from the col- lection. They were based upon lots that had been given tentative manuscript names by Clench in 1937, which Wetherbee assumed uncritically to be undescribed species. 2002. Additions and Corrections to Clench and Turn- er, 1962. New Names Introduced by H. A. Pilsbry. Cambridge, Massachusetts: Department of Mol- lusks, Museum of Comparative Zoology, Harvard University, pp. 1-22 (R. I. Johnson, Ist author) [25 March]. Publications of Ruth D. Turner Publication dates of The Nautilus, al- most always to the exact day of issue, were compiled by: Coan, E. V., and M. G. Har- asewych. 1993. The Nautilus, 106(4): 174— 180. 1942. Editor. Bulletin of New England Bird Life, 6(8—12): 56-104 [September]. 1943. Birding the first year of the war. Bulletin of the Massachusetts Audubon Society, 28(2): 33-42 [March]. 1944. Vassar birds. Vassar Alumnae Magazine, 30(4): 15-17 [March]. 1946. The genus Bankia in the western Atlantic. Johnsonia, 2(19): 1-28, 16 pls. (W. J. Clench, Ist author) [27 April]. 1946. John Gould Anthony, with a bibliography and catalogue of his species. Occasional Papers on Mol- lusks, 1(8): 81-108, 15 pl. [20 July]. 1947. [Review] A List of the Mollusca of the Atlantic Coast from Labrador to Texas, by Johnson, C. W. 1934. Johnsonia, 2(23): 92 [10 March]. 1947. [Review] Fauna of New England, List of Mol- lusca, by C.W. Johnson, 1915. Johnsonia 2(23): 92 [10 March]. 1947. [Republication] Henry Krebs, The West Indian marine shells. Revista de la Sociedad Malacologica “Carlos de la Torre,” 5(1): 23-40 (W. J. Clench, 1st; C. G. Aguayo, 2nd, and R. D. Turner, 3rd authors) [April]. 1947. Collecting shipworms. Limnological Society of America, Special Publication no. 19, pp. 1-8, text figs. [June]. 1947. Procedimientos para recolectar bromas y otros moluscos perforentes marinos. Revista de la Socie- dad Malacologica “Carlos de la Torre,” 5(2): 43-44 [October]. 1947. [Republication] Henry Krebs, The West Indian marine shells. Revista de la Sociedad Malacologica “Carlos de la Torre,” 5(2): 59-80 (W. J. Clench, Ist; C. G. Aguayo, 2nd, and R. D. Turner, 3rd authors) [October]. 1947. [Republication] Henry Krebs, The West Indian marine shells. Revista de la Sociedad Malacologica “Carlos de la Torre,” 5(3): 91-116 (W. J. Clench, Ist; C. G. Aguayo, 2nd, and R. D. Turner, 3rd au- thors) [31 December]. 1948. The genus Truncatella in the western Atlantic. Johnsonia, 2(25): 149-164, pls. 65-73 (W. J. Clench, 1st author) [30 January]. 1948. A new Thais from Angola and notes on Thais haemastoma Linné. American Museum Novitates, (1374): 1-14, 1 pl. (W. J. Clench, Ist author) [27 May]. 1948. [Republication] Henry Krebs, The West Indian marine shells. Revista de la Sociedad Malacologica “Carlos de la Torre,” 6(1): 11-43 (W. J. Clench, 1st; and C. G. Aguayo, 2nd authors) [10 June]. 1948. A catalogue of the family Truncatellidae with notes and descriptions of new species. Occasional Papers on Mollusks, 1(13): 157-212, pls. 22-24 (W. J. Clench, 1st author) [22 June]. 1948. The family Tonnidae in the western Atlantic. Johnsonia, 2(26): 165-192, 11 pls. [30 October]. 1948. [Republication] Henry Krebs, The West Indian marine shells. Revista de la Sociedad Malacologica “Carlos de la Torre,” 6(2): 45-48 (W. J. Clench, 1st, C. G. Aguayo, 2nd, and R. D. Turner, 3rd authors) [ November]. 1948. William Henry Fluck, 1870-1948. Nautilus, 62(2): 69-70 [8 December]. 1949. Sea shells. Life Magazine, 27(7): 72-75 [deter- mination of all shells figured] (W. J. Clench, Ist author) [14 February]. 1949. [Review] W. H. Dall, P. Bartsch and H. A. Reh- der 1938, A manual of the Recent and fossil marine pelecypod mollusks of the Hawaiian Islands. Oc- casional Papers on Mollusks, 1(14): 231 [30 March]. 1949. [Review] H. H. Edmondson 1933, Reef and shore fauna of Hawaii. Occasional Papers on Mol- lusks, 1(14): 231-232 [30 March]. 1949. [Review] S. Hirase 1934, A collection of Japa- nese shells with illustrations in natural color. Oc- casional Papers on Mollusks, 1(14): 232 [30 March]. 1950. The voyage of the “Tomas Barrera.” Johnsonia, 2(28): 220 [6 January]. 1950. The western Atlantic marine mollusks de- scribed by C. B. Adams. Occasional Papers on Mol- lusks, 1(15): 233-403, pls. 29-49 (W. if Clench, Ist author) [26 June]. 1950. Edward Chitty, with a bibliography and a cat- alogue of his species of Jamaican land mollusks. Occasional Papers Museum of the Institute of Ja- maica, (1): 1-12, 1 pl. (W. J. Clench, 1st author) [1 August]. 1950. The genera Sthenorytis, Cirsotrema, Acirsa, MOLLUSCA OF CLENCH AND TURNER ¢ Johnson 39 Opalia and Amaea in the western Atlantic. John- sonia, 2(29): 221-248, pls. 96-107 (W. J. Clench, Ist author) [30 September]. 1950. [Review] M. C. Sullivan 1942, Bivalve Larvae of Malpeque Bay, Prince Edward Id., Bulletin 77, Fisheries Research Board of Canada, pp. 1—36, 22 pls. Johnsonia, 2(29): 248 [30 September]. 1951. [Review] A. H. Verrill, The Shell Collectors Handbook. Natural History, 60(5): 199 | May]. 1951. [Review] C. M. Yonge 1949, The Sea Shore. Occasional Papers on Mollusks, 1(16): 410-411 [July]. 1951. [Review] G. E. and Nettie MacGinitie 1949, Natural History of Marine Animals. Occasional Pa- pers on Mollusks, 1(16): 411-412 [11 July]. 1951. The genus Epitonium in the western Atlantic (Part I), 2(3): 249-288, 23 pls. (W. J. Clench, Ist author) [28 September]. 1952. Some problems in the Pholadidae. Bulletin of the American Malacological Union Annual Report for 1951: 9-10 [January]. 1952. La Rocolte des Tarets. Catalogues VIII, Xylo- phages et Petricoles Ouest Africains. Institut Fran- cais d Afrique Noirre, pp. 130-134, figs. 156-158 (translation of paper published in Special Publica- tion no. 19 of the Limnological Society of America) [March]. 1952. The genera Epitonium (Part I), Depressiscala, Cylindriscala, Nystiella and Solutiscala in the west- ern Atlantic. Johnsonia, 2(31): 289-356, pls. 131— 177 (W. J. Clench, Ist author) [23 July]. 1952. Mesanella, a new genus in the Camaenidae. Nautilus, 66(1): 32 (W. J. Clench, Ist author) [25 July]. 1953. New England malacologists. American Mala- cological Union Annual Report for 1952: 4-6 [Jan- uary]. 1953. [Supplement to] The Genera Epitonium, Opa- lia and Cylindriscala in the western Atlantic. John- sonia, 2(32): 361-363, pl. 180 (W. J. Clench, Ist author) [13 April]. 1953. [Supplement to] The Genus Bankia in the western Atlantic. Johnsonia, 2(32): 357-359 (D. J. Brown, 2nd author) [13 April]. 1953. Recent works on the marine mollusks of Ar- gentina. Johnsonia, 2(32): 380 [13 April]. 1954. [Review] A. R. Carcelles, Catalogo de la Ma- lacofauna Antarctica Argentina. Johnsonia, 3(33): 64 [17 May]. 1954. The family Pholadidae in the western Atlantic and the eastern Pacific, Part I: Pholadinae. John- sonia, 3(33): 1-63, pls. 1-34 [17 May]. 1954. [Review] Frederico Lange de Morretes 1949, Ensaio de Catalogo dos Moluscos do Brasil. Oc- casional Papers on Mollusks, 1(18): 449 [14 Au- gust]. 1954. [Supplement to] John Gould Anthony. Occa- sional Papers on Mollusks, 1(18): 442 [14 August]. 1954. [Supplement to] The Catalogue of the Family Truncatellidae. Occasional Papers on Mollusks, 1(18): 445 (W. J. Clench, Ist author) [14 August]. 1954. [Supplement to] The western Atlantic marine mollusks described by C. B. Adams. Occasional Pa- pers on Mollusks, 1(18): 447 (W. J. Clench, Ist au- thor) [14 August]. 1955. [Abstract] The Genus Melongena. American Malacological Union, Annual Report 1954: 10 [Jan- uary]. 1955. The family Pholadidae in the western Atlantic and the eastern Pacific, Part Il: Martesiinae, Juan- netiinae and Xylophaginae. Johnsonia, 3(34): 65— 100, pls. 35-93 [27 March]. 1955. The North American genus Lioplax in the fam- ily Viviparidae. Occasional Papers on Mollusks, 2(19): 1-20, pls. 1-4 (W. J. Clench, Ist author) [13 August]. 1955. Scaphopods of the Atlantis dredgings in the western Atlantic with a catalogue of the scaphopod types in the Museum of Comparative Zoology. Deep Sea Research, 3(Suppl.): 309-320 [Decem- ber]. 1955. Collecting shipworms, pp. 32—35. [In] How to Collect Shells. American Malacological Union [Special publication]. 1956. [Abstract] The work of Charles B. Adams in the West Indies and Panama. American Malacolog- ical Union Annual Report for 1955: 7-8. 1956. The family Melonginidae in the western Atlan- tic. Johnsonia, 3(35): 161-188, pls. 94-109 (W. J. Clench, Ist author) [30 January]. 1956. Additions to the Pholadidae ia, 3(35): 188 [30 January]. 1956. Melongena corona Gmelin, an excellent marine laboratory mollusk. Turtox News (34): 106-108, pls. it, 2 looney 1956. Notes on Xylophaga washingtona Bartsch and on the genus. Nautilus, 70(1): 10-12 [17 August]. 1956. The eastern Pacific mollusks described by C. B. Adams. Occasional Papers on Mollusks, 2(20): 21-133, pls. 5-21 [22 September]. 1956. Additions to the western Atlantic Marine Mol- lusks described by C. B. Adams. Occasional Papers on Mollusks, 2(20): 134-136, 1 pl. [22 September]. 1956. Freshwater mollusks of Alabama, Georgia and Florida from the Escambia to the Suwannee River. Bulletin of the Florida State Museum (Biological Series), 1(3): 97-239, 9 pls. (W. J. Clench, Ist au- thor) [October]. 1957. Charles Johnson Maynard and his work in mal- acology. Occasional Papers on Mollusks, 2(21): 137-152, 1 pl. [23 January]. 1957. Molluscan wood borers, pp. 10-13. [In] Sym- posium on Wood for Marine Use and Its Protection from Marine Organisms. American Society for Testing Materials. Philadelphia, Pennsylvania: Spe- cial Technical Publication no. 200 [June]. 1957. The family Cymatiidae in the western Atlantic. Johnsonia, 3(36): 189-244, pls. 110-135 (W. J. Clench, Ist author) [20 December]. 1958. [Review] The Museum Boltenianum or the Bolten Catalogue. Johnsonia, 3(37): 283-284 [8 May]. Part I. Johnson- 40 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 1958. The genus Hemitrochus in Puerto Rico. Oc- casional Papers on Mollusks, 2(22): 153-178, pls. 23-30 [29 May]. 1958. [Review] Voyage Aux Iles de Teneriffe, La Trin- ite Saint-Thomas, Saint Croix et Porto Rico by An- dre Pierre LeDru. Occasional Papers on Mollusks, 2(22): 179-180 [29 May]. 1958. The works of Georgius Everhardus Rumphius. Johnsonia, 3(38): 326-327 [28 June]. 1958. The family Pinnidae in the western Atlantic. Johnsonia, 3(34): 283-326, pls. 149-171 (J. Rose- water, 2nd author) [28 June]. 1959. Notes on the genus Taheitia (Truncatellidae) in New Guinea with the description of a new species. Occasional Papers on Mollusks, 2(23): 181-188, pls. 31, 32 [29 April]. 1959. The genera Hemitoma and Diodora in the western Atlantic. Johnsonia, 3(39): 334-344, pls. 176-179 [26 June]. 1959. [Supplement to] The genera Amaea and Epi- tomium in the western Atlantic. Johnsonia, 3(39): 344 [26 June]. 1959. [Introduction] Henry A. Pilsbry. Johnsonia, 3(26): ii-iv, 2 pls. [26 June]. 1959. Notes on the feeding of Melongena corona. Nautilus, 73(1): 11-13 [20 July]. 1959. Melongena egg cases. Nautilus, 73(2): 77 [8 October]. 1959. Two new genera of land mollusks (Papuininae) from the Central Highlands of New Guinea. Jour- nal Malacological Society Australia (3): 4-9, pl. 1, text figs. 1-3 (W. J. Clench, Ist author) [13 Novem- ber]. 1959. The status of systematic work in the Teredini- dae, pp. 124-136. In D. L. Ray (ed.), Symposium on marine boring and fouling organisms. Seattle: University of Washington Press. 1960. Some techniques for anatomical work. Annual Report of the American Malacological Union for 1959: 6-8 [1 January]. 1960. Land shells of Navassa Island, West Indies. Bulletin Museum of Comparative Zoology, 122(5): 233-244, 7 pls. [March]. 1960. Mounting minute radulae. Nautilus, 73(2): 135-137 [4 April]. 1960. A new Meliobba from Schrader Range, New Guinea. Journal Malacological Society Australia (4): 30-31, 1 pl. (W. J. Clench, Ist author) [13 Novem- ber]. 1960. The occurrence of a nematode parasite in the genus Stylodon. Journal Malacological Society Aus- tralia (4): 56-59, text fig. 1, pl. 7 (M. A. Pini, 2nd author) [13 November]. 1960. The genus Calliostoma in the western Atlantic. Johnsonia, 4(40): 1-80, pls. 1-56, 1 text fig. (W. J. Clench, Ist author) [25 November]. 1960. Teredos en de mens. Correspondentiblad van Nederlandse Malacologische Vereniging, (91): 924— 925. [Translated into Dutch by C. O. V. Regteren, Altena] | December] 1961. Helix pomatia Linné, colonized at Plymouth, Massachusetts. Nautilus, 74(3): 122 [11 January]. 1961. Natural history museums of Europe. American Malacological Union Report for 1960: 13-14 [Feb- ruary]. 1961. Report on the American Malacological Union meeting at McGill University. American Malaco- logical Union Report for 1960: 28—32 [March]. 1961. Pleurotomariidae in Bermuda waters. Nautilus, 74(4): 162-163 [6 April]. 1961. [Review] Traité de Zoologie. Vol. 5, fascicule 2. Embranchement des Mollusques, pp. 1625-2164. Occasional Papers on Mollusks, 2(26): 260 [3 May]. 1961. Remarks on WNettastomella and Jouannetia. American Malacological Union Report for 1961: 17-18 [12 December]. 1961. The genus Lignopholas Turner (Mollusca: Pho- ladidae). Mitteilungen aus dem Zoologischen Mu- seum, Berlin, 37: 287-295 [20 December]. 1961. Collecting shipworms, pp. 32-35. [In] How to Collect Shells, 2nd ed. American Malacological Union [Special publication]. 1962. Nettastomella japonica Yokoyama in North America and notes on the Pholadidae. Occasional Papers on Mollusks, 2(18): 289-308, 7 pls. [26 February]. 1962. New names introduced by H. A. Pilsbry in the Mollusca and Crustacea. Academy of Natural Sci- ences of Philadelphia, Special Publication no. 4, pp. 1-218 (W. J. Clench, Ist author) [June]. 1962. Books help beachcombers play the shell game. Natural History, 71(7): 4-7 [August]. 1962. The genus Lithophaga in the western Atlantic. Johnsonia, 4(41): 81-116, 19 pls. (K. J. Boss, 2nd author) [7 September]. 1962. [Review] V. Fretter and A. Graham, British prosobranch molluscs, their functional anatomy and ecology. Johnsonia, 4(41): 116 [7 September]. 1962. James H. Orton, his contributions to the field of fossil and Recent mollusks. Revista Museo Ar- gentino de Ciencias Naturales “Bernardino Riva- davia,” Buenos Aires, 8(7): 89-99 [December]. 1963. Monographs of the genera Papustyla, Forcartia and Meliobba (Papuininae: Camaenidae). Journal Malacological Society of Australia, (6): 3-33 (W. J. Clench, 1st author) [January]. 1963. Nest building in the bivalve mollusk genera, Musculus and Lima. The Veliger, 6(2): 55-59 (A. S. Merrill, Ist author) [1 October]. 1964. The subfamilies Volutinae, Zidoninae, Odon- tocymbiolinae and Calliotectinae in the western At- lantic. Johnsonia, 4(43): 129-180, pls. 80-114 (W. J. Clench, 1st author) [13 February]. 1964. [Review] R. Riedl, Fauna und Flora der Adria. Johnsonia, 4(43): 180 [13 February]. 1964. Anatomical relationships in the Teredinidae. Annual Report American Malacological Union for 1964: 16-17 [1 December]. 1964. Monographs of the genera Megalacron and Rhytidoconcha (Papuininae: Camaenidae). Journal MOLLUSCA OF CLENCH AND TURNER ¢ Johnson 4] Malacological Society of Australia, (8): 36-71 (W. J. Clench, Ist author) [15 December]. 1965. Introduction. Occasional Papers on Mollusks, 2: i-xvi (W. J. Clench, 1st author) [8 November]. (Includes dedication to and notes on the life of J. C. Bequaert, pp. iii—ix.) 1965. Mussel, pp. 1096-1098, 2 figs. [In] Encyclo- paedia Britannica; 1964, ibid. Snail, pp. 848A— 848H, 11 figs. Chicago: Encyclopaedia Britannica. (Other articles by Turner in the Encyclopaedia Bri- tannica include: Mollusk, Periwinkle, Cockle, Qua- hog, Piddock, Teredo, Whelk, Scallop, and Chiton.) 1965. Some results of deep water testing. Annual Re- port American Malacological Union for 1965: 9-11 [1 December]. 1966. Monograph of the genus Rhynchotrochus (Pa- puininae, Camaenidae). Journal Malacological So- ciety of Australia, (9): 59-95, text figs. 1-6, pls. 15— 22 (W. J. Clench, Ist author) [28 January]. 1966. Report to the government of India on system- atic and biological research on marine wood-boring Mollusca. Rome, Italy: FAO (Food and Agriculture Organization of the United Nations) Report TA 2155, pp. 1-30 [February]. 1966. A survey and illustrated catalogue of the Ter- edinidae. Museum of Comparative Zoology, [Spe- cial publication]: 265, 64 pls., 25 text figs (received MCZ Library 23 March). 1966. Implications of recent research in the Teredi- nidae. Beihefte zu Material und Organismen, Ber- lin, (1): 437-446. 1966. Marine borer research in cooperation with the Office of Naval Research. Report of First Inter- American Naval Research Congress [not seen]. 1966. Collecting shipworms, pp. 50-52. [In] How to Collect Shells, 3rd ed. American Malacological Union [Special publication]. 1967. A new species of Lyria (Volutidae) from His- paniola. Nautilus, 80(3): 83-84, figs. 2, 3 (W. J. Clench, Ist author) [24 January]. 1967. A new species of fossil Chlamys from Wright Valley, McMurdo Sound, Antarctica. New Zealand Journal of Geology and Geophysics, 10(2): 446— 455, figs. 1-5 [May]. 1967. Teredo, pp. 861-862. In Encyclopaedia Britan- nica. Chicago: Encyclopaedia Britannica, Inc. 1968. The Xylophagainae and the Teredinidae—a study in contrasts. Annual Report American Mala- cological Union for 1967: 46-48 [20 March]. 1968. Monograph of the genus Letitia (Papuininae: Camaenidae). Journal of Malacological Society of Australia, (11): 32-49, pls. 3-7, text figs. 1, 2 (W. J. Clench, Ist author) [22 March]. 1968. Biological studies in marine wood borers. An- nual Report American Malacological Union for 1968: 14-16 (A. C. Johnson, 2nd author) [27 De- cember]. 1969. | Review] H. Stix, M. Stix and R. T. Abbott, The shell, five hundred million years of inspired design. Natural History, 78(3): 60-62 [March]. 1969. Some problems and techniques in rearing bi- valve larvae. Annual Report American Malacologi- cal Union for 1969: 9-12, 1 pl. (A. C. Johnson, 2nd author) [19 December]. 1969. Pholadacea, pp. 702-742. [In] R. Moore (ed.), Treatise on Invertebrate Paleontology, (N) Mollus- ca, vol. 6, no. 2 of 3, figs. 162-214. 1970. Richard Winslow Foster. Johnsonia, 4: ii—v, 3 figs. [29 June]. 1970. [Supplement to] The family Volutidae in the western Atlantic. Johnsonia, 4(48): 369-372, pls. 172-174 (W. J. Clench, Ist author) [29 June]. 1971. Some anatomical and life history studies of wood boring bivalve systematics. Annual Report American Malacological Union for 1970: 65-66 (J. L. Culliney, 2nd author) [18 February]. 1971. Identification of marine wood boring mollusks of the world, Chap. 1, pp. 17-64, text figs. 1-74. [In] E. B. G. Jones and S. K. Elthringham (eds.), Marine Borers, Fungi and Fouling Organisms of Wood. Paris: Organization for Economic Co-oper- ation and Development. 1971. Biology of the marine boring mollusks of the world, Chap. 13, pp. 259-301, text figs. 1-14. [In] E. B. G. Jones and S. K. Elthringham (eds.), Ma- rine Borers, Fungi and Fouling Organisms of Wood. Paris: Organization for Economic Co-oper- ation and Development (A. C. Johnson, 2nd au- thor). 1971. Australian shipworms. Australian Natural His- tory, Sydney, 17(4): 139-145, 4 pls. [December]. 1971. [Review] Walter Deas and Clarrie Lawler, Be- neath Australian seas. Australian Newsletter N. S., (De &. 1972. Cinephotomicrography: a tool in biological studies. Bulletin of the American Malacological Union for 1971: 30 [February]. 1972. Land and freshwater snails of Savo Island, Sol- omons, with anatomical descriptions (Mollusca, Gastropoda). Steenstrupia, 2(15): 207-232, pls. 1— 13 (W. J. Clench, 2nd author) [20 August]. 1972. Xyloredo, new teredinid-like abyssal wood-bor- ers (Mollusca, Pholadidae, Xylophagainae). Bre- viora, (397): 1-19, pls. 1-6 [6 November]. 1972. Teredicola typicus C. B. Wilson, 1942 (Copepo- da, Cyclopoida) from shipworms in Australia, New Zealand, and Japan. Australian Journal Marine and Freshwater Research, 23(1): 63-72, figs. 1-16 (A. G. Humes, Ist author) [June]. 1972. Results of an international cooperative research program on the biodeterioration of timber sub- merged in the sea. Material und Organismen, 7(2): 93-118 (E. B. G. Jones, Ist, H. Kuhne, 2nd, P. C. Trussell, 3rd, and R. D. Turner, 4th authors). 1972. A new genus and species of deep water wood- boring bivalve (Mollusca, Pholadidae, Xylophagai- nae). Basteria, 36(2—5): 97-104, figs. 1-12 [19 De- cember]. 1973. First report on marine borers in Oyster Creek, Waretown, New Jersey. Museum of Comparative 42 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 Zoology, Harvard University, 30 pp. [3 April] [not seen ].° 1973. Wood-boring bivalves, opportunistic species in the deep sea. Science, 180(4093): 1377-1379, 2 figs., 1 table [29 June]. 1973. The biologists view of the Teredinidae and their control (with a documentary film on the life history of the Teredinidae). Proceedings Third Interna- tional Congress on Marine Corrosion and Fouling, National Bureau of Standards, Gaithersburg, Mary- land, pp. 83-87 (J. L. Culliney, 2nd author) [No- vember]. 1973. Deep water wood-boring mollusks. Proceed- ings Third International Congress on Marine Cor- rosion and Fouling, National Bureau of Standards, Gaithersburg, Maryland, pp. §36—841 [November]. 1974. In the path of a warm saline effluent. American Malacological Union Bulletin for 1973: 36-44, figs. 1-3 [22 May]. 1974. A new blind Physa from Wyoming with notes on its adaptation to the cave environment. Nauti- lus, 88(3): SO-S5, 19 figs. (W. J. Clench, 2nd au- thor) [22 July]. 1974. Fourth report on marine borers in Oyster Creek: the introduction of Teredo furcifera von Martens into Oyster Creek, Waretown, New Jersey. Museum of Comparative Zoology, Harvard Univer- sity, 4 pp. [27 August: not seen].* 1974. Collecting shipworms, pp. 50-53. [In] How to Study and Collect Shells. American Malacological Union. 1974. New approaches and techniques for studying bivalve larvae, pp. 257-271, 2 figs. [In] W. L. Smith and M. H. Chanley (eds.), Culture of Marine In- vertebrate Animals. New York: Plenum Publishing Corp (J. L. Culliney, Ist, and P. J. Boyle, 2nd au- thors). 1975. Studies of bivalve larvae using the scanning electron microscope and critical point drying. Bul- letin American Malacological Union for 1974: 59— 65 (P. J. Boyle, 2nd author) [May]. 1975. [Review] Alan Solem, The Shell Makers, Intro- ducing Mollusks. Journal of the Fisheries Research Board of Canada, 32(5): 719-720 [May]. 1976. Bivalve larvae, their behavior, dispersal and identification. Proceedings of U.S.—U.S.S.R. Work- shop in Biological productivity and biochemistry of the worlds oceans, pp. 23-25. [In] J. Costlow (ed.), Ecology of Fouling Communities. Beaufort, North Carolina: Duke University Marine Laboratory. 1976. Fixation and preservation of marine zooplank- ton, pp. 290-304. [In] H. F. Steedman (ed.), SCOR/UNESCO Handbook Zooplankton Fixation ‘Hoagland and Turmer (1980, Range extension of teredinids .. .) referred to these two reports. None of the four implied reports are again mentioned by Turner and no copies of any were located among her papers, suggesting that they were proprietary to the U.S. Nuclear Regulatory Commission. and Preservation, Chap. 8, Mollusca, Part 11. Paris: UNESCO Press. 1976. Larval development of the deep-water wood boring bivalve Xylophaga atlantica Richards (Mol- lusca, Bivalvia, Pholadidae). Ophelia, 15(2): 149- 161 (J. L. Culliney, Ist author) [November]. 1976. Larval development of the wood boring pid- dock Martesia striata (Linnaeus) (Mollusca: Pho- ladidae). Journal of Experimental Marine Biology and Ecology, 22(1): 55-68, text figs. 14 (P. J. Boyle, 1st author) [April]. 1976. Marine biodeteriogenic organisms, I. Lignicol- ous fungi and bacteria and the wood boring Mol- lusca and Crustacea. International Biodeterioration Bulletin, 12(4): 120-134 (G. Jones, Ist, R. D. Turmn- er, 2nd, S. E. Furtado, 3rd, and H. Kuhne, 4th authors). 1976. Reproductive pattern in an abyssal snail. Amer- ican Zoologist, 16(2): 269 (M. A. Rex. Ist, and C. A. Van Ummersen, 2nd authors) [Spring]. 1976. Search for a weak link, pp. 31-40. In J. D. Bultman (ed.), Proceedings Workshop on Biode- terioration of Tropical Woods. Washington, D.C.: Naval Research Laboratory. 1976. Some factors involved in the settlement and metamorphosis of marine bivalve larvae, pp. 409- 416. [In] Sharpley and Kaplan (eds.), Proceedings 3rd International Biodegradation Symposium, 1975: University of Rhode Island. London: Applied Science Publishers. 1977. Control of marine borer attack on wood. U.S. Patent 4,012,529 (J. D. Bultman, Ist, and L. Jurd, 2nd authors) [not seen]. 1977. Development, metamorphosis and natural his- tory of the nudibranch Doridella obscura Verrill (Corambidae: Opisthobranchia). Journal of Exper- imental Marine Biological Ecology, 27(2): 171-185 (F. E. Perron, lst author) [May]. 1977. Genetic similarities of wood-boring bivalves (Pholadidae and Teredinidae) based on compari- sons of allozymes. Biological Bulletin, 153(2): 420 (T. J. Cole, 1st author). 1977. Analysis of populations of boring and fouling organisms in the vicinity of the Oyster Creek Nu- clear Generating Station over the period: April 30— Nov. 30, 1976. 1st report to the U.S.N.R.C., 61 pp. Washington, D.C.: U.S. Nuclear Regulatory Com- mission Public Document Room (K. E. Hoagland, Ist. R. D. Turmer, 2nd, and M. Rochester, 3rd au- thors) [not seen]. 1977. Analysis of populations of boring and fouling organisms in the vicinity of the Oyster Creek Nu- clear Generating Station over the period: Dec. 1, 1976—Feb. 28, 1977. 2nd report to the U.S.N.R.C., 61 pp. Washington, D.C.: U.S. Nuclear Regulatory Commission Public Document Room (K. E. Hoag- land, Ist, R. D. Turner, 2nd, and M. Rochester, 3rd authors) [not seen]. 1977. Analysis of populations of boring and fouling organisms in the vicinity of the Oyster Creek Nu- clear Generating Station over the period: March 1— MOLLUSCA OF CLENCH AND TURNER * Johnson 43 May. 31, 1977. 3rd report to the U.S.N.R.C., 26 pp. Washington, D.C.: U.S. Nuclear Regulatory Commission Public Document Room (K. E. Hoag- land, Ist, M. Rochester, 2nd, and R. D. Turner, 3rd authors) [not seen]. 1978. The feeding behaviour and diet of Calliostoma occidentale, a coelenterate-associated prosobranch gastropod. Journal of Molluscan Studies, 44(1): 100-103 (F. Perron, 1st author) [16 May]. 1978. Contribution of field and life history studies to an understanding of some cases of opportunism, pp. 241-244. [In] U.S.S.R—U.S.A. Symposium on the Program Biological Productivity and Biochem- istry of the World’s Oceans. Leningrad, November 30—December 4, 1976. 1978. Wood, mollusks, and deep-sea food chains. Bulletin American Malacological Union for 1977: 13-19, figs. 1-3. 1978. Genetic relations of deep-sea wood-borers. Bulletin American Malacological Union for 1977: 19-25 (T. Cole, Ist author). 1979. The role of phytoplankton in the diets of adult and larval shipworms, Lyrodus pedicellatus (Bival- via: Teredinidae). Estuaries, 2(1): 58-60 (J. A. Pechenik, Ist, and F. A. Perron, 2nd authors) [March]. 1979. Bankia neztalia n. sp. (Bivalvia: Teredinidae) from Australia-New Zealand, and its relationships. Journal of the Royal Society of New Zealand, 9(4): 465-473 (J. L. McKoy, 2nd author) [December]. 1979. Mollusks as prey of ariid catfish in the Fly Riv- er, New Guinea. Bulletin of the American Mala- cological Union for 1978: 33-40, pls. 1-6 (T. R. Roberts, 2nd author). 1979. New techniques for preparing shells of bivalve larvae for examination with the scanning electron microscope. Bulletin of the American Malacologi- cal Union for 1978: 17—24, pls. 1-3 (C. B. Calloway, lst author). 1979. [Abstracts] High larval dispersal capability of a deep-sea hydrothermal vent bivalve from the Ga- lapagos Rift. American Society of Zoologists Meet- ing, Dec. 27-30, 1979. American Zoologist, 19(3): 927 (R. A. Lutz, Ist, D. C. Rhoads, 2nd, D. Ja- blonski, 3rd, and R. D. Turner, 4th authors) [April]. 1979. Reproductive pattern in the abyssal snail, Ben- thonella tenella (Jeffreys), pp. 173-188. [In] S. Stancyk (ed.), Reproductive Ecology of Marine In- vertebrates. Belle W. Baruch Library in Marine Science publication 9 (M. A. Rex, Ist, and C. A. Van Ummerson, 2nd authors). Columbia, South Carolina: University of South Carolina. 1979. [Abstract] Biology, life history and relationships of Zachsia zenkewitschi. XTV Pacific Science Con- gress, Khabarovsk. Abstracts, Committee F, Sec. lla, pp. 139-141 (Y. M. Yakovlev, 2nd author) [not seen |. 1979. Galapagos 79: initial findings of a deep-sea bi- ological quest. Oceanus, 22(2): 1-10 (F. Grassle, Ist author, and members of the cruise) [Spring]. 1980. Effects of closed-culture competitive interac- tions on growth of Teredo navalis. Biological Bul- letin, 159(2): 465 (G. A. Tracy, Ist, and C. J. Berg, 2nd authors) [October]. 1980. The giant white clam from the Galapagos Rift, Calyptogena magnifica n. sp. (Bivalvia; Vesicomyi- dae). Malacologia, 20(1): 161-194 (K. J. Boss, Ist author) [22 August]. 1980. Macrobiodegradation of plastics, pp. 117—122. In The Proceedings of the fourth International Bio- deterioration Symposium, Berlin (August-Septem- ber 1978). The Biodeterioration Society (G. J. L. Griffin, 1st author). London: Pitman [for] the Bio- deterioration Society. 1980. Larval dispersal of a deep-sea hydrothermal vent bivalve from the Galapagos Rift. Marine Bi- ology, 57(2): 127-133 (R. A. Lutz, Ist, D. Jablonski, Ind, D. C. Rhoads, 3rd, and R. D. Turner, 4th au- thors) [1 September]. 1980. Range extension of teredinids (shipworms) and polychaetes in the vicinity of a temperate-zone nu- clear generating station. Marine Biology, 58(1): 55— 64 (K. E. Hoagland, Ist author) [2 October]. 1980. Evolution and adaptive radiation of shipworms. Haliotis, 10(2): 68 (K. E. Hoagland, Ist author) [October]. 1981. Evolution and adaptive radiation of shipworms. Malacologia, 21(1—2): 111-148 (K. E. Hoagland, Ist author) [8 December]. 1981. [Abstract] Life cycle of Zachsia zenkewitschi, bivalve mollusk with dwarf males, p. 207. [In] Sixth All-Union Conference on Embryology Abstracts of papers, Nauka, Moscow [in Russian, not seen]. 1981. Physiological aspects of wood consumption, growth, and reproduction in the shipworm, Lyro- dus pedicellatus Quatrefages (Bivalvia: Teredini- dae). Journal of Experimental Marine Biology and Ecology, 52(1): 63-76 (S. Gallager, 1st, R. D. Turn- er, 2nd, and C. Berg, 3rd authors) [7 April]. 1981. Preliminary observations of bacteria and ship- worms (Bivalvia: Teredinidae). Biological Bulletin, 161(2): 332 (A. Wright, lst, C. Cavanaugh, 2nd, S. M. Gallager, 3rd, R. Mann, 4th, and R. D. Turner, 5th authors) [October]. 1981. Wood islands and thermal vents as centers of diverse communities in the deep-sea. Biologiia Morya, 7(1): 3-10 [in Russian]; Soviet Journal of Marine Biology, 7(1): 1-9 [translation by Plenum Publishing]. 1981. Documentation and implications of rapid suc- cessive brooding in the shipworm, Lydorus flori- danus (Mullusca: Bivalva), pp. 172-177, figs. 1-2. In Proceedings XIV Pacific Science Congress, Kha- barovsk, USSR, August 1979. Section Marine Bi- ology 2, Genetics and Reproduction of Marine Or- ganisms (C.B. Calloway, 1st author) [in Russian]. 1981. The ecology and reproduction of Zachsia zen- kewitschi, a teredinid with dwarf males, pp. 215— 219, figs. 1-5. In Proceedings XIV Pacific Science Congress, Khabarovsk, USSR, August 1979. Sec- tion Marine Biology 2, Genetics and Reproduction 44 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 of Marine Organisms (Y. Yakoley, 2nd author) [in Russian]. 1982. [Abstract] Feeding types in vent macro-organ- isms. Abstracts of Papers of the 148th National Meeting American Association for the Advance- ment of Science, 1982: 34 [not seen]. 1983. Dwarf males in the Teredinidae (Bivalvia: Pho- ladacea). Science, 219(4588): 1077-1078 (Y. Ya- kovlev, 2nd author) [4 March]. 1983. Documentation and implications of rapid suc- cessive gametogenic cycles and broods in the ship- worm, Lyrodus floridanus (Bartsch) (Bivalvia: Ter- edinidae). Journal of Shellfish Research, 3(1): 65— 69 (C. B. Calloway, Ist author) [June]. 1983. A cellulolytic nitrogen-fixing bacterium cul- tured from the gland of Deshayes in shipworms (Bivalvia: Teredinidae). Science, 221(4618): 1401— 1403 (J. Waterbury, lst, and C. B. Calloway, 2nd authors) [30 September]. 1983. Some aspects of the life history of a bivalve mollusc, Zachsia zenkewitschi. Biologiia Morya, 9(5): 27-34 (Y. Yakovlev, 2nd, and E. M. Karaseva, 3rd authors) [in Russian]. 1983. Documentation and implications of rapid suc- cessive brooding in the shipworm, Lyrodus flori- danus (Mollusca: Bivalvia), pp. 172-177, figs. 1-2. In Proceedings XIV Pacific Science Congress, Kha- barovsk, USSR, August 1979. Section Marine Bi- ology 2, Genetics and Reproduction of Marine Or- ganisms (C. B. Calloway, 1st author) [in Russian]. 1983. The ecology and reproduction of Zachsia zen- kewitschi, a teredinid with dwarf males, pp. 215— 219, figs. 1-5. In Proceedings XIV Pacific Science Congress, Khabarovsk, USSR, August 1979. Sec- tion Marine Biology 2, Genetics and Reproduction of Marine Organisms (Y. Yakovlev, 2nd author) [in Russian]. 1984. An overview of research on marine borers: past progress and future directions, pp. 3-16. [In] J. D. Costlow and R. C. Tipper (eds.), Marine Biodete- rioration: An Interdisciplinary Study. Annapolis, Maryland: Naval Institute Press. 1984. Growth and distribution of mollusks at deep- sea vents and seeps. Oceanus, 27(3): 55-62 (R. A. Lutz, 2nd author) | Fall]. 1984. Some aspects of the life history of Zachsia zen- kewitschi (Teredinidae, Bivalvia). Soviet Journal of Marine Biology, 9(5): 257-264 [Translation from the Russian by Plenum Publishing, New York); Biologiia Morya (1983) (Y. M. Yakovlev, 2nd, and E. M. Karaseua, 3rd authors) [in Russian]. 1984. [Abstract] Hydrothermal vents, sulfide seeps and mollusks. Program for American Malacological Union—Fiftieth Annual Meeting, Norfolk, Virgin- ia, July 1984: 10. 1984. [Abstract] Larval ecology of mollusks at deep- sea hydrothermal vents. Program for American Malacological Union—Fiftieth Annual Meeting, Norfolk, Virginia, July 1984: 12 (P. Bouchet. Ist, and R. A. Lutz, 2nd authors). 1984, Larval development and dispersal at deep-sea hydrothermal vents. Science, 226(4681): 1451— 1454 (R. A. Lutz, Ist, and D. Jablonski, 2nd au- thors) [21 December]. 1985. [Abstract] Hydrothermal vents, sulfide seeps and mollusks. American Malacological Bulletin, 3(1): 96 [1984 meeting] [March]. 1985. Notes on mollusks of deep-sea vents and re- ducing sediments, pp. 23-34. [In] Perspectives in Malacology, Special edition no. 1 of the American Malacological Bulletin honoring Professor Mel- bourne R. Carriker [July]. 1985. William J. Clench—October 24, 1897—Febru- ary 1984. Malacological Review, 18: 123-124 [De- cember]. 1985. Modes of molluscan larval development at deep-sea hydrothermal vents. [In] M. L. Jones (ed.), Hydrothermal Vents of the Eastern Pacific: An Overview. Bulletin of the Biological Society Washington, (6): 167-184, figs. 1-28 (R. A. Lutz, 2nd, and D. Jablonski, 3rd authors) [30 Decem- ber]. 1985. [Abstract] Description of a hydrocarbon seep community on the Louisiana slope. American Zo- ologist, 25(4) 10A (C. J. Denoux, M. C. Kennicutt, R. R. Bidigare, J. M. Brooks, R. R. Fay, M. L. Jones, R. D. Turner, 7th author) [December]. 1985. [Abstract] Squat lobsters, Munidopsis, associ- ated with mesh enclosed wood panels submerged in the deep-sea. American Zoologist, 25(4): 141A (A. B. Williams, 1st author) [December]. 1986. Larval ecology of mollusks at deep-sea hydro- thermal vents. American Malacological Bulletin, 4(1): 49-54 (R. A. Lutz, Ist, P Bouchet, 2nd, D. Jablonski, 3rd, and A. Waren, 5th authors) [Feb- ruary]. 1986. Squat lobsters (Galatheidae: Munidopsis) as- sociated with mesh-enclosed wood panels sub- merged in the deep sea. Journal Crustacean Biol- ogy, 6(3): 617-624 (A. B. Williams, Ist author) [Au- gust]. 1986. [Abstract] The biology of molluscan hard-sub- strate borers, p. 35. [In] International Conference on Marine Sciences of the Arabian Sea, March 28— April 2, 1986, Karachi, Pakistan [not seen]. 1986. The language of benthic marine invertebrate development patterns: problems and needs, pp. 227-235, figs. 1-10. [In] M.-F. Thompson, R. Sa- rojini, and R. Nagabhushanam (eds.), Biology of Benthic Marine Organisms: Techniques and Meth- ods as Applied to the Indian Ocean. Bombay: Ox- ford and IBH Publishing (J. A. Pechenik, 2nd, and C. B. Calloway, 3rd authors). 1987. Introduction to Symposium on Deep-Sea Hy- drothermal Vents and Cold-Water Seeps, p. 21. In 153rd National Meeting of the American Academy for the Advancement of Science, Chicago, 14—18 February, Abstracts of Papers [not seen]. 1987. Seasonal recruitment of marine invertebrates to hard substrates on Georges Bank and the eastern continental shelf of the United States. Nautilus, 101(1): 19-24 (C. J. Berg, 1st, B. Butman, 2nd, J. MOLLUSCA OF CLENCH AND TURNER * Johnson 45 A Barly sid) and Ro 1D) Turver, Ath authors) [30 January]. 1987. [Abstract] Species pairs in the Teredinidae p. 44. In American Malacological Union Annual Meeting July 19-23 Key West, Florida (C. B. Cal- loway, 2nd author). 1987. Species pairs in the Teredinidae. International research group on wood preservation, document IRG/WP/4142: 1-2 (C. B. Calloway, 2nd author). 1988. Biodeterioration—multidisciplinary, collabora- tive research, pp. 3-12. [In] M.-F. Thompson, R. Sarojini, and R. Nagabhushanam (eds.), Marine Biodeterioration—Advanced Techniques Applica- ble to the Indian Ocean. Bombay, India: Oxford and IBH Publishing. 1988. Biodeterioration—multidisciplinary, collabora- tive research, pp. 215-226. [In] M.-F. Thompson, R. Sarojini, and R. Nagabhushanam (eds.), Marine Biodeterioration—Brooding in the Teredinidae (Mollusca: Bivalvia). Bombay, India: Oxford and IBH Publishing (C. B. Calloway, Ist author). 1988. Cellulolytic nitrogen-fixing bacteria in the Ter- edinidae (Mollusca: Bivalvia), pp. 743-748. [In] D. R. Houghton, R. N. Smith, and H. O. W. Eggins (eds.) Biodeterioration. Vol. 7. London and New York: Elsevier Applied Science. 1988. Recruitment of marine invertebrates to hard substrates at deep-sea hydrothermal vents on the East Pacific Rise and Galapagos spreading center. Deep-Sea Research, 35(10/11A): 1833-1849 (C. L. van Dover, lst, and C. J. Berg, 2nd authors) [No- vember]. 1988. Wood, phytoplankton, dissolved organic mate- rial and nitrogen in teredinid nutrition (Mollusca: Bivalvia: Teredinidae), pp. 585-606. [In] M.-F. Thompson and N. Tirmizi (eds.), Marine Science of the Arabian Sea. Proceedings of the Internation- al Conference. Washington, D.C.: Institute of Bi- ological Sciences. 1989. Bacteria for Cellulose Digestion. U.S. Patent 4861721. Assignee Research Corporation, Woods Hole, Massachusetts (J. B. Waterbury, Ist, and C. B. Calloway, 2nd authors) [29 August] [not seen]. 1989. The Genera Martesia and Lignopholas in the Indo-Pacific. (Mollusca: Bivalvia: Pholadidae) Ophelia, 30(3): 155-186 (L. N. Santhakumaran, 2nd author) [September]. 1990. [Abstract] Bivalves of hydrothermal vents and reducing sediments. Fourth International Congress of Systematic and Evolutionary Biology. University of Maryland: College Park, Maryland. July, 1990 (E. A. Cobabe, 2nd author). 1990. Species richness and diversity of algal-associ- ated micromolluscan communities from Sao Mi- guel, Acores. Acgoreana, (Suppl.): 39-58 (R. C. Bullock, Ist, and R. A. Fralick, 3rd authors). 1990. Xylophile ostracoda in the deep-sea, pp. 307— 319. [In] R. Whatley and R. F. Maddocks (eds.), Ostracoda and Global Events. Proceedings of the Aberystwyth Conference on Ostracods. (P. L. Stei- neck, Ist, R. F. Maddocks, 2nd, R. D. Turner, 3rd, C. Coles, 4th, and R. Whatley, 5th authors). 1992. Characterization and site description of Sole- mya borealis. (Bivalvia; Solemyidae), another bi- valve, bacteria symbiosis. Marine Biology, 112(4): 601-613 (N. M. Conway, Ist, B. L. Howes, 2nd, R. D. Turner, 3rd, J. E. Capuzzo, 4th, and C. M. Ca- vanaugh, 5th authors) [April]. 1992. [Abstract] Deep sea wood borers and ancient wrecks, p. 38. [In] Program and Abstracts for American Malacological Union—58th Annual Meeting, Sarasota, Florida, August 2—7, 1992. 1992. [Abstract] Types and prevention of biodeteri- oration in the deep sea. [In] Indo-United States Meeting on Recent Developments in Biofouling Control. Bangalore (Plenary Session: C) [not seen]. 1998. A new genus and five new species of mussels (Bivalvia: Mytilidae) from deep-sea sulfide (hydro- carbon seeps in the Gulf of Mexico). Malacologia, AQ G22) 16321121 (Re Es Gustatsonslist Re) slum er, 2nd, R. A. Lutz, 3rd, and R. C. Vrijenhoek, 4th authors) [17 December] 1998. Superfamily Pholadiodea, pp. 371-378. [In] P. L. Beesley, G. J. B. Ross, and A. Wells (eds.), Mol- lusca: The Southern Synthesis. Fauna of Australia. Vol. 5, part A. Melbourne: CSIRO Publishing. 2002. On the subfamily Xylophagaine (Family Pho- ladidae, Bivalvia, Mollusca). Bulletin of the Muse- um Comparative Zoology, 157(4): 223-308, 38 pls. [31 October]. Publications to which Clench, Turner, or Clench and Turner Made Contributions Academy of Natural Sciences of Philadelphia. Special Publication (Clench and Turner). Achatina. A newsletter of African non-Marine Mala- cology, Leiden. 1970-83 (1-12) [mimeographed] (Clench). American Journal of Tropical Medicine (Clench). American Malacological Union. Annual Reports and Bulletin (Clench and Turner). American Microscopical Society, Transactions (Clench). American Museum of Natural History, Novitates (Clench). American Society for Testing Materials. Special Tech- nical Publication (Turner). American Zoologist (Turner). Aquasphere. Journal of the New England Aquarium, Boston (Clench). Australian Journal of Marine and Freshwater Re- search (Turner). Australian Natural History (Turner). Australian Newsletter (New series). Malacological Society of Australia (Turner). Basteria. Tijdschrift van de Neederlandse Malacolo- gische Vereniging (Turner). Biological Bulletin (Turner). Biologiia Morya [Kiev] (Turner). 46 Bulletin Museum of Comparative Zoology, Vol. 158, No. 1 Breviora. Museum of Comparative Zoology, Harvard University (Clench). Caribbean Journal of Science (Clench). Carnegie Institution of Washington, D.C. (Clench). Conchological Club of Southern California, Minutes [mimeographed] (Clench). Copeia (Clench). Correspondentiblad van Nederlandse Malacologische Vereniging (Turner). Deep Sea Research. Supplement (Turner). Encyclopedia Americana (Clench). Encyclopaedia Britannica (Clench and Turner). Estuaries (Turner). Florida State Museum, Bulletin (Biological Series) (Clench and Turner). Freshwater Biology, John Wiley and Sons, New York (Clench). Freshwater Research (Turner). Harvard Alumni Bulletin, Cambridge, Massachusetts (Clench). Hawaiian Shell News (Clench). Institut Frangais d Afrique Noirre (Turner). Johnsonia. Museum of Comparative Zoology, Har- vard University, Cambridge, Massachusetts (Clench and Turner). Journal de Conchyliologie (Clench). Journal of Conchology (Clench). Journal of Crustacean Biology (Turner). Journal of Experimental Marine Biological Ecology (Turner). Journal of Fisheries Research Board, Canada (Turn- er). Journal of Molluscan Studies, formerly Journal of the Malacological Society of London (Turner). Journal of Shellfish Research (Turner). Journal of the Royal Society of New Zealand (Turn- er). Life Magazine (Clench and Turner). Limnological Society of America. Special Publication (Turner). Malacologia (Clench). Malacological Review (Turner). Malacological Society of Australia, Journal and News- letter (Clench). Malacological Society of London, Proceedings (Clench). Marine Biology (Turner). Massachusetts Audubon Society, Bulletin (Turner). Michigan Academy of Science, Arts and Letters, Pa- pers (Clench). Mollusca. Paul Reed Publisher, Tavares Florida, 1944-49, 1(1-12) to 2(1-6) [mimeographed] (Clench). Musée Royal d'Histoire naturelle de Belgique, Mé- moires and Bulletin (Clench). Museo Argentino Ciencias Naturales “Bernardino Ri- vadavia,” Revista (Clench). Museum of Comparative Zoology, Harvard Univer- sity, Cambridge, Massachusetts, Bulletin (Clench). Museum of the Institute of Jamaica, Occasional Pa- pers (Clench). Museum of Zoology, University of Michigan, Ann Ar- bor, Michigan, Occasional Papers (Clench). Natural History of Rennell Island, British Solomon Islands. Danish Scientific Press, Ltd., Copenhagen (Clench and Turner). Nautilus, The (Clench and Turner). Nemouria, Occasional Papers of the Delaware Mu- seum of Natural History (Clench). New England Naturalist (Clench). New England Zoological Club, Proceedings (Clench). New Hampshire Fish and Game Commission, Survey Reports (Clench). New Zealand Journal of Geology and Geophysics (Turner). Occasional Papers on Mollusks, Museum of Compar- ative Zoology (Clench and Turner). Oceanus (Turner). Ophelia, International Journal of Marine Biology (Turner). Philippine Journal of Science (Clench). Pittsburgh Shell Club Bulletin (Clench) Revista del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” (Turner) Revue de Zoologie et de Botanique Africaines (Clench). Robert S. Peabody Foundation for Archaeology, An- dover, Massachusetts (Clench). Science (Clench). Shell Case, The Naples (Florida) Shell Club [mim- eographed] (Clench). Shells and Their Neighbors, Redlands, California (Clench). Sociedad Cubana de Historia Natural, “Felipe Poey,” Habana, Memories (Clench). Sociedad Malacologica “Carlos de la Torre,” Revista (Clench and Turner). Soviet Journal of Marine Biology (Turner). St. Petersburg Shell News, St. Petersburg (Florida) Shell Club (Clench). Steenstrupia (Zoological Museum, University of Co- penhagen) (Clench). Sterkiana (Clench). Systematic Zoology (Clench). Torreia (Universidad de la Habana) Publication Oc- casional de Museo Poey (Clench). Turtox News (General Biological Supply House, Inc., Chicago, Illinois (Turner). Vassar Alumnae Magazine (Turner). Veliger, The California Malacozoological Society, Berkeley, California (Turner). Zoological Nomenclature, Bulletin (Clench). Rulletin OF THE Museum of Comparative — Zoology Comments and New Records for the American Genera Gea and Argiope with the Description of New Species (Araneae: Araneidae) HERBERT W. LEVI HARVARD UNIVERSITY VOLUME 158, NUMBER 2 CAMBRIDGE, MASSACHUSETTS, U.S.A. | 17 September 2004 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY BreEviora 1952— BULLETIN 1863— Memoirs 1865-1938 Jounsonia, Department of Mollusks, 1941-1974 OCCASIONAL PAPERS ON Mo.uusks, 1945— SPECIAL PUBLICATIONS. 1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and Evolution of Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere- dinidea (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino-. derms. 284 pp. 4. Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 725 pp. 6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the M. C. Z., Vol. 108.) Reprinted 1971. Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. GC. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter’s Check-list of Birds of the World, vols. 1-16. Proceedings of the New England Zoological Club 1899-1947. (Complete sets only.) Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2004. COMMENTS AND NEW RECORDS FOR THE AMERICAN GENERA GEA AND ARGIOPE WITH THE DESCRIPTION OF A NEW SPECIES (ARANEAE: ARANEIDAE) HERBERT W. LEVI ABSTRACT. ‘There are one species of Gea and seven species of Argiope in the Americas. Distributions of Gea and Argiope species are recorded with new re- cords from South America. Only one new species, A. ericae, from southern Brazil to northern Argentina, was found and is here described. The coloration of the female abdomen of Argiope species differs more than the visible differences of the genitalia. Males are separated by differences in palpi, the attachment of the embolus to stipes, and the shape of the coiled embolus. INTRODUCTION In 1968, I started my araneid revisions with the genera Gea and Argiope. These genera were first because they are the most distinct of the araneid genera, sepa- rated from most others by the presence of a procurved posterior eye row. At the time, I had mostly North American collections from the MCZ and AMNH. The individ- ual variation among specimens was aston- ishing, especially in the structure of the epigynum (these are not again illustrated here). At the time, Willis Gertsch (person- al communication) stressed that species could not be variable: when differences were found, they must indicate two spe- cies. I disagreed because when many spec- imens were examined, there were inter- mediates. These were the taxonomic issues of arachnologists in the 1960s. This is an update of the 1968 revision. Since 1968, many additional South Amer- ican specimens have become available, begging to be examined, especially in view ‘Museum of Comparative Zoology, Harvard Uni- versity, Cambridge, Massachusetts 02138. E-mail: levi@fas.harvard.edu of today’s better understanding of varia- tion. When I started this revision I ex- pected several species to be new, but I found only one. In the 1960s lack of funds limited the amount of data that could be included in the paper, so the localities of specimens were omitted, represented only by dots on a map. Although lengthy, both new localities and disposition of specimens are provided here, much abbreviated for common species, except for southern bor- der specimens. I have not repeated infor- mation published in 1968, except that es- sential for maps and diagnoses. At present, I have revised most Ameri- can araneid genera and have examined the Argiope of the Pacific area (Levi, 1983). The Chinese Argiope have been illustrated by Yin (1997), the African species by Bjgrn (1997), and the Mideastern species by Levy (1998). METHODS The methods used here were the same as in the revisions of other American ara- neid genera (Levi, 1993). Specimens came from the following col- lections: AMNH American Museum of Natural History, New York, United States; N. Platnick, L. Sorkin BMNH Natural History Museum, London, England; J. Becca- loni CAS California Academy of Scien- ces, San Francisco, California, Bull. Mus. Comp. Zool., 158(2): 47-66, Month, 2004 AT 48 Bulletin Museum of Comparative Zoology, Vol. 158, No. 2 United States; C. Griswold, D. Ubick Florida State Collection of Arthropods, Gainesville, Flor- ida, United States; G. B. Ed- wards Instituto Butantan, S4o Paulo, Brazil; A. Brescovit Institut Royal des Sciences Naturelles de Belgique, Brus- sels, Belgium; L. Baert Museo Argentino de Ciencias Naturales, Buenos Aires, Ar- gentina; C. L. Scioscia Museu de Ciéncias Naturais, Fundagao Zoobotanica do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil; E. H. Buckup, M. A. L. Marques Museu de Ciéncias, Pontificia Universidade Catélica do Rio Grande do Sul, Porto Alegre, Rio Grande do Sul, Brazil; A. A. Lise Museum of Comparative Zo- ology, Cambridge, Massachu- setts, United States Museo de Historia Natural, Medellin, Colombia; L. A. Za- mudio Museo de La Plata, La Plata, Angemomes, (Ci. ilar, Ib. vA\, Pereira Museu de Zoologia, Univer- sidade de SA0 Paulo, SAio Pau- lo, SP, Brazil; R. Pinto da Ro- cha Staatliches Museum fiir Na- turkunde Karlsruhe, Karlsru- he, Germany; H. Hé6fer National Museum of Natural History, Smithsonian Institu- tion, Washington, D.C., Unit- ed States; J. Coddington, S. F. Larcher Zoologisk Museum, Copen- hagen, Denmark; N. Scharff FSCA IBSP IRSNB MACN MCN MCP MCZ MHNMC MLP MZSP SMNK USNM ZMUC RESULTS Species Characters. The eye arrange- ment, procurved posterior eye row, and structure of the male palpus place Gea close to Argiope. Females of Argiope are easiest to deter- mine by their coloration as epigyna of many are quite similar (Levi, 1968). This differs from Bjgrn’s (1997) observations of African Argiope, which show much greater differences in genitalia as well as colora- tion. Extracting the broken male embolus tip stuck in the copulatory ducts inside of the epigynum is helpful for determining females in faded specimens. Males of Argiope can be separated by examining the palpus. The palpi of most species are much alike, differing, however, in the attachment of the embolus to the stipes (arrow, Fig. 25) and the shape and structure of the curved embolus. The relatively small differences between Argiope species in morphology of the gen- italia and the consistent differences in col- oration of the abdomen reminds one of similar differences in the theridiid widow spiders, Latrodectus (Abalos, 1980). Both genera also have a long, thread-shaped embolus that breaks off and plugs the epi- gynum. In both genera, relatively common species have been overlooked in well-col- lected areas. Gea heptagon is probably introduced from the South Pacific where it and other species of Gea are found. There are seven species of Argiope in the Americas, one in Europe, 13 in Africa and Madagascar, and about 29 in China. Only one, A. trifasciata, is cosmopolitan. The similarity of genitalia suggests all but A. aurantia are close to A. trifasciata. TAXONOMIC SECTION Gea C. L. Koch Gea C. L. Koch, 1843: 101. Type species Gea spinipes C. L. Koch, 1843: 101, pl. 823, from the East In- dies. Diagnosis. Ged species, together with those of Argiope, differ from most araneid genera by having the posterior eye row strongly procurved, from Mangora by the low thoracic region of the carapace and by lacking trichobothria on the third tibia, and from Mecynogea by the wider cara- pace and different structure of the palpus. Gea specimens are smaller than Argiope and have larger posterior median eyes. In females of Gea, the eyes of the posterior eye row are almost equally spaced (Fig. 5), whereas in Argiope, the median eyes are closer to each other than to the laterals. Gea heptagon (Heniz) Maps 1A; Figures 1-8 Epeira heptagon Hentz, 1850: 20. Type specimens destroyed, from North Carolina and Alabama. Gea heptagon:—Keyserling, 1892: 76, pl. 3, fig. 58. Levi, 1968: 324, figs. 1-24, 2 6. Platnick, 2003. Diagnosis. The small size of females, the shape of the abdomen (Figs. 4, 5), the epi- gynum with a ventral transverse slit and paired posterior openings (Figs. 1-3), and the male palpus with the embolus only gently curved, held by a large conductor (Fig. 8), distinguish the species from oth- ers. Differences of the first tibia are un- reliable. Variation. Total length of females 3.7— 6.4 mm, males 2.6—4.1 mm. The color of the abdomen is variable. There may be transverse lines or a dark folium. The il- lustrations of a female (Figs. 1-5) were made from a specimen from Florida, male (Figs. 6-8) from a specimen from Hon- duras. Natural History. Found in soybean field in Arkansas, in high grass in Florida. The female drops from the web when dis- turbed. Distribution. Pacific, probably intro- duced to America and found from eastern United States and West Indies (Map 1A), probably to Argentina. Additional Records. UNITED STATES, NORTH CAROLINA Dare Co.: Kill Devil Hills, 12 Sep. 1956, 12 (K. V. Krombein, USNM). GEORGIA Bullock Co.: Statesboro, 1984, 2 (E. Dismer, USNM). Chat- ham Co.: Rt. 95, nr. South Carolina border, 1 Aug. 1985, 2d (J. Coddington, USNM). ALABAMA Dal- las Co.: Selma, 2 (USNM). FLORIDA Franklin Co.: Su Neresay2o MarelIols 2 (|. ‘Carico, USNM). Sez Lucie Co.: Fort Pierce, 2 d (M. Mikkelson, USNM): 7 Sep. 1985, 2 (P. Mikkelson, USNM). ARKANSAS Gea and Argiope * Levi AQ Chicot Co.: 29 Aug. 1962, 2? (CAS). Mississippi Co: Big Lake, 23 June 1966, 3 (W. Peck, CAS). LOUI- SIANA Cameron Par: Cameron, 2 (USNM). OKLAHOMA Garfield Co.: Enid, 36°24'N, 97°54’W, 28 July 1989, 2 (L. E. Anhorn, USNM). CALIFOR- NIA San Diego Co.: Del Mar, 2 (M. Martinez, USNM); Los Angeles, 2d (USNM). MEXICO San Luis Potost: Huichihuayan, 98°50'N, 21°19'W, 24 July 1966, 3 Ge W. Ivie, AMNH). HONDURAS Atlan- tida: Lancetilla, 20 July 1929, 5 (A. M. Chickering, MCZ). Copan: Copan, 16 Feb. 1937, 6 (MCZ). COS- TA RICA Turrialba, 25-31 May 1962, 2 (H. Ruckes, AMNH). PANAMA Balboa, 1 Aug. 1943, S (J. B. Duncan, AMNH). BAHAMA ISLANDS Grand Bahama Island, 13 May 1953, 2 (L. Giovannoli, AMNH). DOMINI- CAN REPUBLIC Ciudad Trujillo [Santo Domingo], nest of Sceliphron wasp, 1947, 2d (H. F. Allard, USNM). PUERTO RICO SW Guayama, Pta. Pozue- la, 25 Dec. 1985, ¢ (V., B. Roth, CAS); Aguas Bueno, 5 Nov. 1971, imm. (J. Carico, USNM). VIRGIN IS- LANDS St.Croix, 1-11 Sep. 1966, 2 (A. M. Chick- ering, USNM). COLOMBIA Valle: nr. Cali, 1,000 m elev., 1977, 3 (W. Eberhard, MCZ); Palmira, 27 June 1964, ¢ (R. Hunter, CAS). ECUADOR Guayaquil, 22, Mar. 1942, 3 (Landes, CAS). PERU Piura: Hiquéron, July 1941, 6 (H. E., D. L. Frizzell, CAS). BRAZIL Mato Gros- so: S. Antonio de Levergera, 8 Sep. 1992; 2 (M. E. Marques, MCP 2558). Rio Grande do Sul: Guaiba, 1 Jan. 1989, 26 (A. B. Bonaldo, MCN 18009); 24-27 Dec. 1992, 26 (A. B. Bonaldo, MCN 22643); Mon- tenegro, 7 July 1977, 9 (H. A. Gastal, MCN 6177); Porto Alegre, 21) Sep. 1992, 6 (A. Bonaldo; MCP 2051); Taim, Rio Grande, 18 Mar. 1982, ¢ (J. Grazia, MCN 10247); Vila Nova, 23 May 1991, 2 (F. Garcia, MCP 2729); Xangrila, 5 Jan. 1993, 24 Feb. 1993, 2 (A. A. Lise, MCP 2902, 2974). ARGENTINA Spec- imens have been examined from northern Argentina without recording the data. Argiope Audouin Argyope Audouin, 1826: 121. Type species designat- ed by Thorell, 1869: 51, Argiope lobata from the Mediterranean and Africa. Argiope Audouin,1827: 328. Miranda C. L. Koch 1835: 128, pl. 14. Type species Miranda transalpina C. L. Koch (=Argiope bruen- nichi (Scopoli)). First synonymized by Thorell, 1869: 51. Metargiope F. P.-Cambridge, 1903: 451. Type species by monotypy Argiope trifasciata. Note. The International Commission on Zoological Nomenclature (Opinion 1038, 1975: 105) validated the name Argiope Au- douin, 1826, and placed it on the Official List of Generic Names in Zoology with the 50 Bulletin Museum of Comparative Zoology, Vol. 158, No. 2 Gea heptagon MEXICO \, A. aurantia Map 1. number 2009, and considered Argyope an incorrect spelling. Diagnosis. Argiope and Gea differ from most araneids by having the posterior eye row procurved (Figs. 12, 14), from Man- gora by their low thoracic region of the carapace and by lacking trichobothria on the third tibia, and from Mecynogea by a A. trifasciata Distribution of Gea and Argiope species. wider carapace and a different structure of the palpus. Argiope differs from Gea (Figs. 5, 7) by having the posterior median eyes smaller and closer to each other than to the lat- erals (Figs. 12, 14) and by females being larger (Fig. 12) than those of Gea. Unlike most araneids, the epigynum of A. argeniata A. ericae Gea and Argiope * Levi 51 A. savignyi A. blanda Map 2. Distribution of Argiope species. Argiope lacks a scape; it usually has a bulg- ing hood with a cavity (Figs. 9, 10) or cav- ities below (Figs. 18, 20). The male palpus has a separate sclerite, the stipes, between the radix and embolus (Figs. 161, 171). The palpus lacks a terminal apophysis and has an elaborate conductor (Figs. 16C, 17C) supporting the embolus (Figs. 16E, 17E). A thin curved branch extends from the median apophysis (Figs. 16M, 17M). i KEY TO FEMALES Venter of abdomen with a central transverse white band (Figs. 39, 49) that may be broken iiiice 50) Weasel ie taTee ibe ee het ote ae 5 Venter of abdomen without central transverse band (Figs. 11, 21, 30) or lines (Fig. 67) on black 2(1) Dorsum of abdomen with median black band (Fig. 12); epigynum a longitudinal bar (Figs. 9, 10); North America, Mexico (Map 1C) JN GYG (OL GAY SIT gO} d OVEN aK) ev oe ee 3 Dorsum of abdomen with transverse lines or fo tind sa CLhies B26 GS) fvate a ake Me Pe ea tee 4 Dorsum of abdomen with pair of posterior, lon- gitudinal bands (Fig. 31); SE United States (NA caro A) IE eres eat ES oid ow tae florida Dorsum of abdomen with transverse bands (Fig. 68), abdomen with pair of anterior tubercles and three lateral pairs of tubercles (Fig. 68); Sao Paulo State to NE Argentina (Map 2C) So ee Me ek Ee SUR one eS Sage tae eer aE ericae Dorsum of abdomen with transverse lines (Fig. 22); cosmopolitan (Map 1B) __.... trifasciata Transverse, ventral band wider than length of anterior median black trapezoid (Fig. 49); Texas to WentralvAmenca (Map): = blanda Transverse band narrower than length of black pate: ME 19s GSO). DG) as Slee hn A 6 Transverse band pointing anteriorly and broken in middle (Fig. 58); dorsum with posterior black (Fig. 59); epigynum enlarged posteriorly (Fig. 59, 56); Mexico to Argentina (Map 2B) 52 Bulletin Museum of Comparative Zoology, Vol. 158, No. 2 Pe ee a adem ee eC) RE AB 9 et ae = savignyt = Transverse band straight, rarely broken (Fig. 39); dorsum with posterior having windows in black area (Fig. 40); epigynum smaller posteri- orly (Figs. 36, 37); Florida, Texas, California to southern Brazil (Map 2A) argentata KEY TO MALES il Embolus straight (Fig. 15); North America, WWileseieo, GUMleyo) IC) oe aurantia = Embolus and conductor coiled (Figs. 16, PT 20s AAS ede oh a eee, ne 2) 2(1) Base of sclerotized embolus with jointed to large flat stipes (arrow, Figs. 25, 34) 3 = Base of embolus turning into a screw- shaped base (Figs. 43, 53, 62), stipes small, partly hidden (I in Figs. 16, 17) Curl of embolus small, showing stipes and tegulum on each side (Fig. 26); cosmopoli- tam02 6Z cas oyeula (Il = N) 09% + WLS 16-€8 a GS a) A A sjepneoqns (CR IND) ataan Ol SzleZ0 SLI == oyeule yy (Gil — aN) NO Sateen Gaevall OSI-LYT zoe SLI JPN spequo,A CL = WN) GIRLIE (OI OND Biles CT-SI-SIT CI-SI-¢ SQ[BOS [eBSIOC (SND avail Ga — oN RSI ial — yqee} Areypexe yy (G = N) €10'0 + €60 FS0-1Z-0 wall = ayeula (Ole — IN) SO0I0)==2G.0 830-960 ee 960 ss YA susy peyoYVUpSuyy] [lel (ie Q)aGy (9Z9) L&S a a[euloy JSST] (LPG) SSL = (60) 169 a[eur ysosIe'] (uu) (TAS) Asus] [POL a et 2 ae ee ee a de eS SS ee OPIS Apnys VURT, Oly o[vulot o[eul :(BOIVUIVL T)) TIO I1OpePnoy ‘vloT LOpenoy spa10 spsdic] (G89 NHNW) (GILOL dSNV) odAjojoy ta1puv snyjpusojdaT adAjojoy svaso spsdiq NN eee eee “HLONAT LNAA OL LAONS = TAS ‘SNOILVAYASAO HO SNAWIOddS AO HHANON = NT W1IaVL NI SV GHZIYVNWOAS Hav AIA NVS VNVZ OI AHL WOU VLVG ‘(LHOIW/LAAT) SNATIOS V Ad CaALVUVdAS AYV SHdALOTOH AHL HOA SLNNOO TWHALV ITA ‘(ATAINVS TVLOL GNV SNAWIONdS NVINOGVNOW) SVAYO ‘GT YOU VLVA GHLVINGVL XTALVYVdas WOH T ATAV EL AHS (ALIS AGNLS VNVZ, ON) AYAd WOH SVZYO ‘CG AO HIdINVS NOILLVINdOd V AUNV (QG-6P ‘GS6I ‘NOYWHONM WOW VLIVA -SVaYO “J =) AOVANVS IAYANV SQAHIVNDOOLdA’T UNV (adOD) SVAUO SVSdIG. AO SAdALOTIOH AHL WOH VLVA YAHLO CNV “CNW ) SLNANAYASVAIN “SLNQOO ATVOS (“¢ ATAVL JUL Dipsas oreas Complex in Ecuador and Peru * Cadle ‘Ysue] Apod o1QUS oY} 2[QISIA spuRq SurAry sfenpriArpur A;UO opnzour spueq Apoq Jo syunoy (CIR aN) eGGale eel Apoq uo soyoo[q Gall [Tg mnoqy (are /Spueq [esiop JO 1oquinyy (Giese) nel (Ol = N) GI G7 = IN) IE (tL = NY) OL OL/6 IT/11 seiqeyetyuy CW NE (Omigns 6 =N (G7) 8 6 = N (G-€) 8 I =N (¥-€) 8 8S=N(G¥F)L Ole NG (GaGa T=N(r£) Lz ; I = N 7€) 9 (G-€) LAG-€)2 (G-€)LS-€)L (e490 Suryono}) sperquyedng (VI = N)¥ (61 = N)€ (T=N)@ o/e B/E sjerodure, Arepuosesg (= N) & (Gita) me (ST =N)G (C=N)T L/I LI sjerodure, ABUL g (Si = MN) S (Si = N) (aa G/E C/G S.1e]Nd0}sOg ous Apnys vuez, ony o[euloy oyeul (RorrweleD) nog topenoy “vloy loprnoy spaso svsdiq, _ (S869 NHNI) (SLLOL dSNV) odAqojoy yaupup snyqwousojdaT adAyojoy svaso spsdigg ‘GHANILNOD ‘© ATAV 112 tal relationships might explain a pattern of intraspecific variation in the Dipsas oreas group, whereby loreal patterns | and 2 or, alternatively, loreal patterns 3 and 4 tend to be characteristic of a species (Table 1). All the scutellational differences and geographic trends are minor in view of the overwhelming similarities in both scutel- lation and color patterns between the Pe- ruvian and Ecuadorian specimens of Dip- sas oreas. However, the extensive variation in color pattern within and between pop- ulations of D. oreas in Peru (see discussion of color above) was not observed in spec- imens from Ecuador. It is evident that the status of Dipsas ellipsifera and D. elegans relative to D. or- eas has no bearing on the status of the populations of southern Ecuador and northern Peru here referred to D. oreas. All Peruvian specimens are referable to the nominotypical form of D. oreas, rather than to D. ellipsifera or D. elegans, as di- agnosed herein. Orcés and Almendariz (1987) noted that the range of “D. oreas oreas’ extended farther south than the other subspecies but indicated that its dis- tribution needed clarification. Their gen- eralization appears to hold with the addi- tional material examined for this study (Figs. 8, 23). Dipsas ellipsifera appears to be restricted to the Rio Mira system of northern Ecuador, and I am unaware of specimens of D. elegans from south of central Ecuador (Fig. 8). In contrast, nom- inotypical D. oreas has previously been known south almost to the Peruvian bor- der for more than a century (e.g., the ho- lotype of Leptognathus andrei Sauvage from Loja). Hence, D. elegans and D. el- lipsifera are not germane to evaluating the relationship of the Peruvian populations to nominotypical D. oreas. Distribution and Type Locality Dipsas oreas is distributed from approx- imately 2°10’S (southern Chimborazo Province, Ecuador) to the Rio Zafia in Ca- jamarca Department, Peru (6°51'S; Figs. 8, 23). Because of a copying error, Cadle Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 and Myers (2003:43, fig. 10) referred KU 142803 to Pichincha Province, Ecuador. The coordinates given for the locality are correct, but the province is Loja. Most specimens of Dipsas oreas come from the foothills and slopes of the west- ern cordillera of the Andes (elevational range 1,400—2,900 m). However, a true lowland locality is represented by a series (USNM 60006, 62797—02) from “Guaya- quil, Ecuador (sea level),” a locality that seems out of character not only in terms of elevation but also in terms of habitats at other known localities for this species (see Chapman, 1926, and subsequent dis- cussion of habitats). Cadle and Myers (2003) considered the records from Gua- yaquil “probably in error.” I am no longer convinced that the locality is erroneous, but only that it seems different from other known localities. The series was obtained from Dr. Frederick W. Goding, American Consul General in Guayaquil, and was cat- alogued about 1920. Goding obtained specimens of other species known from western Ecuador for the USNM during the same period. Several specimens of D. oreas from imprecise localities in the Rio Chanchan valley due east of Guayaquil (ANSP, MCZ, and UMMZ specimens) could be from <1,000 m elevation, and it is not possible to unequivocably rule out lowland localities for this species on the basis of present knowledge. At the Rio Zafia Study Site, the docu- mented elevational range of Dipsas oreas is 1,450—-1,800 m. From the distribution of appropriate habitats in this local area (as of 1993), the elevational range may be much greater (1,400—2,500 m). It is very likely that the distribution of D. oreas be- comes increasingly restricted to mesic and humid environments on the Andean slopes toward the southern end of its range be- cause the lowlands along the northern Pe- ruvian coast are primarily occupied by arid to xeric habitats (Chapman, 1926; Koep- cke, 1961). Nonetheless, small parcels of humid lowland tropical forests in Tumbes department could harbor populations of Dipsas oreas Complex in Ecuador and Peru * Cadle 1} s(n MET: Dipsas ellipsifera: 275 _ ty wie Dipsas elegans: kilometers Figure 23. Ecuador and northwestern Peru showing summary distributions of Dipsas ellipsifera and D. elegans (based on locality records in Fig. 8), and locality records for D. oreas in Ecuador and northern Peru. Stippled area, above 1,000 m; hatched area, above 3,000 m. Numbered localities for D. oreas in Ecuador: (1) Guayaquil; (2) Huigra/Rio Chiguancay and Rio Chanchan valley; (3) Alausi; (4) Velacruz; (5) Loja; (6) Rio Catamayo valley. In Peru: (7) Ayabaca and Cerro Aypate; (8) Canchaque; (9) Abra de Porculla; (10) Rio Zana Study Site. 114 D. oreas because another species, D. grac- ilis, has recently been discovered there (see later comments on D. gracilis). Some known or suspected erroneous geographic records of Dipsas oreas in the literature bear some commentary. Parker's (1934: 271, 1938: 444) records of “Dipsas mikanii oreas” from the province of Loja in southern Ecuador seem to be based, at least in part, on misidentified specimens in the The Natural History Museum in Lon- don (BMNH) that I refer to Sibynomor- phus petersi and S. oligozonatus (Cadle, unpublished data). Nonetheless, D. oreas is known from the province of Loja from other specimens (Clee USNM 98923, KU 142803, and the holotype of Leptognathus andrei Sauvage). Fowler (1913: 169) re- ferred three specimens to D. ellipsifera and D. andiana (ANSP 18117, 18120, 18123) that are here referred to D. oreas (Cadle and Myers, 2003: footnote 17, fig. 11). Peters’ (1960a: 94, map 8) record of D. oreas from “Rio Saloya” (Pichincha Province, Ecuador; Fig. 8: locality 7) is based on BMNH 1940.2.20.32, a juvenile here referred to D. elegans. Orcés and Al- mendariz (1987: fig. 1) attributed the same locality to “D. oreas oreas,” probably based on Peters’ record. Boulenger (1896: 454) referred four specimens to “Leptognathus mikanii, variant C (L. oreas, Cope)”; two of these (specimens a and b, BMNH 60.6.16.56, 60.6.16.63; Figs. 14, 22.) from unknown localities in western Ecuador are D. oreas, but specimens c and d are D. elegans (see synonymy and distribution in the D. elegans species account). On the other hand, Despax’s (1911: 36) record of “Leptognathus mikani” from Alausi (upper reaches of the Rio Chan- chan valley, Chimborazo Province, Ecua- dor; Fig. 8: locality 17) probably correctly refers to Dipsas oreas. Despax (1911) stat- ed that the specimen conformed to “Vari- ant C” of Boulenger (1896: 454; =“Lep- tognathus oreas” Cope). In fact, Boulen- gers “Variant C” included specimens of both D. elegans and D. oreas (see referred specimens in the synonymies given above Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 for each species). However, D. elegans is not known from the region of Alausi, whereas D. oreas has been obtained by other collectors from the Rio Chanchan valley (e.g, ANSP 18117, 18120, 18123: MCZ 17083; UMMZ 56491). Thus, it seems most likely that Despax’s specimen is D. oreas. With the above clarifications of litera- ture records, it is clear that the northern limit of the distribution of Dipsas oreas is at approximately 2°S latitude. No con- firmed records exist for D. oreas north of the Rio Chanchan valley in southern Chimborazo Province. Additionally, there are no documented areas of range overlap for any of the three species of the oreas group (Fig. 8; contra Orcés and Almen- dariz, 1987; Peters, 1960a). Dipsas ellip- sifera seems likely to be restricted to the Rio Mira basin and surrounding areas in northwestern Ecuador, whereas D. elegans is found to the south of that, along the western slopes of the Andes in northern and central Ecuador. Nonetheless, D. ele- gans is recorded from Pallatanga in the Rio Chimbo valley (Chimborazo Prov- ince), the river system just north of the Rio Chanchan valley from which D. oreas is known (Fig. 8: localities 14, 16, 17). The nearest localities for D. elegans and D. or- eas (Pallatanga and Alausf, respectively) are separated by only about 26 km straightline distance (but also by a ridge >3,000 m). Thus, if there are areas of sym- patry among any species of the oreas group, it seems most likely to occur be- tween D. elegans and D. oreas in the re- gion encompassing southern Bolivar/ Chimborazo Provinces and adjacent areas of Guayas Province, Ecuador. Similarly, northern Imbabura Province (Fig. 8) is a potential area of contact between D. ele- gans and D. ellipsifera. Most of the localities from which Dipsas oreas is documented are small remnants of once much more extensive humid forests (but see note below on a specimen from a dry semideciduous forest). In at least one instance, the locality near Porculla Pass Dipsas oreas Complex in Ecuador and Peru» Cadle (Piura department, Peru) documented by MVZ 119330 (a road-killed specimen ob- tained in 1969), the population of D. oreas might be extinct. I visited the site in 1994 in an effort to sample remnant humid for- ests previously known to be extensive in this area (Koepcke, 1961). However, the easily accessible area had been completely denuded, except for a few trees remaining in steep ravines; a few small and highly fragmented forest remnants remain in this area (Sagdstegui et al., “2003” [2004)]). Most of the area is now very dry, and slopes are covered with grassland and oc- casional shrubs. I doubt that D. oreas and some other herpetofauna of humid mon- tane forests of northwestern Peru (e.g., Cadle, 1989, 1991: Cadle and McDiarmid, 1990) can survive in such a habitat. The mid-1990s destruction of the forest of Monte Seco was mentioned above (see Materials and Methods). With continuing fragmentation and obliteration of these unique environments, endemics such as D. oreas will survive in increasingly smaller ranges or go extinct. Curiously, despite the work of many herpetologists and other field biologists in western Ecuador, Dipsas oreas is known from relatively few specimens. Orcés and Almendariz (1987) reported that the EPN collection contained no specimens, and Kofron (1982) stated that only eight spec- imens were known at that time. Recent brief biological surveys in southwestern Ecuador (e.g., Almendariz and Carr, 1992) yielded no specimens of D. oreas. There is a distinct distributional gap between spec- imens collected in southern Chimborazo and Guayas Provinces (Rio Chanchan val- ley and Guayaquil, respectively) and Loja Province to the south. However, few com- prehensive collections are available from this area. Clarification of the Type Locality: Or- tons “Valley of Quito.” James Orton was a naturalist and professor who obtained sig- nificant collections on several lengthy ex- peditions to South America, primarily Ec- uador and Peru, in the 1860s and 1870s 115 (xrtonnels7 >= GadlenGdgsow422= 123) briefly summarized the Orton expeditions. The herpetological collections were turned over to Edward D. Cope of the Academy of Natural Sciences in Philadelphia, who catalogued the results of Orton’s expedi- tions in a series of important papers on the herpetology of South America (Cope, 1868, 1874, 1876, 1877). Many species de- scribed by Cope (1868), such as Dipsas or- eas, have the designated type locality “the elevated Valley of Quito,” a vague state- ment that has been narrowly interpreted by most subsequent workers as the valley immediately surrounding the city of Quito, Ecuador (e.g., Peters, 1955: 347). Hence, “Valley of Quito” has often been ques- tioned as a type locality for many reptiles described from Orton’s collections because they are species unlikely to have ever oc- curred in that vicinity. However, Orton himself makes clear that his meaning was much broader, as noted by Orcés and Al- mendariz (1987) and indicated in this quo- tation from Orton (1871: 620) and further amplified in his book (Orton, 1875: 152 fia) The Valley of Quito ... is a remarkably well- defined district, having a uniform temperate cli- mate. It is nearly three hundred and fifty miles in length, stretching from 1°N. to 4°S., and has an average width of forty miles, being walled in by the grandest group of volcanic mountains in the world. These barriers have an average elevation of 12,000 feet above the sea, and are broken at few points, chiefly by the narrow gorges of the Santiago and Pastaza, and the sources of the Mira and Esmer- aldos. The valley is subdivided by ridges into three basins—Quito, Ambato and Cuenca, having the re- spective altitudes of 9500, 8000 and 7500 feet... . Thus, Orton’s “Valley of Quito” com- prised the entire Ecuadorian highlands from the Colombian border to the vicinity of Loja in southern Ecuador. The species described from this “locality,” including Dipsas oreas, could have come from vir- tually anywhere in the Ecuadorian high- lands or their western slopes. Cope (1868: 96) adds the following details, obtained di- rectly from Orton, concerning the collec- tions of 1867-68: “|The expedition] first ILI touched the continent at Payta, Peru [=Paita], and afterwards at Guayaquil; then proceeded inland over the Andes to Quito; thence [eastward to the Rio Napo].... They collected Reptiles chiefly from Gua- yaquil, Pallatanga (on the west slope of the western Cordillera south of Chimborazo); Ambato (in Valley of Quito); western slope of the volcano Antisana, 13000 ft. above sea (a small black frog)... .” A similar route be- tween Guayaquil and Quito was followed by later naturalists (e.g., Samuel N. Rhoads) and resulted in specimens of D. oredas (Oey ANSP IS, 18120) and Wsil23 obtained by Rhoads; MCZ 17083, UMMZ 56491). Fowler (1913: 153) says of the Rhoads collection that it “... was made on the western slope of the Andes, from Gua- yaquil to Quito. ...” Thus, the Orton ex- pedition of 1867-68 crossed that portion of southern Ecuador from which D. oreas is well known (Fig. 8) and followed a route commonly used in the 18th and early 19th centuries. The holotype of D. oreas was most likely obtained en route between Guayaquil and the upper slopes of the Cor- dillera Occidental directly to the east. Of the localities specifically mentioned by Cope (1868) and quoted above, D. oreas is known from subsequent collections at Gua- yaquil, but Pallatanga represents the south- ernmost locality for D. elegans (Fig. 8: lo- cality 14), from which D. oreas is not doc- umented. Apart from species recorded from Gua- yaquil, virtually none of Orton’s specimens from western Ecuador reported by Cope (1868) have locality data more precise than “Valley of Quito.” In a curious paradox, Orton (1871: 619) began his paper with a lengthy discussion of the importance of ac- curate localities and the failure of previous field workers in Ecuador (e.g., Fraser) to supply accurate specimen data: “. . . we are still very far from knowing the precise range of species. This has arisen partly from the failure of collectors to give exact localities.” Thus, although it is clear what Orton himself meant by “Valley of Quito,” that understanding does not further elu- Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 cidate the geographic origin of the species described from this type locality. Natural History: Habitats, Seasonal Activity Patterns, Eggs, and Hatchlings Habitats and Seasonal Activity Patterns. In northern Peru Dipsas oreas seems to be primarily an inhabitant of humid montane forests, although it occurs in secondary forest habitats if these are recent or adja- cent to existing primary forests. MUSM 16750 was collected from an area of sec- ond growth at the base of Cerro Aypate (Piura department), which still had a frag- ment of humid forest on its crest in 1991 when the specimen was collected. Alwyn H. Gentry, who accompanied me to Cerro Aypate in 1991 classified the forest as “bmh-MT” (very humid montane forest) in the Holdridge Life Zone System (Gen- try, 1995: “Cerro Aypate, Piura” in table 1). At the Rio Zafia Study Site, D. oreas was collected from 1,450—1,800 m eleva- tion and always inside or at the edge of primary montane humid forest. The lowest elevation constitutes the lower edge of the humid forest belt as it existed in 1987—90. Appropriate habitat at the Rio Zafia Study Site occurred up to approximately 2,500 m, which might be the upper elevational extent for D. oreas in that local area. During extensive field work in other ar- eas of northern Peru, I did not encounter individuals in subhumid or arid habitats, which are common throughout the mon- tane regions of northwestern Peru. How- ever, a specimen from southern Ecuador was found “in a bromeliad, by day, in dry semi-deciduous thorn forest” (field notes of Linda Trueb for KU 142803). Thus, the habitat breadth of Dipsas oreas may be greater than suggested by my field expe- riences, which primarily derive from the Rio Zafia Study Site. Nonetheless, all the known localities in northern Peru are areas in which fragments of humid montane for- est are known to have occurred, at least historically (see Specimens Examined and Locality Records and Koepcke, 1961). Wet and dry seasons at the Rio Zana Dipsas oreas Complex in Ecuador and Peru ¢ Cadle Study Site are very pronounced. Except for egg clutches found during the early dry season, I found Dipsas oreas only during the rainy season (approximately December or January to March or April) at the Rio Zaha Study Site, although MUSM 16750 was collected 24 September 1991 at a site farther north. The contrast in activity pat- tern is best seen by comparing encounters during two field seasons at the Rio Zana Study Site. In nearly 2 months of contin- uous field work during May and June (ear- ly dry season), no D. oreas (except eggs) were encountered, whereas in only 2 weeks of field work from the same field camp in January (early rainy season), 10 adults and several juveniles were found. Few collecting dates accompany other specimens of D. oreas, but specimens have been obtained in July (KU 142803) and August (MVZ 119330). The climatic con- ditions that affect seasonal activity in D. oreas are probably very localized. The Rio Zana Study Site is the southernmost local- ity known for the species, and it is prob- ably more strongly seasonal than localities farther north. These observations suggest that Dipsas oreas is surface active at the Rio Zana Study Site only when ambient humidity is relatively high. During the dry season (roughly June to September), it may con- ceal itself in inaccessible retreats deep un- derground. Large boulder jumbles offer- ing deep crevices and large surface boul- ders are common at the Rio Zafia Study Site. Anecdotal observations (e.g., the pro- pensity to dry out when kept in cloth bags in the field) suggest that, like some other species of Dipsas, D. oreas is particularly prone to desiccation (personal observa- tions). This probably explains why no spec- imens were encountered during 2 months of field work during the early dry season. Temperature could also be a factor, for al- though average nocturnal (minimum) tem- peratures are only slightly higher in the rainy season compared with the dry sea- son, the dry season was characterized by less night to night temperature equability LILY and greater frequency of lower minimum temperatures than the wet season (see Ma- terials and Methods). The rainy season also corresponds to the period when gastropod prey of Dipsas are most active on the sur- face (see next section). Diel Activity Patterns, Diet, and Defen- sive Behavior. At the Rio Zaiia Study Site, Dipsas oreas is active nocturnally and en- countered in saplings or shrubs. All active adults were found less than 2 m from the ground, and most were encountered on nights of high humidity or light rain. This species possibly never ascends to great heights in vegetation. Inactive individuals were encountered during the day on or below the surface of the ground in the fol- lowing circumstances summarized from my field notes (all except MUSM 16750 from the Rio Zafia Study Site). ANSP 31779-81, 31783-—86 (adult males and one adult female; see subsequent section on aggrega- tion behavior): In well-shaded montane humid for- est on the surface of leaf litter or beneath the sur- face litter at 1015 hr, 25 January 1989. ANSP 31777 (adult female): Under a rock (40 cm diameter) in undisturbed montane humid forest at 1730 hr, 13 January 1989. MUSM 5532 (near-hatchling, 178 mm SVL): Under a rock in the morning at the lower edge of the humid forest, 15 January 1989. MUSM 16750 (adult female; Fig. 16): In a crev- ice in hardened lateritic soil during the day on 24 September 1991 (corresponding to the late dry sea- son). The crevice was in a road cut in disturbed habitats at 1,950 m; it measured 2 cm wide at the ground surface, was >50 cm deep, and was covered by dead brush above. The snake was coiled about 4—5 cm below ground level and about 20 cm from the surface in a horizontal direction. The location within the crevice where the snake was found was quite dry, although it was much wetter deeper within the crevice. In addition to these observations sug- gesting that Dipsas oreas retreats under- ground or under surface objects when in- active, several clutches of D. oreas eggs were found within a crevice in the ground at the Rio Zafia Study Site (see Eggs and Hatchlings). On the other hand, a speci- men from a dry thorn forest in southern Ecuador was found in a bromeliad (KU 142803). 118 Species of Dipsas are well known to prey upon terrestrial gastropods (e.g., Cunha and Nascimento, 1993; Martins and Oliveira, 1998; Peters, 1960a; Sazima, 1989). A single food record is available for D. oreas from the Rio Zana Study Site. The posterior portion of the intestine of ANSP 31777 was packed with slug and/or snail remains. Terrestrial and arboreal gas- tropods were both abundant and diverse at this locality during the rainy season. Dipsas oreas exhibits defensive postur- ing common among species of Dipsadini (Cadle and Myers, 2003), including head triangulation, raising the anterior part of the body, and bringing the head and neck back into an S-shaped loop (Fig. 15). In- dividuals occasionally added an additional bend to the loop and the body was often “anchored” with a loose coil. Although some individuals would occasionally crawl while holding this posture, none could be induced to strike. The posturing and head triangulation de- fenses of Dipsas oreas resemble the defen- sive behaviors of some vipers (Greene, 1988) and thus are potentially a form of behavioral mimicry, and perhaps the band- ing pattern of D. oreas could resemble that of some vipers to certain predators. How- ever, vipers are said by local residents not to occur at the Rio Zafia Study Site, and none were recovered in my herpetofaunal sampling there (residents report “viboras” at lower elevations toward the coast; at least Ole WspeCcies: Bothrops roedingeri, poten- tially occurs there, although this species is documented only farther south in the Pe- ruvian coastal region; cf. Campbell and La- mar, 2004). In other parts of the range of D. oreas, a possibility exists that one or more viperid species is sympatric with D. oreas, judging from distribution maps in Campbell and Lamar (2004); however, such sympatry remains to be demonstrated. The only venomous species potentially occur- ring sympatrically with D. oreas at the Rio Zana Study Site is Micrurus mertensi, which is a coastal/foothill species (Camp- bell and Lamar, 2004) that is recorded in Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 my collections from 1,300 to 1,400 m at the site, just short of the lowest recorded ele- vation for D. oreas (1,450 m). However, there is (to my perception) no particular resemblance between these two species. Comparisons of Behavior with Other Species of Dipsas. Species of Dipsas occur in tropical dry forests and rain forests from central Mexico to the subtropical zone of southern Brazil, thus inhabiting an array of relatively aseasonal to highly seasonal en- vironments. As indicated for the Rio Zaiia Study Site, seasonal activity correlated with the onset of rains or high ambient humidity has been reported in other spe- cies of Dipsas inhabiting seasonal environ- ments. For example, Hartmann et al. (2002) found that adult D. albifrons were active only during the rainy season in the Atlantic forest of southeastern Brazil (ju- veniles seemed less seasonal in their activ- ity). On the other hand, a related snake from the same region, Sibynomorphus neuwiedi, is more active during the cooler, drier season (Marques et al., 2001). Activ- ity patterns of species of Dipsas from less seasonal rain forests in equatorial Amazon- ia do not show strong seasonality (e.g., Martins and Oliveira, 1998). Cadle and Chuna (1995: footnote 5) commented on the seemingly paradoxical behavior of an arboreal snake such as Dip- sas oreas retreating to the ground surface or subsurface during its inactive period, rather than seeking arboreal retreats. However, this behavior could be frequent in both montane and lowland members of the genus, as the following remarks indi- cate for several species. Dipsas catesbyi: At La Selva Lodge (Rio Napo, Ecuador) I found an adult Dipsas catesbyi (not collected) under dead leaves in deeply shaded forest at 1500 hr. At the same locality another individual encoun- tered while active in low vegetation at night dropped to the ground after several minutes of observation and hid under- neath the leaf litter. Comments associated with other specimens include “on ground Dipsas oreas Complex in Ecuador and Peru ¢ Cadle at night” (FMNH 165846; Colombia, Pu- tumayo department) and “Under banana leaves on ground [presumably during the day] (FMNH 39640; Peru, Ayacucho de- partment). Dipsas indica: FMNH 165847 (Colom- bia, Putumayo department) was “on ground at daytime” (remarks in FMNH catalogue entry). Dipsas latifasciata: LSUMZ 45499 (Peru, Pasco department) was “caught on ground of second growth at edge of humid forest in late afternoon.” (remarks on field tag). Dipsas peruana: USNM 299232-34, MUSM-JEC 6750 (Peru, Puno depart- ment). I collected these specimens inac- tive under rocks during the day. This lo- cality was a somewhat dry rain shadow val- ley with xerophytic vegetation and few epi- phytes to create arboreal retreats. Observations of terrestrial activity for other species of Dipsas seem rarely to have been reported. Duellman (1978) re- ported terrestrial and arboreal activity for D. catesbyi in Ecuador. Orcés and Almen- dariz (1987) reported, without stating whether the snakes were active or inactive, that neither D. ellipsifera nor D. elegans seemed to be arboreal snakes. At least one species, D. pavonina in central Amazonia, may be primarily terrestrial when foraging at night (Martins and Oliveira, 1998), al- though Duellman (1978: 240) reported ar- boreal activity for this species in western Amazonia. These examples suggest that diel movement between terrestrial retreats and arboreal microhabitats when noctur- nally active may be common behaviors in some species of Dipsas, at least at some localities. Other species may even be ter- restrial while active.° Conversely, many di- ° Movement between terrestrial retreats and arbo- real active sites may be more widespread in tropical nocturnal, arboreal snakes than is currently recog- nized. In Madagascar, the colubrids Geodipsas la- phystia and G. zeny are similar to species of Dipsas, in being nocturnally active in low vegetation (Cadle, 1996). However, both species are occasionally found hidden in moist leaf litter on the ground during the day (personal observations). EIS) urnal terrestrial snakes ascend vegetation to sleep at night, perhaps in response to carnivorous ants (Martins, 1993). At the Rio Zafia Study Site, ants did not seem es- pecially prevalent compared with my ex- perience in lowland Amazonian rain for- ests; diurnal colubrids of genera known elsewhere to sleep in vegetation were nev- er encountered sleeping in vegetation at the Rio Zafia Study Site (e.g., Chironius, Dendrophidion, Leptophis, and Mastigo- dryads). The factors that might affect where a snake seeks refuge during its inactive pe- riod are numerous, and individual, season- al, geographic, and species-specific pat- terns of behavior are possible. Geographic trends may be related to the availability of arboreal retreats such as epiphytes—epi- phytes are much more abundant in contin- ually wet tropical rain forests than in more seasonal or subhumid forests (e.g., Myers, 1969). Bromeliads and other epiphytes are common at the Rio Zafia Study Site, so the propensity for Dipsas oreas at that site to use terrestrial retreats when inactive is not because of the lack of appropriate arboreal ones. Bromeliads were routinely searched at the Rio Zafia Study Site but yielded only frogs (Eleutherodactylus and Gastrotheca), and these only during the rainy season. The extended dry season at the Rio Zafia Study Site results in low ambient moisture and humidity within the forest, and the surface of the soil, the leaf litter, and even many bromeliads become very dry. During the dry season, some streams at the Rio Zafia Study Site temporarily cease flow, and the humidity is low enough that des- iccation becomes a problem for the aerial egg clutches of centrolenid frogs laid to- ward the end of the rainy season (Cadle and McDiarmid, 1990). The factors com- bined probably explain the terrestrial se- clusion behavior at this locality for D. or eds. Although the observations reported here suggest that terrestrial seclusion for several species of Dipsas occurs with some frequency, other observations suggest that 120 arboreal seclusion also occurs within Dip- sas. Porto and Fernandes (1996) reported an inactive D. neivai found within a bro- meliad on a tree branch close to the ground, and I collected an inactive D. ver- miculata (MVZ 163259) from a bromeliad 2 m from the ground during the day. Park- er (1926) reported that the type of D. var- iegata trinitatis was obtained from a bro- meliad on a fallen tree. Charles W. Myers collected several inactive D. temporalis from bromeliads on the sides of trees 1.2— 3 m aboveground during the day (Cadle and Myers, 2003: 36). Eggs and Hatchlings. Cadle and Chuna (1995: 32-34) reported communal egg-lay- ing in Dipsas oreas (21 total eggs), which in that instance was coincident with a large number of communally laid eggs of the gymnophthalmid lizard, Macropholidus ruthveni. The eggs, discovered 17 June 1987 at the lower edge of the humid forest (1,490 m) at the Rio Zafia Study Site, were underground in a moist crevice within a road embankment. The eggs were 20—30 cm from the entrance to the crevice and 15 cm beneath the surface of the soil (the crevice ran somewhat parallel to the soil surface); dimensions of the Dipsas eggs were 28-30 X 20 mm. On the basis of their clustering within the crevice, the snake eggs probably represented three clutches of eight, seven, and six eggs. One egg was opened at the time from each of the two deepest clutches and contained advanced embryos of slightly different stages. Five of the eggs were transported in plastic bags containing moist paper tow- els back to an air-conditioned office, where they were kept in similar fashion on a ta- ble. These eggs hatched between 23 Sep- tember and 1 October. Hatchlings GIVEN TEIN 232570=725 = VIUSINi G7 Sil 52) were 214-225 mm total length (172-176 mm SVL) and 2.8—3.2 g. A hatchling from this clutch is illustrated in Figure 17, and another was illustrated by Cadle and My- ers (2003: fig. 11). Two hatchlings or near-hatchlings of Dipsas oreas (MUSM 5530, 5532; total Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 lengths 230-243 mm) were collected 13- 15 January 1989 near the site where the above eggs were discovered. It seems like- ly that these hatchlings derived from eggs laid the previous dry season, as indicated by the eggs discovered in June and their slightly larger size than the above series of hatchlings. Two other females of Dipsas oreas from Ecuador were gravid, and clutch sizes were estimated by palpation through the ventral body wall. ANSP 18117 (421 mm SVL) contained four eggs. BMNH 60.6.16.56 (566 mm SVL) contained about 8 eggs. No collection dates are available for these specimens. Aggregation Behavior in Dipsas oreas A peculiar aggregation of Dipsas oreas was discovered on 25 January 1989 at 1,800 m at the Rio Zafia Study Site. At 10: 15 a.m., a field assistant found a ball of four snakes on the surface of the leaf litter adjacent to a small log (10-15 cm diame- ter, 2.5 m long) in a somewhat disturbed patch of cloud forest; all of these snakes were intertwined in a tight ball. Another individual was on the leaf litter surface about 20 cm from the ball. After alerting me to the discovery we returned to the site and found another snake hidden under the leaf litter (15-20 cm deep) where the ini- tial ball was found. At the opposite end of the log (2.5 m from the ball), another in- dividual was hidden under the leaf litter. A search of the immediate area (surface and under logs, litter, etc.) turned up no more snakes. The night before finding the aggregation included a light drizzle for several hours; on the morning of the en- counter, the leaf litter was wet and the day was overcast. Temperature at 11:00 a.m. Weasel Ge All snakes were quiescent and loosely or tightly coiled. The four snakes in the initial ball were completely and tightly inter- twined, and consisted of an adult female and three adult males. The three individ- uals not associated directly with the ball were adult males. The female is ANSP Dipsas oreas Complex in Ecuador and Peru ¢ Cadle 31784 and the males are ANSP 31779-81, 31783, 31785—86 (497-551 mm SVL; see Figs. 18, 19). Ageregations of snakes occur for a va- riety of reasons, including hibernation (in temperate snakes), response to unfavor- able temperature or humidity conditions, feeding, mating, and oviposition (e.g., Amr et al., 1997; Ford and Burghardt, 1993: 139-140; Gillingham, 1987: 197-198; Gregory et al., 1987: 370-373; Halpern, 1992; Noble and Clausen, 1936). The ag- gregation of Dipsas oreas just described seems unlikely to be due to unfavorable environmental conditions, hibernation, or feeding. The rainy season at the Rio Zana Study Sites commences in December or early January (as in 1989) and corresponds to the most active and environmentally fa- vorable period for D. oreas and its gastro- pod prey. On the other hand, several ob- servations suggest an association with re- productive behavior for the aggregation: all individuals are adults and all except one are males. This suggests that the aggrega- tion may have been for mating, as does the morphology of the reproductive tracts de- scribed below. Unfortunately, the aggre- gation was disrupted before specific be- haviors such as copulation or combat could be assessed. The fact that the aggregation was discovered during the day in a snake typically nocturnally active prompts ques- tions that only additional serendipitous field observations could answer: Was this aggregation simply a continuation of court- ship activity that had begun the previous night? Alternatively, do males locate fe- males during the day? Gregory et al. (1987: 370) defined ag- gregations as “any concentration of snakes in a relatively small area such that the den- sity of snakes in the aggregation contrasts sharply with that in the surrounding area.” Thus, aggregations need not involve con- tinuous physical contact among individu- als. In some snakes, males in mating ag- gregations are not necessarily continuously associated with the female, such as three of the males in the aggregation of Dipsas 121 oreas just described. For example, Feio et al. (1999) observed six male Chironius fla- volineatus courting a female, but only two were ever directly associated with the fe- male, the other males being nearby on the ground or low bushes. Thus, even though three males were not directly associated with the tight cluster of D. oreas, it seems very unlikely that they were only coinci- dentally close by. That the aggregation of Dipsas oreas de- scribed was associated with reproduction is also suggested by the morphology of the reproductive tracts of the female and males. The female (ANSP 31784) has small ovarian eggs (2-4 mm) in her left ovary. These are somewhat yellowish, per- haps indicating the initiation of yolk supply to the ova. Her left oviduct is extremely dilated (3.5—4 mm broad and thrown into many pleats), which is the condition ob- served in female snakes of many species prior to ovulation and fertilization (review in Fox, 1977). The middle portion and cra- nial end of the left oviduct, corresponding to the tube portion of the oviduct (Guil- lette et al., 1989), are particularly dilated and pleated. The right oviduct is less di- lated than the left and not at all pleated. Except for a single ovum similar in size to those in the left oviduct, eggs in the right ovary are smaller and paler than those in the left ovary. These observations suggest that the left reproductive tract of the fe- male was gearing up for reproduction. Moreover, the size of the ova in the left ovary correspond to follicle size classes I (“early growth”) to III (“hydration and ini- tial vitellogenesis”) recognized for D. ca- tesbyi by Zug et al. (1979: 2-3), which cor- roborates the other indications of repro- ductive competence of this female. The males in the aggregation have swollen and highly convoluted vasa deferentia, which are characteristic of male snakes during the breeding season. Because so little is known of reproduc- tive cycles and behavior of tropical snakes, it is worth exploring the details known for Dipsas oreas at the Rio Zafia Study Site 122 and attempting to paint a scenario of what seems to occur. At the Rio Zafa site, eggs of D. oreas with well developed embryos were discovered in mid-June in a well-se- cluded, relatively cool location deep within a crevice (details reported above). The ad- vanced developmental stages of two em- bryos suggest that the eggs were laid some time prior to their discovery in mid-June. This observation, combined with the ob- served aggregation of several adult males and an adult female in late January, sug- gests that a mating period in late January may not be unreasonable. Moreover, the condition of the oviducts in the female and the vasa deferentia in the males suggest that these individuals were reproductively competent. The delay between the pre- sumed mating period and egg laying would give ample time for fertilization and yolk- ing of the ova. In a related colubrid, Sibynomorphus ventrimaculatus, mating apparently takes place several months (perhaps as many as seven) before vitellogenesis commences and egg-laying occurs (Cechin and Oliv- eira, 2003). A similar sequence in Dipsas oreas would explain the small follicles in the female described above well before the inferred oviposition season. The strong seasonal activity pattern of D. oreas at the Rio Zafia Study Site, as well as the few observations on eggs and hatchings, imply that reproduction is strongly seasonal in this species, at least in the southern part of its range. Other species of Dipsas that inhabit less seasonal environments of Amazonia or the Atlantic Forest of south- eastern Brazil are apparently aseasonal in their reproductive patterns (Zug et al. [1979] for D. catesbyi; Porto and Fernan- des [1996] for D. neivai), and some other members of the Dipsadini have extended periods of oviposition (e.g., two species of Sibynomorphus; Laporta-Ferreira et al., 1986). Aggregation behavior is best known in north temperate species of natricine col- ubrids and viperids, although scattered ob- servations of scolecophidians, elapids, and Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 other groups of colubrids suggest that ag- gregation behavior is taxonomically wide- spread. Aggregations have previously been reported in three other species of Neo- tropical colubrids. An aggregation of Rhadinaea calligaster (Myers, 1974: 221) in a pile of decomposing thatch consisted of six females and a male. Females were apparently attracted to the site for ovipo- sition because a communal nest was in the same pile of debris. The other cases in- volve aggregations for mating functions in snakes of the clade Colubrinae (sensu Ca- dle, 1984c, 1985): Drymobius margariti- ferus (Campbell, 1998: 213) and Chironius flavolineatus (Feio et al., 1999). The ob- servations reported here for Dipsas oreas constitute the first report of aggregation behavior in any species of Dipsadini (Dip- sas, Sibon, Tropidodipsas, Sibynomor- phus) and one of only three cases of mat- ing aggregations in Neotropical colubrids. Because communal nests are now known in two distantly related members of the larger Neotropical clade Dipsadinae (sensu Cadle, 1984b; Zaher, 1999), D. oreas and Rhadinaea calligaster, aggregation behav- ior for oviposition may be widespread within this group if particular oviposition sites offer favorable conditions for egg de- velopment. TAXONOMIC AND GEOGRAPHICAL NOTES ON DIPSAS GRACILIS, D. LATIFASCIATUS, AND D. LATIFRONTALIS While studying the systematics of spe- cies in the Dipsas oreas complex and oth- ers in western Ecuador and Peru (e.g., Ca- dle and Myers, 2003) I had occasion to examine specimens of other species re- ported from the eastern and western ver- sants of the Andes in this region. In a few cases, the specimens examined amplified distributional knowledge of species or raised questions concerning systematic and nomenclatural issues. I here comment on these points for three species. Dipsas gracilis has been well known from the western lowlands of Ecuador, but new in- Dipsas oreas Complex in Ecuador and Peru « Cadle formation extends its range into Peru. Dip- sas latifasciatus and D. latifrontalis have been reported from eastern Ecuador and/ or Peru, but my examination of specimens disclosed some systematic and nomencla- tural problems relative to the treatment of these species in the literature. Detailed lo- cality data on specimens of these taxa I examined are given in the Specimens Ex- amined and Locality Records. See Note Added in Proof, p. 136. Dipsas gracilis Dipsas gracilis (Boulenger, 1902) was not recorded south of Guayaquil, Ecuador (Peters, 1960a) until Tello (1998) reported it from a lowland (620 m) evergreen rain forest in extreme northwestern Peru on the basis of MUSM 17589 (Quebrada de los Naranjos, Tumbes department). I pro- visionally refer two additional specimens from northern Peru to D. gracilis, MUSM 2700-01 (Rio Quiroz valley, Piura depart- ment). Although D. gracilis thus far has been reported in detail only from western Ecuador (Peters, 1960a; Peters and Ore- jas-Miranda, 1970), Perez-Santos and Mo- FenOmelsss lg)» neported) the ispecies from Colombia without providing docu- mentation. A specimen of Dipsas that I ex- amined from northwestern Colombia near the Panama border (FMNH 74376) had been previously identified as D. gracilis, but I tentatively refer this specimen to D. viguieri; see comments in the following section (key to species) concerning some difficulties in distinguishing D. gracilis and D. viguieri. | here comment on character- istics of the Peruvian specimens of D. gracilis, whose scutellation and other char- acters are summarized in Table 4. Unfor- tunately, my examination of MUSM 17589 was brief, and basic data were not com- pleted, although the specimen is well pre- served. All three Peruvian specimens are males, so the most relevant comparison would be males of Ecuadorian D. gracilis (Table 4). MUSM 17589 was obtained during a bi- ological inventory of the lowland tropical 123 forests of Tumbes department (Wust, 1998a). It is an adult male in excellent con- dition with well-everted hemipenes. The specimen has 22 broad black bands (en- compassing 6—8 dorsal scale rows) that ex- tend completely across the venter. Inter- spaces are whitish (3-4 dorsal rows in breadth) with brownish stippling concen- trated in the central part of each inter- space. The head is white with many irreg- ular black markings on the top and sides. Quebrada de los Naranjos, whence MUSM 17589 comes, is within a region of lowland humid tropical forest extending southward from Ecuador into far north- western Peru (Chapman, 1926; Koepcke, 1961). This forest is the only truly lowland humid forest in western Peru and has never been extensive: remaining fragments are now a protected reserve (Wust, 1998a). Tel- lo (1998:79) described the vegetation of Quebrada de los Naranjos as “an ephemeral thicket composed mostly of small plants of the Rubiaceae and ferns.” Wust (1998b:22) stated that “In the hills of Quebrada de los Naranjos the forest is humid and evergreen, reaching more than 20 m in some parts, and with a more or less uniform canopy domi- nated by [Ficus, Centrolobium, Vitex, Pou- teria, Ocotea, Bombax, and Gallesia].” MUSM 2700-01 are assigned only pro- visionally to Dipsas gracilis because they are in poor condition (soft, poorly pre- served, and with pattern elements ob- scure). The MUSM catalogues indicate that the specimens were collected 17 March 1966 (collector not listed) from the imprecise locality “Valle del Rio Quiroz.” Additional geographical indications in the catalogue (perhaps added later when the specimens were catalogued in Lima) in- clude “Piura [department], Pv. [Provincia] Ayabaca.” The Rio Quiroz flows north- westward across extreme northern Piura department (western versant of the Andes) north of the Cordillera Guamani and even- tually joins the Rio Chira, which drains the southern part of Loja Province, Ecuador. Ayabaca Province is at the northern ex- tremity of Piura department on the Ec- Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 124 (L = N)€ (iia) ac = C/G S/S sperodute, Arepuooas (6 =N)€ (ST = N)@ (it = ND) it =a V/G C/G speaodute, Areurtig (FI =N)€ (as NING — C/G C/G sie[no0\sog (Sii—oN) RO. a 1A V/I sae[Nd0e1g (Gie—w NO eG — T ii utoyed [eo10'] a[suls a] SUIS Q[SuIS a] surS ayeos yeuy (¢ = N) 9I€-86G a ce = ayeula,y (9 = N) €€E-CTE 98G S66 16G ed it S}UNOS [e}USUIBES [ROL (€ = N) €6°9 = 0'601 LIt-SOT a = = ayeulo yy (9 = N) G6'S = L061 ScIFII V6 601 POL STP sjepneoqns (¢ = N) $9°% = 0961 (sjerzuoaoid ¥-G) 66I-F6IL — = — oyeulo (9 = N) LOE = 80S (sjeazuoaoid G-L) OLG-66IL (jeaquoaoid 1) G6IL (jeaquoAoid I) 68ST (sjerzuosoid G) LET [PIN (speazuoAoid pur) speque,, SGT-ST-ST CGI-SI-ST GT-SI-ST GI-ST-ST SopBos [eSIOd (S = N) ¥% (I = N) 61 (IT = N) LI = aan ST yq090} AIeTEXe | (€¢ = N) 100 ¥ 0€°0 [e066 0 oyeule J (9 = N) 10°0 ¥ GE0 Se'0-08'0 = LGO 6a 0 PWN ysuey peyoyysug] [eL (STS) LoL a[euray ysosiey] (VES) OL = Girarcue (ILr) S99 ofl 4SOBIE'] (uIuL) (TAS) YAsuey [ROL IOpenoy o[eul oyeul o[elu syyovis spsdiq, 68SLT SAW 10L@ WSO 00L% WSAIN “HLONAT LNYA OL LOONS = TAS ‘SNOILVAYHSAO YO SNHWIOddS AO HATNAN = N (LHOI/LAAT) SACTTIOS V AT GHLVUVdAS AYV SLNNOO TWHALVIIG “[T HIAVL NI GHLVOIGNI SV GHHOOS AYV SATVOS dvaH ‘YOdVNOW WOW AIdWNVS V HLIM GaYVdNOO SITIOVYUD SVSdIq. OL GHYYAAHY SNAWIONdS NVIANYAd WOH SHALOVYVHO GYVAGNVLS “fF H1AVL lor Dipsas gracilis Ecuac male MUSM 17589 male TABLE 4. CONTINUED. MUSM 2701 MUSM 2700 male Dipsas oreas Complex in Ecuador and Peru ¢ Cadle 5 uadorian border. The general region is arid in lower reaches of the valley to the west and subhumid toward the east. Relictual ae a a A iad DISA patches of humid forest still existed in the . it a Ses er i, ‘ Fk ® 2 : 1990s at higher elevations in montane re- PEIN aR Sheree eo gions in the eastern part of the valley (per- Sn Botts ne “ sonal observations; see above notes on A AIGIGXOT Ie Te habitats of D. oreas in northern Peru). Dipsas gracilis is otherwise known only from the lowlands (<1,000 m), whereas the only parts of the Rio Quiroz valley presently having humid forest habitats are at higher elevations in the eastern part of the river system (>2,000 m). s The pattern elements visible in MUSM are 2700-01 are similar to other specimens of aa Dipsas gracilis. The dorsal pattern consists of a series of dorsal blotches or bands, which, when viewed in lateral aspect, are oval on the anterior body and round pos- teriorly. The anterior bands nearly meet a midventrally; posterior ones encroach onto YF the outer edges of ventrals. The top of the = fl A head is mainly dark blackish with light 2 i S StS patches and flecks (more light patches are i rz a present in the smaller specimen, MUSM ee 2701). The vertebral scale row is greatly mA expanded (1.5-2 the width of the para- vertebral rows). The three Peruvian specimens referred a to Dipsas gracilis conform generally to a other males of D. gracilis | examined, but a is ‘ there are differences from Ecuadorian © = = a specimens as well (Table 4). The Peruvian ak q q specimens have lower ventral (187-192) i and subcaudal (94—109) counts than males from Ecuador (ventrals 199-210, subcau- dals 114-128). MUSM 2700-01 have shorter tails and a different loreal pattern from Ecuadorian specimens. In D. gracilis from Ecuador, all the dorsal bands are complete across the venter (anterior body) or encroach very broadly onto the ventral scutes, nearly meeting midventrally (pos- terior body). This is the case in MUSM 17589, but in MUSM 2700-01, the pos- terior bands end on the outer edges of the ventral scutes. Although some scutellation characteris- tics of the three Peruvian specimens re- contact behind mental Number of bands or blotches Supralabials (touching eye) on body Tertiary temporals Number of infralabials in Infralabials 126 ferred to Dipsas gracilis are outside the range of scale counts seen in Ecuadorian specimens of D. gracilis, they are likewise dissimilar to other species of Dipsas known from western South America in scutellation and/or color pattern. Dipsas oreas is also known from the “valley of the Rio Quiroz” (Fig. 16, Fig. 23: locality 7), the locality for MUSM 2700-01. However, compared with characters of D. oreas males (Table 1 for total sample, Table 3 for Peruvian populations), MUSM 2700— Ol have significantly greater numbers of ventrals, subcaudals, and supralabials. Moreover, the number of maxillary teeth of MUSM 2700 (18) is much greater than the number in a large sample of D. oreas (12-14: Table 1). Collection of additional material might permit a resolution of the systematic status of these Peruvian popu- lations, particularly if new specimens of D. gracilis from the geographic gap between northern Peru and the vicinity of Guaya- quil, Ecuador, can be obtained. Biogeo- graphically, the presence of Dipsas gracilis in lowland humid forests of Tumbes de- partment (MUSM 17589) was perhaps ex- pected on the basis of other geographic and biological data from this region (e.g., Chapman, 1926; Koepcke, 1961), and MUSM 17589 is well documented in the context of a biological inventory (Wust, 1998a). On the other hand, MUSM 2700—O1 are more problematic because they are not ac- companied by specific locality data or col- lector, and there may be reason to ques- tion their origin. Appropriate habitats do not apparently exist presently in the low- lands of the Rio Quiroz valley, and Dipsas gracilis is otherwise known only from low- land localities. Moreover, D. latifasciata is recorded from inter-Andean valleys im- mediately east of the Rio Quiroz (e.g., by MCZ 17404; see comments below). Most characteristics of the color pattern, scale counts, body proportions, loreal scale pat- tern, and maxillary tooth counts of MUSM 2700-01 are similar to those I recorded for D. latifasciata from the Amazonian versant Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 of Peru and Ecuador (unpublished data). One characteristic of the patterns of MUSM 2700-01 is more typical of D. la- tifasciata than of D. gracilis: The posterior bands fail to encroach broadly onto the ventral scutes or to meet midventrally. If, in fact, MUSM 2700-01 were obtained from east of the continental divide or if D. latifasciata occurs on both versants of the Andes in this region, then the identity of these specimens should be reconsidered. Dipsas latifasciatus and D. latifrontalis in Eastern Ecuador and Peru Dipsas latifasciata (Boulenger, 1913) was described from the type locality “Up- per Marafion, eastern Peru,” and D. latif- rontalis (Boulenger, 1905) has the type lo- cality “Aricaqua” [Venezuela]; Peters (1960a: 110) identified the latter locality as the town of that name on the Amazonian slopes of the Andes in western Venezuela south of Mérida. These names and their applicability to populations in eastern Ec- uador require further study. Without ex- amining the holotype of either species, Pe- ters (1960a) assigned specimens from east- ern Ecuador to each. However, Peters’ (1960a) characterizations reveal similar scutellation and few distinctions between them, such as whether some or none of the dorsal bands fail to meet midventrally, as stated in Peters’ key (1960a: 33, couplet 35) or by the number of infralabial pairs in contact behind the mental. The last character is now known to be highly vari- able within species of Dipsas; for exam- ples, see descriptions of the three species discussed in detail in this report or Cadle and Myers (2003) for D. nicholsi and D. andiana. There is broad overlap in the scu- tellation characters of these two taxa as construed by Peters (1960a). This in itself is not necessarily grounds for questioning their validity, but in combination with the other similarities noted by Peters and some inconsistencies in his own discus- sions (see below), it begs the question of how these nominal taxa are distinguished Dipsas oreas Complex in Ecuador and Peru * Cadle and how the names should apply to Ec- uadorian and Peruvian specimens. Moreover, Peters (1960a) had little ma- terial of each species, particularly from near the type localities. He referred only three specimens from extreme southern Ecuador and northern Peru to D. latifas- ciata. More problematic was his referral of only one specimen from Venezuela to D. latifrontalis but, in addition, 44 others from eastern Ecuador. Thus, the distribu- tion of “Dipsas latifrontalis,” as Peters (1960a: 99) conceived it, left a broad gap across eastern Colombia, from which no specimens referred to either of these spe- cies were reported. Peters himself seems to take great pains to justify his applica- tions of these names to samples available to him based only on the type descriptions. Similarly, he justified his referral of the names Leptognathus palmeri Boulenger (1912) (type locality, “El Topo, Rio Pasta- za, Ecuador”) and L. praeornata Werner (1909) (type locality, “Venezuela”) to the synonymy of Dipsas latifrontalis by ex- tended discussion of relatively few char- acters in the original descriptions (Peters, 1960a: 103, 109-110). The brevity of the original descriptions of all these taxa and the poor comprehension of intraspecific variability within Dipsas at the time of their descriptions mean that reference to the types is needed to resolve the system- atics of these snakes. These peculiarities and my interpreta- tions of some material that both Peters and I examined lead me to question the proper name of specimens from eastern Ecuador and Peru. MCZ 17404, a speci- men that Peters (1960a) assigned to D. la- tifasciata, has the pattern by which he characterized D. latifrontalis (all bands fail to meet midventrally). In fact, Peters (1960a: 103) stated that this specimen was “intermediate in some characters between what might be called typical latifrontalis and latifasciata. ...” In three specimens that Peters assigned to D. latifrontalis from the same general area of Ecuador (“Llanganates Area” [probably southwest- 127 ern Napo Province]), two (FMNH 23530— 31) show the pattern characteristic as- cribed to D. latifrontalis, whereas FMNH 23532, although it has a predominantly black venter, seems to have bands meeting midventrally, the pattern that Peters as- cribed to D. latifasciata. Peters (1960a: 103) suggested the possibility that addi- tional collections might “demonstrate that the relationship between [latifrontalis and latifasciata| is at best on the subspecific level.” Interpretive problems such as these sug- gest that a new look at snakes with the requisite characteristics from Venezuela, Ecuador, and Peru is warranted. As with many similar problems concerning species identities in Dipsas, adequate resolution will require renewed study of the types and other geographically proximate speci- mens. Consequently, Peters’ assignment of specimens from eastern Ecuador to D. la- tifrontalis is unconvincing, as is his sug- gestion of a possible subspecific relation- ship between D. latifrontalis and D. lati- fasciatus. The specimens here referred to D. latifasciata (see Specimens Examined and Locality Records) are done so provi- sionally, pending further study of the Ec- uadorian, Peruvian, and Venezuelan ma- terial (including types) referred to D. la- tifasciata and D. latifrontalis. Full resolu- tion of the systematics of this group should also review the validity and proper syn- onymy of the names Leptognathus palmeri Boulenger (1912) and L. praeornata Wer- ner (1909), which Peters considered syn- onyms of D. latifrontalis on the basis solely of the type descriptions. KEY TO SPECIES OF DIPSAS IN WESTERN SOUTH AMERICA I provide the following key to assist in identifying species and individual speci- mens of Dipsas in western South America. Six species of Dipsas are known from the western slopes of the Cordillera Occiden- tal and adjacent Pacific lowlands of Ecua- dor and Peru: D. andiana, D. elegans, D. ellipsifera, D. gracilis, D. oreas, and D. 128 temporalis. All of these are known from Ecuador, but only D. oreas and D. gracilis have been reported from Peru. The key also includes three additional species known or expected in the Choco region of Colombia: Dipsas sanctijohannis, D. vi- guieri, and (possibly) D. nicholsi; the last species, previously known only from cen- tral Panama (Cadle and Myers, 2003), has recently been recorded in eastern Panama near the Colombian border (Myers et al., unpublished data). Characters of D. ni- cholsi and D. andiana in the key are based on diagnoses given in Cadle and Myers (2003) and another female of D. nicholsi reported by Myers et al. (ms.). The differ- ential characteristics of D. sanctijohannis in the key are based on data from Peters (1960a) and on data kindly provided by Charles W. Myers from specimens he ex- amined. Characters of all other species are based on discussions in this paper (D. ele- gans, D. ellipsifera, D. gracilis, D. oreas) or unpublished data (D. temporalis) in conjunction with some data from Peters (1960a). Without additional study, I am unable to adequately differentiate Dipsas viguieri (eastern Panama and northern Chocé6, Co- lombia) and D. gracilis (western Ecuador and extreme northern Peru). Geography currently seems to be the only reliable means of assigning names to specimens of these species. The characters that Peters (1960a) used to distinguish D. viguieri and D. gracilis (e.g., preocular presence/ab- sence, number of postoculars) are known to be highly variable intraspecifically with- in Dipsas. Peters (1960a: 52) considered the two perhaps only “subspecifically dis- tinct” and otherwise noted (Peters, 1960a: 48) that “The characters separating [Dip- sas gracilis] and D. viguieri are compara- tively weak ones... .” Thus, specimens of D. viguieri will key out to D. gracilis with this key. The differential characters (if any) of these two species and the disjunction (or lack thereof) between their ranges in western Colombia require further re- search. Nonetheless, if these two taxa are Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 closely related or conspecific, then they would add a third example within Dipsas of a biogeographic connection between northwestern (Chocoan) South America and eastern Panama. The other examples are D. andiana—D. nicholsi (Cadle and Myers, 2003) and D. temporalis, which has scattered records from northwestern Ec- uador into eastern Panama. In addition to illustrations provided herein, illustrations and discussions of some of the relevant taxa are found in Pe- ters (1960a; D. gracilis, pl. Ib, c and D. ellipsifera, pl. 1Va), Kofron (1982; D. ele- gans, D. ellipsifera, and D. oreas), and Ca- dle and Myers (2003; D. andiana, D. ni- cholsi, and D. oreas). The key is dichoto- mous except for the last numbered sec- tion, a triplet. 1. Dorsal bands very broad anteriorly and poste- riorly, covering 8 or more dorsal scale rows in longitudinal dimension; bands continuous, or nearly so, across the venter at least ante- riorly. Subcaudals >90, often >100 (up to 132). Tail =29% of total length. Either no infralabials in contact behind the mental scale on one woain imicontach ea = aes Y — Dorsal bands narrower, covering 10 or fewer dorsal scale rows in longitudinal dimension (usually <7); anterior bands usually much broader than posterior bands, and all bands end on outer edges of ventrals or on low- ermost dorsal rows. Subcaudal counts vari- able, but usually <100 (except male D. an- diana, up to 106). Tail =28% of total length. At least one pair of infralabials in contact behind the mental scale . 2. No infralabials in contact behind the mental scale. Anterior pair of chin shields very small (in contact with mental), followed by a pair of much larger chin shields. —~------ Sa) BE a Le? Li cena Dipsas temporalis (Werner) (eastern Panama, western [Chocoan] Co- lombia, and western Ecuador; primarily in low uplands in Panama, 800—1,000 m; ap- parently also in the lowlands in Colombia and Ecuador) — One pair of infralabials in contact behind the mental scale. Anterior pair of chin shields about the same size as, or only slightly smaller than, the second pair, -—---------------- Pires ee recs Nos. £ W) Dipsas gracilis (Boulenger) (lowlands [150-620 m] of western Ecuador from just north of the equator to extreme northwestern Peru [Tumbes and, less cer- Dipsas oreas Complex in Ecuador and Peru ° Cadle tainly, Piura departments; see discussion herein]) [D. viguieri will also key here] 3. A distinct V- or U-shaped mark with well-de- fined, regular borders on top of the head (apex anteriorly at frontal/prefrontal bor- der); head otherwise pale brown or grayish brown and relatively unmarked. Ventrals =185 in males and females. Subcaudals = Jim males, =S2 im temales, 4 — No distinct stereotypic inverted V- or U- shaped mark on the top of the head.” Head usually with many dark markings, including darkened suture lines; entire head may be dark. Ventrals <190 in males and females. Subcaudal counts variable, but often much lessMtlnars GO tees aeeet eye. Oe od. NS le 5 4. Ventrals 185-196 in males, 185-191 in fe- males. Subcaudals 91—106 in males, 82—83 in females. Branches of the V-shaped mark on top of head generally not connected to the first pair of neck blotches. Dorsal blotches at midbody taller than wide. -------- See MSS Se ee Dipsas andiana (Boulenger) (lowlands and lower montane slopes of western Ecuador from approximately the equator to latitude 2°S; 5-1,140 m) — Ventrals 198-208 in males, 200—206 in two females. Subcaudals 92—98 in males, 87—95 in females. Branches of the V-shaped mark on top of head connected to the first pair of neck blotches. Dorsal blotches at mid- body wider than tall. __ Dipsas nicholsi (Dunn) (lowlands <200 m in the Rio Chagres basin of central Panama; Darién highlands, 875 m, of extreme eastern Panama; Myers et al., ms.) 5. Ventrals 153-164 in males, 157-158 in two females. Subcaudals 72-78 in males, 62-63 in two females _. Dipsas ellipsifera (Boulenger) (known only from the valley of the Rio Mira in extreme northwestern Ecuador, 570— 2,600 m) — Ventrals >165 in males and females. Subcau- dals >80 in males, >70 in females 6. Subcaudals 82-91 in males, 70—83 in females. Top of head usually with a pair of large elongate irregular blotches centered on the parietal region. Head otherwise with many dark spots and flecks and darkened suture lines. Thirty or fewer bands on the body, with anterior bands covering more than 5 dorsal rows in longitudinal dimension. Each band in adults usually with a somewhat pale central portion, but bands in most speci- ‘The elongate blotches on the parietal region in Dipsas oreas occasionally fuse to form an irregular V- shaped mark that is very different from the marking in D. andiana. See Cadle and Myers (2003: 24, fig. TWD); 129 mens without distinct borders enclosing a highly contrasting pale central area. Ante- rior bands end on outer edges of ventrals. Venter dirty whitish to grayish, usually with many small dark irregular spots and streaks oN A Gy LL OS. een ee Dipsas oreas (Cope) (western Ecuador and northwestern Peru, approximately latitude 2°S to nearly 7°S; primarily Andean foothills and_ slopes, 1,400-2,600 m; lowlands in the vicinity of Guayaquil, Ecuador) — Subcaudals 94-105 in males, 68—88 in fe- males. Top of head with many dark brown irregular marks on a pale brown ground col- or, but generally not forming paired irreg- ular blotches and occasionally nearly solid dark brown. Twenty-five to 46 bands on body. Each band in adults comprising a pair of bold blackish vertical edges enclosing a contrasting pale brown central portion. An- terior bands cover no more than 5 dorsal scale rows and end on outer edges of ven- trals. Venter grayish with many small dark flecks and squarish markings, which some- times form longitudinal arrays ---.-- DREN ACN e «PE Meera Se he Dipsas elegans (Boulenger) (western Ecuador from just north of the equator to approximately latitude 1°10'S, 500—2,650 m) — Subcaudals 86—94 in males, 70-83 in females. Top of head in adults relatively uniform me- dium brown (dark-spotted in juveniles). Twenty-one to 28 bands on body; anterior bands complete or nearly complete across venter; each band solid, without pale cen- tral portion. Venter brown without numer- ousedankelecks* and: spots y-2--2--te Bt See ee See Dipsas sanctijohannis (Boulenger) (Colombia: western slopes of the Cordillera Occidental in Chocé and the inter-Andean valley of the Rio Cauca) ACKNOWLEDGMENTS For loans of specimens and other assis- tance I thank Linda S. Ford and Charles W. Myers (AMNH); Edward B. Daeschler and Edward Gilmore (ANSP); E. Nicholas Arnold, Colin J. McCarthy, and Mark Wil- kinson (BMNH); Robert F. Inger, Jamie Ladonski, Alan Resetar, and Harold Voris (FMNH); Hugo Alamillo, Juan M. Guay- asamin, John E. Simmons, Omar Torres, and Linda Trueb (KU); Christopher J. Austin (LSU); James Hanken and José P. Rosado (MCZ); César Aguilar, Nelly Car- rillo de Espinoza, and Jestis Cordova (MUSM); Harry W. Greene and Barbara 130 Stein (MVZ): Ronald Nussbaum and Gregory Schneider (UMMZ); and Steve Gotte, W. Ronald Heyer, Roy W. Mc- Diarmid, Robert Wilson, and George R. Zug (USNM). I especially thank Jests Cérdova and César Aguilar for kindnesses on visits to the MUSM. Several individuals helped locate or clar- ify localities for some specimens: Steve Gotte, Roy W. McDiarmid, Charles W. Myers, and Thomas S. Schulenberg. My- ers also provided the basemap used to pre- pare Figure 8, provided data and photo- graphs for Dipsas sanctijohannis, and shared ideas and information about Dip- sas; I am grateful for these exchanges and for his generosity. Martin Henzl translated a portion of Koepcke (1961). The Chicago Zoological Society, especially through its Director Emeritus George B. Rabb, has supported my biodiversity studies, partic- ularly during completion of this research and visits in 2004 to ANSP, MCZ, MUSM, and USNM. Field work was supported by the Field Museum of Natural History (1987 expe- dition to the Rio Zafa Study Site), the American Philosophical Society, the Put- nam Fund of the Museum of Comparative Zoology, and a grant from Faculty of Arts and Sciences of Harvard University. The David Rockefeller Center for Latin Amer- ican Studies (Harvard University) provided funds for attendance at the symposium Es- tratégias para Bioconservacion en el Norte del Pert, held at the Universidad Antenor Orrego, Trujillo (1995), at which some of this work was presented; invitation to that conference was extended by Abtindio Sa- sdstegui Alva, whose enthusiastic moral and logistical support also made much of the field work possible. The field work would have been impos- sible without the efforts and camaraderie of my field companions: Pablo Chuna Mo- gollon, Camilo Diaz, Michael O. Dillon, the late Alwyn H. Gentry, Rosa Ortiz de Gentry, José Guevarra Barreto, Segundo Leiva, Pedro Lezama, Raul Quiroz, Abtin- dio Sagastegui Alva, José Santisteban, and Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 Helena Siesniegas. The family of Francis- co Quiroz was especially supportive during two field seasons at the Rio Zafia Study Site, and the support and friendship of many strangers during field work in north- ern Peru is much appreciated. I am grateful to the Direcci6n General Forestal y de Fauna del Pert and the Mu- seo de la Universidad Nacional de San Marcos in Lima for their many years’ sup- port of my efforts to understand the bio- diversity of Peru. For discussions of plant diversity, I thank Michael O. Dillon, the late Alwyn H. Gentry, and Abundio Sagas- tegui Alva. For very valuable comments on the manuscript, I thank Harry W. Greene, Charles W. Myers, and Jay M. Savage. SPECIMENS EXAMINED AND LOCALITY RECORDS A few literature records are included herein when there is reason to be assured of the identity of the specimens, which is rarely the case with South American spe- cies of Dipsas. Some of these records are discussed in the above species accounts (see sections on Distribution). In addition to specimens of the three species of the oreas group, this list also includes material of D. gracilis, D. latifasciata, and D. tem- poralis examined during the course of this study. This should aid future workers on this genus because many of the identifi- cations in the literature are questionable and much material has accumulated since the group was last reviewed (Peters, 1960a). Discussion of some apparent prob- lems concerning the systematics of D. gracilis and D. latifasciata is given in the text. Other material relevant to the iden- tity of species in western Ecuador was list- ed in Cadle and Myers (2003) (D. andiana, D. nicholsi, and D. variegata). Institutional abbreviations for museums are as follows. AMNH American Museum of Natural History, New York ANSP Academy of Natural Sciences of Philadelphia Dipsas oreas Complex in Ecuador and Peru ¢ Cadle BMNH The Natural History Muse- um, London Field Museum of Natural History, Chicago Museum of Natural History, University of Kansas, Lawr- ence Museum of Natural Science, Louisiana State University, Baton Rouge Museum of Comparative Zo- ology, Harvard University, Cambridge Museo de Historia Natural de San Marcos, Lima, Peru Muséum National d Histoire Naturelle, Paris Museum of Vertebrate Zool- ogy, University of California, Berkeley Museum of Zoology, Univer- sity of Michigan, Ann Arbor FMNH KU LSUMNS MCZ MUSM® MNHN MVZ UMMZ Dipsas elegans ECUADOR: “Tehuantepec” [Oaxaca, Mexico], in error (see text), BMNH 1946.1.21.77 (holotype). “Western Ecuador,’ BMNH 60.6.16.73. [PROv. CHIMBORAZO]: Pallatanga [1,500 m; 01°59’S, 78°57'W], BMNH 80.12.5.267. PROV. COTOPAXI!: Corazé6n [900—1,200 m; 01°08'S, 79°04’W], USNM 210938—39.° PROV. PICHINCHA: Way to Mindo [0°2'S, 78°48’W], USNM 210934, 210961 (see foot- note 9). 2.9 km SW Tandayapa on Mindo Road, 1,820 m [0°1’S, 78°46’W], USNM 285957. Nanegal Grande [about 1,400 m; 0°7’N, 78°40’W], USNM 210936. Below Pacto, [1,200 m; approximately 0°8'N, 78°45'W], USNM 210937. Rio Saloya, 4,000 feet [1,220 m; O°1’N, 78°57'W], BMNH 1940.2.20.32. Santo Domingo de los Colorados [500 m; 0°15’S, 79°9’W], USNM 210935. Perucho [1,830 m; 0°6’N, *MUSM-JEC refers to Cadle field numbers for which the MUSM catalogue numbers are not yet available. °USNM specimens from “Corazén” (USNM 210938—39) and “Way to Mindo” (USNM 210934, 210961) are probably equivalent in whole or in part tom Ee Ne (ina S le Corazones). and PN 719 (“camino a Mindo”) cited by Peters (1960a: 92) as “Dipsas ellipsifera.”. According to USNM records, these specimens were obtained by Peters from Gus- tavo Orcés-V. and recorded originally as “GOV 717— 719” (Steve Gotte, personal communication, October 2004). Orcés-V. was, at the time, curator at the EPN. 131 78°25'W] (Orcés and Almendariz, 1987]. E] Quinche, 2,640 m [0°6’S, 78°17’W] (Orcés and Almendariz, 1987). Cumbayé, 2,354 m [0°12’S, 78°26’W] (Orcés and Almendariz, 1987). Chiriboga, 1,800 m [0°15’S, 78°44'W] (Orcés and Almendariz, 1987). Tumbaco [2.400 m; 0°13’S, 78°24'W] (Orcés and Almendariz, 1987). PROv. IMBABURA: Near Pefiaherrera-Intag, 1,500 m [0°21'N, 78°31’W], UMMZ 92073. Dipsas ellipsifera ECUADOR: [PROv. IMBABURA]: Ibarra |=Ybar- ra; 2,211 m; 0°21’'’N, 78°7’W], BMNH 1946.1.21.26 (formerly 98.4.28.87) (syntype), BMNH 1946. 1.21.27 (formerly 98.4.28.89) (syntype), BMNH 1946.1.21.28 (formerly 98.4.28.88) (syntype), BMNH 1946.1.21.29 (formerly 98.4.28.90) (syntype), MCZ 8431. Lita, 572 m? [0°52’N, 78°28’W] (Orcés and Almendariz, 1987). Chachimbiro, 2,600 m [0°27’N, 78°13'W] (Orcés and Almendariz, 1987). Pimampiro, 2,000 m [0°30’N, 77°56’W], UMMZ 83697—700. Dipsas gracilis ECUADOR: PROV. ESMERALDAS [?PROV. PI- CHINCHA]:!! Hacienda Equinox, 30 km NNW Santo Domingo de los Colorados, 1,000 ft. [305 m; approx- imately 0°5'’N, 79°8’W], USNM 210945. PRov. GUAYAS: near Guayaquil [approximately 2°10’S, 79°50’W], USNM 210947. PROv. PICHINCHA: 47 km S. Centro Cientffico Rio Palenque, 150-220 m |ap- proximately 0°50'S, 79°30’W], USNM 286478-80. Centinela, 14.1 km SE Patricia Pilar by road, 570— 600 m [0°37’S, 79°18’W], MCZ 156894. Santo Do- mingo de los Colorados [0°15’S, 79°9'W], USNM 286477. Centro Cientifico Rio Palenque [200 m: 0°33'S, 79°22'W], MCZ 147183. Estacién Bioldgica Rio Palenque, 56 km N Quevedo, 220 m [0°33’S, 79°22'’W], KU 152604. PERU: DEPTO. TUMBES: Quebrada Los Naranjos, 3°50’S, 80°14’W, 620 m, MUSM 17589. T confirmed the elevation of Lita given by Orcés and Almendariz (1989) on two maps (“Republica del Ecuador” 1:1,000,000 published by the Instituto Geo- grafico Militar [Quito], 1991; and “Ecuador” 1: 1,000,000 published by ITMB Publishing, Vancouver, 1994-1996). Paynter (1993) gave 900 m as the ele- vation. My maps place this locality, as given by distance from Santo Domingo de los Colorados, in Pichincha Province. The specimen was obtained by James A. Peters, who used “Esmeraldas Province” in his field notes and as registered in the USNM catalogues (S. Gotte, personal communication, September 2004). Data on a paper label within the jar differ slightly from the catalogue data: 32 km NNW Santo Domin- go de los Colorados and “James Brown Farm,” rather than “Hacienda Equinox.” 132 Dipsas gracilis(’?) PERU: DEPTO. PIURA: Provincia Ayabaca, Valle del Rio Quiroz [approximately 4°45'S, 79°50'W], MUSM 2700-01. Dipsas latifasciata ECUADOR: PROV. NAPO: Coca_ [0°28'S, 76°58'W], MCZ 166589-90. El] Reventador [approx- imately 0°40'S, 77°40'W; locality assumed to refer to lower eastern slopes of Volcan El] Reventador, a high peak in the eastern cordillera], MCZ 164510-11. 21 km SSW Reventador, 1,700 m [approximately 0°7'S, 77°38'W; Lynch and Duellman, 1980: 85], KU 164212. Lumbaqui OZStINE ie2 ONE NACo 4674 US. PERU: [DEPTO. CAJAMARCA]: Tabaconas [1,892 m; 5°19'S, 79°18’'W], MCZ 17404. [DEPTO. SAN MARTiN/LORETO]: Cumbre Ushpayacu-Mishquiya- cu, 3,200 feet [975 m; 6°57'S, 76°3’W],!2 AMNH 52444. DEPTO. PAsco: Playa Pampa, about 8 km NW Cushi on trail to Chaglla, 2,100 m [9°57’S, 75°42'’W?3], LSUMZ 45499. Dipsas oreas ECUADOR: “The elevated Valley of Quito” (probably southern Ecuador, as discussed in the text), ANSP 10115 (holotype). “Western Ecuador,” BMNH 60.6.16.56, 60.6.16.63. PROV. CHIMBORAZO: Alausi, 2,390 m [2°12’S, 78°50'W] (Despax, 1911: 36, as “Leptognathus mikani”’). PROV. CHIMBORAZO/ CANAR:" Huigra to Rio Chiguancay [2°13’S, 79°3’W], ANSP 18117, 18120, 18123. [Rio] Chanchan Valley [approximately 2°17’S, 79°24'W], MCZ 17083, UMMZ 56491. Prov. GuayAs: Guayaquil [2°10'S, 79°50’W], USNM 60006, 62797—802. PROv. Loja: Catamayo Valley, 9,780 feet [2,982 m; approximately 4°5'S, 79°35’W], USNM 98923. “Loja, Nouvelle-Gre- nade” [inferred to be the city of Loja, 2,200 m; 4°S, 79°13'W], MNHN 6285 (holotype of Leptognathus andrei Sauvage, 1884; see Kofron, 1982: 48-50]. 13 km E Velacruz (20 km NE Catacocha), 2,250 m [3°58’S, 79°33’W], KU 142803. The specimen is part of the Harvey Bassler col- lections from eastern Peru (see Myers, 2000: 139- 141). The locality is along a ridge separating the headwaters of the Rio Mishquiyacu (Rio Huallaga drainage) and the Rio Uchpiyacu (Rio Ucayali drain- age) and forming the border between San Martin and Loreto departments. The Mishquiyacu flows west- ward into the Rio Huallaga near the town of Pilluana. The Uchpiyacu flows eastward into the Rio Cusha- batay, a tributary of the Rio Ucayali. '5 Coordinates from Dr. Thomas S. Schulenberg, who collected the specimen. '4 Tower reaches of the Rio Chanchan form the border between Chimborazo and Cafar provinces. Huigra itself is in Chimborazo province. Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 PERU: DEPTO. PIURA: Immediate vicinity of Ay- abaca, approximately 2,600 m [4°38’S, 79°43’W], MUSM-JEC 10347. Toronche (town at base of Cerro Aypate), approximately 16 km (airline) SE Ayabaca, 1,950 m [4°35'S, 79°32'W], MUSM 16750. 2 km W Porculla [Pass] [5°51'S, 79°31'W], MVZ 119330. About 15 km (by road) E Canchaque on road to Huancabamba [about 5°24'S, 79°36’W], LSUMNS 27373. DEPTO. CAJAMARCA: Rio Zafia Study Site, 1,450-1,800 m [6°51'S, 79°6’W; see Materials and Methods for interpretation of this locality], ANSP Oli (Ol. all (63-502 0F MINE eZ an0= 7 ae2 ooo a MUSM 3361, 3452, 5530, 5532-33, 16751—52. Dipsas temporalis COLOMBIA: DEPTO CHOCO: Agua Clara, Rio Tamana [approximately 4°53’N, 76°45’W], USNM 267244, PANAMA: [PROv. PANAMA]: Pequeni-Cha- gres ridge, head of Rio Limpio and Quebrada Las Tres Honeras (Panama snake census) [9°18’N, 79°27'W], MCZ 50214. South slope of Cerro La Campana, 850 m [8°43’N, 79°54'W], KU 110293. PROV. DARIEN: Ridge between Rio Jaque and Rio Imamad6, 800-890 m [7°35'N, 77°57'W; see Cadle and Myers, 2003: fig. 16], KU 110294-96. North ridge of Cerro Cituro, Serrania de Pirré, 900—1,000 m [approximately 8°5’N, 77°46’W], KU 110298— 110301. LITERATURE CITED ALMENDARIZ, A., AND J. L. CARR. 1992. Herpetofau- na, pp. 62 + appendices 11-12 (pp. 128-132). In T. A. Parker III and J. L. Carr (eds.), Status of Forest Remnants in the Cordillera de la Costa and Adjacent Areas of Southwestern Ecuador (RAP Working Papers 2). Washington, D.C.:: Conservation International. AMARAL, A. DO. 1926. 2nd nota de nomenclature Ophiologica. S6bre o emprego do nome generico Sibynomorphus em vez de “Leptognathus”, “Stremmatognathus”, “Anholodon”, etc. Revista do Museu Paulista. 14: 7-9. . 1929a. Estudos sobre ophidios neotropicos. XVII—valor systematico de varias formas de ophidios neotropicos. Memorias do Instituto Bu- tantan. 4: 3-68. . “1929"b [1930]. Estudos sobre ophidios neo- tropicos XVIII. Lista remissiva dos ophidios da Regiao Neotropica. Memorias do Instituto Bu- tantan. 4: 127-128 + i-vii + 129-271. AMR, Z. S., R. M. AL-ORAN, AND W. N. AL-MELHIM. ‘5 Another specimen from near Canchaque collect- ed in the early 1990s is in the Museo de Historia Natural de la Universidad Privada Antenor Orrego (Trujillo, Peru). It is referable to Dipsas oreas on the basis of color pattern (I examined the specimen brief- ly in 1994 but did not complete details such as scale counts). Dipsas oreas Complex in Ecuador and Peru * Cadle 1997. Aggregation behavior in two Jordanian snakes: Coluber rubriceps and Typhlops vermi- cularis. Herpetological Review. 28(3): 130-131. BAILEY, J. R. 1967. The synthetic approach to colu- brid classification. Herpetologica. 23(2): 155— 161. . 1981. Notes on the Genus Thamnodynastes. Abstracts: 1° Simpésio Internacional sobre Ser- pentes em Geral e Artrépodes Peconhentos, 16 a 18 de Novembro de 1981 (Comemoragées do 80° aniversdrio do Instituto Butantan, 1901— 1981). Sdo Paulo, Brazil: Instituto Butantan. BOULENGER, G. A. 1896. Catalogue of the Snakes in the British Museum (Natural History). Vol. 3. London: British Museum (Natural History). i—xiv + 727 pp. + pls. 1-25. . 1898. An account of the reptiles and batra- chians collected by Mr. W. F. H. Rosenberg in western Ecuador. Proceedings of the Zoological Society of London. 1898: 107-126. . 1902. Descriptions of new batrachians and reptiles from northwestern Ecuador. Annals and Magazine of Natural History. 9(Ser. 7): 51-57. . 1905. Description of a new snake from Ve- nezuela. Annals and Magazine of Natural His- tory. 15(Ser. 7): 561. . 1912. Descriptions of new reptiles from the Andes of South America, preserved in the British Museum. Annals and Magazine of Natural His- tory. 10(Ser. 7): 420-424. . 1913. Description of a new snake discovered by Mr. A. E. Pratt in eastern Peru. Annals and Magazine of Natural History. 12(Ser. 8): 72. CADLE, J. E. 1984a. Molecular systematics of Neo- tropical xenodontine snakes: I. South American xenodontines. Herpetologica. 40(1): 8-20. . 1984b. Molecular systematics of Neotropical xenodontine snakes: IJ. Central American xeno- dontines. Herpetologica. 40(1): 21-30. . 1984c. Molecular systematics of Neotropical xenodontine snakes: III. Overview of xenodonti- ne phylogeny and the history of New World snakes. Copeia. 1984(3): 641-652. . 1985. The Neotropical colubrid snake fauna (Serpentes: Colubridae): lineage components and biogeography. Systematic Zoology. 34(1): 1— 20. . 1989. A new species of Coniophanes (Ser- pentes: Colubridae) from northwestern Peru. Herpetologica. 45(4): 411-424. . 1991. Systematics of lizards of the genus Stenocercus (Iguania: Tropiduridae) from north- ern Peru: new species and comments on rela- tionships and distribution patterns. Proceedings of the Academy of Natural Sciences of Philadel- phia. 143: 1-96. . 1996. Systematics of snakes of the genus Geodipsas (Colubridae) from Madagascar, with descriptions of new species and observations on natural history. Bulletin of the Museum of Com- parative Zoology. 155(2): 33-87. 133 CADLE, J. E., AND P. CHUNA M. 1995. A new lizard of the genus Macropholidus (Teiidae) from a re- lictual humid forest of northwestern Peru, and notes on Macropholidus ruthveni Noble. Bre- viora. 501: 1—39. CADLE, J. E.. AND R. W. MCDIARMID. 1990. Two new species of Centrolenella (Anura, Centrolen- idae) from the western slope of the Andes in northern Peru. Proceedings of the Biological So- ciety of Washington. 103: 746-768. CADLE, J. E., AND C. W. MYERS. 2003. Systematics of snakes referred to Dipsas variegata in Panama and western South America, with revalidation of two species and notes on defensive behaviors in the Dipsadini (Colubridae). American Museum Novitates. 3409: 1-47. CAMPBELL, J. A. 1998. Amphibians and Reptiles of Northern Guatemala, the Yucatan, and Belize. Norman, Oklahoma: University of Oklahoma EIess. XIX, 1.360) pp: CAMPBELL, J. A., AND W. W. LAMAR. 2004. The Ven- omous Reptiles of the Western Hemisphere. 2 vols. Ithaca, New York: Comstock Publishing As- sociates. xviii + 1-476; xiv + 477-870 (+ index, 1-28) + 1,500 pls. on 127 pp. CECHIN, S. Z., AND J. L. OLIVEIRA. 2003. Sibyno- morphus ventrimaculatus (southern snail-eater): mating. Herpetological Review. 34(1): 72. CHAPMAN, F. M. 1926. The distribution of bird-life in Ecuador, a contribution to a study of the origin of Andean bird-life. Bulletin of the American Museum of Natural History. 55: 1-784. Cope, E. D. 1868. An examination of the reptilia and batrachia obtained by the Orton expedition to Equador and the upper Amazon, with notes on other species. Proceedings of the Academy of Natural Sciences of Philadelphia. 20: 96-140. . 1874. On some batrachia and nematognathi brought from the upper Amazon by Prof. Orton. Proceedings of the Academy of Natural Sciences of Philadelphia. 26: 120-137. . 1876. Report on the reptiles brought by Pro- fessor James Orton from the middle and upper Amazon, and western Peru. Journal of the Acad- emy of Natural Sciences of Philadelphia. 8(Ser. 2): 159-183. . 1877. Synopsis of the cold blooded Verte- brata procured by Prof. James Orton during his exploration of Peru in 1876-77. Proceedings of the American Philosophical Society. 17: 33-49. CUNHA, O. R., AND F. P. NASCIMENTO. 1993. Ofidios da AmazOnia. As cobras da regiaio leste do Para [2nd edition]. Boletim do Museu Paraense Em- flio Goeldi, série Zoologia. 9(1): 1-191. DEsPAx, R. 1911. Reptiles et batraciens de lEquateur recueillis par M. le Dr. Rivet, pp. 17— 44 + 3 pls. In Ministére de Instruction Pub- lique (ed.), Mission du Service Géographique de l’Armée pour la mesure d’un Arc de Méridien Equatorial en Amérique du Sud sous le Contréle Scientifique de Académie des Sciences, 1899— 134 1906, Tome 9 (Zoologie), Fascicule 2 (Reptiles- Poissons-Batraciens). Paris: Gauthier-Villars. DILLON, M. O., A. SAGASTEGUI ALVA, I. SANCHEZ VEGA, S. LLATAS QUIROZ, AND N. HENSOLD. 1995. Floristic inventory and biogeographic anal- ysis of montane forests in northwestern Peru, pp. 251-269. In S. P. Churchill, H. Balslev, E. For- ero, and J. L. Luteyn (eds.), Biodiversity and Conservation of Neotropical Montane Forests. Bronx, New York: The New York Botanical Gar- den. DODSON, C. H., AND A. H. GENTRY. 1991. Biological extinction in western Ecuador. Annals of the Missouri Botanical Garden. 78(2): 273-295. DUELLMAN, W. E. 1978. The biology of an equatorial herpetofauna in Amazonian Ecuador. Miscella- neous Publications, Museum of Natural History, University of Kansas. 65: 1-352. FEIO, R. N., P. SANTOS, R. FERNANDES, AND T. S. DE FREITAS. 1999. Chironius flavolineatus (NCN) courtship. Herpetological Review. 30(2): 99. FERNANDES, R., D. S. FERNANDES, AND P. PASSOS. 2002. Leptognathus latifasciatus Boulenger, 1913, a junior synonym of Dipsas polylepis (Bou- lenger, 1912) (Serpentes, Colubridae). Boletim do Museu Nacional (Rio de Janeiro), nova série, Zoologia. 493: 1-7. FORD, N. B., AND G. M. BURGHARDT. 1993. Percep- tual mechanisms and the behavioral ecology of snakes, pp. 117-164. In R. A. Seigel and J. T. Collins (eds.), Snakes: Ecology and Behavior. New York: McGraw Hill. FOWLER, H. W. 1913. Amphibians and reptiles from Ecuador, Venezuela, and Yucatan. Proceedings of the Academy of Natural Sciences of Philadel- phia. 65: 153-173. Fox, H. 1977. The urinogenital system of reptiles, pp. 1 eal 1 Cansvand Mies Sesarsonse eds» aelbi= ology of the Reptilia. Vol. 6 (Morphology E). New York: Academic Press. GALLARDO, J. M. 1972. Observaciones bioldgicas so- bre una falsa yarara, Tomodon ocellatus Duméril, Bibron et Duméril (Reptilia, Ophidia). Neotro- pica (Buenos Aires). 18: 58-63. GENTRY, A. H. 1995. Patterns of diversity and floristic composition in Neotropical montane forests, pp. 103-126. In S. P. Churchill, H. Balslev, E. For- ero, and J. L. Luteyn (eds.), Biodiversity and Conservation of Neotropical Montane Forests. Proceedings of the Neotropical Montane Forest Biodiversity and Conservation Symposium, The New York Botanical Garden, 21-26 June 1993. Bronx, New York: New York Botanical Garden. GILLINGHAM, J. C. 1987. Social behavior, pp. 184— 209. In R. A. Seigel, J. T. Collins, and S. S. Novak (eds.), Snakes: Ecology and Evolutionary Biolo- gy. New York: MacMillan Publishing Company. GREENE, H. W. 1988. Antipredator mechanisms in reptiles, pp. 1-152. In C. Gans and R. B. Huey Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 (eds.), Biology of the Reptilia, vol. 16B: Defense and Life History. New York: Alan R. Liss. GREGORY, P. T., J. M. MACARTNEY, AND K. W. LAR- SEN. 1987. Spatial patterns and movements, pp. 366-395. In R. A. Seigel, J. T. Collins, and S. S. Novak (eds.), Snakes: Ecology and Evolutionary Biology. New York: MacMillan Publishing Com- pany. GUILLETTE, L. J., JR., S. L. FOX, AND B. D. PALMER. 1989. Oviductal morphology and egg shelling in the oviparous lizards Crotaphytus collaris and Eumeces obsoletus. Journal of Morphology. 201: 145-159. GUNTHER, A. 1872. Seventh account of new species of snakes in the collection of the British Muse- um. Annals and Magazine of Natural History. 9(Ser. 4, no. 49): 13-37. PUSS a— 902s Biologia Centrali-Americana, Reptilia and Batrachia [1987 facsimile reprint]. Ithaca, New York: Society for the Study of Am- phibians and Reptiles. lxix + xx + 326 pp. + 76 pls. HALPERN, M. T. 1992. Nasal chemical senses in rep- tiles: structure and function, pp. 423-523. In C. Gans and D. Crews (eds.), Biology of the Rep- tilia. Vol. 18. New York: Alan R. Liss, Inc. HARTERT, E. 1898. On a collection of birds from north-western Ecuador collected by Mr. W. F. H. Rosenberg. Novitates Zoologicae. 5: 477-505. HARTMANN, M., M. L. DEL GRANDE, M. J. DA C. GONDIM, M. C. MENDES, AND O. A. V. MARQUES. 2002. Reproduction and activity of the snail-eating snake, Dipsas albifrons (Colubri- dae), in the southern Atlantic Forest in Brazil. Studies on Neotropical Fauna and Environment. Bi7(Dye WII zb. HARVEY, M. B., AND A. MUNOZ. 2004. A new species of Tomodon (Serpentes: Colubridae) from high elevations in the Bolivian Andes. Herpetologica. 60(3): 364—372. KOEPCKE, H. W. 1961. Synédkologische studien an der Westseite der peruanischen Anden. Bonner Geographische Abhandlungen. 29: 1—320. KOFRON, C. P. 1982. The identities of some dipsadine snakes: Dipsas elegans, D. ellipsifera and Lepto- gnathus andrei. Copeia. 1982(1): 46-51. LAPORTA-FERREIRA, I. L., M. G. SALOMAO, AND P. SAWAYA. 1986. Biologia de Sibynomorphus (Co- lubridae—Dipsadinae )—reprodugao e habitos alimentares. Revista Brasileira de Biologia 46(4): 793-799. LAURENT, R. 1949. Notes sur quelques reptiles ap- partenant a la collection de l'Institut Royal des Sciences Naturelles de Belgique, [I1—formes américaines. Bulletin de Institute Royal des Sci- ences Naturelles de Belgique. 25(9): 1-20. LYNCH, J. D., AND W. E. DUELLMAN. 1980. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae). Lawrence, Kansas: University of Kansas Museum Dipsas oreas Complex in Ecuador and Peru * Cadle of Natural History, Miscellaneous Publication 69. 86 pp. MARQUES, O. A. V., A. ETEROVIC, AND I. SAZIMA. 2001. Serpentes da Mata Atlantica, guia ilustrado para a Serra do Mar. Ribeirao Preto, Sao Paulo, Brazil: Holos Editora. 184 pp. MARTINS, M. 1993. Why do snakes sleep on the veg- etation in central Amazonia? Herpetological Re- view. 24(3): 84—85. MARTINS, M., AND M. E. OLIVEIRA. 1998. Natural history of snakes in forests of the Manaus region, Central Amazonia, Brazil. Herpetological Natu- ral History. 6(2): 78-150. MtyAaTA, K. 1982. A check list of the amphibians and reptiles of Ecuador with a bibliography of Ec- uadorian herpetology. Smithsonian Herpetologi- cal Information Service. 54: 1—70. Myers, C. W. 1969. The ecological geography of cloud forest in Panama. American Museum Nov- itates. 2396: 1-52. . 1974. The systematics of Rhadinaea, a genus of New World snakes. Bulletin of the American Museum of Natural History. 153(1): 1-262. . 2000. A history of herpetology at the Amer- ican Museum of Natural History. Bulletin of the American Museum of Natural History. 252: 1— YI . 2003. Rare snakes—five new species from eastern Panama: reviews of northern Atractus and southern Geophis (Colubridae: Dipsadinae). American Museum Novitates. 3391: 1—47. MYERS, C. W., AND J. E. CADLE. 1994. A new genus for South American snakes related to Rhadinaea obtusa Cope (Colubridae) and resurrection of Taeniophallus Cope for the “Rhadinaea” brevi- rostris group. American Museum Novitates. 3102: 1-47. . 2003. On the snake hemipenis, with notes on Psomophis and techniques of eversion: a re- sponse to Dowling. Herpetological Review. 34(4): 295-302. MYERS, C. W., R. IBANEZ D., AND J. E. CADLE. ms. On the fragmented distribution of a rare Pana- manian snake, Dipsas nicholsi (Colubridae: Dip- sadinae). American Museum Novitates. NOBLE, G. K., AND H. J. CLAUSEN. 1936. The aggre- gation behavior of Storeria dekayi and other snakes, with especial reference to the sense or- gans involved. Ecological Monographs. 6: 269— 316. ORCES V, G., AND A. ALMENDARIZ. 1987. Sistematica y distribucién de las serpientes Dipsadinae del grupo oreas. Politécnica (Revista de Informacién Tecnico-Cientifica, Quito). 12(4): 135-144. ORTON, J. 1871. Contributions to the natural history of the Valley of Quito—No. I. American Natu- ralist. 5: 619-626; No. II: 693-698. . 1875. The Andes and the Amazon; or Across the Continent of South America. 3rd ed. New York: Harper and Brothers. 645 pp. + 2 foldout maps. 135 PARKER, H. W. 1926. Description of a new snake from Trinidad. Annals and Magazine of Natural History. 18(Ser. 9): 205-207. . 1934. Reptiles and amphibians from south- ern Ecuador. Annals and Magazine of Natural History. 14(Ser. 10): 264-273. . 1938. The vertical distribution of some rep- tiles and amphibians in southern Ecuador. An- nals and Magazine of Natural History. 2(Ser. 11): 438—450. PAYNTER, R. A., JR. 1993. Ornithological Gazetteer of Ecuador. 2nd ed. Cambridge, Massachusetts: Museum of Comparative Zoology. xi + 247 pp. . 1997. Ornithological Gazetteer of Colombia. Cambridge, Massachusetts: Museum of Compar- ative Zoology. ix + 537 pp. PEREZ-SANTOS, C., AND A. G. MORENO. 1988. Ofi- dios de Colombia. Torino, Italy: Museo Region- ale di Scienze Naturali. 517 pp. . 1991. Serpientes de Ecuador. Torino, Italy: Museo Regionale di Scienze Naturali. 538 pp. PETERS, J. A. 1955. Herpetological type localities in Ecuador. Revista Ecuatoriana de Entomologia y Parasitologia. 2: 335-352. . 1960a. The snakes of the subfamily Dipsa- dinae. Miscellaneous Publications of the Muse- um of Zoology, University of Michigan. 114: 1— 224. . 1960b. The snakes of Ecuador, a checklist and key. Bulletin of the Museum of Comparative Zoology. 122: 489-541. . 1965. Liste der rezenten Amphibien und Reptilien: Colubridae (Dipsadinae). Das Tier- reich. 81: i-viii + 1-18 pp. PETERS, J. A., AND B. OREJAS-MIRANDA. 1970. Cat- alogue of the Neotropical Squamata. Part I. Snakes. Bulletin of the United States National Museum. 297: 1—347. PORTO, M., AND R. FERNANDES. 1996. Variation and natural history of the snail-eating snake Dipsas neivai (Colubridae: Xenodontinae). Journal of Herpetology. 30(2): 269-271. ROSSMAN, D. A., AND D. A. KIZIRIAN. 1993. Variation in the Peruvian dipsadine snakes Sibynomorphus oneilli and S. vagus. Journal of Herpetology. 27(1): 87-90. SAGASTEGUI ALVA, A., AND M. O. DILLON. 1991. In- ventario preliminar de la flora del bosque Mon- teseco. Arnaldoa 1(1): 35-52. SAGASTEGUI ALVA, A., I. SANCHEZ VEGA, M. ZAPATA CRUZ, AND M. O. DILLON. “2003” [2004]. Div- ersidad Floristica del Norte del Peri, Tomo II: Bosques Montanos. Trujillo, Peru: Universidad Privada Antenor Orrego. 305 pp. SAUVAGE, H. E. 1884. Sur quelques reptiles de la collection du Muséum d’Histoire Naturelle. Bul- letin de la Société Philomathique de Paris. 8(Ser. 1) AEA SAVAGE, J. M. 1973. A revised terminology for plates in the loreal region of snakes. British Journal of Herpetology. 5(1): 360-362. 136 SAZIMA, I. 1989. Feeding behavior of the snail-eating snake, Dipsas indica. Journal of Herpetology. 23(4): 464-468. SHINE, R. 1993. Sexual dimorphism in snakes, pp. 49-86. In R. A. Seigel and J. T. Collins (eds.), Snakes: Ecology and Behavior. New York: Mc- Graw Hill. SMITH, H. M., AND E. H. TAYLOR. 1945. An anno- tated checklist and key to the snakes of Mexico. Bulletin of the United States National Museum. 187: 1-239. STEPHENS, L., AND M. A. TRAYLOR. 1983. Ornitho- logical Gazetteer of Peru. Cambridge, Massachu- setts: Museum of Comparative Zoology. Vill Teo. G. 1998. Herpetofauna de la Zona Reservada de Tumbes, pp. 79-87. In W. H. Wust (ed.), La Zona Reservada de Tumbes, Biodiversidad y Diagnostico Socioecondémico. Lima: F ondo Na- cional por las Areas Naturales Protegidas por El Estado (PROFONANPE). VIDAL, N., S. G. KINDL, A. WONG, AND S. B. HEDG- ES. 2000. Phylogenetic relationships of xenodon- tine snakes inferred from 12S and 16S ribosomal RNA sequences. Molecular Phylogenetics and Evolution. 14(3): 389-402. WERNER, F. 1909. Uber neue oder seltene Reptilien des Naturhistorischen Museums in Hamburg. I. Schlangen. Mitteilungen aus dem Naturhisto- rischen Museum in Hamburg. 26: 205-247. Bulletin Museum of Comparative Zoology, Vol. 158, No. 3 1922. Synopsis der Schlangenfamilie der Amblycephaliden und Viperiden nebst Ubersicht iiber die kleineren Familien und die Colubriden der Acrochordinengruppe. Auf Grund des Bou- lengerschen Schlangenkatalogs (1893-1896). Ar- chiv fur Naturgeschichte [Berlin]. 8: 185-244. Wust, W. H. (ed.) 1998a. La Zona Reservada de Tumbes, Biodiversidad y Diagnéstico Socioecon- 6mico. Lima: Fondo Nacional por las Areas Na- turales Protegidas por El Estado (PROFON- ANPE). xiv + 180 pp. + pls. 1-6. . 1998b. Aves de la Zona Reservada de Tum- bes, pp. 21-41. In W. H. Wust (ed.), La Zona Reservada de Tumbes, Biodiversidad y Diagnés- tico Socioeconémico. Lima: Fondo Nacional por las Areas Naturales Protegidas por El Estado (PROFONANPE). ZAHER, H. 1999. Hemipenial morphology of the South American xenodontine snakes, with a pro- posal for a monophyletic Xenodontinae and a re- appraisal of colubroid hemipenes. Bulletin of the American Museum of Natural History. 240: 1— 168. ZUG eE, RES. 2B EDGES) AND Om SUNKE Es 81979) Variation in reproductive parameters of three Neotropical snakes, Coniophanes fissidens, Dip- sas catesbyi, and Imantodes cenchoa. Smithson- ian Contributions to Zoology. 300: 1—20. NOTE ADDED IN PROOF: Paulo Passos kindly called my attention to Fernandes et al. (2002), in which Dipsas latifasciatus is synonymized with D. polylepis. These authors could not resolve the status of D. polylepis vis-a-vis D. latifrontalis. ; ; f > se eae eh oe, = a 7 , we . =a «ex ae & , oa! brute 7 t] AY ‘ 7 » F Mint ste a | | * cl ie oD ae. itt » shed yt Dimi an = nh | . - | pee ‘\aale Maconies. ase 4 niin Mat ba ey * ¥ i ae: : wi tat | : «3 , | Ni as ee | ie, 1 ef wy: eis Geel Palaces asd 7 (as aa died Pee Rey aati ; Ba di af ys pi qanta m. § nie poe | | frp ¢ q Adie x atl eae ae Ses ayuee ad Rulletin OF THE , Museum of” Compare Loology The Orb-Weaver Genus Mangora of Mexico, Central America, and The West Indies (Araneae: Araneidae) HERBERT W. LEVI : HARVARD UNIVERSITY | VOLUME 158, NUMBER 4 _ CAMBRIDGE, MASSACHUSETTS, U.S.A. 28 July 2005 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY BREVIORA 1952— BULLETIN 1863— Memoirs 1865-1938 Jounsonia, Department of Mollusks, 1941-1974 OCCASIONAL PAPERS ON MO .uusks, 1945— SPECIAL PUBLICATIONS. i 2. 3. 6. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and Evolution of Crustacea. 192 pp. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere- dinidea (Mollusca: Bivalvia). 265 pp. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. | . Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. . Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ermest E. Williams. 725 pp. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the M. C. Z., Vol. 108.) Reprinted 1971. Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter’s Check-list of Birds of the World, vols. 1-16. Proceedings of the New England Zoological Club 1899-1947. (Complete sets only.) Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2004. THE ORB-WEAVER GENUS MANGORA OF MEXICO, CENTRAL AMERICA, AND THE WEST INDIES (ARANEAE: ARANEIDAE) HERBERT W. LEVI' ABSTRACT. Thirty-two species of Mangora are found in the region under consideration. Three tem- perate species extend their ranges into northern Mex- ico, three Mexican species extend into the Southwest, and four species extend into South America. Only one species, M. melanocephala from Central America and South America, is both widespread and common. One species is found in the West Indies, including Trinidad. Of the 32 species, 15 are new. Six new syn- onyms are found. A few synapomorphies place Man- gora close to Araneus. INTRODUCTION At present, Mexican and Central Amer- ican Mangora can be determined only with the 1894-1904 publications of O. P.- and F. O. P.-Cambridge in the Biologia Centrali-Americana (on Guatemalan spi- ders mostly, but including some from Ta- basco, Mexico), and Chickering’s two pa- pers (1954, 1963) on Panamanian Man- gora. They deal with only a small part of the region. Mangora of most of the region remain unknown and undetermined col- lections have accumulated in museums. The inability to recognize species of the region has limited needed work on ecolo- gy, behavior, and physiology of the genus. Mangora pia was used by Craig (1987a,b, 1988, 1989) for her observations on be- havior and structure of silk. Distinctiveness of most South American Mangora species make it advisable to pub- lish on the species of Mexico, Central America, and the West Indies (excluding Trinidad) and to deal with the many spe- cies of Trinidad and South America sepa- ‘Museum of Comparative Zoology, Harvard Uni- versity, 26 Oxford Street, Cambridge, Massachusetts 02138-2902. E-mail: levi@fas.harvard.edu Bull. Mus. Comp. Zool., 158(4): 139-182, July, 2005 rately. The astonishing abundance of trop- ical South American Mangora species, mostly undescribed, may make this the most species-rich genus of araneids in the Americas. METHODS Preserved Mangora specimens are more fragile than other araneid specimens. It is nearly impossible to avoid breaking leg macrosetae when handling. For examina- tion, specimens were placed on a substrate such as black paraffin with suitable de- pressions, black silicon carbide crystals (80 grit), or black Velcro tape glued to a dish. The black background minimizes reflec- tions. Description. Alcoholic specimens were used to describe coloration even though many have some bright coloration that washes out. No color slides were available to show the color when alive. Eyes. Eyes in araneids are difficult to measure because the curved cornea ex- tends beyond the eye as seen from above. Thus, eyes seen from the side appear larg- er than when viewed from above. There also is individual variation in eye place- ment. Eye measurements are rough esti- mates. The line of posterior eyes was checked against an imagined line behind the eyes when viewing the eyes dorsally. An imag- inary line outside of the eyes, when seen from above, defined the median eye trap- ezoid. Despite the difficulty in measuring, numerical sizes and distances are given. Approximations are easier to describe in numbers than in words. 139 140 Measuring. Total length was measured without stretching the animal, whose elon- gate abdomen hangs at an angle with the prosoma (Levi, 1975, fig. 59; 2002, fig. 2). Length of body parts and leg articles are accurate only to 0.1 mm because legs were not amputated and leveled for measure- ments. Figures. Figures of holotypes of species north of Mexico were made in 1970 (Levi, 1975). Here, the tips of palpi are addition- ally illustrated, perhaps making them eas- ier to determine without amputating the structure. The apical view also helps to dis- tinguish relationship of the complicated sclerites without expanding the palpus. The ventral view of epigyna is often il- lustrated from slightly anterior. Lifting the epigynum with a needle or folding it back exposed the posterior view. The epigyna were cleared with Hoyer’s medium. Indi- vidual specimens of Mangora less than 2.5 mm in total length were submerged in the clearing medium. In larger specimens, the epigynum was removed and cleared for ex- amination and later stored in a small vial with the rest of the specimen. The directions for locating structures in the illustrations (h) refer to the numbers in the face of a clock. Maps. Maps were made with a comput- er. Collecting sites were located with var- ious automobile maps and the gazetteers of the United States Board on Geographic Names (1956, 1965), Office of Geography, Department of the Interior. A gazetteer of the localities of the Cambridges’ types is given by Selander and Vaurie (1962). Col- lectors of specimens often published maps of their collecting sites (e.g., Goodnight and Goodnight, 1953; Petrunkevitch, 1909: Vaurie and Vaurie, 1949). Specimens Used. Specimens used came from the following collections and individ- uals: AD A. Dean, Texas A&M Univer- sity, Collége Station, Texas, United States AMNH American Museum of Natural BMNH CAS CNC CUC ECOSUR FSCA HEF JM MCZ MIUP MNHN NMB PAN REL SMF USNM Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 History, New York United States; W. J. Gertsch, J. A. L. Cooke, N. Platnick Natural History Museum, Lon- don, United Kingdom; P. Hill- yard, J. Beccaloni California Academy of Scienc- es, San Francisco, California, United States: C. Griswold, D. Ubick Canadian National Collections, Ottawa, Ontario, Canada; C. Dondale Cornell University Collection kept in the AMNH ECOSUR Collection, Chetu- mal, Quintana Roo, Mexico; G. Alayo6n Florida State Collection of Ar- thropods, Gainesville, Florida, United States: G. B. Edwards The late Harriet Exline Friz- zell, W. Peck, kept in CAS J. M. Maes, Leon, Nicaragua Museum of Comparative Zo- ology, Cambridge, Massachu- setts, United States: G. Giribet Museo de Invertebrados, Univ- ersidad de Panama, Panama Panama; D. Quintero A. Museum National d’ Histoire Naturelle, Paris, France; M. Hubert, C. Rollard Naturhistorisches Museum, Basel, Switzerland: E. Sutter, A. Hanggi Polish Academy of Science, Warsaw, Poland; J. Proszynski, A. Slojewska Robin E. Leech, Edmonton, Alberta, Canada Senckenberg Museum, Frank- furt am Main, Germany; M. Grasshoff National Museum of Natural History, Washington, D.C., United States; J. Coddington TAXONOMIC SECTION Mangora O. P.-Cambridge Mangora O. P.-Cambridge, 1889: 13. Type species: Mangora picta O. P.-Cambridge from Guatemala, designated by Simon, 1895: 793. Levi, 1975: 116. Diagnosis. Mangora differs from all other araneid genera by the presence of feathered trichobothria on the tibia of the third leg (unfortunately often broken off in poorly handled specimens) (Levi, 1975, fig. 59; 2002, fig. 2). Easier to observe is the unique shape of the carapace: it is bald and narrow in the eye region, usually only one-half or less of the total width of the carapace, and the thoracic region of the carapace is high, equal to or greater than the width of the carapace in the eye region. The high thoracic region slopes evenly toward the eye region (Levi, 1975, fig. 59; 2002, fig. 2). Unlike most araneids, the abdomen is always oval, longer than wide (Figs. 3, 4, 10). The thin legs have long, strong macrosetae, which stand up, not appressed against the appendage (Levi, 1975, fig. 59; 2002, fig. 2). First legs are subequal in length with the fourth. Discrete black frames of the posterior median eyes (Levi 1975, fig. 58), and the posterior eye row being procurved in some species, further distinguish Mango- ra. Description of Female. The venter of the female abdomen is often unmarked. The posterior eye row can be straight, re- curved, or procurved. The anterior median or posterior median eyes are the largest, the laterals smallest, the posterior laterals always slightly smaller than the anterior. The clypeus height is usually equal to the diameter of the anterior median eyes. The first and fourth legs are subequal in length; the fourth is often thicker than the first. Individual specimens of a species vary in darkness of their coloration. Description of Male. The male has sim- ilar but lighter coloration than the female. The eyes are slightly closer to each other than in the female. The endite tubercle is minute, often hard to see or just a tiny CENTRAL AMERICAN MANGORA * Levi 141 swelling. The hook on the first coxa is al- ways present; it may be very small and shifted posteriorly. The male genitalia are surprisingly com- plex. There is always a radix (R), a median apophysis (M), and a soft conductor (C) holding the tip of the embolus (E) (Figs. 89, 90). The embolus is of variable shape, rarely filiform. The terminal apophysis (A) is complex and of enormous size, with soft and hard lobes, difficult to figure out un- less expanded (Figs. 89, 90). The palpal patella has always one strong seta. Sizes. Many of the tropical Mangora are small, females 2.0 to about 7.0 mm total length, males 1.5-4.5 mm. The largest here is Mangora pia, found in Panama and observed by Lubin (1978) and used for ex- periments by Craig (1987-1989). Relationship. Relationship with Ara- neus is suggested by the frequent pres- ence of a scape with a distal pocket in the epigynum of the female (Figs. 62, 97, 110), and in the palpus of the male, the conductor close to the rim, median apophysis with spines (Figs. 89, 90), with- out a paramedian apophysis, and the pres- ence of a large terminal apophysis. This agrees with the cladogram of Scharff and Coddington (1997). Distribution. Thirty-two species are found in Mexico and Central America, 15 of them new. Of these 32, 3 temperate species extend their range into northern Mexico (M. gibberosa, M. placida, and M. spiculata) and 3 Mexican species extend into the Southwest (M. calcarifera, M. fas- cialata, and M. passiva). Four species also are found in South America (M. amchick- eringi, M. falconae, M. melanocephala, and M. pia; Maps 1, 2). Mangora melanoce- phala, the most common Mangora, is also the most widespread, from the Isthmus of Tehuantepec to Peru and Brazil (Map IE). Only one species, M. fascialata, is found in the West Indies excluding Trinidad (Map 2A). Names. An attempt was made to not use specific names used before in the family. 142 Distinguishing Characters of Species. For separating Mangora species, exami- nation of the color pattern is useful, and it is essential to examine not only the ventral view of the epigynum, but also the poste- rior view (Figs. 2, 9, 15), by lifting the epi- gynum with a needle or folding it over. In- ternal genitalia do not provide good char- acters to separate species. Males generally are similar to females in color pattern. Their palpi are so complex that only a slight twist of the palpus pro- vides a different image (Figs. 5-7). Unrecognizable Species. 1 start system- atic revisions by checking catalogs (Plat- nick, 2004; Roewer, 1942). According to these catalogs, Zilla decolorata Keyserling, 1893, is described from Brazil. I borrowed the holotype from BMNH and found it to be a male collected in Bahia, Brazil, and the type is marked Zilla decolorata (C. L. Koch). Koch described a Zilla decora, list- ed in Volume 3 of Bonnet (1961) as de- colorata, but it comes from Germany and is a synonym of Mangora acalypha (Wal- ckenaer, 1802). Although Zilla decolorata Keyserling had been placed in Araneus and Zygiella, the male examined was a Mangora. When working on the Brazilian Man- gora and checking original literature, I found to my surprise that Zilla decolorata was described from a female from Gua- temala. Now that I have seen most Central American Mangora, I do not recognize it. It is relatively large, total length 5.9 mm, but in Central America, only Mangora with a pair of posterior rectangular black spots are this large. Most Mangora are smaller. It might be Chrysometa lancetilla Levi (Tetragnathidae) but Chrysometa has shorter fourth legs than those Keyserling measured for Zilla decolorata. Bonnet (1957) cites Petrunkevitch (1911) as writing that Z. decolorata also appears in Brazil, but Petrunkevitch (1911) indicated that it is found in Gua- temala. (A line below cites a spider from Brazil.) I consider the name unrecogniz- able. Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 KEY TO FEMALE MANGORA Female of one species, M. sufflava, is not known I A black longitudinal line on venter of femora one and two (Fig. 100); abdo- men dorsum with two or three lines (Figé¢ 98409) ain Ose ee deal 2 Legs without black longitudinal lines; dorsum at times with lines In posterior view, epigynum with median plate upside-down T-shaped (Fig. 92); southeastern Canada to Tamaulipas (Mayo Sli) Giese eset eh eee eee gibberosa Epigynum with median plate heart- shaped (Fig. 98); Mexico to Honduras, NWestalnidiesn (lay 2).) sae fascialata Abdomen with circular gray or black dots and/or posterior rectangles on whitish (in alcohol) dorsum (Figs. 139, 178, LSD Wyte ae Wy a, Se eee ee aes 4 Without circular spots or posterior, black réctamgles, Bats Wu a Ee ee 12 One pair of gray spots, plus black rec- tangles or bands on posterior of whitish (in alcohol) abdomen (Figs. 139, 143) Abdomen with spots only or posterior black rectangles only (Figs. 178, 203) Epigynum with borders around lateral plates (Fig. 140); Chiapas (Map 2E) - SH Sod Yah ale A a ee ee goodnightorum Epigynum with lateral plates indistinct (Fig. 136); Campeche (Map 2E) i mene Cashel, ara, SACS 3 TTA Hien campeche Abdomen dorsum with circular, black Cotes (mies WHS), WES)) 5 te 7 Abdomen without black dots, posterior with gray or black rectangles, some- times pairs of lighter rectangles more eimgeralore (Carte MEP) oo a ave oa 8 Abdomen dorsum with a series of black dots (Fig. 178); epigynum as in Figure 175; Yucatan Peninsula to Chiapas GAY Fey opal Niaase PSR ae pS Sle chicanna Abdomen with only one pair of black dots (Fig. 185); epigynum as in Figure 182; Guatemala to Panama (Map 2H) xt A gens tA 6 oY OE Se purulha Epigynum with a posterior, median, oval or angular notch (Figs. 193, 200) ___.. 9 Epigynum with a median posterior scape Notch oval with a median anterior tuber- cle (Fig. 193); central Mexico to Costa Rica (Miaph2 eae a ee bimaculata Notch triangular (Fig. 200); Panama to northern South America (Map 2]) —. Epigynum scape flanked by a semicircu- lar notch on each side (Fig. 189); Costa Rica, Panama (Map 2H) -........... schneirlai 12(3) 14(13) 15(12) 16(15) 21(15) No such notches present Scape a semicircular, short lobe (Fig. 215); in posterior view, median plate flanked by narrow lateral plates (Fig. 216); Costa Rica (Map 2K) corcovado Scape with base narrower than tip (Fig. 208); posterior median plate upside- down T-shaped (Fig. 209); Panama to Venezuela (Map 2K) falconae Abdomen (in alcohol) all white (Fig. 160), or with a median longitudinal white band flanked by darker lateral bands (Figs. 134, 147) Abdomen otherwise Abdomen all white (Fig. 160); México, Michoacan (Map 2G) ixtapan Abdomen with median band flanked by dark lateral bands (Figs. 134, 147) Epigynum with a median oval scape (Fig. 131); Costa Rica, western Panama (Map 2D) fortuna Epigynum with a small median posterior lobe (Fig. 144); Costa Rica (Map 2E) Venter of abdomen posterior to epigyn- um with a median patch with white pigment spots (Figs. 53, 61, 66, 153) Venter of abdomen behind epigynum without a median white patch Abdomen dorsum with a pair of upside- down, comma-shaped marks on black (Fig. 152); epigynum as in Figures 148-151; Arizona to Nicaragua (Map 2F) passiva Abdomen and epigynum otherwise Posterior of epigynum with median plate longer than wide (Fig. 63); Mexico to lomeumrasn (Vitara gil seen eens mobilis Median plate wider than long (Figs. 50, DO) aie Wiens FE eve Pm ee AVR BONO ET Nee Posterior median plate a narrow trans- verse band (Fig. 50); Yucatan Penin- Sulay GMa i hte set ent oh eh eine Posterior median plate not band-shaped (Bigs Shane) les Oe wee Posterior median plate with an anterior lobe on each side (Fig. 72); Honduras to Costa Rica (Map 1H) distincta Median plate without anterior lobes (ERs Ge 5S Silo) create Varet oEae oh 2/1 a Lateral plates in ventral view with a lat- eral, dark mark on median side of scape (Fig. 57); Nayarit (Map 1F) acaponeta Lateral plates with dark marks anteriorly (Fig. 80); Costa Rica to northern South America (Map 11) amchickeringi Epigynum with diagonal duct marks showing through cuticle on each side anterior to median depression (Figs. 18 itza 19 CENTRAL AMERICAN MANGORA ° Lewi 27(26) 28(22) 30(29) 1438 20, 22); Texas to Costa Rica (Map 1D) ie SO aL.” a oe, ee ee calcarifera Epigynum lacking these diagonal marks DD Epigynum with a median scape or pos- terior lobe or sphere (Figs. 104, 110, GRE Gale) Epigynum without scape or median lobe (Figs. 1, 8, 14, 30) Scape flanked by large depressions (Fig. 164); Panama (Map 2G) candida Epigynum lacking depressions next to scape Scape flanked by diagonal sclerotized bars (Fig. 110); southeastern Canada to Nuevo Léon (Map 2B) placida Epigynum otherwise Scape flanked at some distance by curved, parallel bars (Fig. 104); eastern United States to probably Tamaulipas (Map 1K) spiculata Epigynum otherwise In posterior view, epigynum sclerotized, and lateral plates with a dorsal lobe (Fig. 117); Oaxaca (Map 2C) oaxaca In posterior view, lightly sclerotized, lat- eral plates without a dorsal lobe (Figs. eT) In posterior view, median plate with a pair of ventral, lateral lobes (Fig. 122); México, Veracruz (Map 2C) nahuatl Posterior median plate without lateral lobes (Fig. 127); western Panama (Map 9] Bp| lalate 2 SOL er ee OS Se ret Ro volcan Epigynum with indistinct median oval cavity, but with distinct, large, oval spermathecae, closer than their length from the margin (Fig. 14); Mexico to Honduras (Map 1C) Epigynum otherwise Spermathecae more than two diameters from posterior border and a flattened, posterior chevron, usually with a me- dian notch or seam (Figs. 30, 34, 39); Veracruz to Brazil and Peru (Map 1E) staadeass UP MON en ee ties Int melanocephala Spermathecae closer to border, epigyn- um with different border (Figs. 1, 8) Epigynum with a posterior, sclerotized, slightly wavy border (Fig. 8); Costa Rica (Map 1B) craigae Epigynum with a straight border (Fig. 1); Costa Rica, Panama (Map 1A) —_ montana KEY TO MALE MANGORA Males of nine species (M. acaponeta, M. campeche, M. corcovado, M. fortuna, M. goodnightorum, M. nahuatl, M. oaxaca, M. schneirlai, and M. volcan) 1 are not known A black longitudinal line on venter of 144 Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 SES : ae , ; wok SOE, hed “COLOMBIA melanocephala calcarifera SeAz area @acaponeta ® jtza Map 1. Distribution of Mangora species. CENTRAL AMERICAN MANGORA * Levi 145 UNITED STATES fascialata volcan e fortunaa @ campeche 8 goodnightorum vito e nahuatl go Oaxaca Th i Wi Vo mee ao @ candida ® jxtapan 8 sufflava e chicanna 1S a purulha @ schneirlai a corcovado ~® falconae Map 2. Distribution of Mangora species. 146 femora one and two (Fig. 100); abdo- men dorsum with two or three lines (Rips/93 400) Mae a tees ee 2 = Legs without black longitudinal lines ___ 3 Palpus in apical view with a blunt trian- gular projection (Fig. 96); southeastern Canada to Tamaulipas (Map IJ) £cfb Seb iad an Oe dee MN ce, SE gibberosa = Palpus, in apical view, with a wide band (Fig. 101); Mexico to Honduras, West Iinchiesa(Mape2/\) ee ee eee fascialata Macroseta ventrally on proximal end of Oude wcerontoe CS, 87, LOW) ea 4 = Without such macrosetae 6 Small size, less than 2.5 mm total length; palpus with filiform embolus (Figs. 43— 45), Veracruz to Brazil and Peru (Map 1h] Dy eee eh Shae ee SOR tn melanocephala = Large size, more than 3 mm total length; palpus otherwise, Panama to northern South America (Maps 2], K) Palpus, in apical view, with a sclerotized, mG tdavorern (Imes, ALD) falconae = Palpus with only a small thorn (Fig. 204; Map 2]) Abdomen dorsum with black, circular spots or rectangles posteriorly (Figs. SESS 192) ees een ee Tl = Abdomen otherwise 10 Abdomen dorsum with black or gray cir- @ullare Crous (anws, WHS, WSS) ae 8 = Abdomen with black or gray rectangles jomstwesetord yyy (TEMG NOR) ee 9 Abdomen dorsum with a series of pairs of dots (Fig. 178); palpus as in Figures 179-181; Yucatan Peninsula to Chiapas (Miaro: 251) ioe 2s oie ae eee eek shames chicanna ~ Abdomen dorsum with one pair of dots (Fig. 185); palpus as in Figures 186— 188; Guatemala to Panama (Map 2H) CE Bean ee ed tina! Petit St purulha Palpus, in apical view, with sickle-shaped structure (Fig. 197); central Mexico to Costa Rica (Mia) 2) ee bimaculata _ Palpus, in apical view, with drawn-out point (Fig. 173); western Panama (Map 2G) sufflava Left palpus, in apical view, showing em- bolus with counterclockwise filament (Figs: 5 aileli26). 0 eee a eee ul = AN BICAll VIO OWNEINIISS cccnecaccceeecceenneeneenese 14 11(10) Embolus turns clockwise before revers- ing direction (Fig. 26); Texas to Costa Rica (Map 1D) calcarifera = Embolus turns only counterclockwise (Higs. Social) tes ace ian Si 12 12(11) Terminal apophysis, in apical view, with single, large, sclerotized thorn adjacent to a truncate tip (Fig. 11); Costa Rica craigae Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 = Terminal apophysis with only thorn or only truncate tip 13(12) Terminal apophysis, in apical view, with only a thorn (Fig. 5); Costa Rica, Pan- ama (Map 1A) montana = Terminal apophysis, in apical view, with only a truncate tip (Fig. 17); Mexico to Honduras (Map 1C) picta 14(10) Terminal apophysis, in apical view, with a sclerotized, pointed rod (Figs. 67, 76, SS, SOL ASA? 6. Sok 2 Oh A) dace! ee 15 Terminal apophysis otherwise 20 15(14) Rod sword-shaped with one straight and one curved edge (Fig. 154); Arizona to Nicaragua (Map 2F)) =o) =) passiva = Terminal apophysis tip otherwise _______ 16 16(15) Rod with right angle (Fig. 161); México, Michoacan (Map 2G) __ ixtapan = Rod otherwise (Fig. 168) 17(16) Sword-shaped rod with both sides evenly curved (Fig. 168); Panama (Map 2G) TSR ee a a ele candida = Terminal rod first curved in one direc- tion, distally in another (Figs. 67, 76, 85) ek ee a A oe ee 18 18(17) In mesal view, median apophysis with evenly curved “top” (Fig. 79); Hondu- ras to Costa Rica ( Map iE) == distincta - Median apophysis “top” not so curved CHigs. <0, 88) 2) Ae Saas eee ee 19 19(18) Both spines of median apophysis almost equal in size; “top” flattened (Fig. 88); Costa Rica to northern South America (Mays 2. oe a ee amchickeringi “= One spine much larger than other; “top” skewed (Fig. 70); México to Honduras (Map: LG) Ss 00: (eae ae eee mobilis 20(14) Terminal apophysis, in apical view, with thorn and lobe, separated by a notch as in Figure 54; Yucatan Peninsula CMA ain SIGE) coe. ole die EN itza = Terminal apophysis otherwise (Figs. 107, iLa]RS) eaten 2 ey RCT 2 a ee Ae ee Mh 21(20) Median apophysis with two spines (6h in Fig. 108); southeastern Canada to Nueva Léon (Map 1K) __. spiculata = Median apophysis with one spine (6 h in Fig. 114); eastern United States to Nuevo Kéonk (Map) 23) oss placida Mangora montana Chickering Figures 1-7; Map 1A Mangora montana Chickering, 1954: 204, figs. 15-17, 26. Male holotype and female allotype from El Voledn, Panama, in MCZ, examined. Platnick, 92004. Description. Female paratype from El Voleén. Carapace, legs, sternum grayish orange-white. Abdomen dorsum with black, gray, and white pigment spots (Fig. 3); venter well marked (Fig. 4). Posterior eye row procurved. Ocular trapezoid long- er than wide, widest posteriorly. Posterior median eyes 1.5 diameters of anterior me- dians; lateral eyes 0.8 diameter. Anterior median eyes 0.9 diameter apart, 0.9 from laterals. Posterior median eyes 0.7 diame- ter apart, 0.7 from laterals. Total length 3.4 mm. Carapace 1.5 mm long, 1.1 wide in thoracic region, 0.7 wide behind eyes, 0.7 high. First femur 1.4 mm, patella and tibia 1.7, metatarsus 1.4, tarsus 0.7. Second pa- tella and tibia 1.7 mm, third 1.0, fourth 1.6. Male from Osa Peninsula, Costa Rica. Lighter than female, with only faint col- oration on dorsum of abdomen. Posterior eye row procurved. The median eye trap- ezoid is longer than wide, slightly wider posteriorly. Posterior median eyes 1.3 di- ameters of anterior medians; lateral eyes 0.6 diameter. Anterior median eyes 0.8 di- ameter apart, 0.8 from laterals. Posterior median eyes 0.4 diameter apart, 0.4 from laterals. Height of clypeus 1.5 diameters of anterior median eye. Total length 2.3 mm. Carapace 1.3 mm long, 1.0 wide in tho- racic region, 0.4 wide behind lateral eyes, 0.7 high. First femur 1.3 mm, patella and tibia 1.4, metatarsus 1.2, tarsus 0.6. Sec- ond patella and tibia 1.2 mm, third 0.7, fourth 1.2. Variation. Total length of females 2.6— 3.6 mm, males 2.1—2.6 mm. The illustra- tions were made from female paratypes and males from El Volc4n, Panama. Diagnosis. Mangora montana is darker (Figs. 3, 4) than M. craigae (Fig. 10), and the epigynum has a sclerotized, almost posterior straight edge (Fig. 1), whereas that of M. craigae is less sclerotized (Fig. 8) and the posterior edge has slight dents (Figs. 8, 9). The male is distinguished from that of M. craigae by a differently shaped terminal apophysis (compare 12 h in Figs. 5-7 and 12 h in Figs. 11-13). Distribution. Costa Rica, western Pan- ama (Map 1A). CENTRAL AMERICAN MANGORA * Levi 147 Paratypes. PANAMA Chiriqut: Boquete, July 1939, 12 (A. M. Chickering, MCZ); El Volcan, 20 Feb. 1936, many 26 (W. J. Gertsch, AMNH); Aug. 1950, 29 (A. M. Chickering, MCZ); Cerro Punta, 3 Feb. 1936, 12 (W. J. Gertsch, AMNH). Specimens Examined. COSTA RICA Puntarenas: Butlers Finca, 9°15’N, 83°47'W, 28 Jan. 1976, 29 (Roth-Schropfer, AMNH); San Isidro del General, 660—1,300 m, 22 (D. Round, MCZ); Osa Peninsula, 4 km SW Rincon, 08°42'N, 83°29'W, Feb. 1967, 1d (Organization for Tropical Studies, MCZ). Mangora craigae new species Figures 8-13; Map 1B Holotype. Male holotype from Monteverde Cloud Forest, Bosque del Rio, 1,580 m, Puntarenas Prov- ince, Costa Rica, 13 July 1977, female paratype, 4 Aug. 1977 (C. L. Craig) in MCZ. The species is named after the collector, arachnologist Cay Craig. Description. Female paratype. Prosoma orange, with legs lighter. Abdomen or- ange-white with a pair of dorsal, longitu- dinal bands of white spots (Fig. 10), sides with similar bands, joining anteriorly; ven- ter with bands between epigynum and spinnerets. Posterior eye row procurved. Ocular trapezoid longer than wide, widest posteriorly. Posterior median eyes 1.3 di- ameters of anterior medians; anterior lat- eral eyes 1.0 diameter, posterior laterals 0.7. Anterior median eyes their diameter apart, 1.0 diameter from laterals. Posterior median eyes their diameter apart, 1.0 di- ameter from laterals. Total length 3.5 mm. Carapace 1.6 mm long, 1.3 wide in tho- racic region, 0.7 wide behind eyes, 0.8 high. First femur 1.7 mm, patella and tibia 1.8, metatarsus 1.7, tarsus 0.8. Second pa- tella and tibia 1.7 mm, third 1.3, fourth ily Male holotype. Lighter than female. Eye arrangement similar to that of female. Posterior median eyes 1.0 diameter of an- terior medians; lateral eyes 0.6 diameter. Anterior median eyes 1.0 diameter apart, 0.8 from laterals. Posterior median eyes 0.8 diameter apart, 0.8 from laterals. Total length 2.7 mm. Carapace 1.4 mm long, 1.2 wide in thoracic region, 0.5 wide behind eyes, 0.8 high. First femur 1.5 mm, patella ang) tibia 16, metatarsus 123, tarsus 0,7. 148 Second patella and tibia 1.5 mm, third 0.8, fourth 1.5. Variation. Total length of females 2.8— 3.8 mm, males 2.6—2.7 mm. The illustra- tions were made from female paratypes, and male from the holotype. Diagnosis. Mangora craigae is lighter than M. montana and has the epigynum lightly sclerotized with an undulating pos- terior edge (Fig. 8), unlike that of M. mon- tana (Fig. 1); in posterior view the median plate is smaller (Fig. 9) than that of M. montana. In the palpus of the male, the terminal apophysis (12 h in Figs. 11-13) is different in shape from that of M. montana (Figs. a Natural History. Specimens have been found in cloud forest at Monteverde, and roadside in forest in Braulio Carrillo Na- tional Park. Distribution. Costa Rica (Map 1B). Paratypes. COSTA RICA Cartago [?]: Orosi [“cen- tral flat land,’ Reimoser, 1940], 12 (Picado, MC7Z). San José: Braulio Carrillo Natl. Park, 1,400-1,600 m, 26 July 1983, 22 (H., L. Levi, W. Eberhard, MCZ); Bajo Hondura, 1,250 m, Mar. 1986, 32; 26 July 1983, 3¢; Dec. 1989, 22 (W. Eberhard, MCZ): Rio Hon- dura, 30 Mar. 1991, 1,200 m, 22 (W. Eberhard, MCZ); Zurqui, 1,600 m, May 1992, 1d (W. Eber- hard, MCZ). Alajuela: Pefias Blancas, headwaters of Rio Pefias Blancas, 13 July 1980, 12 (J. Coddington, MCZ). Puntarenas: Monteverde Biol. Reserve, 17 Mar. 1979, 12; 3 Apr. 1979; 1¢; 5 june 1980, 12 (J. Coddington, MCZ, USNM); Bosque del Rio, 1 May 1977, 19; 11 May 1977, 19; 3 July 1977, 19; 23 July 1977, 19; 3 Aug. 1977, 12; 24 May 1978, 16 (C. L. Craig, MCZ); Bosque Nuboso, 22 July 1978, 1¢ (C. L. Craig, P. Klass, MCZ); Las Cruces nr. San Vito, 1,100 m, Jan. 1987, 22 (W. Eberhard, MCZ). Mangora picta O. P.-Cambridge Figures 14-19; Map 1C Mangora picta O. P.-Cambridge, 1889: 14, pl. 3, fig. 5, 3. Male lectotype here designated from Chac- Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 tum [?], Guatemala in BMNH, examined. F. P- Cambridge, 1904: 479, pl. 45, fig. 14, ¢. Platnick, 2004. Mangora trilineata O. P.-Cambridge, 1889: 14, pl. 3, fig. 7, 2. Female lectotypes, here designated from Chiacam [coffee plantation, Alta Verapaz, 26 km NE Coban (Selander and Vaurie, 1962)], Guate- mala, in BMNH, examined. F. O. P.-Cambridge, 1904: 480; pl. 45, fig. 20, 2. Chickering, 1954: 213, figs. 27, 28, 2. Platnick, 2004. NEW SYNONYMY. Note. Chickering (1954) noted, correct- ly, that the female described with Mangora picta is actually the female of M. melano- cephala (=M. spinula), but overlooked the correct match of the female of M. trilinea- ta with the male of M. picta. Males and females are commonly collected together. Description. Female from Xilitla, San Luis Potosi, Mexico. Carapace orange, with a dusky patch on each side of thoracic region. Legs light orange. Abdomen or- ange-white, with posterior median black ladder, and areas with white pigment spots (Fig. 16); venter orange-white. Posterior eye row procurved. Ocular trapezoid lon- ger than wide, widest posteriorly. Posterior median eyes 1.5 diameters of anterior me- dians; lateral eyes 0.7 diameter. Anterior median eyes 0.6 diameter apart, 0.6 from laterals. Posterior median eyes 0.4 diame- ter apart, 0.5 from laterals. Total length 2.7 mm. Carapace 1.2 mm long, 1.0 wide in thoracic region, 0.5 wide behind eyes, 0.7 high. First femur 1.5 mm, patella and tibia 1.7, metatarsus 1.4, tarsus 0.6. Second pa- tella and tibia 1.5 mm, third 0.9, fourth LZ, Male lighter than female. Posterior me- dian eyes 1.5 diameters of anterior medi- ans; lateral eyes 0.7 diameter. Anterior me- dian eyes 1.0 diameter apart, 0.8 from lat- erals. Posterior median eyes 1.3 diameters apart, 0.8 from laterals. Total length 1.8 —s Figures 1-7. Mangora montana Chickering. 1-4, female. 1, 2, epigynum. 1, ventral. 2, posterior. 3, abdomen, dorsal. 4, ab- domen, ventral. 5-7, male, left palpus. 5, apical. 6, mesal. 7, ventral. Figures 8-13. Mangora craigae new species. 8-10, female. 8, 9, epigynum. 8, ventral. 9, posterior. 10, abdomen, dorsal. 11— 13, male, palpus. 11, apical. 12, mesal. 13, ventral. Figures 14-19. Mangora picta O. P.-Cambridge. 14—16, female. 14, 15, epigynum. 14, ventral. 15, posterior. 16, abdomen, dorsal. 17-19, male, palpus. 17, apical. 18, mesal. 19, ventral. CENTRAL AMERICAN MANGORA °* Levi 149 Figures 20-29. Mangora calcarifera F. O. P.-Cambridge. 20-25, female. 20-23, epigynum. 20, 22, ventral. 21, 23, posterior. 24, abdomen, dorsal. 25, abdomen, ventral. 26—29, male, palpus. 26, apical. 27, mesal. 28, ventral. 29, lateral. Scale lines: 1.0 mm, genitalia 0.1 mm. 150 mm. Carapace 1.1 mm long, 0.9 wide in thoracic region, 0.4 wide behind eyes, 0.6 high. First femur 1.2 mm, patella and tibia 1.5, metatarsus 1.2, tarsus 0.7. Second pa- tella and tibia 1.4 mm, third 0.8, fourth 18 Variation. Total length of females 2.5— 3:4 mom, males: 1:8—24 nm: DheMllistra tions were made from the female lecto- types of M. trilineata, and from the male lectotype of M. picta. Diagnosis. The epigynum of M. picta is lightly sclerotized; the depression is vari- able in shape, and the rim barely distinct. It is distinguished from all other species by the large, distinct pair of seminal re- ceptacles, less than their length from the posterior edge (Fig. 14). Males have a distinct terminal apophysis (12 h in Figs. 17-19) differing from that of M. montana (Figs. 5-7) and M. craigae (Figs. 11-13). Natural History. Specimens have been collected from a bromeliad at Fortin, Ve- racruz, and from tropical, deciduous forest in Jalisco, Mexico. Distribution. Nuevo Léon, Mexico to western Honduras (Map IC). Specimens Examined. MEXICO Nuevo Léon: Tier- an, 20 km NW Laredo, 27 July 1945, 12 (H. Wagner, AMNH). San Luis Potost: Fortin, 25 July 1956, 1°, 2d (V. Roth, AMNH); 26 June 1944, 2° (L. I. Davis, AMIN ED)? 322, IN) ontinyn Se 95.61 (| eu aelzies AMNH); Huachinango, 7 Oct. 1947, 12 (H. M. Wag- ner, AMNH); Huichichuyan, 19 May 1952, many 2 6 (W. NE Gertsch, AMNH): Picolo, 21 May 1952, 2 (W. J. Gertsch et al, AMNH); Tamazunchale, 6, 7 July 1941, 19 (A. M., L. I. Davis, AMNH); 23 Nov. 1946, 12 (CAS); 20 May 1952, 19, 16 (W. J. Gertsch, AMNH); 19 Apr. 1963, 12 (W. J. Gertsch, W. Ivie, AMNH): Valles, 1961, 12 (L. Steude, AMNH); 13 km NNW Xilitla, 23 July 1954, 12, 2d (R. Dreisbach, MCZ); Xilitra, 16 km NE Xilitla, 19 Apr. 1963, 12 (W. J. Gertch, W. Ivie, AMNH); Cueva de Salitre, ca, 21°23'N, 98°23 W, 13 june 1983719 (W. Maddison, MCZ). Nayarit: Compostela, 16 Sep. 1957, 12 (R. Dreisbach, MCZ); Tepic, 22-24 Sep. 1947, 22 (B. Malkin, AMNH); 30 km SW Rio Cafnas, 4 Nov. 1987, 12 (V. F. Lee, CAS). Jalisco: Municipio El] Limon, 3 Sep. 1997, 19 (F. Alvarez, MCZ); 26 km S Autlan, 8 July 1984, 1? (J. Woolley, AD); Esta. Biol. Chamela, 100 m, Sep. 1987, 12 (W. Eberhard, MCZ). Colima: 14 km N Comala, 12 July 1984, 2d (J. Woolley, AD). Veracruz: Cérdoba, 1909, 42, 1d Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 (A. Petrunkevitch, AMNH); 13-15 May 1946, 49 (J. C., D. L. Pallister, AMNH); 3.2 km E Cordoba, 6 Aug. 1966, 12 (J., W. Ivie, AMNH); Fortin, 20 May 1947, 12 (Bordas, AMNH); 5 Aug. 1966, many ? d (J., W. Ivie, AMNH); 4.2 km NE Huatusco, 22 July 1985, many 2d (J. Woolley et al., AD); Jalapa, 22 May 1946, many ¢¢ (J. C., D. L. Pallister, AMNH); Roma nr. Jalapa, Aug. 1948, 1d (C., M. Goodnight, AMNH); July 1981, 32, 16 (C. Gold, CAS); La Plan- ta de Moctezuma, nr. Fortin; 7 July 1947, 12 (C., M. Goodnight, AMNH); 17 Dec. 1948, 12 (H. B. Leech, CAS); Mirador, Zacuapam, 20 Aug. 1933, 12 (W. James, AMNH); Orizaba, 12 Feb. 1954, 12 (R. R. Dreisbach, MCZ); Penuela, 18°53’N, 96°48’W, 26 Apr. 1963, 12 (W. J. Gertsch, W. Ivie, AMNH); Po- trero, 24 June 1936, 19 (L. I. Davis, AMNH); Tla- pacoyan, 300 m, 7-8 July 1946, 12 (H. Wagner, AMNH); 24 June 1947, 29S Gaal. DaviswANUNIED)? Hidalgo: Chapulhuacan, 20 May 1952, 12 (W. J. Gertsch et al., AMNH); 27 July 1966, 1° (J., W. Ivie, AMNH). Puebla: Huauchinango, 7 Oct. 1947, many 23 (H. Wagner, AMNH); Villa Juarez, 10 Oct. 1947, many ¢d6 (H. Wagner, AMNH); 1.5 km. W of Hwy. 130, bypass of Xicotepec de Juarez, ca. 20°17’'N, 97°59'W, 17 June 1983, 1d (W. Maddison, MCZ); 7.5 km. SW La Cumbre, 1,730 m, 23 July 1987, 32, 4d (Kopvarik, Schaffner, AD); 6.2 km NE San Gabriel Mixtepec, 16 July 1985, 12 (J. Woolley, AD). Guer- rero: 3.2 km E Ocotito, 11 July 1985, 1d (J. Woolley et al, AD). Oaxaca: 7.5 km S San Gabriel Mixtepec, 16 July 1985, 2d (J. Woolley et al., AD); 30.4 km S San Miguel Suchixtepec, 17 July 1985, 12 (J. Wool- ley, AD); Soledad, 7 May 1947, 12 (H. Wagner, AMNH); Porto Escondido, 15 July 1985, 2d (J. Woolley et al., AD); 12.8 km NE EI Punto, 18 July 1985, 1d (J. Woolley et al., AD). Chiapas: Finca Cuauhtemoc nr. Cacahuatan, 3-13 Aug. 1950, 2°, 2d (C., M. Goodnight, AMNH); Finca Cusuhtemoc, 6 Aug. 1950, 12 (C. Goodnight, AMNH); Finca Lu- beca, 20 km N Huixtla, 11 Jan. 1945, 22, 1d (T. C. Schneirla, AMNH); Lagunas de Montebello, 48 km Comitan, 23 Aug. 1977, 42, 1d (T. C. Meikle, C. E. Griswold, CAS); Ocosingo, 24, 25 June 1950, 12, 1d (C., M. Goodnight et al, AMNH); rim of canyon 5 km S Sumidero, 16°48’N, 93°05'W, 17 Aug. 1966, 1° (J., W. Ivie, AMNH). GUATEMALA Yepocapa, Chi- maltenango, 30 Aug. 1948, 12 (H. T. Dalmat, USNM): 1,400 m, 27 July 1949°426, “Ci “Rar: AMNH); Moca, June 1947, many 2 6; 31 Aug. 1947, 12 (C., P. Vaurie, AMNH); Tumahu, 1,000 m, 10-11 July 1947, 22 (C., P. Vaurie, AMNH). Quixchaya: Suchitepequez, 9 Oct. 1944, 1d (H. Elishewitz, AMNH). HONDURAS Copdn: Copan, 16 Feb. 1937, sweeping weeds, 12 (Roys, MCZ). Mangora calcarifera F. O. P.-Cambridge Figures 20-29; Map 1D Mangora calcarifera F. O. P.-Cambridge, 1904: 479, pl. 45, figs. 12, 13, 2d. Two female, two male syn- types from Petexbattin, Guatemala, in BMNH, ex- amined. Levi, 1975: 132, figs. 131-144. Platnick, 2004. Description. Description is found in Levi (1975). Variation. Total length of females 3.2— 4.4 mm, males 2.2—2.7 mm. Diagnosis. The epigynum of M. calcar- ifera is lightly sclerotized and variable; it is distinguished from that of other species by the one or two pairs of diagonal marks made by ducts that show through the transparent cuticle, anterolateral to the median depression (Figs. 20, 22). Unlike other species with a lightly sclerotized epi- gynum, in M. calcarifera the venter of the abdomen has two pairs of areas containing silver spots (Fig. 25). The male has a longer filamentous em- bolus in the palpus (Figs. 27, 28) than have other Mexican and Central American Mangora, and has a distinct distal flap like a rabbit’s ear on the tip of the palpus (Figs. 26-29), similar to the palpus of M. melan- ocephala (Fig. 45). However, the male lacks the macroseta on the fourth femur present in the male of M. melanocephala (Fig. 47). Natural History. A specimen was found in deep weeds at Salto Falls, San Luis Po- tosi. Distribution. Texas to Costa Rica, most- ly in eastern regions, the Gulf drainage (Map 1D). Specimens Examined. TEXAS Cameron Co.: 16 km SE Brownsville, 8 Oct. 1937, 12 (Davis, Fones, MCZ). MEXICO Tamaulipas: 1 km N Gomez Farias, ZO Deca Sila: 2a (Crap Durden, WISNIM) ole kimes Ciudad Victoria, 29 Jan. 1947, 12 (AMNH). San Luis Potost: 10 km N Naranjo Salto, 5 May 1973, 1d (B. Vogel, USNM); nr. Ciudad del Maiz, 760 km, 28 July 1953, 12 (C. J. Goodnight, AMNH); Huichichuyan, 19 May 1952, 2° (W. J. Gertsch, AMNH); Salto Falls, 5 Feb. 1967, 12 (W. Peck, CAS): Tamanzuchale, 15 Apr. 1946, 22 (L. Davis, M. Johnston, AMNH); 20 May 1952, 13 (W. J. Gertsch et al., AMNH); 20 July 1956, 1d (W. J. Gertsch, V. Roth, AMNH); 1.6 km SW Tamazunchale, 25 July 1966, 12 (J., W. Ivie, AMNH); Xilitla, 23 July 1954, 1¢ (R. Dreisbach, MCZ); 16 km NE Xilitla, 19 Apr. 1963, 1d (W. ip Gertsch, W. Ivie, AMNH): Valles, 1961, 22, 16 (L. Steude, AMNH). Veracruz: Catemaco, Playa Azul, 9 Aug. 1966, 12, 1d (J., W. Ivie, AMNH); 7.5 km W of Catemaco, 23 June 1982, 12 (F. Coyle, MCZ); 4 CENTRAL AMERICAN MANGORA ¢ Levi Salk km E Huatusco, 23 July 1984, 1? (J. B. Woolley, AW) =) nie ela aealmas IN (of Catemaco, WNSrs6eN- 95°07'W, 29 June-1 July 1983, 29: 1, 2 Aug. 1983, 22 (W. Maddison, MCZ); Jesus Carranza, 13 July 1947, 12 (C. Goodnight, AMNH); Los Tuxtlas Biol. Sta., July 1990, 12 (B. Traw, MCZ); Martinez de la Torre, 4 July 1946, 1d (H. Wagner, AMNH); Papan- tla, 12 Oct. 1947, 2d (H. Wagner, AMNH); Poza Rica, 13 Apr. 1946, 12 (C. Bolivar, AMNH); Teco- lutla, 13 Oct..1947, 12 (H. Wagner, AMNH); Tla- pacoyan, 300 m, 1, 8 July 1946, 1d (H. Wagner, AMNH). Hidalgo: SW Progreso, March 1936, 19, 16 (L. I. Davis, AMNH). Puebla: Huachinango, 7 Oct. 1947, 12, 26 (H. M. Wagner, MCZ). Oaxaca: Tolosa, 1-12 Apr. 1947, 12 (B. Malkin, AMNH). Campeche: 6 km W Francisco Escarcegan, 18°37'N, 90°48’W, 11, 12 July 1983, 12 (W. Maddison, MCZ). Quintana Roo: 31 km NE of Felipe Carrillo, 19°48'N, 87°52'W, 17 July 1983, 22 (W. Maddison, MCZ). Chiapas: Palenque Ruins, 9 July 1949, 1d (C. Good- night, AMNH): Ocosingo, 900 m, 24, 25 June 1950, 12, 26 (C., M. Goodnight et al, AMNH). GUA- TEMALA Petén: Tikal, 7 July 1975, 1¢ (W. Sedg- wick, MCZ); Uaxactin, Mar., Apr. 1931, 1° (H. H. Bartlett, MCZ). HONDURAS nr. Lancetilla, 19 July 1929, 1d (A. M. Chickering, MCZ). COSTA RICA San José: Cerro Zurqui, 1,600 m, May 1992, 12 (W. Eberhard, MCZ). Mangora melanocephala (Taczanowski) Figures 30-47; Map 1E Linyphia melanocephala Taczanowski, 1874: 70. Male and female syntypes from Cayenne, French Gui- ana, in PAN, examined. Zilla melanocephala:—Keyserling, 1881: 552, pl. 16, fig. 4, 93; 1893: 302, pl. 15, fig. 223, 22. Mangora picta:—O. P.-Cambridge, 1889: 14, pl. 3, fig, 6, 2; F. O. P.-Cambridge, 1904: 479, pl. 45, ° (female only, not male holotype). Error first re- ported by Chickering, 1954. Mangora spinula F. O. P.-Cambridge, 1904: 480, pl. 45, fig. 18, d. Male holotype from Teapa, [Tabas- co], Mexico, in BMNH, examined. Chickering, ele HIE figs. 93-26, 2. Platnick, 2004. NEW SYNONYMY. Mangora dentembolus Chamberlin and Ivie, 1936: 59, pl. 12, figs. 114-116, ¢. Male holotype from Barro Colorado Island, Panama, vial present in AMNH, but specimen lost. Synonymized with spi- nula by Chickering, 1954. NEW SYNONYMY. Zygiella melanocephala:—Roewer, 1942: 887. Mangora aragarcensis Soares and Camargo, 1948: 372, figs. 27, 28, 2. Female holotype from Aragar- cas, Goids, Brazil, in MZSP no. 1215, examined. Platnick, 2004. NEW SYNONYMY. Mangora melanocephala:—Caporiacco, 1948: 659. Platnick, 2004. Caporiacco first recognized that Linyphia melanocephala is a Mangora. Mangora pozonae Schenkel, 1953: 20, fig. 18, 2. Fe- male holotype from Conwarook (Potaro), Poz6n, 152 Depto. Acosta, Falc6én, Venezuela, in NMB, ex- amined. Platnick, 2004. NEW SYNONYMY. Note. Keyserling (1881) had apparently examined the original specimens of Tac- zanowski and made good illustrations of the genitalia, which had been ignored. Description. Female from Panama. Pro- soma orange, with a dusky patch on each side of thoracic region of carapace (Fig. 38), legs lighter. Abdomen orange-white, with posterior median black ladder, and ar- eas with white pigment spots (Fig. 38); venter orange-white. Posterior eye row procurved. Ocular trapezoid longer than wide, widest posteriorly. Posterior median eyes 1.5 diameters of anterior medians; lateral eyes 0.7 diameter. Anterior median eyes 0.6 diameter apart, 0.6 from laterals. Posterior median eyes 0.4 diameter apart, 0.5 from laterals. Total length 2.7 mm. Carapace 1.2 mm long, 1.0 wide in tho- racic region, 0.6 wide behind eyes, 0.7 high. First femur 1.5 mm, patella and tibia 1.7, metatarsus 1.4, tarsus 0.6. Second pa- tella and tibia 1.5 mm, third 0.9, fourth Lf. Male from Panama lighter than female. Posterior median eyes 1.3 diameters of an- terior medians; lateral eyes 0.8 diameter. Anterior median eyes 1.0 their diameter apart, 0.8 from laterals. Posterior median eyes 1.3 diameters apart, 0.8 from laterals. Total length 1.8 mm. Carapace 1.1 mm long, 0.9 wide in thoracic region, 0.4 wide behind eyes, 0.6 high. First femur 1.2 mm, patella and tibia 1.5, metatarsus 1.2, tarsus 0.7. Second patella and tibia 1.4 mm, third 0:8, fourth 1.3. Variation. Total length of females from Central America 2.3-3.5 mm, males 1.7— 2.1 mm. The epigynum is lightly sclero- tized and the posterior rim quite variable (Figs. 30, 34, 39); the posterior swelling varies in angles and in thickness; in pos- terior view the median borders of the lat- eral plates are usually, but not always, par- allel and close to each other (Figs. 31, 35). Diagnosis. The epigynum of M. melan- ocephala is lightly sclerotized and distin- Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 guished from that of other Mangora in the region by having the spherical spermathe- cae of the epigynum placed far anterior from the chevron-shaped posterior swell- ing (Figs. 30, 34); the swelling is variable in thickness and shape, rarely sclerotized (Fig. 30). The median plate, in posterior view, is narrow and bordered by parallel edges of the lateral plates (Fig. 31). Males are the only tiny (1.7—-2.1 mm) Mangora in Central America with a ma- croseta on the ventral face of the fourth femur (Fig. 47); the others with macroseta on the fourth femur are the larger-sized M. pia and M. falconae. Another distinctive character is the projection on the terminal apophysis of the palpus, and the ragged- tipped embolus (Figs. 43-46). Natural History. The species has been collected in various habitats: river bottom; in lowland forest south of Volcan, Costa Rica; in understory of secondary forest, in open vegetation and cultivated areas at La Selva, Costa Rica; in low vegetation, forest edge and low vegetation, dense jungle near Turrialba, Costa Rica; in vegetation along ditch at Finca la Laja Forest in Costa Rica. Also, it has been found as prey of Trypoxylon saussurei wasps in Costa Rica (no locality, R. Coville collection) and can- opy fogging of monsoon forest in Panama City and humid forest in Colon. Distribution. From the Isthmus of Te- huantepec to Peru and Brazil (Map 1E). (Additional South American records will be published in a separate paper.) Specimens Examined. MEXICO Veracruz: 61 km S Acayucan, 17°57'N, 94°54'W, 2 March 1976, 2 (V. Roth, AMNH); Catemaco, 9 Aug. 1966, 26 (J., W. Ivie, AMNH); La Buena Ventura, July 1909, 2d (A. Petrunkevitch, AMNH); 16 km S San José del Car- men, 16 Apr. 1953, 3d (L. I. Davis, AMNH). Chiapas: Cacahuatan, 9 Aug. 1950, 2 (C., M. Goodnight, AMNH); El Real, 6, 7 July 1950, 2 (C., M. Good- night, AMNH); Escuintla, 2¢ (N. Banks, MCZ); Finca Santa Marta, nr. Huehuetan, 31 July—1 Aug. 1950, 2 (C., M. Goodnight, AMNH); Ocosingo, 900 m, 25 June 1950, ¢ (C., M. Goodnight et al., AMNH); Las Ruinas Palenque, July 1948, 2d (C., M. Goodnight, AMNH); 22 July 1949, d (C. Good- night, AMNH); Pichuacalco, 17 July 1947, 2 (C. J. Goodnight, AMNH); La Zacualpa, Aug. 1909, 2d (A. CENTRAL AMERICAN MANGORA * Levi 153 ine Figures 30-47. Mangora melanocephala (Taczanowski). 30-42, female. 30-37, 39-41, epigynum. 30, 34, 39, ventral. 31, 35, 40, posterior. 32, 36, ventral cleared. 33, 37, 41, posterior, cleared. 30-34, syntype. 35-37, Costa Rica, unusual specimen. 38, carapace and abdomen. 39-42, Nicaragua, doubtful determination. 42, abdomen, dorsal. 43-46, male, left palpus. 43, apical. 44, mesal. 45, ventral. 46, left palpus, expanded. 47, macroseta on fourth femur. Scale lines: 1.0 mm, genitalia 0.1 mm. Abbreviations: A, terminal apophysis; C, conductor; E, embolus; M, median apophysis. 154 Petrunkevitch, AMNH):; Selva del Ocoto, 32 km NW Ocozocoautla, 27 Aug. 1973, d (C. Mullinex et al., CAS). BELIZE Mt. Pine Ridge, Feb.—Mar. 1931, 2 (H. H. Bartlett, MCZ); nr. Rio Frio Cave, 5 June 1974, d (C. J. Goodnight, AMNH). GUATEMALA Los Petén: Tikal Ruins, 1 July 1980, 2 (J. Coddington, MCZ); Tikal, July 1975, @ (W. Sedgwick, MCZ); 7— 12 Aug. 1979, 2 (C. E. Griswold, T. C. Meikle, CAS). Izabal: Los Amates, 1908, 26 (Kellerman, MCZ). Alta Verapaz: Lanquin nr. Gruta, 5 Feb. 1980, 2 (V., B. Roth, AMNH). Suchitepéquez: Moca, June 1947, 3 (C., P. Vaurie, AMNH); Variedades, 1-4 July 1947, 2 (C., P. Vaurie, AMNH). HONDURAS Copan: Co- pan, March 1939, d (AMNH). Atlantida: Lancetilla, 1929, 2 (A.M. Chickering, MCZ). NICARAGUA 50 km E Matagalpa, El Coyolar, 800 m, 20 Nov. 1991, 23 (J. Maes, JM); Musawas, Waspuc River, Sep., Oct. 1955, 2d (B. Malkin, AMNH). Granada: Volcan Mombacho, 700-800 m, 15 July 1998, 2 (J. M. Maes, JM). COSTA RICA Limon: 20 km N Siquirres, 100 m, July 1980, 2 (W. Eberhard. MCZ); Hamburg Farm, 2 (C. R. Dodge, MCZ); Finca La Laja, 190 m, 2d (K. A. Arnold, MCZ). Heredia: Puerto Viejo, La Selva, many records (CAS, MCZ, USNM). Guan- acaste: Orosi, 2 3 (Picado, MCZ). Cartago: Turrialba, many records (AMNH, CUC); Finca Sinfonia, 9°50'N, 83°05’W, 26 Jan. 1976, 2 (Roth, Schroepfer, AMNH). San José: Bajo La Hondura (Braulio Car- rillo Natl. Park), 1,400-1,600 m, 26 July 1983 (W. Eberhard, MCZ): Rio Hondura, 10°N, 84°W, 1,200 m, 30 Mar. 1991, do (W. Eberhard, MCZ): nr. Villa Colén, 800 m, Nov. 1990, (W. Eberhard, MCZ):; San José, 3 (E. Schmidt). Puntarenas: nr. Quepas Ma- nuel-Antonio, 9°23’N, 84°09’W, 15-21 Feb. 1976, @ (Roth-Schoepfer, AMNH); 24 km S Volcan, 12 July 1970, 2 (S. Riechert, AMNH): Osa Peninsula, 3.2 km SW Rincon, 21-28 Feb. 1967, 2 (J. Nelson, MCZ); Corcovado Natl. Park, Sirena, 25 Feb. 1979, @ (J. Coddington, MCZ); Santa Elena nr. Monteverde, 20 June 1998, ¢ (K. J. Ribardo, CAS); Quizarra, 6 km E San Isidro, May 1989, 2 (W. Eberhard, MCZ); San Isidro del General, 600—1,200 m, 2 (D. Rounds, MCZ); nr. Tocales, Reserva Carara, Aug. 1983, 2 (W. Eberhard, MCZ); Golfito, 25 Jan. 1995, 2 (W. Eber- hard, MCZ). PANAMA Chiriqut: Bugaba, July 1939, Aug. 1940, (A. M. Chickering, MCZ); 1, 2 Nov. 1985, 2 (D. Quintero, MIUP); Cerca Cementerio, Bugaba, Rio Mulo, 2 Nov. 1985, 2¢ (D. Quintero, MIUP); La Fortuna, 1,100—1,200 m, 5 Apr. 1984, 26 (W. Eberhard, MCZ); El Volcan, 20 Mar. 1936, 2¢ (W. J. Gertsch, AMNH). Veraguas: Alto Limon (La Yeguado), 11 Aug. 1984, 2 (D. Quintero, MIUP); San Martin, Santiago, 21 July 1983, 2 (M. E. Mendoz, Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 MIUP). Colén: Colén, July 1979, 2 (Broadhead et al., USNM). Panamd: El Valle, several records, 2 3 (AMNH, MCZ); Arraijan, 2d (A. M. Chickering, MCZ); Canal area, very common (MCZ, USNM); Cermenio, Jan.—Feb. 1940, d (Zetek, MCZ); Panama City, July 1979, 23 (E. Broadhead et al., USNM). Darién: Villa Darién, 12-18 Feb. 1984, ¢ (M. N. Garcia, MIUP). TRINIDAD nr. Port of Spain, 1913, 2d (R. Thax- ter, MCZ); 9 Feb. 1926, 29 ¢ (W. S. Brooks, MCZ): Sangre Grande, 1913, 2¢ (R. Thaxter, MCZ); Cu- muto, Feb. 1926, 2 (W. S. Brooks, MCZ). St. Andrew Co.: Valencia Ward at Oropuche River, 17 Aug. 1986, 6 (G. B. Edwards, FSCA). Mangora itza new species Figures 48-56; Map 1F Holotype. Female holotype from Chichen Itza, Yu- catan, Mexico, 12 Sep. 1964 (J. C. Pallister), in AMNH. The name is a noun in apposition after the type locality. Description. Female holotype. Carapace yellowish, with a narrow, median longitu- dinal gray line. Labium, endites, sternum, legs yellowish; sternum grayish along bor- ders. Abdomen with median, longitudinal dark band that is narrower anteriorly, con- taining posteriorly three darker bands, sides with scattered white spots (Fig. 52); venter light yellowish, with three white patches and two indistinct darker, longi- tudinal bands. A pair of white patches on sides (Fig. 53). Posterior eye row procur- ved. Ocular trapezoid longer than wide, rectangular. Posterior median eyes 1.2 di- ameters of anterior medians; anterior lat- eral eyes 1.0 diameter, posterior 0.8. An- terior median eyes their diameter apart, 1.2 from laterals. Posterior median eyes 0.7 diameter apart, 1.2 from laterals. Height of clypeus equals 0.8 diameter of anterior median eyes. Total length 3.5 mm. Carapace 1.7 mm long, 1.3 wide in tho- racic region, 0.7 wide behind lateral eyes, 0.7 high. First femur 1.7 mm, patella and tibia 2.1, metatarsus 1.7, tarsus 0.8. Sec- =— Figures 48-56. Mangora itza new species. 48-53, female. 48-51, epigynum. 48, ventral. 49, ventral, scape torn. 50, posterior. 51, posterior, cleared. 52, abdomen, dorsal. 53, abdomen, ventral. 54-56, male, left palpus. 54, apical. 55, mesal. 56. ventral. Figures 57-61. 60, abdomen, dorsal. 61, abdomen, ventral. Mangora acaponeta new species, female. 57-59, epigynum. 57, ventral. 58, posterior. 59, posterior, cleared. CENTRAL AMERICAN MANGORA °* Levi 155 mobilis Figures 62-70. Mangora mobilis (O. P.-Cambridge). 62-66, female. 62-64, epigynum. 62, ventral. 63, posterior. 64, posterior, cleared. 65, abdomen, dorsal. 66, abdomen, ventral. 67—70, male, palpus. 67, apical. 68, mesal. 69, ventral. 70, median apoph- ysis, conductor and embolus. Scale lines: 1.0 mm, genitalia 0.1 mm. 156 ond patella and tibia 1.8 mm, third 1.2. Fourth femur 2.0 mm, patella and tibia 1.8, metatarsus 1.8, tarsus 0.7. Male paratype. Coloration similar to that of female. Posterior eye row straight. Ocular trapezoid longer than wide, widest anteriorly. Posterior median eyes 0.8 di- ameter of anterior medians; lateral eyes 0.5 diameter. Anterior median eyes 0.7 di- ameter apart, 0.7 from laterals. Posterior median eyes 0.3 diameter apart, 1.0 from laterals. Height of clypeus equals 1.0 di- ameter of anterior median eyes. Total length 2.1 mm. Carapace 1.2 mm long, 0.8 wide in thoracic region, 0.3 wide behind lateral eyes, 0.4 high. First femur 1.4 mm, patella and tibia 1.5, metatarsus 1.3, tarsus 0.6. Second patella and tibia 1.3 mm, third 0.8, fourth patella and tibia 1.8. Variation. Total length of females 3.5— 3.7 mm, males 2.1—2.2 mm. The specimen from Uxmal is darker with a black sternum and black spots on legs. The epigynum is variable: the scape varies in width; in ven- tral view the lateral sclerites vary in amount of sclerotization; and in posterior view the transverse sclerite varies in scler- otization. One specimen had the scape torn off the epigynum (Fig. 49). The pal- pus of the male has a large concave me- dian apophysis (4 h in Figs. 55, 56). The illustrations are a composite from various specimens. Diagnosis. The epigynum of M. itza has a wide scape flanked by sclerotized lobes (Fig. 48), and is distinguished from other species by the wide transverse posterior band (Fig. 50). The male is separated from others by the large concave median apophysis in the palpus with a spine pointing toward the cymbium (4 h in Figs. 55, 56). Distribution. Yucatan Peninsula. Paratypes. MEXICO Campeche: Campeche, 27, 28 Oct. 1946, 12, 1d (H. Wagner, AMNH). Yucatan: Chichen Itza, 16 July 1952, 12 (J. ‘G) ‘Pallister; AMNH); Uxmal, July 1981, 12 (C. Gold, CAS); Val- ladolid, 13 Sep. 1952, 1d (J., D. Pallister, AMNH). Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 Mangora acaponeta new species Figures 57-61; Map 1F Holotype. Female holotype from 12.8 km NW Aca- poneta, Nayarit, Mexico, 25 Nov. 1948 (H. B. Leech) in CAS. Description. Female holotype. Carapace light orange, with a median gray line and gray patch on each side of thoracic region. Chelicerae with gray patch. Labium, en- dites gray. Sternum gray anteriorly and on sides. Legs with fine black rings and black spots at origin of macrosetae. Abdomen dorsum orange-white, with posterior lon- gitudinal bands (Fig. 60); venter with a white patch surrounded by black behind epigynum, lateral diagonal bands, and ar- eas with white pigment spots (Fig. 61). Posterior eye row straight. Ocular trape- zoid longer than wide, rectangular. Poste- rior median eyes 1.0 diameter of anterior medians; lateral eyes 0.8 diameter. Ante- rior median eyes 0.8 diameter apart, 1.0 from laterals. Posterior median eyes 0.8 di- ameter apart, 1.2 from laterals. Total length 3.8 mm. Carapace 1.3 mm long, 1.1 wide, 0.6 wide behind lateral eyes, 0.7 high. First femur 1.8 mm, patella and tibia 1.8, metatarsus 1.6, tarsus 0.7. Second pa- tella and tibia 1.7 mm, third 1.1, fourth femur 1.8. The male is not known. Diagnosis. The epigynum of M. acapo- neta is distinguished from that of other species by the lateral plates which in ven- tral view have a median, dark mark and in posterior view a heart-shaped median plate, pointed behind (Fig. 58). Distribution. Nayarit (Map 1F). Specimens Examined. No other specimens have been found. Mangora mobilis (O. P.-Cambridge) Figures 62—70; Map 1G Epeira mobilis O. P.-Cambridge, 1889: 30, pl. 6, fig. 1, 2. Female specimens from Ciudad in Durango [Mexico] and Motagua Valley, Chicoyoto [P Chi- quito], Guatemala in the BMNH, examined, prob- ably not syntypes. Keyserling, 1892: 240, pl. 12, fig. VS eh Mangora mobilis:—F. O. P.-Cambridge, 1904: 479, pl. 45, figs. 16, 17, 2d. Additional locality from Teapa in Tabasco, Mexico. Platnick, 2004. Note. The syntypes are lost. The speci- men surviving and labeled mobilis is the one from Teapa, examined by F. O. P.- Cambridge. Description. Female from Jalisco. Pro- soma light orange, with a gray longitudinal line through middle of carapace. Dorsum of abdomen with three gray lines posteri- orly, and posterior black laterally; white pigment spots in light areas (Fig. 65). Ven- ter with patch of white pigment spots be- hind epigynum on black venter (Fig. 66). Posterior eye row straight. Median eye trapezoid square. Posterior median eyes 1.0 diameter of anterior medians; lateral eyes 0.8 diameter. Anterior median eyes 0.8 diameter apart, 1.0 from laterals. Pos- terior median eyes 0.8 diameter apart, 1.5 from laterals. Height of clypeus equals 0.8 diameter of anterior median eye. Total length 4.2 mm. Carapace 1.7 mm long, 1.4 wide, 0.7 wide behind eyes, 0.8 high. First femur 1.9 mm, patella and tibia 2.3, meta- tarsus 1.8, tarsus 0.8. Second patella and alone, 2 rearan, tdavireol ILO, iteoadn 2), Male from Jalisco lighter than female, abdomen with two dorsal bands of white pigment spots; venter without marks. Pos- terior eye row slightly recurved. Median eye trapezoid wider anteriorly than long. Posterior median eyes 0.8 diameter of an- terior medians; lateral eyes 0.6 diameter. Anterior median eyes 0.6 diameter apart, 0.6 from laterals. Posterior median eyes 0.5 diameter apart, 1.3 from laterals. Total length 2.8 mm. Carapace 1.5 mm long, 1.3 wide in thoracic region, 0.4 wide behind lateral eyes, 0.7 high. First femur 1.8 mm, patella and tibia 2.0, metatarsus 1.6, tarsus 0.9. Second patella and tibia 1.6 mm, third 0.9, fourth 1.5. Variation. Total length of females 3.2— 4.3 mm, males 2.4-2.8 mm. Rarely the sternum is black and the legs have small, black patches at the insertion of macrose- tae. The illustrations (Figs. 67-70) were from a male from Teapa, Tabasco. CENTRAL AMERICAN MANGORA °* Levi Wa Diagnosis. The epigynum of M. mobilis, in posterior view, has the median plate longer than wide (Fig. 63), unlike the re- lated species M. acaponeta, M. distincta, and M. amchickeringi, which have the plate wider than long (Figs. 58, 72, 81). It differs from M. fascialata, which has a sim- ilar epigynum, by lacking the ventral black lines on the venter of the femora as seen in M. fascialata (Fig. 100). The palpus of the male has a short scler- otized, triangular median apophysis, wider on one end than the other (Fig. 70). Natural History. Specimens came from tropical, deciduous forest in Jalisco; pine forest in Chiapas. Distribution. Mexico to western Hon- duras (Map 1G). Specimens Examined. MEXICO Tamaulipas: Ono km E Nuevo Morelos, 2 Dec. 1939, 12 (L. I. Davis, AMNH): Naciemente del Rio Frio, nr. Gomez Farias, 23°1'N, 99°L'W, 6, 7 June 1983, 12 (W. Maddison, MCZ); 9.6 km E Villa Juarez, 7 July 1941, 7@ (L. I. Davis, AMNH); ridge betw. Antiguo Morelos and Nuevo Morelos, 18 Nov. 1948, 12 (H. B. Leech, CAS). Coahuila: 19.8 km S Saltillo, 4 July 1985, 19 (J. Woolley et al., AD). Sonora: 13 km W Alamos, 23 Aug. 1965, 1d (W. J. Gertsch et al.. AMNH). San Luis Potost: Ciudad del Maiz, 760 m, 28 July 1953, 1d (C. J. Goodnight, AMNH); El Salto, 21-23 June 1955, 12 (C., P. Vaurie, AMNH); Huichichuyan, 19 May 1952, 12, 1d (W. J. Gertsch et al., AMNH); [FMON OOM OMe NUS He LOOAe SER IS (pe Wie livies AMNH); Picolo, 21 May 1952, 12, 16 (M. Cazier et al, AMNH); Pujal, 19 May 1952, 12 (W. J. Gertsch, AMNH); 13 km W San Joaquin, 19 Apr. 1963, 12, 1d (W. J. Gertsch, W. Ivie, AMNH); Tamazunchale, § July 1944, 2° (L. IL. Davis, AMNH); 23 Nov. 1946, (E. S. Ross, CAS); 20 May 1952, many 2d (W. J. Gertsch et al., AMNH); 20 July 1956, 12, 2d (W. Gertsch, V. Roth, AMNH): 13 km NNW Tamazun- chale, 19 Apr. 1963, 1d (W. J. Gertsch, W. Ivie, AMNH); 1.6 km SW Tamazunchale, 29 July 1966, 12 (J., W. Ivie, AMNH); Valles, 1961, 1d (L. Staude, PING) Eos fully OS Ore ea oom (Waa). Gextschy nV: Roth, AMNH); 15 Aug. 1964, 1d (Tes W. Ivie, AMNH); Valles El Bafito, 27 June, 1940, 12 (H. Hoogstraal, MCZ); 8 km N Valles, 17 Apr. 1938, 12 (L. I. Davis, AMNH); 32 km S Valles, 14 Apr. 1946, 1d (A. M., L. I. Davis, AMNH); Volcan San Martin, nr. San Andrés, 1,520 m, 14 July 1953, 12 (C. J. Goodnight, AMNH). Durango: 56 km N Villa Juarez, 17 Apr. 1938, 1 (L. I. Davis, AMNH); 1 km S Villa Jwairez, I eyo IDSs eh SGN Ms Ia Dennis, AMNH). Sinaloa: Mazatlan, 6 Sep. 1956, 1d (A. F. Archer, AMNH); 3.2 km S Elota, 23°55'N, 106°48’W, 158 11 Sep. 1966, 1d (J., W. Ivie, AMNH). Nayarit: La Libertad, 6 Aug. 1947, 12, 1d (J. Goodnight, B. Mal- kin, AMNH); San Blas, Manbauchen Beach, 9 Sep. 1966, 1d (J., W. Ivie, AMNH); 16 km E San Blas, 12 Aug. 1954, 12 (R. E. Ryckman et al., AMNH). Jal- isco: El Limon, 720 m, 19°47’N, 104°03’W, 6 Dec. 1996, 12 (Alvarez, Castello, MCZ); 2 Aug. 1997, 16; 31 Aug. 1997, 32 (F. Alvarez, MCZ); Esta. Biol. Cha- mela, 100 m., Sep. 1988, 12 (W. Eberhard 3515, MCZ); Sep. 1988, 12 (W. Eberhard SAE-12B, MCZ): Sep. 1989, 16d; Sep. 1990, 12, 36 (W. Eber- hard, MCZ); Chamela Station, Aug. 1992, 1d (W. Eberhard, MCZ); Puerto Vallarta, Sep. 1957, 12 (J. A. Comstock, CUC); hillside above Plan de‘ Barran- ces, 21°O1'’N, 104°05’W, 8 Sep. 1966, Pe) qs W. Ivie, AMNH). Colima: Potrero Grande, 15 Jan. 1943, 1° (F. Bonet, AMNH); Orilla Rio America, Tecoman, 18 Jan. 1943, 12 (F. Bonet, AMNH); Tecolapa, 31 July 1954, 2°, 1d (W. J. Gertsch, AMNH). Veracruz: Aca- yucan, 25 Oct. 1957, 12 (R. R. Dreisbach, MCZ); 6.4 km NE Acayucan, 27 Apr. 1963, 1d (W. J. Gertsch, W. Ivie, AMNH); Alamo, 17 Oct. 1947, 12, 16 (H. Wagner, AMNH); Catemaco, Playa Azul, 9 Aug. 1966, many $4 (J., W. Ivie, AMNH); Fortin, 25 July 1956, many °d (V. Roth, W. J. Gertsch, AMNH); 5 Aug. 1966, 32 (J. W. Ivie, AMNH); Cérdoba, 11-13 May 1946, many 26 (J. C., D. L. Pallister, AMNH); 21 Sep. 1984, 12, 1d (C. W. Agnew et al., AD); 4.8 km NE Huatusco, 22 July 1985, 12 (J. Woolley et al., AD); La Buena Ventura, July 1909, many 2? d (A. Petrunkevitch, AMNH); Lake Catemaco, 330 m, 26 July 1955, 12 (C., P. Vaurie, AMNH); Los Tuxtlas, 15 km N Catemaco, Aug. 1986, 12 (W. Eberhard, MCZ); July 1990, 12 (B. Traw, MCZ); Mantla, 1, 2 July 1946, many ? d (H. Wagner, AMNH); Martinez de la Torre, 4, 5 July 1946, 22, 26 (H. Wagner, AMNH); Papantla, 12 Oct. 1947, many ? d (H. Wag- ner, AMNH); 24 Aug. 1946, 32 (J. Goodnight et al., AMNH); Plan del Rio, 26 July 1956, 12 (V. Roth, W. J. Gertsch, AMNH); Potrero, 24 June 1936, 1@ (L. I. Davis, AMNH); San Rafael [Carretore, Mantla Martinez], 3 July 1946, 22, 1d (H. Wagner, AMNH); 4 km N Sontecomapan, 21 Aug. 1967, 12 (R. E. Leech, CNC); Tecolutla, 13 Oct. 1947, many ? 3 (H. Wagner, AMNH); Tierra Colorado, 26 July 1956, 1d (W. J. Gertsch, V. Roth, AMNH); Tlapacoyan, 300 m, 7, 8 July, 1946, many 2d (H. Wagner, AMNH). Mi- choacan: 78 km SE Aquila, 13 July 1984, 1d (J. Wool- ley, AD); 16 km S Uruapan, 6, 7 July 1985, 29 (J. Woolley et al., AD). Puebla: Acatlan, 24-27 Sep. 1946, 12 (H. Wagner, AMNH). Guerrero: Arcelio, 400 m, 2 Nov. 1947, 2° (H. Wagner, AMNH): Iguala, 730 m, 27 Oct. 1947, 16 (H. Wagner, AMNH); 1.6 km NE La Laguna, 17 July 1984, DO is (J. Woolley, Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 AD). Oaxaca: Huajuapan, 27 Sep.—1 Oct. 1946, 19 (H. Wagner, AMNH); Jalapa, 28 Aug. 1947, 32, 1d (H. Wagner, AMNH); San Felipe, nr. Oaxaca City, 1,700 m, 6 Oct. 1946, 32 (H. Wagner, AMNH1); Pa- paloapan, 24, 25 July 1946, 12 (H. Wagner, AMNH); Paso Real, Rio Tonto, 28, 29 July 1946, many ? 6 (H. Wagner, AMNH); Playa Hati, Rio Tonto, 27 July 1946, many 2d (H. Wagner, AMNH); Puerto Escon- dido, 15 July 1985, 1d (J. Woolley et al., AD); 7 km NE San Pedro Mixtepec, 16 July 1985, 1d (J. Wool- ley et al, AD); Soyaltepec, 1, 2 Aug. 1946, many ? d (H. Wagner, AMNH); Tehuantepec, 26 Aug. 1947, many 2d (H. Wagner, AMNH); Tuxpan, 15 Oct. 1947, many 2d (H. Wagner, AMNH); Tuxtila, 23 July 1946, 3°, 3d (H. Wagner, AMNH); 22 km W Zanatepec, 28 Aug. 1967, 2 (R. E. Leech, REL). Ta- basco: Villahermosa (La Venta), 18°00'N, 92°53’W, 13 Aug. 1966, 12 (J. W. Ivie, AMNH). Campeche: Cam- peche, 27, 28 Oct. 1946, 22 (H. Wagner, AMNH); San José, Dec. 1946, 2°, 1d (H. Wagner, AMNH). Yucatan: Piste, 4-8 June 1959, 12 (C., P. Vaurie, AMNH); 4 km N of Xocenpich, 12 km N of Piste, ca. 20°47'N, 88°34’W, 20 July 1983, 12, 1d (W. Mad- dison, MCZ); Uxmal, 18 Aug. 1949, 1d (C. J. Good- night, AMNH). Chiapas: Arriaga, low coast, 1, 2 Sep. 1947, 32,463 (H. Wagner, AMNH); 24 km NW Arri- aga, 27 Aug. 1966, 52 (J., W. Ivie, AMNH); Chiapa, 6 Sep. 1947, 12, 36 (H. Wagner, AMNH); Cintalapa, 17 Sep. 1947, 32, 1d (H. Wagner, AMNH); Escuin- tla, 12, 36 (N. Banks, MCZ): 36 (Crawford, MCZ): Las Cruces, 15-18 Sep. 1947, 22 (H. Wagner, AMNH); Puerto Madero [Puerto de San Benito], 2 Aug. 1950, 2d (C., M. Goodnight, AMNH); Ocos- ingo, 24, 25 June 1950, 42 (C., M. Goodnight et al., AMNH); Rio de Las Flores, 15 Sep. 1947, 3d (H. Wagner, AMNH); Rio San Gregorio, km 1,327, betw. Comitan and Ocotal, 18 July 1950, 1d (C., M. Good- night, AMNH); nr. Simojovel, 4 Aug. 1964, 12 (J. Shatterly, MCZ); Tonala, Aug. 1909, 12, 1d (A. Pe- trunkevitch, AMNH). GUATEMALA Baja Verapaz: San Jer6nimo, 24, 26 July 1947, 12 (C., P. Vaurie, AMNH). Chiquimula: Chiquimula, 375 m, 21-23 July 1947, 1° (C., P. Vaurie, AMNH). Suchitepequez: Variedades, 1-4 July 1947, 12 (C., P. Vaurie, AMNH). EL SALVADOR Santa Tecla [13°41’'N, 89°17'W], 15 Oct. 1949, 12 (J. Boursot, AMNBHE). HONDURAS Copdn: Copan, El Cedro, March 1939, 1d (AMNH); Copan ruins, 7 March 1939, 12 (R. V. Chamberlin, AMNH). Mangora distincta Chickering Figures 71-79; Map 1H Mangora distincta Chickering, 1963: 6, figs. 8-11, d. Male holotype from Escuela Agricultura Panamer- —S Figures 71-79, Mangora distincta Chickering. 71-75, female. 71-73, epigynum. 71, ventral. 72, posterior. 73, posterior, cleared. 74, abdomen, dorsal. 75, abdomen, ventral. 76-79, male, left palpus. 76, apical. 77, mesal. 78, ventral. 79, median apophysis, conductor and embolus. CENTRAL AMERICAN MANGORA ° Levi 159 Figures 80-90, Mangora amchickeringi new species. 80-84, female. 80-82, epigynum. 80, ventral. 81, posterior. 82, posterior, cleared. 83, abdomen, dorsal. 84, abdomen, ventral. 85-90, male, palpus. 85, apical. 86, mesal. 87, ventral. 88, median apoph- ysis, conductor and embolus. 89, 90, left palpus, expanded. Figures 91-95. Mangora gibberosa (Hentz). 91-93, female. 91, 92, epigynum. 91, ventral. 92, posterior. 93, abdomen, dorsal. 94, 95, male, palpus. 94, mesal. 95, ventral. Scale lines: 1.0 mm, genitalia 0.1 mm. Abbreviations: A, terminal apophysis; C, conductor; E, embolus; M, median apophysis; R, radix. 160 icana, San Antonio de Oriente, 27 km S [east] of Tegucigalpa, Honduras in MCZ, examined. Plat- nick, 2004. Note. Chickering (1954, 1963) did not mention M. mobilis in his long description of the very similar M. distincta and gives no differential characters. He illustrated the palpus of M. mobilis from one side, and that of M. distincta from another. Description. Female from Nicaragua. Prosoma light orange. Abdomen orange- white with white pigment spots and with posterior median three indistinct longitu- dinal lines (Fig. 74); venter with small area of white pigment spots behind epigynum, and two areas of white pigment spots on each side (Fig. 75). Posterior eye row slightly recurved. Ocular trapezoid longer than wide, about rectangular. Posterior median eyes 1.3 diameters of anterior me- dians; anterior lateral eyes 0.6, posterior 1.0. Anterior median eyes 1.0 diameter apart, 1.5 from laterals. Posterior median eyes 0.5 diameter apart, 1.2 from laterals. Height of clypeus equals 0.8 diameter of anterior median eyes. Total length 4.2 mm. Carapace 1.7 mm long, 1.4 wide in tho- racic region, 0.7 wide behind eyes, 0.7 high. First femur 2.0 mm, patella and tibia 2.2, metatarsus 1.8, tarsus 0.8. Second pa- tella and tibia 1.9 mm, third 1.3. Fourth femur 2.0 mm, patella and tibia 2.0, meta- tarsus 1.7, tarsus 0.7. Male holotype. Coloration as in female. Posterior eye row slightly recurved. Ocular trapezoid longer than wide, widest anteri- orly. Posterior median eyes 1.0 diameter of anterior medians; laterals 0.7. Anterior median eyes 0.7 diameter apart, 0.7 from laterals. Posterior median eyes 0.4 diame- ter apart, 1.2 from laterals. Total length 2.4 mm. Carapace 1.2 mm long, 0.9 wide be- hind eyes, 0.4 wide behind lateral eyes, 0.5 high. First femur 1.4 mm, patella and tibia 1.5, metatarsus 1.3, tarsus 0.6. Second pa- tella and tibia 1.3 mm, third 0.9, fourth thg)z Variation. Total length of females 3.2— 4.5 mm, males 2.3-2.8 mm. The illustra- tions were made from females from Nic- Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 aragua, and the palpus from the male ho- lotype. Diagnosis. The epigynum of M. distinc- ta is distinguished from epigyna of M. mobilis and M. amchickeringi by the wide, transverse median posterior plate having two lateral lobes (Fig. 72). Males differ from these two species by the curved “upper” edge of the median apophysis (Fig. 79). Natural History. Specimens have been found in dry forests on Volcan Mombacho, in malaise traps; from undergrowth in cof- fee plantation; in tropical dry forest rem- nants and organic coffee plantation in Nic- aragua. Distribution. From southeastern Hon- duras to Costa Rica (Map 1H). Specimens Examined. NICARAGUA Islas de So- lentiname, 30 July 1989, 12 (J. Maes, JM); Apanas, 15 July 1989, 1d (F. Reinbold, JM); Masaya Las Flo- res, 1 Aug. 1994, 1d (J. M. Maes, JM); 50 km E Metagalpa, El Coyolar, 800 m, 20 Nov. 1991, 3d (J. Maes, JM); San Ramon demlinessnOcmocpns Oolemec (|. M. Maes, JM). Granada: Volcan Mombacho, Mar.—Aug. 1998, 22, 14d (J. M. Maes, MCZ). COS- TA RICA Guanacaste: Bagaces, Palo Verde, 16-22 Jan. 1978, 32, 1d (W. Eberhard 1316, MCZ); Rio Portrero, Bagaces, 7 July 1966, 12 (S. Peck, AMNH); ca. Cannas, 100 m, Nov. 1982, 12 (W. Eberhard, MCZ); 9.5 km W Cajfias, Taboga, 10°19'N, 85°09'W, 1967, 12 (J. Nelson, MCZ). San José: San José, 1d (E. Schmidt, MCZ). Mangora amchickeringi new species Figures 80-90; Map 11 Holotype. Female holotype, 17 male and 39 female paratypes from Madden Dam, Canal Zone, Pana- ma, July 1950 (A. M. Chickering) in MCZ. The species is named after the collector, arachnologist A. M. Chickering. Mangora mobilis:—Chickering, 1954: 202, figs. 10- 14, 26 (misidentification). Description. Female holotype. Carapace yellow. Abdomen speckled white except midline, with posterior pair of longitudinal black bands (Fig. 83); venter with a patch of white spots behind epigynum and white spots on sides (Fig. 84). Posterior eye row straight. Ocular trapezoid longer than wide, rectangular. Posterior median eyes 1.3 diameters of anterior medians; lateral eyes 0.8 diameter. Anterior median eyes 0.8 diameter apart, 1.1 from laterals. Pos- terior median eyes 0.5 diameter apart, 1.1 from laterals. Height of clypeus equals 0.8 diameter of anterior median eyes. Total length 3.9 mm. Carapace 1.7 mm long, 1.4 wide in thoracic region, 0.7 wide behind lateral eyes, 0.7 high. First femur 2.2 mm, patella and tibia 2.3, metatarsus 1.9, tarsus 0.8. Second patella and tibia 2.0 mm, third 1.2. Fourth femur 2.1 mm, patella and tib- ia 2.2, metatarsus 1.8, tarsus 0.8. Male paratype. Coloration as in female, except abdomen has a pair of longitudinal, dorsal bands with white pigment spots. Posterior eye row slightly procurved. Oc- ular trapezoid longer than wide, rectan- gular. Posterior median eyes 1.2 diameters of anterior medians; lateral eyes 0.6 di- ameter. Anterior median eyes 0.7 diameter apart, 1.0 from laterals. Posterior median eyes 0.6 diameter apart, 1.3 from laterals. Height of clypeus equal to 0.8 diameter of anterior median eye. Total length 2.6 mm. Carapace 1.3 mm long, 1.1 wide in tho- racic region, 0.4 wide behind lateral eyes, 0.5 high. First femur 1.6 mm, patella and tibia 1.8, metatarsus 1.4, tarsus 0.7. Sec- ond patella and tibia 1.5 mm, third 0.8, fourth 1.4. Variation. Total length of females 3.2— 4.2mm, males 2.3-2.8 mm. Mangora am- chickeringi has less black pigment than M. mobilis, but coloration is variable. The fe- male dark, abdominal band is present in most males. The illustrations were made from female holotype and male paratype. Diagnosis. Ventrally the epigynum of M. amchickeringi is like that of M. mobilis; however, in posterior view it differs by having a transverse V-shaped, median plate without folds or lobes (Fig. 81), whereas that of M. mobilis is longer than wide with many folds (Fig. 63) and the median plate of M. disticta has lateral lobes (Fig. 72). The male differs by having the two teeth of the median apophysis large and directed toward the cymbium, and by the straight CENTRAL AMERICAN MANGORA * Levi 161 “upper edge of the median apophysis (Fig. 88). Natural History. Fogging trees in mid- dle savanna in Calabozo, Venezuela, has collected the species. Distribution. Panama to Venezuela, Trinidad (Map 11). Paratypes. PANAMA Bocas del Toro: Rio Chan- guinola, Campamente Irie, 13-17 Mar. 1980, 12 (R. Ibanez, MCZ). Cocle: Cermefio, Feb. 1940, 12 (A. M. Chickering, MCZ). Panamd: Barro Colorado Isl., several collections (A. M. Chickering, MCZ); Mad- den Dam, many collections (A. M. Chickering, MCZ); Summit, July 1950, many 26; Aug. 1950, 12 (A. M. Chickering, MCZ); Chilibre, July 1950, 1d (A. M. Chickering, MCZ); Experimental Gardens, 29 July 1954, 12; 12 Aug. 1954, 19° (A. M. Chickering, MCZ). WEST INDIES. TRINIDAD St. Augustine Uni- versity, April 1964, 12, 36 (A. M. Chickering, MCZ); Port of Spain, 1913, 12, 3d (R. Thaxter, MCZ); Gas- parce, 3 Nov. 1944, 12 (R. H. Montgomery, AMNH). St. George Co.: Diego Martin Ward, Edith Falls, Trail, 18 Aug. 1986, 1d (G. B. Edwards, FSCA); San Rafael Ward, E side of Talparo River, 20 Aug. 1986, 1d (G. B. Edwards, FSCA). St. Andrew Co.: Valencia Ward, at Oropuche River, 17 Aug. 1986, 1d (G. B. Edwards, FSCA). VENEZUELA Bolivar: San Felix, Oct.—Dec. 1947, 12 (AMNH). Guarico: Estacion Biolégica de los Lla- nos, Calabozo, 280 m, 18 Jan. 1985, 12 (J. Palmer, MCZ); Hato Masaquarai, 45 km S of Calabozo, 17 Mar. 1980, 12 (K. Rabenold, MCZ). COLOMBIA Magdalena: 10 km E Santa Marta, Oct. 1985, 12 (H.- G. Miiller, SMF). Mangora gibberosa (Hentz) Figures 91-96; Map 1J Epeira gibberosa Hentz, 1847: 477, pl. 31, fig. 20, 2. Types from Alabama, destroyed. Mangora gibberosa:—Levi, 1975, figs. 118-130, ° d, map 4. Platnick, 2004. Description. Description is found in Levi (1975). Diagnosis. Unlike other Mangora, ex- cept M. fascialata (Fig. 100), specimens have the venter of the first and second femora with a black line. The female differs from M. fascialata by having, in posterior view, rectangular lat- eral plates (Fig. 92). The male differs, in apical view of the palpus (Fig. 96), by lacking the wide band present in M. fascialata (Fig. 101). 162 Distribution. Southeastern Canada, eastern United States, only one record from Mexico (Map 1J). Additional record. MEXICO Tamaulipas: Nueva Karedor 22) iullyael O46N when (| ea@e ID penleeeeeralllastere AMNH). Mangora fascialata Franganillo Figures 97-103; Map 2A Mangora fascialata Franganillo, 1936: 83. Specimens came from Cuba. Levi, 1975: 128, figs. 110-117, 26. Platnick, 2004. Mangora striatipes Bryant, 1945: 375, figs. 17, 18, 2d. Male holotype from Puerto Plata, Dominican Republic, in MCZ, examined. Synonymized by Levi, 1975. Mangora conspicua Chickering, 1963: 191, pl. 2, figs. 1-7, 2d. Female holotype from El Potosi, Nuevo Léon, Mexico, in MCZ, examined. Synonymized by Levi, 1975. Description. Female from Nuevo Leon. Carapace whitish with a median dusky line. Legs whitish with a median black line under the first and second femora (Fig. 100); other articles with black patches. Ab- domen light with marks as in Figures 99 and 100. Posterior eye row straight. Ocular trapezoid longer than wide, widest anteri- orly. Posterior median eyes 0.9 diameter of anterior medians; lateral eyes 0.7 diameter. Anterior median eyes 1.0 diameter apart, 1.0 from laterals. Posterior median eyes 0.8 diameter apart, 1.0 from laterals. Total length 3.6 mm. Carapace 1.4 mm long, 1.0 wide in thoracic region, 0.6 wide behind lateral eyes, 0.7 high. First femur 1.7 mm, patella and tibia 1.5, metatarsus 1.4, tarsus 0.7. Second patella and tibia 1.4 mm, third 0.9, fourth 1.5. Male from Haiti. Coloration lighter than in female. Posterior eye row straight. Oc- ular trapezoid longer than wide, widest an- teriorly. Posterior median eyes 1.0 diame- Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 ter of anterior medians; lateral eyes 0.6 di- ameter. Anterior median eyes 1.0 diameter apart, 1.0 from laterals. Posterior median eyes 0.4 diameter apart, 1.5 from laterals. Total length 2.2 mm. Carapace 1.2 mm long, 1.0 wide in thoracic region, 0.4 wide behind lateral eyes, 0.6 high. First femur 1.2 mm, patella and tibia 1.2, metatarsus 1.0, tarsus 0.5. Second patella and tibia 1.1 mm, third 0.6, fourth 1.1. Variation. Total length of females 2.8— Gay mmm, males 2253) manta Diagnosis. Mangora fascialata is the only species known from the West Indies (except Trinidad). As in M. gibberosa, it has a distinct black line along the venter of the first and second femora (Fig. 100). The lightly sclerotized epigynum is dis- tinguished from others by the presence of a scape and distinctly curved, lateral plates surrounding the median plate in posterior view (Fig. 98). The black line on the ven- ter of the femora of M. fascialata prevents confusion with the similar M. mobilis. Males have four or five macrosetae on the venter of the second tibia, and are fur- ther distinguished by the numerous scler- ites in the palpus (Figs. 101-103). In api- cal view the palpus has a structure that looks like a wide, tucked-in band (Fig. 101). Natural History. Specimens came from cane and mesquite along river in Texas; on low shrubs and herbs in Bocatoma, Ta- maulipas, and in creek bed in Nuevo Leon. Distribution. Texas, Mexico to Hondu- ras, Greater Antilles (Map 2A). Specimens Examined. TEXAS Comal Co.: New Braunfels, 7 June 1942, 12 (E. S. Ross, CAS). Brew- ster Co.: Big Bend National Park, Boquillas Camp, 24, 25 May 1967 (E. Sabath, MCZ). MEXICO Ta- Figure 96. Mangora gibberosa (Hentz), male, left palpus, apical. Figures 97-103. Mangora fascialata Franganillo. 97-100, female. 97, 98, epigynum. 97, ventral. 98, posterior. 99, abdomen, dorsal. 100, abdomen, sternum, and proximal parts of left legs, ventral. 101-103, male, palpus. 101, apical. 102, mesal. 103, ventral. Figures 104-109. Mangora spiculata (Hentz). 104-106, female. 104, 105, epigynum. 104, ventral. 105, posterior. 106, abdo- men, dorsal. 107-109, male, palpus. 107, apical. 108, mesal. 109, ventral. CENTRAL AMERICAN MANGORA ° Levi GS Figures 110-115. Mangora placida (Hentz). 110-112, female. 110, 111, epigynum. 110, ventral. 111, posterior. 112, abdomen, dorsal. 113-115, male, palpus. 113, apical. 114, mesal. 115, ventral. Figures 116-120. Mangora oaxaca new species, female. 116-120, epigynum. 116, ventral. 117, posterior. 118, posterior, cleared. 119, abdomen, dorsal. 120, abdomen and sternum, ventral. Scale lines: 1.0 mm, genitalia 0.1 mm. 164 maulipas: Bocatoma, 26 Feb. 1976, 12 (W. B. Peck, CAS); 1.6 km S Antiguo Morelos, 21 July 1954, 19 (Chilcott, CNC); Padilla, 17 May 1952, 2d (W. J. Gertsch et al., AMNH); NE of Padilla, 24°03’N, 98°03'W, 21 July 1966, 1d (J., W. Ivie, AMNH); Vic- toria, 17 May 1952, 1d (W. J. Gertsch et al., AMNH); Villagran, 19 July 1956, 1d (V. Roth, W. Gertsch, AMNH). Nuevo Léon: Linares, 3 July 1941, 22, 1d (L. I. Davis, MCZ); Los Cristales, Aug. 1972, 12 (A. F. Archer, AMNH); Sabinas Hidalgo, 13 June 1940, 292 (H. Hoogstraal, MCZ); Villa de Santiago, Haci- enda Vista Hermosa, 19 June 1940, 12, 1d (H. Hoogstraal, MCZ); El Potosi, Cerro Potosi, 13 June 1938, 12 (Hoogstraal, MCZ); Montemorelos, 23 May 1952, 12 (W. J. Gertsch et al, AMNH). Coahuila: Saltillo, 23 Aug. 1947, 12 (W. J. Gertsch, AMNH); 20 km S Saltillo, 4 July 1985, 12 (J. Woolley et al., AD). Baja California Sur: 10 km S San Antonio, 31 Dec. 1976, 16 (C. Griswold, L. Vincent, CAS). Sierra San Lazaro, 12 (N. Banks, MCZ). San Luis Potost: 4 2 km E Ciudad del Maiz, 29 Nov. 1950, 12 (A. M. Davis, AMNH); Valles, 45.6 km S Huizache, 4 July 1985, 1d (J. Woolley et al., AD); 19 July 1956, 1°, 1d (V. Roth, W. J. Gertsch, AMNH); 11 km S Tam- azunchale, 16 Feb. 1961, 12 (D., H. Campbell, CAS). Durango: Santa Maria del Oro, 1,700 m, 28 July 1947, 12 (W. J. Gertsch, AMNH). Veracruz: Fortin, 25 July 1956, 12 (V. Roth, W. Gertsch, AMNH); 4 km NE Huatusco, 22 July 1985, 19, 2d (J. Woolley et al., AD); Plan del Rio, 26 July 1956, 12 (V. Roth, W. Gertsch, AMNH); Mirador, Zacual- pan, 25 Aug. 1933, 1d (W. James, AMNH). Hidalgo: Apulco, 6 Oct. 1947, 12 (H. Wagner, AMNH); Ixmi- quilpan, 16-18 Aug. 1947, 22, 1d (H. Wagner, AMNH); 3 km SW Jacala, 12 Aug. 1972, 1d (G. F. Hevel, USNM). Guerrero: 37 km N Chilpancingo, 31 July 1956, 135 (V. Roth, W. Gertsch, AMNH); 8 km NE Tixtla de Guerreo, 16 July 1984, 12 (J. B. Wool- ley, AD). Oaxaca: 13 km NE El Punto, 18 July 1985, 12,16 (J. Woolley et al., AD); Huajuapan, 27 Sep.— 1 Oct. 1946, 35 (H. Wagner, AMNH); Juan Garcia, WEBILAN,, GErZiy wy, il Sep. 1964, 2 en YE IN, AMNH); Oaxaca, 1,550 m, 2 Oct. 1946, 12, 26 (H. Wagner, AMNH); 22 Aug. 1947, 16 (H. Wagner, AMNH); San Felipe, 24 Aug. 1947, 22 (H. Wagner, AMNH). Yucatan: Dolores Otero, 13 July 1952, 1°, 1d (J., D. Pallister, AMNH). Chiapas: Ocosingo, 900 m, 25 June 1950, 32, 1d (C., M. Goodnight et al., AMNH); Ocozocoautla, 3 Sep. 1947, 12 (H. Wagner, AMNH); Tuxtla Gutierrez, 9 Sep. 1947, 12, 16 (H. Wagner, AMNH). HONDURAS 27 km S Tegucigal- pa, San Antonio del Oriente, 17 Nov. 1945, 39, Id (A., M. Carr MCZ). CUBA Pinar del Rio: Sierra del Rosario, nr. Insti- tute of Botany research area, May 1976, 12 (R. Lev- ins, MCZ). Oriente: Santiago, June 1967, 1d (P. Ala- yon, MCZ); Loma “La Farola” Baracoa, June 1967, 12 (P. Alayon, MCZ). HAITI Las Cayes, Les Platons, Nov. 1971, 22 (T. Moermond, MCZ); Diquini, 2d (W. M. Mann, MCZ);: hills nr. Port of Prince, 2 Oct. 1934, 62 (P. Darlington, MCZ); nr. Kenscoff, 1,300 Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 m, 1 May 1935, 12 (Roys, MCZ). DOMINICAN REPUBLIC Puerto Plata, Apr—May 1941, 292 (D. Hurst, MCZ). TRINIDAD Port of Spain, 1913, 19 (R. Thaxter, MCZ); St. Augustine, 1 Apr.1964, 12 (A. M. Chickering, MCZ). Mangora spiculata (Hentz) Figures 104-109; Map 1K Epeira spiculata Hentz, 1847: 475, pl. 31, fig. 13, @. Female holotype destroyed. Mangora spiculata:—Levi, 1975: 125, figs. 82-89, 26, map 4. Platnick, 2004. Description. Description is found in Levi (1975). Diagnosis. The female differs from M. placida by having, in ventral view of the epigynum, longitudinal, sclerotized bars at a distance from the scape (Fig. 104). The median apophysis of the palpus of M. spiculata differs from that of M. pla- cida by having two spines (6 h in Fig. 108). Distribution. Eastern United States to southern Texas (Map IK). No records are known from Mexico, although specimens can be expected there. Mangora placida (Hentz) Figures 110-115; Map 2B Epeira placida Hentz, 1847: 475, pl. 31, fig. 12, 9. Female holotype from Alabama, destroyed. Mangora placida:—Levi, 1975: 126, figs. 80, 81, 90— 101, 2d; map 4. Platnick, 2004. Description. Description is found in Levi (1975). Diagnosis. In ventral view of the epi- gynum, the female differs from M. spicu- lata by having diagonal bars flanking the scape (Fig. 110). The median apophysis of the palpus of the male has one spine (6 h in Fig. 114), whereas that of M. spiculata has two spines (Fig. 108). Distribution. Southeastern Canada, eastern United States to Nuevo Léon, Mexico (Map 2B). Additional Record. MEXICO Nuevo Léon: Lina- res, 8 July 1941, 32 (L. I. Davis, AMNH). Mangora oaxaca new species Figures 116-120; Map 2C Holotype. Female holotype from Finca Patichuiz, SE of Oaxaca, Mexico, 22 Oct. 1962 (Gardner) in MCZ. Description. Female holotype. Carapace light orange. Large black circles around secondary eyes. Chelicerae dusky orange. Labium, endites black. Sternum black. Legs light orange, distally darker. Abdo- men with median dorsal gray band, black- ish posteriorly with four black spots (Fig. 119); venter black posteriorly, sides with a gray band (Fig. 120). Posterior eye row procurved. Ocular trapezoid longer than wide, rectangular. Posterior median eyes 1.2 diameters of anterior medians; lateral eyes 0.8 diameter. Anterior median eyes their diameter apart, 1.0 from laterals. Posterior median eyes 0.8 diameter apart, 1.3 from laterals. Total length 3.0 mm. Carapace 1.3 mm long, 1.0 wide in tho- racic region, 0.6 wide behind lateral eyes, 0.7 high. First femur 1.5 mm, patella and tibia 1.8, metatarsus 1.4, tarsus 0.7. Sec- ond patella and tibia 1.6 mm, third 1.0, fourth 1.5 mm. The male is not known. Diagnosis. The abdomen of M. oaxaca (Fig. 119) resembles that of M. placida but the epigynum of the former is a semicircle having a dark, sclerotized, posterior, lateral black frame (Fig. 116), and the posterior view is heavily sclerotized with two lateral, dorsal lobes (Fig. 117). Distribution. Oaxaca (Map 2C). Specimens Examined. No other specimens have been found. Mangora nahuatl new species Figures 121-125; Map 2C Holotype. Female holotype from Jalapa, Veracruz, Mexico, July 1981 (C. Gold) in CAS. The specific name is a noun in apposition after the Indians in- habiting the area of the type locality. Description. Female holotype. Carapace yellowish with median eye area and clyp- eus black. Endites, labium, sternum black. Legs yellowish. Abdomen orange-white, CENTRAL AMERICAN MANGORA * Levi 165 with posterior median black area, and ar- eas with white pigment spots and gray chevron marks (Fig. 124); sides with a black patch; venter with a median gray area (Fig. 125). Posterior eye row straight. Ocular trapezoid square. Posterior median eyes 1.5 diameters of anterior medians; lateral eyes 1.0 diameter. Anterior median eyes 1.3 diameter apart, 1.0 from laterals. Posterior median eyes 0.5 diameter apart, 0.7 from laterals. Total length 2.8 mm. Carapace 1.3 mm long, 0.9 wide in tho- racic region, 0.5 wide behind lateral eyes, 0.7 high. First femur 1.3 mm, patella and tibia 1.4, metatarsus 1.1, tarsus 0.5. Sec- ond patella and tibia 1.3 mm, third 0.8, fourth 1.3 mm. The male is not known. Diagnosis. The dorsal abdominal mark- ings of M. nahuatl (Fig. 124) differ from those of M. passiva (Fig. 152). Also, in posterior view of the epigynum, the me- dian plate is wider (Fig. 122), with antero- lateral lobes, than that of M. passiva (Fig. 149). Distribution. Veracruz, México, Mexico. Specimen Examined. MEXICO México: Tenango de Valle, 2,400 m, 26-29 Aug. 1946, 12 (H. Wagner, AMNH). Mangora volcan new species Figures 126-130; Map 2D Holotype. Female holotype and an immature para- type from El] Volcan, Chiriqui, Panama, 24 Feb. 1936 (W. J. Gertsch) in AMNH. The name is a noun in apposition after the type locality. Description. Female holotype. Carapace pale yellow with eye region black. Labium, endites, sternum black. Coxae pale yellow- ish, distal leg articles gray. Abdomen whit- ish with a dorsal, posterior black band that fades anteriorly (Fig. 129); sides each with a black patch; venter with a gray area in middle and black around epigynum (Fig. 130). Posterior eye row straight. Ocular trapezoid slightly wider than long, widest anteriorly. Posterior median eyes 1.0 di- ameter of anterior medians; lateral eyes 0.7 diameter. Anterior median eyes 1.0 di- ameter apart, 1.0 diameter from laterals. 166 Posterior median eyes 0.8 diameter apart, 1.0 from laterals. Total length 2.2 mm. Carapace 1.0 mm long, 0.8 wide in tho- racic region, 0.4 wide behind lateral eyes, 0.4 high. First femur 1.0 mm, patella and tibia 1.2, metatarsus 1.0, tarsus 0.5. Sec- ond patella and tibia 1.1 mm, third 0.6, fourth 1.0. The male is not known. Diagnosis. Mangora volcan has the scape of the epigynum almost spherical (Fig. 126), and in posterior view, has a wide trapezoid median plate (Fig. 127). In coloration it is unlike any other Central American species (Figs. 129, 130). Distribution. Chiriqui, Panama (Map 21))) Specimens Examined. No other specimens have been found. Mangora fortuna new species Figures 1381-135; Map 2D Holotype. Female holotype from El] Fortuna, 1,100— 1,200 m, Chiriqui, Panama, 5 May 1984 (W. Eber- hard) in MCZ. The name is a noun in apposition after the name of the type locality. Description. Female holotype. Prosoma yellowish, darkest in eye region, legs with distal articles dusky. Sternum with a dusky frame, light in center. Dorsum of abdomen with two gray longitudinal bands, most dis- tinct posteriorly (Fig. 134); venter with a V-shaped gray patch anterior to spinnerets and a dusky mark covering epigynum and on each side of the structure (Fig. 135). Posterior eye row slightly procurved. Oc- ular trapezoid longer than wide, rectan- gular. Posterior median eyes 1.0 diameter of anterior medians; lateral eyes 0.8 di- ameter. Anterior median eyes 1.0 diameter apart, 0.4 from laterals. Posterior median eyes 0.8 diameter apart, 0.8 from laterals. Height of clypeus equals 0.8 diameter of Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 anterior median eyes. Sternum bulging. Total length 2.4 mm. Carapace 1.2 mm long, 0.8 wide in thoracic region, 0.4 wide behind lateral eyes, 0.6 high. First femur 1.2 mm, patella and tibia 1.3, metatarsus 1.1, tarsus 0.6. Second patella and tibia 1.2 mm, third 0.7, fourth 1.2. The male is not known. Variation. Total length of females 2.3— 2.4 mm. Females may be lighter colored than illustrated (Fig. 134), and the ventral coloration is variable. The illustrations were made from the holotype. Diagnosis. The epigynum of M. fortuna is lightly sclerotized, small, and difficult to study in ventral view (Fig. 131); the oval scape is indistinct unless viewed laterally. The posterior view is more distinct and shows a wide median plate (Fig. 132) more curved than that of M. volcan (Fig. 127). Distribution. Costa Rica, western Pan- ama (Map 2D). Paratypes. COSTA RICA Heredia: La Selva nr. Puerto Viejo, Feb. 1981, 12 (W. Eberhard, MCZ); 26 March 1979, 12 (J. Coddington, MCZ); 22 Nov. 1981, 12 (J. Coddington, MCZ). Mangora campeche new species Figures 136-139; Map 2E Holotype. Female holotype from Reserva de la Bios- fera, Calakmul, km 32, Campeche, Mexico, 20 July 1998 (F. Alvarez) in MCZ. The specific name is a noun in apposition after the type locality. Description. Female holotype. Cara- pace, sternum, legs, golden yellow, except for black eye rings and a gray ring at distal ends of tibiae and tarsi. Abdomen yellow- ish white, with a posterior pair of longi- tudinal black bands, a pair of black spots, and pairs of white patches (Fig. 139). Ven- ter without marks but sides of spinnerets blackish. Posterior eye row recurved. Oc- ular trapezoid longer than wide, widest an- — Figures 121-125. Mangora nahuatl new species, female. 121-123, epigynum. 121, ventral. 122, posterior. 123, posterior, cleared. 124, abdomen, dorsal. 125, abdomen, ventral. Figures 126-130. Mangora volcan new species, female. 126-128, epigynum. 126, ventral. 127, posterior. 128, posterior, cleared. 129, abdomen, dorsal. 130, abdomen, ventral. CENTRAL AMERICAN MANGORA ° Levi 167 137 campeche <8 Figures 131-135. Mangora fortuna new species, female. 131-133, epigynum. 131, ventral. 132, posterior. 133, posterior, cleared. 134, abdomen, dorsal. 135, abdomen, ventral. Figures 136-139. Mangora campeche new species, female. 136-138, epigynum. 136, ventral. 137, posterior. 138, posterior, cleared. 139, abdomen, dorsal. Figures 140-143. Mangora goodnightorum new species, female. 140-142, epigynum. 140, ventral. 141, posterior. 142, pos- terior, cleared. 143, abdomen, dorsal. Figures 144-147. Mangore vito new species, female. 144-146, epigynum. 144, ventral. 145, posterior. 146, posterior, cleared. 147, abdomen, dorsal. Scale lines: 1.0 mm, genitalia 0.1 mm. 168 teriorly. Posterior median eyes 1.0 diame- ter of anterior medians; lateral eyes 0.8 di- ameter. Anterior eyes 1.0 diameter apart, 1.2 from laterals. Posterior median eyes 0.5 diameter apart, 1.5 from laterals. Total length 3.8 mm. Carapace 1.7 mm long, 1.3 wide in thoracic region, 0.7 wide behind lateral eyes, 0.7 high. First femur 1.7 mm, patella and tibia 1.8, metatarsus 1.5, tarsus 0.7. Second patella and tibia 1.7 mm, third DO omnes The male is not known. Diagnosis. The epigynum of M. campe- che is lightly sclerotized and distinguished from others, in posterior view, by the nar- row, bowling-pin—shaped median plate Clie, 137). Distribution. Campeche (Map 2E) Specimens Examined. No other specimens have been found. Mangora goodnightorum new species Figures 140-143; Map 2E Holotype. Female holotype from Finca Monte Liba- no [12 mi E El Real], Chiapas, Mexico, “7.5.50” (C., M. Goodnight, J. Stannard), in AMNH. Description. Female holotype. Prosoma light orange, black between median eyes; distal leg articles gray. Abdomen grayish, light orange with a pair of anterior dorsal white spots and two posterior black bands tipped by a black dot (Fig. 143); venter and spinnerets gray. Posterior eye row straight. Ocular trapezoid longer than wide, widest anteriorly. Posterior median eyes same diameter as anterior medians; lateral eyes 0.8 diameter. Anterior median eyes 1.0 diameter apart, 1.0 from laterals. Posterior median eyes 0.6 diameter apart, 1.3 from laterals. Total length 2.6 mm. Carapace 1.1 mm long, 0.8 wide in tho- racic region, 0.4 wide behind lateral eyes, 0.6 high. First femur 1.3 mm, patella and tibia 1.5, metatarsus 0.9, tarsus 0.6. Sec- ond patella and tibia 1.3 mm, third 0.8, fourth 1.3. The male is not known. Diagnosis. The epigynum of M. good- nightorum is sclerotized and is distin- Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 guished, in posterior view, from all other Central American Mangora by the trian- gular outline (Fig. 141). Unlike other spe- cies, the eye region is slightly projecting above the clypeus. Distribution. Chiapas (Map 2E). Specimens Examined. No other specimens have been found. Mangora vito new species Figures 144-147; Map 2E Holotype. Female holotype from San Vito, Las Cru- ces, 1,300 m, Puntarenas, Costa Rica, Jan. 1987 (W. Eberhard) in MCZ. The specific name is a noun in apposition after the name of the type locality. Description. Female holotype. Prosoma yellowish. Large black circles around sec- ondary eyes. Legs distally brown. Abdo- men white, dorsally with a narrow gray band on each side (Fig. 147), posteriorly wrapping around spinnerets on venter; sides with white pigment spots; venter with gray band anterior of spinnerets. Spinnerets gray. Posterior eye row procur- ved. Ocular trapezoid longer than wide, widest posteriorly. Posterior median eyes 2.0 diameters of anterior medians; anterior lateral eyes 0.8 diameter, posterior laterals 1.0. Anterior median eyes their diameter apart, 1.0 from laterals. Posterior median eyes 0.5 diameter apart, 0.6 from laterals. Height of clypeus equals 1.5 diameters of anterior median eyes. Total length 3.8 mm. Carapace 1.8 mm long, 1.2 wide in tho- racic region, 0.7 wide behind lateral eyes, 0.7 high. First femur 1.5 mm, patella and tibia 1.7, metatarsus 1.3, tarsus 0.6. Sec- ond patella and tibia 1.5 mm, third 0.8, fourth 1.6. The male is not known. Diagnosis. In ventral view, unlike other species, the epigynum is short, having a median lobe flanked by narrow areas that are lobed laterally, and have a wedge- shaped sclerotization (Fig. 144); in poste- rior view, the lateral plates overlap the less sclerotized narrow posterior median plate (Fig. 145). Distribution. Costa Rica (Map 2E). Specimen Examined. COSTA RICA San José: Zur- qui, 1600 m, May 1992, 12 (W. Eberhard 3632, MCZ) [not mapped]. Mangora passiva (O. P.-Cambridge) Figures 148-156; Map 2F Epeira passiva O. P.-Cambridge, 1889: 20, pl. 5, fig. 4, 2. Female syntypes from Cahabon [Alto Vera- paz], San Juan [? Alto Verapaz], Chamelco [? Alto Verapaz], Tactic [Alto Verapaz], Chichochoe [?], Laguna de los Coheteros [?], Coban [Alto Verapaz], all from Guatemala in BMNH, examined. Epeira rostrata Keyserling, 1893: 230, pl. 11, fig. 171, 2. Female holotype from Guatemala, in BMNH, examined. Placed by Levi, 1991, in Mangora. NEW SYNONYMY. Mangora passiva:—F. O. P.-Cambridge, 1904: 480, pl. AD5, fig. 19 oO WevialOna: 124 figs. 6919. Platnick, 2004. Note. The holotype of Epeira rostrata is labeled Epeira rostralipes and is shriveled and faded. Description. Female from Arizona. Car- apace, legs yellow-white. Abdomen with a median black band containing large com- ma-shaped marks formed by white pig- ment spots and smaller marks (Fig. 152), _ venter whitish with median white spots be- hind epigynum, and a pair of white marks anterior of spinnerets (Fig. 153). Posterior eye row recurved. Ocular trapezoid longer than wide, widest anteriorly. Eyes sube- qual. Anterior median eyes 1.0 diameter apart, 1.5 from laterals. Posterior median eyes 0.9 diameter apart, 2.0 from laterals. Total length 4.7 mm. Carapace 2.0 mm long, 1.7 wide in thoracic region, 0.7 wide behind posterior eye row, 0.8 high. First femur 2.3 mm, patella and tibia 2.5, meta- tarsus 1.9, tarsus 0.8. Second patella and tibia 2.4 mm, third 1.4, fourth 2.3. Male from Arizona lighter than female. Eyes subequal. Anterior median eyes 1.0 diameter apart, 1.5 from laterals. Posterior median eyes their diameter apart, 1.5 from laterals. Height of clypeus 0.6 diameter of anterior median eye. Total length 2.7 mm. Carapace 1.3 mm long, 1.2 wide in tho- racic region, 0.5 wide behind posterior eyes, 0.7 high. First femur 1.7 mm, patella and tibia 1.9, metatarsus 1.6, tarsus 0.7. CENTRAL AMERICAN MANGORA * Levi 169 Second patella and tibia 1.8 mm, third 0.9, ieyonecay IL.5). Variation. Total length of females 3.7— 4.7 mm. Diagnosis. The abdomen of M. passiva is recognized by the two upside-down, comma-shaped light marks on dark back- ground (Fig. 152). The epigynum is usu- ally lightly sclerotized (Figs. 148, 149); when more heavily sclerotized, it loses wrinkles (Figs. 150, 151). It differs from other lightly sclerotized epigyna by the presence of the short scape and the vase- shaped posterior median plate. Mangora passive differs from M. nahuatl (Figs. 121— 125) by the distinct coloration of the ab- domen (Figs. 152, 153) and by being larg- er in size. The palpus of the male has a dagger- shaped rod, visible in apical view (Fig. 154), that is heavier than that of M. suffla- va (Fig. 171). Natural History. Specimens have been found in short tropical rain forest at Cha- canna Ruins. Distribution. Arizona, New Mexico to Nicaragua (Map 2F; the map includes data from Levi, 1975, map 4). Specimens Examined. ARIZONA Cochise Co.: Rucker Canyon, Chiricahua Mts., 11 July 1973, 19 (V. Roth, E. Wilkins, CAS); above Herb Martyr Dam, 11 Aug. 1971, 12 (V. Roth, CAS).. Manzanita Co.: Oak Creek Canyon, 27 July 1950, 32, 1d (M. A. Cazier, MCZ). MEXICO Tamaulipas: 4.5 km NW Gomez Farias, 750 m, 28 Dec. 1971, 2 (C. J. Dur- den, USNM). Nuévo Léon: Santa Rosa Canyon, 29 km W Linares, nr. 24°8'N, 99°8’W, 30 June 1983, 12 (W. Maddison, MCZ). San Luis Potost: El Pujal, 18— 21 July 1939, 1? (R. Haag, MCZ). Jalisco: Rio Ru- dioso [Rio Ruidosa], 22 (N. Banks, MCZ). Veracruz: 22.5 km S Catemaco on Rt. 180, 23 June 1982, 19 (F. Coyle, MCZ); Jalapa, July 1981, 52 (C. Gold, CAS); 4.8 km NE Huatusco, 22 July 1985, 29 (J. Woolley et al., AD). Hidalgo: Jacala, 1,370 m, 30 June—3 July 1939, 12 (B. Wathall, AMNH). Puebla: Huauchinango, 9 Oct. 1947, many 2 ¢ (H. Wagner, AMNH). Oaxaca: Llanos de la Flores, 24 km NE Ixtlan de Juarez, 21 July 1985, 1d (J. Woolley, et al., AD); base San Filipe Mt., 16, 17 Sep. 1947, 1d (B. Malkin, AMNH); Oaxaca, 19 July 1947, 12 (B. Mal- kin, AMNH); San Juan Quiotepec, July 1962, 12 (W. S. Miller, AMNH). Campeche: Chicanna Ruins, 8 km W Xpujil, 18°32'N, 89°31'W, 12-14 July 1983, 12 (W. Maddison, MCZ). Chiapas: (Van Patten, MCZ); San 170 Cristobal de Las Casas, 22 July 1947, 1d (C., M. Goodnight, AMNH). GUATEMALA Capetillo, 20— 23 Aug. 1947, 12 (C., PB Vaurie, AMNH). NICA- RAGUA Jinotega: Jinotega, 15 Aug. 1989, 1d (F. Reinboldt, JM); 5 km Jinotega, Los Pinares, 29 May 1992, 12 (J. Maes, C. Pineda, JM); Matagalpa, 4 Oct. 1952, 22 (R. B. Swain, AMNH); 1 Nov. 1991, 12 GE Maes, JM). Mangora ixtapan new species Figures 157-163; Map 2G Holotype. Female holotype, male paratype, and two female paratypes from 6.5 km W Uruapan, 2,300 m, Michoacan, Mexico, 15 Aug. 1967 (R. E. Leech) in MCZ. The name is a noun in apposition after the locality of a paratype. Description. Female holotype. Prosoma light orange, except black circles around secondary eyes; distal articles of legs gray. Abdomen orange-white, with spinnerets dark gray (Fig. 160). Posterior eye row straight. Ocular trapezoid longer than wide, rectangular. Posterior median eyes 1.0 diameter of anterior medians; lateral eyes 0.8 diameter. Anterior median eyes 1.0 diameter apart, 1.0 from laterals. Pos- terior median eyes 0.9 diameter apart, 1.2 from laterals. Height of clypeus equals 0.8 diameter of anterior median eyes. Total length 3.1 mm. Carapace 1.2 mm long, 1.1 wide in thoracic region, 0.6 wide behind lateral eyes, 0.6 high. First femur 1.4 mm, patella and tibia 1.5, metatarsus 1.3, tarsus 0.6. Second patella and tibia 1.4 mm, third 0.8, fourth 1.4. Male paratype. Prosoma yellow, abdo- men lighter yellowish without marks. Pos- terior eye row recurved. Ocular trapezoid longer than wide, widest anteriorly. Pos- terior median eyes 1.2 diameters of ante- rior medians; lateral eyes 0.8 diameters. Anterior median eyes 0.8 diameter apart, 1.0 from laterals. Posterior median eyes 0.3 diameter apart, 1.0 from laterals. Total length 2.3 mm. Carapace 1.2 mm long, 0.9 wide in thoracic region, 0.4 wide behind lateral eyes, 0.4 high. First femur 1.2 mm, patella and tibia 1.3, metatarsus 0.9, tarsus 0.5. Second patella and tibia 1.2 mm, third Oo7 fountin ale Variation. Total length of females 2.7— Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 3.1 mm. The illustrations were made from female holotype and male paratype. Diagnosis. The epigynum of M. ixtapan is lightly sclerotized and distinguished from others by the semicircular scape (Fig. 157) and a posterior median plate widen- ing toward the scape (Fig. 158). The palpus of the male, in apical view, differs from that of M. passiva and M. can- dida by having the dagger-shaped rod with a right-angle bend (Figs. 161, 163). Distribution. Michoacan, México, Mex- ico (Map 2G). Paratype. MEXICO Mexico: Ixtapan de la Sal, 21— 28 Aug. 1946, 12 (HA. Wagner, AMNH). Mangora candida Chickering Figures 164-170; Map 2G Mangora candida Chickering, 1954: 198, figs. 6-9, 263. Male holotype from Barro Colorado Island, Gatun Lake, Panama, in MCZ, examined. Platnick, 2004. Description. Female from Madden Dam, Panama. Prosoma yellow-white. Ab- domen whitish with white pigment spots, and several pairs of black lines posteriorly (Fig. 167). Posterior eye row slightly re- curved. Ocular trapezoid longer than wide, widest anteriorly. Posterior median eyes 0.9 diameter of anterior medians; lateral eyes 0.8 diameter. Anterior median eyes 1.0 diameter apart, 0.8 from laterals. Pos- terior median eyes almost touching, 1.3 di- ameters from laterals. Height of clypeus equals 0.8 diameter of anterior median eyes. Total length 3.8 mm. Carapace 1.7 mm long, 1.3 wide in thoracic region, 0.6 wide behind lateral eyes, 0.8 high. First femur 1.8 mm, patella and tibia 2.3, meta- tarsus 1.8, tarsus 0.8. Second patella and tibia 1.8 mm, third 1.2, fourth 2.0. Male from Madden Dam. Coloration as in female (Fig. 167). Posterior eye row slightly recurved. Ocular trapezoid longer than wide, widest anteriorly. Posterior me- dian eyes 0.8 diameter of anterior medi- ans; lateral eyes 0.5 diameter. Anterior me- dian eyes 0.6 diameter apart, 0.5 from lat- erals. Posterior median eyes almost touch- ing, 1.0 diameter from laterals. Total CENTRAL AMERICAN MANGORA * Levi WL NN SS = ‘em DS. aes Vari \ Figures 148-156. Mangora passiva (O. P.-Cambridge). 148-153, female. 148-151, epigynum. 148, 150, ventral. 149, 151, posterior. 152, abdomen, dorsal. 153, abdomen, ventral. 154-156, male, left palpus. 154, apical. 155, mesal. 156, ventral. Figures 157-163. Mangora ixtapan new species. 157-160, female. 157-159, epigynum. 157, ventral. 158, posterior. 159, posterior, cleared. 160, abdomen, dorsal. 161—163, male, palpus. 161, apical. 162, mesal. 163, ventral. Figures 164-170. Mangora candida Chickering. 164—167, female. 164-166, epigynum. 164, ventral. 165, posterior. 166, pos- terior, cleared. 167, abdomen, dorsal. 168-170, male, palpus. 168, apical. 169, mesal. 170, ventral. Scale lines: 1.0 mm, genitalia 0.1 mm. length 2.7 mm. Carapace 1.3 mm long, 0.9 wide in thoracic region, 0.4 wide behind lateral eyes, 0.7 high. First femur 1.5 mm, patella and tibia 1.8, metatarsus 1.5, tarsus 0.6. Second patella and tibia 1.4 mm, third 0.8, fourth 1.5. Variation. Total length of females 3.3— SS name maless252=2 a7 Simmel ine ellnistra tions were made from the holotype and male allotype. Diagnosis. The epigynum of M. candida is lightly sclerotized and distinguished from others by having the long scape flanked by large depressions (Fig. 164), and a triangular, pointed, median posterior plate (Fig. 165). The pattern of the abdo- men, with several pairs of black streaks (Fig. 167), is distinctive. Unlike that of the males of M. passiva and M. sufflava, the palpus of the male in apical view has a gently curved dagger- shaped rod (Fig. 168) with a distal flat- tened tip (ie. Vea) Natural History. Specimens have been collected in forest in Panama. Distribution. This species is known only from along the Panama Canal area (Map 26); Paratypes. PANAMA Panaméd: Barro Colorado Is- land, 2 Aug. 1954, 1d (A. M. Chickering, MCZ); 1975, 1d (W. Eberhard, MCZ); Madden Dam, Aug. 1939, 12, 2d (A. M. Chickering, MCZ); Forest Re- serve, numerous collections (A. M. Chickering, MCZ). Mangora sufflava Chickering Figures 171-174; Map 2G Mangora sufflava Chickering, 1963: 9, figs. 12-15, ¢. Male holotype from Boquete, Chiriqui, Panama, in MCZ, examined. Platnick, 2004. Note. Chickering did not mention the similarity of this species to M. passiva or how they can be separated. Description. Male holotype. Carapace Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 with a median dusky line, median eye re- gion and clypeus gray. Legs yellowish. Sternum yellowish, gray toward borders. Abdomen yellowish white, dorsally with posterior pair of black rectangles (Fig. 174). Posterior eye row recurved. Ocular trapezoid longer than wide, widest anteri- orly. Posterior median eyes 1.0 diameter of anterior medians; lateral eyes 0.8 diameter. Anterior median eyes 0.8 diameter apart, 0.5 from laterals. Posterior median eyes 0.3 diameter apart, 1.0 from laterals. Height of clypeus equal to 0.8 diameter of anterior median eye. Total length 2.6 mm. Carapace 1.3 mm long, 1.1 wide in tho- racic region, 0.5 wide behind lateral eyes, 0.7 high. First femur 1.5 mm, patella and tibia 1.6, metatarsus 1.2, tarsus 0.7. Sec- ond patella and tibia 1.4 mm, third 0.8. Fourth femur 1.3 mm, patella and tibia 1.2, metatarsus 0.9, tarsus 0.5. The female is not known. Diagnosis. The apical view of the palpus differs from that of M. passiva (Fig. 154) by having a more gracefully shaped curved rod (Fig. 171), and in mesal view, a heavily sclerotized median apophysis spine (4 h in Fig. 172), Distribution. Chiriqui, Panama (Map ey. Specimens Examined. No other specimens have been found. Mangora chicanna new species Figures 175-181; Map 2H Holotype. Female holotype from Chicanna Ruins, ca. 8 km W of Xpujil, ca. 18°32'N, 89°31"W, Campe- che, Mexico, 12-14 July 1983 (W. Maddison) in MCZ. The species is named after the type locality as a noun in apposition. Description. Female paratype from Honduras. Carapace golden yellow, with small black eye rings. Sternum, legs yellow. = Figures 171-174. Mangora sufflava Chickering, male. 171-173, left palpus. 171, apical. 172, mesal. 173, ventral. 174, abdo- men, dorsal. Figures 175-181. Mangora chicanna new species. 175-178, female. 175-177, epigynum. 175, ventral. 176, posterior. 177, posterior cleared. 178, abdomen, dorsal. 179-181, male, palpus. 179, apical. 180, mesal. 181, ventral. CENTRAL AMERICAN MANGORA ° Levi 173 sufflava — é, 172 Qn 173 pal TSS S oa Figures 182-188. Mangora purulha new species. 182-185, female. 182-184, epigynum. 182, ventral. 183, posterior. 184, posterior, cleared. 185, abdomen, dorsal. 186-188, male, palpus. 186, apical. 187, mesal. 188, ventral. Figures 189-192. Mangora schneirlai Chickering, female. 189-191, epigynum. 189, ventral. 190, posterior. 191, posterior, cleared. 192, abdomen, dorsal. Scale lines: 1.0 mm, genitalia 0.1 mm. 174 Abdomen whitish, with pairs of white pig- ment patches and four pairs of round, black spots (Fig. 178); venter without marks. Posterior eye row procurved. Oc- ular trapezoid longer than wide, widest an- teriorly. Posterior median eyes 0.8 diame- ter of anterior medians; lateral eyes 0.7 di- ameter. Anterior median eyes 0.5 diameter apart, 0.5 from laterals. Posterior median eyes 0.3 diameter apart, 1.2 from laterals. Total length 2.4 mm. Carapace 1.2 mm long, 0.8 wide in thoracic region, 0.3 wide behind lateral eyes, 0.5 high. First femur 1.4 mm, patella and tibia 1.6, metatarsus 1.3, tarsus 1.1. Second patella and tibia 1.4 mm, third 0.7, fourth 1.3 mm. Male paratype from Campeche. All light yellowish, abdomen lighter with four pairs of black spots on abdomen, but no white pigment. Posterior eye row slightly pro- curved. Eye trapezoid widest in front, as long as wide in front. Posterior median eyes 0.7 diameter of anterior medians; lat- eral eyes 0.5 diameter. Anterior median eyes 0.7 diameter apart, 0.4 from laterals. Posterior median eyes 0.4 diameter apart, 1.0 from laterals. Total length 2.2 mm. Carapace 1.1 mm long, 0.7 wide in tho- racic region, 0.3 wide behind lateral eyes, 0.5 high. First femur 1.3 mm, patella and tibia 1.6, metatarsus 1.2, tarsus 0.5. Sec- ond patella and tibia 1.3 mm, third 0.7, fourth 1.2 mm. Variation. Total length of females 2.2— 3.0 mm, males 1.8—2.2 mm. The illustra- tions of a female were made from a para- type from Honduras, and male allotype from Campeche, Mexico. Diagnosis. The four pairs of circular, black dots on the abdomen (Fig. 178) eas- ily characterize M. chicanna. The epigyn- um is lightly sclerotized and distinguished from others by the short, wide posterior median plate (Fig. 176). The palpus of the male, in apical view, differs by having a strongly curved rod (Fig. 179) and the median apophysis has two spines (Figs. 179, 180). Natural History. Specimens came from dry, deciduous forest in Campeche. Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 Distribution. Yucatan Peninsula, Chia- pas to Honduras (Map 2H). Paratypes. MEXICO Campeche: Reserva de la Biosfera, Calakmul, km 32, 20 July 1998, 12, 16 (F. Alvarez, MCZ); 26 July 1998, 22, 1d (F. Alvarez, MCZ). Yucatan: Colonia Yucatan, 22 Aug. 1964, 19 (J. C. Pallister, AMNH); Progreso, 25 July 1952, 19 (J., D. Pallister, AMNH). Quintana Roo: Chancanah, Cozumel, 8 Aug. 1949, 12 (C. Goodnight, AMNH); San Felipe de Bacalor, 8 July 1993, 12 (G. Alayén, ECOSUR); Rancho Las Palmas, 30 km S Felipe Ca- rrillo Puerto, 10 June 1993, 1d (G. Alay6n, ECO- SUR). Chiapas: El Real, 6 July 1950, 12 (C., M. Goodnight, AMNH). HONDURAS Bouacca Island [ PI}, aus Apr. 1935, 12 (M. Bates, MCZ). Mangora purulha new species Figures 182-188; Map 2H Holotype. Female holotype with male paratype from Purulha, Baja Verapaz, Guatemala, 2,300 m, 23 May 1979 (J. Coddington) in MCZ. The name is a noun in apposition after the type locality. Description. Female holotype. Prosoma pale orange. Abdomen light orange-white, with a pair of posterior spots and white pigment anterior on sides (Fig. 185); ven- ter orange-white, sides of spinnerets gray. Posterior eye row straight. Ocular trape- zoid longer than wide, rectangular. Poste- rior median eyes 1.2 diameters of anterior medians; lateral eyes 1.0 diameter. Ante- rior median eyes 1.0 diameter apart, 1.0 diameter from laterals. Posterior median eyes 0.6 diameter apart, 1.1 from laterals. Height of clypeus equals 0.5 diameter of anterior median eyes. Total length 3.2 mm. Carapace 1.4 mm long, 1.2 wide in tho- racic region, 0.6 wide behind lateral eyes, 0.7 high. First femur 1.5 mm, patella and tibia 1.8, metatarsus 1.5, tarsus 0.7. Sec- ond patella and tibia 1.7 mm, third 1.0. Fourth femur 1.6 mm, patella and tibia 1.6, metatarsus 1.3, tarsus 0.6. Male paratype. Similar color as female, but white on sides of abdomen more ex- tensive than in female. Posterior eye row straight. Ocular trapezoid longer than wide, widest anteriorly. Posterior median eyes 0.8 diameter of anterior medians; lat- eral eyes 0.7 diameter. Anterior median eyes 1.2 diameters apart, 1.0 from laterals. Posterior median eyes 1.0 diameter apart, 1.8 from laterals. Total length 2.7 mm. Carapace 1.6 mm long, 1.3 wide in tho- racic region, 0.5 wide behind lateral eyes, 0.7 high. First femur 1.6 mm, patella and tibia 1.8, metatarsus 1.6, tarsus 0.7. Sec- ond patella and tibia 1.7 mm, third 1.0, fourth 1.7. Diagnosis. Mangora purulha is distin- guished by two circular, black dots on the abdomen (Fig. 185). The epigynum differs by a lightly sclerotized, slender scape on a curved, transverse plate (Fig. 182), and in posterior view, by lateral sclerotization (Fig. 183). Males also have two black spots on the abdomen and the palpus has a median apophysis with one curved spine visible in apical view (Fig. 186), a soft, large embo- lus (Figs. 187, 188), and a sclerotized ter- minal apophysis (Fig. 186). Because only one male was available, the sclerites were not carefully examined nor was their ho- mology checked. Distribution. Guatemala (Map 2H). Specimens Examined. No other specimens have been found. Mangora schneirlai Chickering Figures 189-192; Map 2H Mangora schneirlai Chickering, 1954: 209, fig. 22, °. Female holotype from Barro Colorado Island, Ga- tun Lake, Panama, in AMNH, examined. Platnick, 2004. Description. Female paratype. Prosoma yellowish. Abdomen yellowish, with four white spots and a pair of black rectangles posteriorly (Fig. 192); venter with a pair of white spots lateral to genital furrow and lateral to spinnerets. Posterior eye row re- curved. Ocular trapezoid slightly longer than wide, widest anteriorly. Posterior me- dian eyes 1.0 diameter of anterior medi- ans; lateral eyes 0.6 diameter. Anterior me- dian eyes 0.8 diameter apart, 0.8 from lat- erals. Posterior median eyes 0.4 diameter apart, 1.0 from laterals. Legs heavy. Total length 4.8 mm. Carapace 1.7 mm long, 1.0 wide in thoracic region, 0.6 wide behind lateral eyes, 0.7 high. First femur 1.8 mm, patella and tibia 2.0, metatarsus 1.5, tarsus CENTRAL AMERICAN MANGORA * Levi WS) 0.7. Second patella and tibia 1.8 mm, third 1.3. Fourth femur 1.7 mm, patella and tib- ia 2.0, metatarsus 1.6, tarsus 0.7. The male is not known. Variation. Total length of females 3.5— 4.8 mm. The illustrations were made from the female holotype. Diagnosis. Mangora schneirlai belongs to the group of species with two black rectangles on the posterior of the abdo- men (Fig. 192). The epigynum differs from that of other species with this pattern by having a median scape flanked by a semicircular notch on each side (Fig. 189). The posterior view of the epigynum of M. schneirlai (Fig. 190) differs from others, except M. corcovado, by having the wide, median plate covered by the narrow lateral ones on each side (Fig. 190). Distribution. Costa Rica, Panama (Map 2H). Paratype. PANAMA Panama, Apr., May 1946, 1° (T. C. Schneirla, MCZ). Specimens Examined. COSTA RICA Puntarenas: Corcovado Natl. Park, 11 Aug. 1979, 12 (J. Cod- dington, MCZ). PANAMA Panamd: Barro Colorado Isl., Gatun Lake, 30 July, 1964, 12 (D. M. Rees, AMNH); nr. Gamboa, Sep. 1975, 22 (W. Eberhard, MCZ). Mangora bimaculata (O. P.-Cambridge) Figures 193-199; Map 2l Epeira bimaculata O. P.-Cambridge, 1889: 21, pl. 6, figs. 12, 13, 2d. Specimens from between Petet [Petén, El Petén] and Chicoyoito [?], Guatemala, Cahabon [Alto Verapaz], Quirigua [Izabal], and Veragua, Panama, in BMNH, examined. Keyser- ling, 1893: 260, pl. 13, fig. 19, 26. Mangora bimaculata:—F. O. P.-Cambridge, 1904: 479, pl. 45, figs. 10, 11, 2d. Chickering, 1954: 197, figs. 1-5, 96. Platnick, 2004. Description. Female from Petén, Gua- temala. Prosoma yellowish. Abdomen whitish with white pigment spots and a pair of dorsal posterior black rectangles (Fig. 196); venter with white pigment spots except in midline. Posterior eye row recurved. Ocular trapezoid as long as wide in front, widest anteriorly. Posterior me- dian eyes 1.0 diameter of anterior medi- ans; lateral eyes 0.7 diameter. Anterior me- 176 dian eyes 0.4 diameter apart, 1.2 from lat- erals. Posterior median eyes 0.2 diameter apart, 1.5 from laterals. Total length 6.3 mm. Carapace 2.2 mm long, 1.8 wide in thoracic region, 0.8 wide behind lateral eyes, 1.2 high. First femur 2.7 mm, patella and tibia 3.0, metatarsus 2.6, tarsus 1.1. Second patella and tibia 2.7 mm, third 1.7, fourth 2.9. Male from Hidalgo, Mexico. Prosoma light orange. Abdomen as in female. Pos- terior eye row procurved. Ocular trapezoid longer than wide, widest anteriorly. Pos- terior median eyes 0.9 diameter of anterior medians; lateral eyes 0.8 diameter. Ante- rior median eyes 1.0 diameter apart, 0.3 from laterals. Posterior median eyes al- most touching, 0.8 from laterals. Total length 2.3 mm. Carapace 1.2 mm long, 0.9 wide in thoracic region, 0.4 wide behind lateral eyes, 0.7 high. First femur 1.3 mm, patella and tibia 1.5, metatarsus 1.2, tarsus 0.7. Second patella and tibia 1.3 mm, third ON fount de?) Variation. Total length of females 5.3— 6.3 mm. A female from Chiapas has indis- tinct black bands rather than patches on the abdomen. The illustrations were made from female and male syntypes; Figure 197 was from a Hidalgo male. Diagnosis. Mangora bimaculata differs from sympatric species in the north by the coloration of the abdomen and from those with similar abdomen patches by the epi- gynum, which is heavily sclerotized and has an oval posterior opening with an an- terior median tubercle (Fig. 193); in pos- terior view the median plate forms a groove (Fig. 194). The palpus of the male differs from that of M. sufflava by the different shape of the terminal apophysis, having a shorter, distal dagger-shaped rod (Fig. 197). Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 Natural History. Specimens from Bar- rancas came from a palm forest; the Chia- pas specimens were collected by sweeping. Distribution. Central Mexico to Costa Rica (Map 21). Specimens Examined. MEXICO Hidalgo: 3.2 km SW Jacala, 18 Aug. 1964, 1d Ge W. Ivie, AMNH). Chiapas: Palenque Ruins, 9 July 1949, 12 (C. J. Goodnight, AMNH). GUATEMALA Petén: Uaxac- tan, Mar—Apr. 1931, 12 (H. H. Bartlett, MCZ). COSTA RICA San José: San Isidro del General, 600— 1,200 m, 12 (D. Rounds, MCZ). Puntarenas: Barran- cas, 13 June 1979, 12 (J. Coddington, MCZ). Mangora pia Chamberlin and Ivie Figures 200-207; Map 2J Mangora pia Chamberlin and Ivie, 1936: 58, pl. 12, fig. 112, 2. Female holotype from Barro Colorado Island, Panama, in AMNH, examined. Chickering, 1954: 208, figs. 18-21, 2d. Platnick, 2004. M. belligerens Chamberlin and Ivie, 1936: 60, pl. 12, fig. 113, 5. Male holotype from Barro Colorado Island, Panama, in AMNH, examined. First syn- onymized with pia by Chickering, 1954. M. wiedenmeyeri Schenkel, 1953: 18, fig. 15, 2. Fe- male holotype from El Pozén, Depto. Acosta, Fal- con, Venezuela, in NMB, examined. Platnick, 2004. NEW SYNONYMY. Description. Female from Barro Colo- rado Island. Prosoma light orange. Abdo- men whitish, with a pair of posterior black rectangles and indistinct longitudinal lines with white pigment spots (Fig. 203); ven- ter whitish with a pair of white spots, one on each side anterior to spinnerets, and two indistinct longitudinal lines of white pigment spots. Posterior eye row straight. Ocular trapezoid longer than wide, widest anteriorly. Posterior median eyes 0.8 di- ameter of anterior medians; lateral eyes 0.6 diameter. Anterior median eyes 0.7 di- ameter apart, 1.0 from laterals. Posterior median eyes 0.4 diameter apart, 1.4 from laterals. Total length 5.5 mm. Carapace 2.4 mm long, 2.0 wide in thoracic region, 0.8 —s Figures 193-199. Mangora bimaculata (O. P.-Cambridge). 193-196, female. 193-195, epigynum. 193, ventral. 194, posterior. 195, posterior, cleared. 196, abdomen, dorsal. 197-199, male, left palpus. 197, apical. 198, mesal. 199, ventral. Figures 200-207. Mangora pia Chamberlin and lvie. 200-203, female. 200-202, epigynum. 200, ventral. 201, posterior. 202, posterior, cleared. 203, abdomen, dorsal. 204-207, male. 204—206, palpus. 204, apical. 205, mesal. 206, ventral. 207, proximal end of fourth leg, ventral. CENTRAL AMERICAN MANGORA ° Levi Wa © corcovado Figures 208-214. Mangora falconae Schenkel. 208-211, female. 208-210, epigynum. 208, ventral. 209, posterior. 210, pos- terior, cleared. 211, abdomen, dorsal. 212—214, male, palpus. 212, apical. 213, mesal. 214, ventral. Figures 215-218. Mangora corcovado new species, female. 215-217, epigynum. 215, ventral. 216, posterior. 217, posterior, cleared. 218, abdomen, dorsal. Scale lines: 1.0 mm, genitalia 0.1 mm. 73 wide behind lateral eyes, 1.4 high. First femur 3.0 mm, patella and tibia 3.2, meta- tarsus 3.0, tarsus 1.1. Second patella and tibia 3.1 mm, third 2.1. Fourth femur 3.2 mim, patella and tibia 3.3, metatarsus 2.8, (resus IL JL. Male from Barro Colorado Island, Pan- ama. Coloration as in female. Posterior eye row straight. Eye trapezoid longer than wide, widest anteriorly. Posterior median eyes 0.8 diameter of anterior medians; an- terior lateral eyes 0.7 diameter, posterior 0.6. Anterior median eyes 0.8 diameter apart, 0.8 from laterals. Posterior median eyes 0.2 diameter apart, 1.2 from laterals. Fourth femur with a proximal, ventral ma- croseta (Fig. 207). Total length 3.8 mm. Carapace 2.0 mm long, 1.8 wide in tho- racic region, 0.7 wide behind lateral eyes, 1.0 high. First femur 2.3 mm, patella and tibia 2.6, metatarsus 2.3, tarsus 0.9. Sec- ond patella and tibia 2.2 mm, third 1.5, fourth 2.4. Variation. Total length of females 5.0— 6.7 mm, males 3.3-4.3 mm. The illustra- tions were made from the female holotype of M. pia: Figures 205 and 206 were from the holotype of M. belligerens; and Figure 204 was from a specimen from Barro Col- orado Island, Panama. Diagnosis. Mangora pia differs from fe- males of other species with a pair of black rectangles on the abdomen by the distinc- tive heavily sclerotized epigynum, which has a median, triangular notch, framed by a sclerotized fold (Fig. 200). Males differ from males of M. melano- cephala and M. falconae (which also have a macroseta on the venter of the proximal end of the fourth femur; Fig. 207), by be- ing much larger in size than M. melano- cephala and lacking the filamentous em- bolus in the palpus (Figs. 205-206), and from M. falconae, in apical view, by the thick spine of the terminal apophysis (Fig. 204). Natural History. Living specimens are green (Craig, personal communication). Specimens have been found in forest in Panama. Mangora pia “is a wet season, Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 forest understory spider and the ones I worked on lived in vegetation near streams but not on the stream edge. They build very delicate and beautiful webs, densely spun, and the viscid silk is extremely stretchy and densely coated with glue” (Craig, personal communication). Lubin (1978) writes that the species is primarily diurnal but complete webs can be found at night. Distribution. Panama, northern Vene- zuela and eastern Colombia (Map 2]). Specimens Examined. PANAMA Taboga Isl., 23 Aug. 1946, 12 (N. L. H. Krauss, AMNH). Panama: Araijan, Cerro Galero, very common (W. Eberhard, MCZ); Barro Colorado Island, Gatun Lake, very common (AMNH, MCZ, USNM); Cocoli area, July— Sep. 1954, 29 (W. E. Lundy, AMNH); France Field, Aug. 1939, 1d (A. M. Chickering, MCZ); Pedro Mi- guel, July 1950, 12 (A. M. Chickering, MCZ); Ex- perimental Gardens, 11—13 July 1954, 12, 2d (A. M. Chickering, MCZ); Forest Reserve, Aug. 1939, 2d; July 1950, 1d (A. M. Chickering, MCZ); Parque Nac. Soberiana, 7 Aug. 1983, 29 (R. J. Raven, AMNH); Pipe line Road nr. Gamboa, 6 Jan. 1977, 12 (H. Levi, M. Robinson, MCZ); Chilibre, July 1950, 1d (A. M. Chickering, MCZ); Madden Dam, Aug. 1939, 19 (A. M. Chickering, MCZ); Fort Knobbe, 3 Aug. 1983, 19 (H., L. Levi et al., MCZ); Summit, Nov. 1946, 192 (N. L. H. Krauss, AMNH); Summit Gardens, July 1954, 12 (A. M. Chickering, MCZ); El Valle, 32 (A. M. Chickering, MCZ); Villa Darién, 12-28 Feb. 1984, 22 (M. N. Garcia, MIUP). COLOMBIA Valle: nr. Cali, 1,000 m, 32 (W. Eberhard 1128, 1131, 1132, MCZ); Rio Jamundi nr. Jamundi, 9 Dec. 1969, 19 (W. Eberhard, MCZ). Mangora falconae Schenkel Figures 208-214; Map 2K Mangora falconae Schenkel, 1953: 19, fig, 17, 2. Fe- male holotype from El Pozén, Depto. Acosta, Fal- con, Venezuela, in NMB, examined. Platnick, 2004. Description. Female from Magdalena, Colombia. Prosoma light orange. Abdo- men orange-white, with two posterior black rectangles and streaks of white pig- ment spots (Fig. 211); venter orange- white. Posterior eye row straight. Ocular trapezoid longer than wide, slightly widest anteriorly. Posterior median eyes 0.8 di- ameter of anterior medians; lateral eyes 0.7 diameter. Anterior median eyes 0.5 di- ameter apart, 0.8 from laterals. Posterior median eyes 0.3 diameter apart, 1.2 from laterals. Total length 4.3 mm. Carapace 1.8 mm long, 1.5 wide in thoracic region, 0.7 wide behind lateral eyes, 1.3 high. First femur 2.1 mm, patella and tibia 2.1, meta- tarsus 1.6, tarsus 0.6. Second patella and tibia 1.8 mm, third 1.3. Fourth femur 2.1 mim, patella and tibia 2.2, metatarsus 1.7, tarsus 0.7. Male from Magdalena, Colombia. Col- oration as in female, except venter of ab- domen with a large, median patch of white pigment spots. Posterior eye row slightly recurved. Ocular trapezoid longer than wide, widest anteriorly. Posterior median eyes 0.8 diameter of anterior medians; lat- eral eyes 0.6 diameter. Anterior median eyes 0.4 diameter apart, 0.4 from laterals. Posterior median eyes 0.3 diameter apart, 1.0 from laterals. Height of clypeus equal to 0.8 diameter of anterior median eye. Fourth femur with proximal, ventral ma- croseta (Fig. 207). Total length 2.8 mm. Carapace 1.8 mm long, 1.2 wide in tho- racic region, 0.5 wide behind lateral eyes, 0.7 high. First femur 1.6 mm, patella and tibia 1.7, metatarsus 1.3, tarsus 0.6. Sec- ond patella and tibia 1.4 mm, third 0.9, fourth 1.5. Variation. Total length of females 3.1— 4.4 mm, males 2.6—2.8 mm. The illustra- tions were made from the female holo- type, and from a male specimen from Co- lombia. Diagnosis. The epigynum of M. falconae is distinct from that of all species in having a pair of black rectangles on the abdomen, by the scape having a narrow base (Fig. 208), and in posterior view having the me- dian plates upside-down-T-shaped (Fig. 209). Males are distinguished from those of M. melanocephala and M. pia, which also have a macroseta on the fourth femur (F ig. 207), by a palpus with the terminal apoph- ysis having a thorn-shaped hook seen in apical view (Fig. 212). Natural History. Specimens have been collected on plants in Colombia, and from a coastal thorn-scrub in Venezuela. CENTRAL AMERICAN MANGORA * Levi 179 Distribution. Panama, northern Vene- zuela, Colombia (Map 2K). Specimens Examined. PANAMA Chiriqut: 1938, 12 (AMNH#). VENEZUELA Sucre: Cartpano, OS ort July 1987, 1d (S., J. Peck, AMNH). COLOMBIA Magdalena: Tayrone Park, Gairaca, 8 km NE Santa Marta, 13 June 1985, 12 (H.-G. Miiller, SMF); Tayrone Park, 16 km NE Santa Marta, 16 June 1985, 92, 1d, 2 imm. (H.-G. Miiller, SMF). Mangora corcovado new species Figures 215-218; Map 2K Holotype. Female holotype from Llocona [?]. Cor- covado National Park, Puntarenas, Costa Rica, 11 Aug. 1979 (J. Coddington), in MCZ. The name is a noun in apposition after the name of the type locality. Description. Female holotype. Prosoma yellowish. Abdomen yellowish white with a pair of black rectangles posteriorly, and areas of white spots anteriorly (Fig. 218); venter without marks. Posterior eye row procurved. Ocular trapezoid longer than wide, widest anteriorly. Posterior median eyes 1.0 diameter of anterior medians; lat- eral eyes 0.6 diameter. Anterior median eyes 0.3 diameter apart, 0.5 from laterals. Posterior median eyes 0.2 diameter apart, 1.0 from laterals. Total length 3.2 mm. Carapace 1.7 mm long, 1.3 wide in tho- racic region, 0.6 wide behind lateral eyes, 1.2 high. First femur 1.6 mm, patella and tibia 2,0) metatarsus 1245 tarsus O47. Sec= ond patella and tibia 1.8 mm, third 1.3. Fourth femur 1.8 mm, patella and tibia 1.9, metatarsus 1.5, tarsus 0.7. The male is not known. Diagnosis. The scape of the epigynum of M. corcovado is a short lobe without constriction (Fig. 215) compared to the longer, constricted scape of M. falconae (Fig. 208) and in posterior view the epi- gynum has a wider median plate (Fig. 216) than in M. falconae. Distribution. Costa Rica (Map 2K). Specimens Examined. No other specimens have been found. 180 ACKNOWLEDGMENTS Without the collections mentioned in the Methods, this revision would not have been possible and I am grateful to the cu- rators and collectors mentioned there for their help. I also thank Robin Leech and J. Beatty each for the gift of a specimen and W. Eberhard, G. Ibarra N., and J. M. Maes for information on obscure localities. W. Piel provided instructions for mapping. Lorna Levi and Laura Leibensperger read the manuscript and helped with wording; C. Dondale made helpful comments. Na- tional Science Foundation grant GB- 36161 supported the start of the revision in the 1970s, enabling the borrowing and illustrating of old types. LITERATURE CITED BONNET, P. 1957. Bibliographia Araneorum, Tou- louse, 2: 1927-3026. . 1961. Bibliographia Araneorum, Toulouse, 3: 1-591. BRYANT, E. 1945. The Argiopidae of Hispaniola. Bul- letin of the Museum of Comparative Zoology, 95: 357-418. CAMBRIDGE, F. O. P.-. 1904. Arachnida, Araneidea and Opiliones, pp. 465—545. In Biologia Centrali- Americana, Zoologia. Vol. 2. London, United Kingdom. CAMBRIDGE, O. P.-. 1889. Arachnida, Araneidea and Opiliones, pp. 1-56. In Biologia Centrali-Amer- icana, Zoologia. Vol. 1. London, United King- dom. CAPORIACCO, L. 1948. Arachnida of British Guiana collected by Prof. Beccari. Proceedings of the Zoological Society of London, 118: 607-747. CHAMBERLIN, R. V., AND W. IvIE. 1936. New spiders from Mexico and Panama. Bulletin of the Uni- versity of Utah, Biological Series, 27(5): 1-103. CHICKERING, A. M. 1954. The genus Mangora (Ar- giopidae) in Panama. Bulletin of the Museum of Comparative Zoology, 111: 193-215. . 1963. Three new species of Mangora (Ara- neae, Argiopidae) from Central America. Bre- viora, Museum of Comparative Zoology, 191: 1—- ee CRAIG, C. L. 1987a. The significance of spider size to the diversification of spider-web architectures and spider reproductive modes. The American Naturalist, 129: 47-68. . 1987b. The ecological and evolutionary in- terdependence between web architecture and web silk spun by orb web weaving spiders. Bio- logical Journal of the Linnaean Society, 30: 135— 162. Bulletin Museum of Comparative Zoology, Vol. 158 No. 4 . 1988. Insect perception of spider orb webs in three light habitats. Functional Ecology, 2: LN PD . 1989. Alternative foraging modes of orb web weaving spiders. Biotropica, 21: 257-264. FRANGANILLO BALBOA, P. 1936. Los Ardcnidos de Cuba Hasta 1936. La Habana, Cuba: Cultural S. A. 183 pp. GOODNIGHT, C. J., AND M. L. GOODNIGHT. 1953. The opilionid fauna of Chiapas, Mexico and ad- jacent areas (Arachnoidea, Opiliones). American Museum Novitates, 1610: 1-81. HENTZ, N. M. 1847. Descriptions and figures of the Araneides of the United States. Boston Journal of Natural History, 5: 443-478. KEYSERLING, E. 1881. Neue Spinnen aus Amerika. Verhandlungen der K. K. Zoologisch-Botanisch- en Gesellschaft in Wien, 30: 547-582. . 1892-1893. Die Spinnen Amerikas, Epeiri- dae. Verlag von Bauer und Raspe, Niirnberg, 4: L877. LEvI, H. W. 1975. The American orb-weaver genera Larinia, Cercidia and Mangora north of Mexico. Bulletin of the Museum of Comparative Zoology, VAT NO 35: . 1991. The Neotropical and Mexican species of the orb-weaver genera Araneus, Dubiepeira and Aculepeira (Araneae: Araneidae). Bulletin of the Museum of Comparative Zoology, 152: 167— Cilia: . 2002. Keys to the genera of araneid orb- weavers (Araneae, Araneidae) of the Americas. Journal of Arachnology, 30: 527-562. LUBIN, Y. 1978. Seasonal abundance and diversity of web-building spiders in relation to habitat struc- ture on Barro Colorado Island, Panama. Journal of Arachnology, 6: 31-51. PETRUNKEVITCH, A. 1909. A trip to southern Mexico for spiders. American Museum Journal, 9: 245— 256. . 1911. Synonymic index—catalogue of spiders on North, Central and South America with all adjacent islands, Greenland, Bermuda, West In- dies, Terra del Fuego, Galapagos etc. Bulletin of the American Museum of Natural History, 29: t=79 PLATNICK, N. I. 2004. The World Spider Catalog, Version 4.5. American Museum of Natural His- tory, hhttp://research.amnh.org/entomology/spiders/ catalog/index. html. REIMOSER, E. 1940 (1939). Wissenschaftliche Ergeb- nisse der 6sterreichischen biologischen Expedi- tion nach Costa Rica. Annalen des Naturhisto- rischen Museum in Wien, 50: 328-386. ROEWER, C. F. 1942. Katalog der Araneae von 1758 bis 1940. Kommissions-Verlag von “Natura,” Bremen, 1: 1—1040. SCHARFF, N., AND J. CODDINGTON. 1997. A phylo- genetic analysis of the orb-weaving spider family Araneidae (Arachnida, Araneae), Zoological Journal of the Linnean Society, 120: 355-434. SCHENKEL, E. 1953. Bericht tiber einige Spinnenti- ere aus Venezuela. Verhandlungen der naturfor- schenden Gesellschaft, Basel, 64: 1—57. SELANDER, R. B., AND P. VAURIE. 1962. A gazetteer to accompany the “Insecta” volumes of the “Biol- ogia Centrali-Americana.” American Museum Novitates, 2099: 1—70. SIMON, E. 1895. Histoire Naturelle des Araignées. Libraire Encyclopédique de Roret, Paris, 1 (fase. 4): 761=1084. SOARES, B. A. M., AND H. F. DE ALMAIDA DA CA- MARGO. 1948. Aranhas coligidas pela Funda¢ao INDEX CENTRAL AMERICAN MANGORA ¢ Lewi 181 Brasil-Central (Arachnida-Araneae). Boletim do Museu Paraense EF. Goeldi, 10: 355—409. TACZANOWSKI, L. 1874. Les Aranéides de la Guyane frangaise. Horae Societatis Entomologicae Ros- sicae, St.-Petersbourg, 9: 64—150. UNITED STATES BOARD ON GEOGRAPHIC NAMES. 1956, 1965. Gazetteer. United States Depart- ment of the Interior, Office of Geography, Wash- ington, DC. VAURIE, C., AND P. VAURIE. 1949. Insect collecting in Guatemala 65 years after Champion. Journal of the New York Entomological Society, 57: 1-18. Valid names are printed in italics. Page numbers refer to the main references, starred page number to illustrations. acalypha, 142 acaponeta, 155*, 156 amchickeringi, 159*, 160 aragarcensis, 151 belligerens, 176 bimaculata, 175, 177* calcarifera. 149*, 150 campeche, 166, 167* candida, 170, 171* chicanna, 172, 173* conspicua, 162 corcovado, 177*, 179 craigae, 147, 149* decora, 142 decolorata, 142 dentembolus, 151 distincta, 158, 159* falconaé, 177*, 178 fascialata, 162, 163* fortuna, 166, 167* gibberosa, 159*, 161, 163 goodnightorum, 167*, 168 tizad, LOA, Lob* ixtapan, 170, 171* Mangora, 141 melanocephala, 151, 153* mobilis, 155*, 156 mobilis, 160 montana, 146, 149* nahuatl, 165, 167* oaxaca, 163*, 165 passiva, 169, 171* Pig. WiC. Mie picta, 141, 148, 149* picta, 151 placida, 163*, 164 pozonae, 15] purulha, 173*, 174 rostrata, 169 schneirlai, 173*, 175 spiculata, 163*, 164 spinula, 151 striatipes, 162 sufflava, 172, 173* trilineata, 148 vito, 167*, 168 volcan, 165, 167* wiedenmeyeri, 176 rae ae ee Walco, atin ata kl + Crepe) at ‘i ie 4 aed al ih ieee Fa — > Toure Ramet 1 “OL ThbeooeD & attvete Leet entail ut Ae 5) (ap weet weil = o* SPA phe iM ‘Were mnt 4% an ‘ aoe oll ney Jee 4) 7 may! 1 Tae iy 7 ne apn a Mere) Livi Re Ty af ah WY pe or ere y 7 ow ‘a! ‘eebdynghntec esa ie heed wall torsional dell enn Deeg) «Cea gta CRINGE le | oF A gal gm i? = , Gaertn! ihe 4 on : ro ay | “1 . Lr . i) pede maaan 2 el ra Werte’ iy ae Wt wey ate cata ih manly re mln i beniird | . a Preéicipefh aye aunt ’ \F asia | | Ba fhe of ie Dargie Seon, | uae Firma e444 athe A a hype Ssineewl ae ame : ail teri, cae iv ‘elgg I agian nepeat A st i) apa Amani oe ; ji Sint, : oh ies whit Bintan . 5 J ,f | D os 1 = Fo ae y Ait if 2 ~| i ‘ ky date sls Right Ae ‘ 2 ss sania ve rh" ot f eS ‘ i 4 ob tal a | = ey 7a ya ’ Ott a Ah 7 De par vr a bat x. 4 a M0) as hyphae Rulletin or 1He : Museum of — Comparative a Loology — The Snake Genus Sibynomorphus (Colubridae: Dipsadinae: Dipsadini) in Peru and Ecuador, with Comments on the systematics of Dipsadini JOHN E. CADLE HARVARD UNIVERSITY VOLUME 158, NUMBER 5 CAMBRIDGE, MASSACHUSETTS, U.S.A. 14 June 2007 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY BREVIORA 1952— BULLETIN 1863— Memoirs 1865-1938 JounsoniA, Department of Mollusks, 1941-1974 OCCASIONAL PAPERS ON MOLLUusks, 1945— SPECIAL PUBLICATIONS. 1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and Evolution of Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere- dinidea (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4. Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology and Evolutionary Biology: Essays in Honor of Ernest E. Williams. 725 pp. : 6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the M. C. Z., Vol. 108.) Reprinted 1971. Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter’s Check-list of Birds of the World, vols. 1-16. Proceedings of the New England Zoological Club 1899-1947. (Complete sets only.) Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2004. THE SNAKE GENUS S/BYNOMORPHUS (COLUBRIDAE: DIPSADINAE: DIPSADINI) INPERU AND ECUADOR, WITH COMMENTS ON THE SYSTEMATICS OF DIPSADINI JOHN E. CADLE' CONTENTS S| ORS EER OIG yo Re A i ape ea AS ee Aa ees a ae 183 TNC SUMING Wipreiata re creak. ese Meh IG teal pale! open ot Oa 184 rarROre CIOL tae A ee ne oy he eso ES 185 IMIGEDOGIS eee sae erator’ cn ei ny AS Wied Bae Le 187 Key to Species of Sibynomorphus in Ecuador STING | «J Vera eae oe oon ee Vit ait le ape 189 Sibynomorphus oligozonatus Orcés and NMC NC aAniz yates Mow wn a a eee Gl es 191 Sibynomorphus oneilli Rossman and Thomas __. 205 Sibynomorphus petersi Orcés and Almendariz bgp | £obs Vela AS os ooh ny ee erat baer aiinnlt 216 Sibynomorphus vagrans (Dunn) 239 Sizomomonplews oaeus, (an) ps ee JAD, Sibynomorphus williamsi Carrillo de Espinoza iy tg 004 pest > Ot ta tr lil tent is 248 Leptognathus atypicus Cope, 1874: Incertae SECIS eee NE eee ee NE ee ee DSS Hemipenial Morphology in Sibynomorphus aime! Otlner IDivoseelinh ae. ee el 257 Comparison of Hemipenes of SUDUMOIORD (US eae te can ee eee DST Sibynomorphus and the Dipsadini: Directions form uctnem Research) ae hese ee 261 The Systematics of Dipsadini—An Outline OLSOMENRTOMlenism eset le as ey 262 Research Needs on the Northern Species of Sibymomonrpnus ast ne a) Lees 270 JNO SIO WAS CKATNCTINS ee Diss Specimens Examined and Locality Records __ 276 Paiterature (Gite Get sted ed eas A ey 278 ABSTRACT. Sibynomorphus, an assemblage of about a dozen species of South American gastropod-eating colubrids, has a peculiar distribution. Six species oc- cur in northern Peru and southwestern Ecuador. The others are distributed south of the Amazon basin in Brazil, Bolivia, Paraguay, Uruguay, and Argentina. ‘ Associate, Department of Herpetology, Museum of Comparative Zoology. Address for correspondence: Department of Herpetology, California Academy of Sciences, 875 Howard Street, San Francisco, Califor- nia 94103. jcadle@calacademy.org. Bull. Mus. Comp. Zool., 158(5): 183-284, May, 2007 Species of Sibynomorphus known from Ecuador and Peru are reviewed. Sibynomorphus oligozonatus and S. petersi are reported from Peru for the first time. Study of existing collections considerably amplifies understanding of the geographic ranges and character variation within S. oligozonatus, S. oneilli, S. petersi, and S. williamsi. Sibynomorphus vagrans is known only from its type locality (Bellavista, Cajamarca De- partment, Peru), and S. vagus (type locality unknown) is known only from near the single historically re- ported locality for the species (Huancabamba, Piura Department, Peru). Sibynomorphus oligozonatus (four specimens known) is distributed in southwest- ern Ecuador (Azuay and Loja Provinces) and north- ern Peru (Piura Department). Sibynomorphus oneilli is distributed in the Cordillera Oriental and Cordil- lera Occidental of northern Peru from southern An- cash Department to southern Cajamarca and Ama- zonas Departments. Available specimens of S. petersi extend the known range from Azuay and Loja Prov- inces in southwestern Ecuador, along the western An- dean slopes to southern Ancash Department, Peru (Pacific versant); a single specimen is also known from the upper Rio Chotano in central Cajamarca Department, thus documenting that this species oc- curs on the Amazonian versant. Sibynomorphus wil- liamsi is known from central Peru in Lima and An- cash Departments from near sea level to at least 2,900 m in the Andes (and perhaps as high as 3,600 m)—an elevational range unmatched in any other species of the genus. Two species (S. vagrans and S. vagus ) are restricted to the Amazonian versant, two are restricted to the Pacific versant (S. oligozonatus and S. williamsi), and two are found on both versants (S. oneilli and S. petersi). A key to the species of Sibynomorphus in Peru and Ecuador is provided. Hemipenes are described for Sibynomorphus oli- gozonatus, S. petersi, S. vagrans, S. vagus, and S. wil- liamsi. Their organs are similar to hemipenes of other Dipsadini in having a well-defined capitulum orna- mented with papillate calyces, several rows of large spines encircling the organ proximal to the capitulum, an exceptionally large basal nude pocket, and the di- 183 184 vision of the sulcus spermaticus within the capitulum. In Sibynomorphus and in some other Dipsadini the branches of the sulcus spermaticus are centrolineal, but their tips pass somewhat to the lateral surface of the hemipenial lobe(s). The exceptional size of the nude pocket might be a synapomorphy of Dipsadini but further comparative studies are needed, espe- cially because the pocket has probably been over- looked in many species. The natural history of all northern species of Si- bynomorphus is summarized, but very little is known. Most localities are in relatively dry to arid habitats, such as dry deciduous forests or thorn forests, on the western Andean slopes or inter-Andean valleys. A few localities for S. petersi in northern Peru are in areas of mesic to humid forests, although specimens are from forest edge or ecotones in this area. Sibyno- morphus williamsi is characteristic of the lomas for- mations on the central Peruvian coast, but it also oc- curs on the Andean slopes at higher elevations. Sibynomorphus shares with three other genera of Neotropical Dipsadinae (Dipsas, Sibon, Tropidodip- sas) a behavioral propensity and associated morpho- logical characteristics for consuming gastropods. In accordance with some previous authors, I suggest that the name Dipsadini be applied inclusively to this clade of four genera of Neotropical molluscivorous colubrids, notwithstanding previous arguments for ex- cluding Tropidodipsas. Several external characters in- dicate that Sibynomorphus is most closely related to Dipsas within Dipsadini. Nonetheless, the monophyly of Sibynomorphus has not been demonstrated and the possibility remains that Dipsas is paraphyletic with respect to Sibynomorphus, which itself is con- ceivably polyphyletic. The peculiar distribution of Si- bynomorphus is somewhat similar to other compo- nents of the herpetofauna of southern Ecuador and northern Peru (Andes, Pacific versant, and coast). However, Sibynomorphus as presently understood lacks some geographic components (e.g., Amazonia, Guayana) associated with these other taxa. Some investigations have suggested that Sibyno- morphus be synonymized with Dipsas, but analytical issues such as taxon and character sampling are prob- lematic with previous studies. None of the northern species of Sibynomorphus have been included in broader comparative studies. I advocate a broader survey of nominal taxa of Dipsadini before making further changes in the generic nomenclature in this group. Some outstanding questions concerning the systematics and natural history of Sibynomorphus are outlined. Especially needed are more detailed studies of microhabitat occurrence and behavior of all spe- cies and field studies to further delineate their dis- tributions. The significance of exceptional color pat- tern variation that occurs within S. oneilli, S. petersi, and S. vagus needs to be explored, perhaps in con- junction with their role as potential mimics of sym- patric species of Bothrops (Viperidae). Similarly, the extensive variation in scutellation that occurs gener- Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 ally in many Dipsadini needs functional, mechanistic, and evolutionary explanation. RESUMEN. Sibynomorphus, un grupo de aproxima- damente una docena de especies de colubridos sud- americanos que comen gasterépodos, tiene una dis- tribucién extrafia. Seis especies occuren en el norte del Pert y el Ecuador sudoccidental y los demas son distribuidas al sur de la cuenca Amazonica en Brazil, Bolivia, Paraguay, Uruguay, y Argentina. Se revisan las especies de Sibynomorphus de Ecuador y Pert. Se reportan para la primera vez S. oligozonatus y S. petersi del Pert. El estudio de colecciones que existe aumenta considerablemente el conocimiento de los rangos geograficos y la variacién de cardcteres de S. oligozonatus, S. oneilli, S. petersi, y S. williamsi. To- davia se conoce S. vagrans solamente de su localidad tipica (Bellavista, Departamento de Cajamarca, Pert), y se conoce S. vagus (localidad tipica descon- ocida) solamente cercana a la tinica localidad hist6- rica para la especie (Huancabamba, Departamento de Piura, Pert). Sibynomorphus oligozonatus (cono- cido de cuatro especimenes) se distribuido en el sur- oeste de Ecuador (provincias Azuay y Loja) y el norte de Pert (departamento de Piura). Sibynomorphus oneilli se distribuido en la Cordillera Oriental y la Cordillera Occidental del norte del Perti desde el sur del departamento de Ancash hasta el sur de los de- partamentos Cajamarca y Amazonas. Los especi- menes disponibles de S. petersi se extiende el rango conocido desde las provincias Azuay y Loja en el sur- oeste de Ecuador, a lo largo de las vertientes andinas occidentales hasta el sur del departamento de An- cash, Pert (vertiente Pacifico); un espécimen tnico se conoce también del alto Rio Chotano en el centro del departamento de Cajamarca, que documenta que esta especie también se encuentra en la vertiente Amazénica. Sibynomorphus williamsi se conoce del Perti central en los departamentos Lima y Ancash desde cerca del nivel del mar hasta a menos 2,900 m en los Andes (y quizds tan alto como 3,600 m)—un rango elevational incomparable en algiin otra especie del género. Dos especies (S. vagrans y S. vagus) son restringidas a la vertiente Amazdnica, dos especies son restringidas a la vertiente Pacifica (S. oligozonatus y S. williamsi), y dos especies se encuentran en las dos vertientes (S. oneilli y S. petersi). Se presenta un clave para las especies de Sibynomorphus en Pert y Ecuador. Se describe los hemipenes de Sibynomorphus oli- gozonatus, S. petersi, S. vagrans, S. vagus, y S. wil- liamsi. Sus 6rganos son similar a los hemipenes de otros Dipsadini al tener un capitulo bien definido y ornamentada con célices que llevan papilas, una serie de espinas grandes que rodean el 6rgano proximal al capitulo, un excepcionalmente grande bolsillo des- nudo basal, y un surco espermatico que divide dentro del capitulo. En Sibynomorphus y algunas otras es- pecies de Dipsadini las ramas del surco espermatico son centrolineal pero sus puntas pasan un tanto al lado lateral del l6bulo(s) del hemipene. El tamafo SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle excepcional del bolsillo desnudo puede ser una sin- apomorfia de Dipsadini pero se necesitan mas estu- dios comparativos, especialmente dado que el bolsillo se ha pasado por alto en muchas especies. Se resumen la historia natural de todas especies septentrionales de Sibynomorphus pero se conoce muy poca de estas especies. La mayoria de las local- idades son en habitats relativamente secos o dridos, tal como bosques caducifolios o bosques espinales, sobre las vertientes occidentales andinas o en valles interandinos. Unas pocas localidades para S. petersi en el norte del Peri son en dreas de bosques ht- medos o muy htimedos, aunque los ejemplares son del borde de los bosques o los ecotonos en esta re- gion. Sibynomorphus williamsi es caracteristico de las lomas en la costa central del Peri pero también se ocurre en las vertientes andinos a elevaciones mas altas. Sibynomorphus comparte con tres otros géneros neotropicales de Dipsadinae (Dipsas, Sibon, y Tro- pidodipsas) una propension conductual y caracteris- ticas morfolégicas asociadas para consumir gaster6- podos. De acuerdo con algunos autores anteriores, sugero que el nombre Dipsadini sea aplicado inclu- sivamente a este clado de cuatro géneros de colubri- dos neotropicales moluscivoros, no obstante razones anteriores para excluir Tropidodipsas. Varias carac- teristicas externales indican que Sibynomorphus es relacionado mas cercano a Dipsas entre los Dipsadi- ni. Sin embargo, la monofilia de Sibynomorphus no se ha sido demostrado y queda la posibilidad que Dipsas es parafilético respecto a Sibynomorphus, que si mismo puede ser polifilético. La distribucién ex- traiia de Sibynomorphus es algo similar a otros com- ponentes de la herpetofauna del sur de Ecuador y el norte del Perti (Andes, vertientes Pacifficas, y la cos- ta). Sin embargo, la distribucién de Sibynomorphus, en cuanto al entendimiento actual, carece de algunos componentes geograficos (p.e., Amazonia, Guayana) asociados con estos otros taxones. A pesar de algunos estudios que sugeren que Si- bynomorphus pase al sinonimia de Dipsas, algunos asuntos analiticos, inclusive cuestiones de muestreo de caracteres y taxones, son problematicos con estu- dios anteriores. Ninguno de las especies septentrion- ales de Sibynomorphus han sido incluidos en estudios comparativos amplios. Recomendo un estudio mas amplio de especies nominales de Dipsadini antes que iniciar mas cambios en la nomenclatura genérica de este grupo. Se explica algunas cuestiones pendientes acerca de la sistematica y la historia natural de Si- bynomorphus. Especialmente se necesitan estudios mas detallados de los microhabitas y comportamiento de todas especies y estudios de campo para refinar sus distribuciones geogrdficas. Se necesita explorar la importancia de la variacién excepcional de patrones de coloracién en S. oneilli, S. petersi, y S. vagus, tal vez conjunto con su rol como mimicos potenciales de especies simpatricas de Bothrops (Viperidae). Igual- mente, la variacién extensa en escamaci6n que ocurre 185 generalmente en Dipsadini necesita explicacién en términos funcionales, mecanicistos, y evolutivos. INTRODUCTION Recent investigations clarified the sys- tematics and distributions of several spe- cies of snakes of the genus Dipsas (Colu- bridae: Dipsadinae: Dipsadini*) in Panama and northern South America (Cadle, 2005; Cadle and Myers, 2003; Fernandes et al., 2002). Nonetheless, despite Peters’ com- prehensive review of Dipsadini (Peters, 1960), much additional work is needed on the systematics of this group. This is no- where more apparent than in the endemic South American genus Sibynomorphus, which comprises about a dozen currently recognized species (Franco et al., 2002, 2003; Hoge et al., “1978/1979” [1981]; Pe- ters and Orejas-Miranda, 1970; Scrocchi et al., 1993). Sibynomorphus has a peculiar distribu- tion. Six or seven species are distributed south of the Amazon in Argentina, Brazil, Paraguay, Uruguay, and Bolivia.* The other six species are found along the coast of central Peru; the western Andean foothills of northern Peru and southwestern Ecua- dor; or the Andes and inter-Andean valleys of the Rio Maranén, the Rio Santa, and their tributaries in northern Peru (Ama- zonian and Pacific versants, respectively). Thus, the range of the genus is disjunct > Dipsadini is here used for snakes of the genera Dipsas, Sibon, Tropidodipsas, and Sibynomorphus, essentially equivalent to the Dipsadinae of Peters (1960) with the addition of Tropidodipsas. See later herein: Sibynomorphus and the Dipsadini: Directions for Further Research. > These include Sibynomorphus inaequifasciatus, S. lavillai, S. mikanii, S. neuwiedii, S. turgidus, and S. ventrimaculatus (Hoge et al., “1978/1979” [1981]; Peters, 1960; Scrocchi et al., 1993). Additionally, S. garmani (Cope), previously considered a synonym of S. mikanii, was apparently resurrected in an unpub- lished thesis that I have not seen (Franco, 1994): the name appears in two online abstracts (Franco et al., 2002, 2003). Sibynomorphus inaequifasciatus Du- méril, Bibron, and Duméril, is known only from the original description. The holotype, said to be “doubt- fully from Brazil but certainly from South America,” has not been located (Peters, 1960: 146-148). 186 and the northern and southern species oc- cupy distinct physiographic and climatic zones. Scrocchi et al. (1993) briefly re- viewed the species of Sibynomorphus from southern South America, but species from Peru and Ecuador remain poorly known and have not received comprehen- sive review because most were described subsequent to Peters’ (1960) work on the group. The lack of a modern review has led to inaccurate identifications of muse- um material and misunderstandings con- cerning their distributions. As recently as the 1960s, when the sys- tematics of Sibynomorphus was last re- viewed (Peters, 1960; Peters and Orejas- Miranda, 1970), the genus was unreported from Ecuador, and only S. vagus (Jan, 1863) and S. vagrans (Dunn, 1923) were known from Peru (but see later comments on the name Leptognathus atypicus Cope).* Jan (1863) had described Leptog- nathus vagus from a specimen said to be from “Hong Kong.” This name has been applied to a species from near the town of Huancabamba in northern Peru (Piura Department), known primarily from a se- ries in the Museum of Comparative Zool- ogy collected by G. K. Noble (Dunn, 1923) and a later series from near the same locality (Rossman and Kizirian, 1993). Dunn (1923) described Pseudopa- reas vagrans from specimens collected by Noble in the inter-Andean valley of the upper Rio Maranon of northern Peru (Bel- lavista, Cajamarca Department), and the type series comprises the only known specimens. Both of these earliest de- scribed northern species of Sibynomor- phus are from inter-Andean valleys on the Amazonian versant of the Andes at the western limits of the Amazon basin. Since 1970, two additional species of Si- bynomorphus from Peru and two from ‘Peters (1960: 57-58) synonymized Sibynomor- phus macedoi Prado and Hoge (1947; type locality: Pucallpa, Ucayali Department, Peru) with Dipsas ca- tesbyi, a conclusion I accept without having seen the type : Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 southern Ecuador have been described: Sibynomorphus williamsi Carrillo de Es- pinoza (1974) from coastal Lima Depart- ment, Peru (type locality: Jicamarca); S. oneilli Rossman and Thomas (1979) from Amazonas Department, Peru (type locali- ty: between Balsas and Abra Chanchillo); S. oligozonatus Orcés and Almendariz (1989) from Azuay Province, Ecuador (type locality: Zhila); and S. petersi Orcés and Almendariz (1989) from Azuay and Loja Provinces, Ecuador (type locality: Zhila, Azuay Province). Few specimens have been reported for most of the north- ern species of Sibynormorphus. Sibyno- morphus oligozonatus and S. oneilli were described only on the basis of their holo- types and until now S. oligozonatus has been known only from the type. Rossman and Kizirian (1993) reported two addition- al specimens of S. oneilli from Cajamarca Department, Peru, and additional speci- mens of S. vagus from near the previously known locality. Sibynomorphus petersi has been known only from three specimens re- ported in the original description. Of the species of Sibynomorphus described since 1970, three (S. williamsi, S. oligozonatus, and S. petersi) have heretofore been known only from the Pacific versant, and S. oneilli only from the Amazonian versant. Even considering the recently described species, it is clear that the species limits and distributions of Sibynomorphus in Ec- uador and Peru are incompletely under- stood. As part of biological surveys in northern Peru (Cadle, 1989, 1991, 1998, 2005; Cadle and Chuna, 1995; Cadle and McDiarmid, 1990; Cadle and Myers, 2003), I obtained a few specimens of Si- bynomorphus, prompting me to review the species known from Peru and Ecuador. The results of that review form the basis of this report. Although the accumulated material im- proves our understanding of variation and the distributions of some taxa, some out- standing problems remain. Nonetheless, a comprehensive review of these species is needed to facilitate future systematic work SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle on this group. In this report, I document the occurrence of S. oligozonatus and S. petersi in Peru and the presence of S. pe- tersi and S. oneilli on both Amazonian and Pacific versants, and I amplify knowledge of variation in all species. Scrocchi et al. (1993) provided a brief review of the southern South American species and lit- erature subsequent to Peters (1960, 1965). METHODS General Methodology and Scale Termi- nology. This review is based on study of most specimens of Sibynomorphus from Ecuador and Peru in collections, including holotypes of all species except S. petersi and S. vagus. My own field experience with the genus is limited to a few speci- mens from northern Peru, here referred to S. petersi. Systematic methods and characters used herein follow previous studies (e.g., Cadle, 1996, 2005; Cadle and Myers, 2003). I used the Dowling (1951) method for ven- tral counts (beginning the count with the first plate bordered on each side by the first dorsal row), but for most specimens, I also recorded preventral scales (unpaired plates anterior to the ventrals that are wid- er than long) for reasons detailed by Myers (1974: 37; 2003: 6-8). Scale terminology follows Peters (1960), but amplification of the scoring of some scales is useful. Scales on the ventral surface of the head are highly variable in Sibynomorphus and other dipsadines. One or two pairs of in- fralabials are in contact posterior to the mental scale. Posterior to these contacting infralabials, I counted as chin shields those scales along the midline (no distinct men- tal groove in Sibynomorphus) that were about equal in length and width consid- ering each scale individually. These scales are squarish or polygonal and occur in pairs with the occasional exception of a posterior shield that is unpaired (resulting in a chin shield count of 3.5 pairs, for ex- ample). Another one to three pairs of gu- lar scales were often present posterior to the chin shields; gular scales are distin- Si guished from the chin shields because each gular scale is wider than long. Gular scales were not scored for this study. The gular scales are followed by the unpaired preventrals (when present). For the Dipsas oreas group (D. oreas, D. elegans, and D. ellipsifera) Cadle (2005: 71-73, fig. 1) described patterns of covari- ation between the shape of the loreal scale and whether the preocular was fused with the prefrontal or not. These scales are more uniform in species of Sibynomor- phus, and a few specimens were interme- diate between some of the six “loreal pat- tern” states defined by Cadle (2005). As in Dipsas, the loreal in Sibynomorphus was either squarish or polygonal, or much lon- ger than tall, and the preocular was fused or not with the prefrontal. In addition, some specimens of Sibynomorphus have other patterns, such as multiple preocu- lars, which were not observed in the D. oreas group; these patterns are discussed in the species accounts. Because of great variability in scutella- tion and color patterns in Sibynomorphus, I present detailed discussion of individual specimens where pertinent to facilitate fu- ture work on Sibynomorphus. For testing sexual dimorphism in ventral and subcau- dal counts and color pattern characteris- tics, I used Student’s t test on the sample means after testing for homogeneity of variances (Sokal and Rohlf, 1981). When variances were unequal (only in the case of color pattern characters for Sibynomor- phus petersi), | used an approximation to the ¢ test for two samples described by So- kal and Rohlf (1981: 408—412). Sexual di- morphism was also evident in relative tail lengths. However, because relative tail lengths are ratios (tail length/total length), the assumptions of the ¢ test are unlikely to be met without tranforming the data, a process not undertaken here. However, the frequency distributions of relative tail lengths between the sexes of species of Si- bynomorphus considered herein over- lapped minimally or not at all. Snout—vent lengths (SVLs) and _ total 188 lengths were measured with a meter stick or a plastic ruler to the nearest millimeter. For a few comparisons, I used head di- mensions, which were measured with dial vernier calipers to the nearest 0.05 milli- meter. Head lengths were measured in two ways: (1) obliquely from the tip of the snout to the posterior tip of the mandible (HL—mandible) and (2) from the tip of the snout to the posterior end of the interpa- rietal suture (HL-—parietal). Eye diameter (ED) was measured with dial vernier cal- ipers under a dissecting microscope as the greatest horizontal dimension of the eye. Figure 3 was drawn with a camera lucida mounted in a dissecting microscope. Hemipenial Techniques and Descrip- tions. Everted hemipenes of Sibynomor- phus petersi and S. williamsi described herein are from field-everted organs. Hemipenes of S. oligozonatus, S. vagrans, and S. vagus were manually everted from the inverted organs of museum specimens according to methods detailed by Myers and Cadle (2003). Hemipenial terminolo- gy follows Myers (1974: 30-33), Myers and Campbell (1981), and Zaher (1999). Some remarks about the terminology of general orientation is useful because of some dif- ferences in the way inverted and everted organs are described. The position of in- verted hemipenes is fixed relative to the snake’s body and they can be described by standard anatomical terminology (e.g., dorsal/ventral, medial/lateral, proximal/dis- tal) for orientation with reference to the snake’s body. In addition, the number of subcaudal scales subtending portions of the inverted hemipenis is commonly used as a relative measure of proportions for in- verted hemipenes. On the other hand, everted hemipenes are frequently described once they have been separated from the snake’s body and subjected to further preparation—eversion in the case of manual eversions, inflation, injection with colored jelly, etc. (Myers and Cadle, 2003). In descriptions of evert- ed organs, structures on the hemipenis it- self provide the best markers for orienta- Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 tion. The sulcus spermaticus provides a convenient landmark and the terms sulcate (side with the sulcus) and asulcate (side opposite the sulcus) were defined by My- ers and Trueb (1967) to denote these sur- faces of the hemipenial body. In Sibyno- morphus and many other snakes the hemi- penial body bears a nude pocket approxi- mately midway between the sulcate and asulcate surfaces, a position referred to as lateral (e.g., Cadle, 2005; Cadle and My- ers, 2003; and herein).® In this context, lat- eral is used with respect to the orientation provided by the sulcate/asulcate sides of the hemipenial body only, and not with re- spect to the snake’s body. A laterally posi- tioned nude pocket thus has a sulcate lip or edge (toward the sulcate side) and an asulcate lip or edge (toward the asulcate side). Terms such as proximal (toward the base of the organ) and distal (toward the apex) are still useful in describing everted hemipenes because their meaning is the same with respect to both inverted or everted organs. Localities and Geographical Data. Ref- erence to the “Rio Zafia Study Site” in the account for Sibynomorphus petersi refers toa locality in western Cajamarca Depart- ment that is better known herpetologically than other localities in northern Peru; see Cadle (2005: 70—71) for discussion and ad- ditional references. Elevations for my col- 5 Current use of the terms sulcate, asulcate, and lateral differs from definitions in Dowling and Sav- age’s (1960) classic work. Dowling and Savage used the terms sulcate and asulcate to refer to the two primary tissue layers of the hemipenis: the sulcate layer referred to the entire external surface of the everted organ, which bears the sulcus spermaticus on one side; the asulcate layer referred to the internal tissue layer of the everted organ (Dowling and Sav- age, 1960: 19 and pl. I). Dowling and Savage (1960: 21) used the term lateral to refer to what is now called the asulcate surface of the organ and medial for what is now referred to as the sulcate surface. Myers and Trueb (1967) redefined sulcate and asul- cate to their current usage because those terms seem most useful and least confusing as descriptors for the gross topology of hemipenes (the usual mode of study), rather than as names of tissue layers that are best studied in histological section. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR ° Cadle lecting localities were determined in the field with a Thommen© altimeter cali- brated in meters, sometimes in conjunc- tion with topographic maps. Coordinates and elevations for other localities, unless otherwise stated, were derived from Payn- ter (1993); Stephens and Traylor (1983); Peruvian departmental maps published by the Instituto Geografico Militar (IGM; now the Instituto Geografico Nacional), Lima; or online versions of the gazetteers of the U.S. Board on Geographic Names at the GEOnet© Names Server: http:// earth-info.nga. mil/gns/html/index.html. Stiglich (1922) is indispensable in locating obscure Peruvian localities, such as small haciendas, and I have also found occasion- al use for the Global Gazetteer (version 2.1, 26 January 2006) at www-fallingrain. com/world. Bracketed data in the list of specimens examined are inferences not supplied with the original collections data. Museum abbreviations used in the text are identified at the beginning of the Speci- mens Examined and Locality Records. Geographical discussions follow Simp- son (1975) for the names of the principal chains of the Andes in northern Peru. There is little cause for confusion except that some references and atlases (e.g., Hopkins, 1997) more finely divide the no- menclature of some of the ranges. In par- ticular, I follow Simpson (1975) in using the name “Cordillera Oriental” for all of the Andean ranges lying east of the inter- Andean portion of the Rio Marafion. Oth- er authors use that term for the relatively low ridges east of the Rio Huallaga and designate the higher mountains between the Rio Marafion and Rio Huallaga as “Cordillera Central” (see Fig. 7 and Simp- son, 1975: 276). As used herein, the Cor- dillera Blanca, Cordillera Negra, and Cor- dillera de Huancabamba are parts of the Cordillera Occidental. KEY TO SPECIES OF SIBYNOMORPHUS IN ECUADOR AND PERU Because of the variability of color pat- terns and scale characters within Sibyno- 189 morphus, individual specimens must be identified by a combination of characters and with careful attention to sexual differ- ences. The six northern species of Siby- nomorphus are rare or absent in most col- lections, and accurately identified compar- ative material is generally unavailable. As a consequence, misidentification of muse- um specimens is frequent. The most com- mon misidentifications of northern Siby- nomorphus in collections that I surveyed involved S. petersi, S. vagus, and S. oneilli (which is not saying much because S. oli- gozonatus has been known only from the holotype and S. vagrans only from the type series; S. williamsi is seemingly ab- sent from non-Peruvian collections). Ec- uadorian specimens of S. petersi were commonly misidentified as Dipsas varie- gata (following Peters, 1960) or, in older collections, as Dipsas oreas (see Cadle, 2005; Cadle and Myers, 2003, for discus- sion), whereas Peruvian specimens or &. petersi and S. oneilli were commonly mis- identified as the better known S. vagus. A previous key to all species of Siby- nomorphus (Scrocchi et al., 1993) was ad- equate on the basis of variation within spe- cies known at the time. However, S. oli- gozonatus and S. oneilli were known only from their holotypes, and this report doc- uments additional variation within all spe- cies that was not available to Scrocchi et al. (1993). For example, in their key, color pattern characteristics (with or without de- finitive bands, number of dorsal bands) were critical to differentiating some spe- cies (S. petersi, S. oneilli, S. vagus), but on the basis of the new material examined for this report, I infer extremely variable color patterns within these species (e.g., “defin- itive bands” present in some individuals of S. petersi and S. vagus and individuals without bands in S. oneilli). Indeed, the variation of color patterns and scutellation within most of the northern species of Si- bynomorphus and overlap of meristic characters between species made the con- struction of a workable key difficult. I have not studied species of Sibynomorphus that 190 occur south of the Amazon basin in any detail, but the following key should permit identification of species known from Peru and Ecuador. Considering present under- standing of variation within these species and their geographic distributions, not to mention the possibility of new species oc- curring in this area, the key should be used in conjunction with detailed descriptions and comparisons of specimens when pos- sible. A few comments concerning the pres- ence or absence of a separate preocular used in the first couplet of the key should be made. The key considers only the “nor- mal” conditions of the preocular(s) in Si- bynomorphus—that is, cases in which a single preocular superior to the loreal is present (Cadle 2005; Peters, 1960: 26; Figs. 11, 15, 23 herein), cases in which this preocular is fused with the prefrontal scale and a separate preocular is absent (see Fig. 3), or cases in which more than one preoc- ular is present, in which case the loreal is separated from the eye by the inferior preocular(s). Less commonly, an irregular preocular can be formed by, for example, a suture across the posterior corner of the loreal. In the key, these irregular condi- tions would not be considered in deter- mining presence or absence of a preocular. That said, I should enter a caveat: the presence or absence of a separate preoc- ular seems to be highly variable intraspe- cifically within some species of Dipsas (e.g., Cadle, 2005; Cadle and Myers, 2003). Although Sibynomorphus seems less variable in this character than Dipsas, some of the species in the key are repre- sented by few specimens, and this char- acter could prove more variable once sam- ple sizes are larger. For this reason, use of the key in conjunction with the detailed discussions and illustrations in the species accounts will provide more accurate iden- tifications. To facilitate the identification of a spec- imen in hand, Table 1 provides summaries of systematic characters for all six species of Sibynomorphus occurring in Ecuador Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 or Peru. The summary distributions in the key are documented herein. KEY 1. Supralabials 6 or 7, with 34 or 4-5 touching the eye. No separate preocular (preocular fused with prefrontal). Venter usually only lightly spotted with dark pigment, if at all, or dusky and without discrete spots. Dis- crete crossbands on body present at least anteriorly, and anteriorly at least twice the width of the pale interspaces 2, Supralabials 8 or more, with 4-5 touching the eye (rarely fewer than 8). One or more preoculars usually present (some individ- uals of S. oneilli lack separate preoculars). Venter usually heavily pigmented, often with half-moon—shaped or squarish spots. Distinct dorsal crossbands present or not, wider than interspaces or not ___.--. 3 2. Distinct bold crossbands present only on an- terior half of body, broken into series of paler vertebral and lateral blotches poste- riorly; dorsal blotches on posterior body distinctly narrower than pale interspaces. Head only slightly broader than neck. Known variation in ventrals: 177 (one fe- male), 145 (two males), and 163 (one Tale) ae ee Sibynomorphus oligozonatus Orcés and Almendariz (Andes of Azuay and Loja Provinces, southwestern Ecuador, Pacific versant; one specimen without specific locality from Piura Department, Peru) Distinct bold crossbands present the whole length of the body (often broken into mid- dorsal and offset lateral blotches posteri- orly, but these maintain their boldness); dorsal blotches on posterior body equiva- lent to, or (usually) slightly wider than, pale interspaces. Head distinctly broader than neck. Known variation in ventrals: 181-188 (females), 173-182 (males) ----- shh Sn ss OY GREAT oid Sibynomorphus williamsi Carrillo de Espinoza (coastal lomas formations and Andean foothills and slopes in Lima and Ancash Departments, Peru) 3. Anterior body with 5-10 discrete crossbands that are much broader than the pale in- terspaces (bands encompass 5-9 dorsal rows middorsally, 3-6 rows laterally, ex- cluding the nape band, which is always wider than any other bands; interspaces encompass 0.5—2 scale rows middorsally). Top of head very ornately patterned, in- cluding a relatively broad irregular dark band across the posterior edge of the pre- frontals (sometimes interrupted at one or more points). Ventrals 149-155 in males, d. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR ° Cadle 191 154-159 in females. Subcaudals 80-87 in Mees, “WDQ=W@ sia wSTMASS eee ee oS. ye Sibynomorphus vagrans Dunn (known only from the vicinity of Bella- vista, Cajamarca Department, in the in- ter-Andean valley of the upper Rio Ma- ranon of northern Peru; Amazonian ver- sant). Discrete crossbands on anterior body pres- ent or not; if present anteriorly, bands are only slightly wider than, or equivalent to, the pale interspaces (usually narrower than interspaces). Top of head ornately patterned or not, but without a discrete band across the posterior edge of the pre- COTA S.-i ef ee ne a mn ee eget oes 4 Ventrals <155 (known range 144-152) in males, <160 (known range 151—158) in fe- males. Subcaudals =65 in males, <60 in females. Maximum known size: 368 mm SVL (males), 326 mm SVL (females) —----- Ak eae Sibynomorphus vagus (Jan) (known only from the vicinity of Huan- cabamba, Piura Department, Peru; Am- azonian versant) Ventrals >150 in males, >160 in females. Subcaudals >60 in males, >55 in females. Maximum size to =470 mm SVL in males, =O HEU PS Vleet ale Smee ee 5 A discrete dark spot or irregular marking, usually outlined with a narrow pale edge, on each prefrontal; head generally with many dark markings. Dorsum usually with narrow, often zigzag, crossbands (bands may be reduced to an indistinct series of irregular vertebral and lateral spots). In- fralabials usually 10-12 (rarely 9). Ven- trals: males, 164-183; females, 173-188. Subcaudals: males, 75-87; females, 65—77 pes A Seal 7 ee ce aoe Sibynomorphus peterst Orcés and Almendariz (Azuay and Loja Provinces, southwestern Ecuador, south along the western slope of the Cordillera Occidental to southern Ancash Department, Peru; Rio Chotano valley, Amazonian versant, in Cajamarca Department, Peru) No discrete dark spots outlined with a pale edge on prefrontals; top of head unpattemmed or with a few dark markings, primarily on parietals and frontal. Discrete narrow blackish crossbands usually present the length of the body; anterior bands about equal to interspaces; posterior bands much narrower than interspaces (crossbands may be broken up on posterior body and some individuals lack bands, having in- stead an obscure network or reticulations). Infralabials 8 or 9. Ventrals: males, 152— 168; females, 163-177. Subcaudals: males, 62277 temaless 6-04.25 2) a ee Rossman and Thomas (Cordillera Oriental and Cordillera Oc- cidental from southern Ancash to south- ern Cajamarca and Amazonas Depart- ments, Peru, in the Rio Maranon and Rio Santa drainages) Sibynomorphus oligozonatus Orcés and Almendariz Figures 1-6 Dipsas mikanii oreas, part: Parker, 1938: 444 (a fe- male from the “Catamayo valley 30 km west of Loja,” here inferred to be BMNH 1935.11.3.108, which Parker thought possibly represented a “dis- tinct species” from the rest of the series) (misiden- tification). The rest of this series discussed by Park- er is identified here as S. petersi. Sibynomorphus oligozonatus Orcés and Almendariz (1989). Type locality: “Zhila (2,250 m) 79°17'26’W, 03°39'45"S [see footnote 19], parroquia Ona, can- ton Girén, Provincia Azuay |Ecuador].” Holotype, EPN 3612 (Fig. 1). Almendariz and Orcés (2004). Notes on the Holotype of Sibynomorphus oligozonatus With a few minor exceptions, my data for the holotype of Sibynomorphus oligo- zonatus (EPN 3612) are virtually identical to those reported in the original descrip- tion (Orcés and Almendariz, 1989) (see Table 2). The difference in ventral counts is probably due to different methods of counting (i.e., whether the preventrals were included in the ventral count or not—see Methods). My notes for the ho- lotype indicate that the “tail tip is damaged and may include one or two additional subcaudals.” The holotype (Fig. 1) has broad dark brown crossbands on the anterior half of the body, alternating side to side. Posteri- orly, these crossbands break up into nar- row zigzag lateral bars alternating with an irregular series of middorsal blotches. The dorsal ground color (interspaces) is whitish anteriorly, medium brown posteriorly. The venter is immaculate white posterior to the head, with fine dark speckling displaced laterally and increasing posteriorly. The only substantive difference be- tween my assessment of the holotype and that in the original description concerns the maturity of the specimen. Orcés and Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 192 (6 = N) (ND) (© = N) (Gin) (Ge wn) SoS * 919 OFT = CUE 79G = SEL 69'S + O'EL GOs = O19 99-9G 09-9¢ WG) LI=S9 VO-LG (G = N) (vy = N) (SL = N) (§ = N) (6 = N) GSS + 0°69 COG = SiG SVG + FES 99'S + O'T8 LOG z= ie) €L-LO ¢9-09 zL8-08 SSP NPC (Ol ND) (Gia) (9 = N) (Gite Ni) (= N)) 96° + O'FST G6G + OFST OL = OLGI SCize GS IS = OLIT SSI-ISI XG UG 6SI-FST SSI-ELT SLI-€9l (G = N) (G = N) (Gia ND) (UE = 1K) (OL = N) IO = OLA PVE = OSPF 99'T + FIST APG s GPO TE 66S = PO GSl-SLI CSI-PrI CGI-6FT SSI-F9T S9I-GST (Cl = A) SIRS CI-SI-GI Fi-Gay CT=GlsSil ClSSisGil (OL = N) SI-SI-ST Cl = WN) AT (6 = N) OL Ol = WN) Al (6 = N) OL (Ga ND ea (6 = N) OT (aN Cn (Gee) neal (Goa) sei (G = N) OT (aw NO Ril (G = N) FI (aN CE = IND) FAL (G= WN) SI (ew apalt (G =) Sl (Giga) (vy = N) (9 = N) (il = iN”) (G = N) 10'0 + 020 GO00'0 + 020 10'0 + ¥6'0 10'0 + G20 800'0 = 130 GS 0-610 1Z'0-06'0 9G 0-83'0 PS 0-060 GS 0-06'0 (G = N) (vy = N) (Sip) (S = N) (GEN) LOMOE=RSGuO GO00'0 = F2'0 10'°0 + LZ°0 GOO) == Soi F100 = F6'0 SG 0-160 PS O-SS'0 6 0-SZ'0 8G 0-GS'0 9 0-GS'0 (19S) STL (9GE) SOF (SGP) LSS (G6S) TLL (FEO) O6L (GET) S9G (S9E) SSP (L8€) SPS (609) S92 (OLVF) S69 $999 LLY Ge eNeeon (@ = N) SFI Giles teak G60 960 VE 0 (SGP) OSS (SPS) GOF ayewloy ITP sfepneoqns a[eulay aTPIN spe.Uo,, Saeos [es1od Y9e} AreypEXey oyewloy IPN yysue] [RIOVYsug] [rel a[eulay Sasser] Q[BUl ySOBIe'T (wu) (TAS) tASuey [POL ee a ee ee ee ee ISUUDYIN a a Ee a ae snapa suDispDa isuajad ‘ds snydiowouhqis 0 aeaeaea>aégsa ‘TISHUALAd “§ ANVW “ITIANO '§ “SALYNOZODITO ‘S HOH SINNOOOV AHL NI NHAID AYV AHAVHYOOAD YO NAWIONdS AM SAIMVININAS V.LVG TVNOILIGGY ‘(HTVNAA T “SHTIVN €) SAZIS ATIdINVS TIVWS AO ASOVOUd NAATID FAV SHONVY YALOVUVHO XINO “SALVNOZODITO ‘§ HOA ‘GS Tx AUV SNVAW ‘HLONAT LNAA-LOONS = "TAS ‘SNOILVAYHSAO YO SNAWIONdS 4O YAPNON = N “AULAWWASVY LNHNOFYA AO ASAVOA NOLLVAYASAO LNAGNAGACNI NV SV CAYOOS SVAA NAWIONdS HOVE AO AVaAH HHL AO ACIS HOVE ‘SATVOS GV4H YO “AYAd HO YOAVNOF NI YAOOO LVHL SQHdYHOWONKAIS AO SHIOAdS XIS AHL WOU VLVG OUNONOXVL GUVANVLS JO AUVWNNAS “TT AIPVE 1]]/12U0 snypuozosyo Single OxCNE ae) williamsi vagus Single (Naso) vagrans Single Sibynomorphus sp. CONTINUED. petersi Single TABLE lL. oneilli Single oligozonatus Single Anal scale Preoculars SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle 193 Nw NN © 4A tt ao asantano ih i al 6 ih ff i i fh ae low wl dl ZEZZ 2S SEB FF eee AIG GQEG N oO Ga) Sa! Ne) GANGS) COC) Oe Oo — = Pe ond ot CD) SEG = aN iy SS aa eae BES Se I Meh AOaa hele ev lll CU WwW BSS ae i i m 0 S285 24 26229595 ie Haq HO naa SHaqc av cys COS RS st -T- -- © © ECO OO oan aA 64 10 ww aS i telah wat lil 224444 if oo 0 2S 224 2S Ze 9 in In & 2242644 Nc =A AO 62) Sai ona OonNnN od wa | on A ee eB | LO CO) Ay a & r= C8) x Box oa aN om FoHOM COAN ATA No nas Yu TN Me th Ieee Te Za eS Ma AN AN 2255 255 S55 266 | Bue AoxdA Ol cd st CECI Hi ce ND OD DRoHA | on Hen HL | KKOO o—— — a I~ a SOCH Boaack ae Sh OE a Os ESS ees too ow Doo i he re Be rere rere ip bette ih ail 2222 2222222 YFHYHyooo 2222 eS arabe slea Tega No 4a HNodt aod BOUOVYYVV wamonan | on fl ee Ol ce PO | Or DMDMAS toa (se) Oa oO mee + tea all 216 (i i il I 22 e lit SS ae S oS te ae aon co + cD ey 2) Sy ao OK ia i?) See omen e = © ° SI S ~ a, 3S oy ® i = od = a2) ist) n re oO a oO 6 = as) re 4 6 is So aes e "39 4 S g 6 = Ss See “3 D = Oo 8 = Os oe © = @) or oO 3 = al a N om DN ao 194 CONTINUED TABLE l. Sibynomorphus sp. Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 williamsi 35-48 Al.9 = 3.7 vagrans vagus petersi Male: 29-35 SLY, 22 IY) oneilli onatus oligo y (Often irregular with well-defined bands) / Only anterior bod ( Number of dorsal (Generally only an- 38-63 when present; flecks or reticula- tions: bands, sometimes absent or fragmented terior body with well-defined bands) bands or blotches on body (N = 10) Female: 31-59 when present, are discrete only an- teriorly) 85200255, 0 (bands sometimes absent) ' The largest male is the holotype (Orcés and Almendariz, 1989), which was not examined in this study. The largest male examined by me was 657+ mm total length, 520 mm SVL. On the basis of measurements in the original description, the holotype has a shorter tail (20% of total length) than males I examined. 2 Peters (1960) reported a lower bound of 74 subcaudals for male S. vagrans, which could be a count obtained from one of the paratypes not examined for this study (see text footnote 25). > Extensively fragmented or fused tertiary temporals, making scoring somewhat arbitrary. Almendariz (1989) reported that the ho- lotype of Sibynomorphus oligozonatus was an adult male. However, several character- istics indicate that it is more likely a sub- adult (the hemipenis of the holotype is de- scribed subsequently in this account). Ca- dle (1996: 43-44), following a suggestion of Charles W. Myers, reviewed observa- tions indicating that the extent of miner- alization of hemipenial spines could be used to infer the sexual maturity of colu- brids. Immature male snakes have non- mineralized hemipenial spines and_ spi- nules, whereas in adults, these ornaments are mineralized, a characteristic that can be assessed with a fine needle (some ex- perience and comparisons of juveniles and adults is helpful);° the mineralization of spines in adult hemipenes compared with juveniles was also noted by Cope (1895: 189). A specific pattern of progressive mineralization of individual spines (tip to base) and on the hemipenis overall (prox- imal to distal) is apparent. The proximal edge of the capitulum of the hemipenis of the holotype of S. oligo- zonatus is fringed with nonmineralized or poorly mineralized spinules (rather than mineralized spines). In addition, spines on the hemipenial body proximal to the ca- pitulum are mineralized only at their tips. These characteristics suggest that the specimen is not fully mature, and the ho- lotype is also relatively small (SVL 265 mm, total length 351 mm) compared with the other referred males, AMNH 110587 (SVL 330 mm, total length 446 mm) and MUSM 2196 (SVL 348 mm, total length 462 mm). These characters are only sug- gestive that the holotype is not fully adult and more definitive internal examination ° If sufficient material, including juveniles to adults, is available, a useful technique for visualizing hemi- penial spine mineralization is a simple adaptation of methods for clearing and alizarin staining to demon- strate mineralized tissue such as bone (e.g., Dinger- kus and Uhler, 1977). Cadle (1996) used this ap- proach to examine the ontogeny of spine minerali- zation in Geodipsas. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle 195 TABLE 2. SCALE COUNTS, MEASUREMENTS, AND OTHER DATA FOR SPECIMENS OF SIBYNOMORPHUS OLIGOZON- ATUS. BILATERAL COUNTS ARE SEPARATED BY A SOLIDUS (LEFT/RIGHT). SVL = SNOUT-VENT LENGTH. BRACK- ETED DATA FOR THE HOLOTYPE ARE FROM THE ORIGINAL DESCRIPTION (ORCES AND ALMENDARIZ, 1989) AND ARE GIVEN ONLY IN CASES OF DISCREPANCY FROM MY OWN DATA (SEE TEXT). EPN 3612 AMNH BMNH MUSM Holotype 110587 1935.11.3.108 2192 Subadult (?) male Adult male Adult female Adult male Total Length Sol (26>)) 446 (330) 550 (428) 462 (348) (SVL) (mm) [BSA (27S) Tail length (mm) 86 [85] 116 22 114 Tail length/total length 0.24 0.26 O22 0.25 Maxillary teeth 12 1S LY 15 Dorsal scales LSM IFN L5NSFAMS LH=MS=MS Ventrals (+preventrals) ASE) 145 (41) bia (G2) 163 (preventrals not [148; preventrals recorded not recorded | Subcaudals 66 [67 | 68 66 68 Anal scale Single Single Single Single Preoculars (loreals) 0/0 (1/1) 0/0 (1/1) 0/0 (1/1) 0/0 (1/1) Postoculars 2/2 2/2, 3/3 2/2, Temporals Primary Dy 2/2, 2/2. 2/2. Secondary 3/3 3/3 3/4 3/3 Tertiary B18; 3/3 [fragmented] fie Supralabialss (touching 16\(8icr4)/ON) ce 4) (4 ee o)/7 (4 ceo) 6u(3 4/7 (3 a 4) GIG + 4)/? (3 + 4) eye) Infralabials 10/9 [10] 10/10 10/10 lee of reproductive organs was not undertak- en. New Material and Comparison of Specimens Referred to Sibynomorphus oligozonatus Sibynomorphus oligozonatus has previ- ously been known only from the holotype, EPN 3612. I refer three additional speci- mens to this taxon: AMNH 110587, BMNH 1935.11.3.108, and MUSM 2192. I was able to make side-by-side compari- sons of BMNH 1935.11.3.108 and the ho- lotype and subsequently made side-by-side comparisons of AMNH_ 110587 and BMNH 1935.11.3.108. Later I determined that MUSM 2192 should be referred to this species. Standard characteristics of each specimen are given in Table 2. BMNH_ 1935.11.3.108 (Fig. D) iS sin adult female from approximately 70 km SSW of the type locality. It was part of a Figure 1. Sibynomorphus oligozonatus. Holotype in dorsal : and ventral views (EPN 3612; Azuay Province, Ecuador). Sub- collection from southern Ecuador ob- adult male, 265 mm SVL. tained for the British Museum by Clodov- 196 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 2. Sibynomorphus oligozonatus. Dorsal and ventral views of BMNH 1935.11.3.108 (Loja Province, Ecuador). Female, 428 mm SVL. The specimen illustrates the typical pattern in the known specimens: broad anterior dorsal crossbands, fragmented posterior bands, and relatively unpatterned venter. eo Carrién (Parker, 1930a, 1930b, 1932, 1934, 1938). Parker (1938) referred sev- eral specimens in this collection to “Dipsas mikanii oreas” (= Dipsas oreas; Cadle, 2005). Rather peculiarly for a moderately sized collection from this region, none of the snakes that Parker referred to D. oreas appear to represent that species. Instead, they comprise the single specimen here identified as Sibynomorphus oligozonatus and a larger series I identify as S. petersi (see subsequent species account).’ Parker (1938: 444) did not use BMNH collection numbers, but he noted that one of the fe- males “differs from other specimens in having a smaller eye, a higher and narrow- er rostral, and the first labial in contact with the loreal; it may possibly represent a distinct species.” BMNH 1935.11.3.108 7 Although Dipsas oreas is known from the same region as Carrién’s collections (Cadle, 2005), the presence of Sibynomorphus in his collection and the apparent absence of D. oreas suggest that his collec- tions were from drier, more open habitats (as op- posed to humid forested habitats). Dipsas oreas is restricted to humid forests (Cadle, 2005), whereas species of Sibynomorphus in Ecuador and Peru gen- erally occur in subhumid to arid environments, such as dry deciduous forests or thorn forests, which pre- dominate in this region. matches the locality and characteristics given by Parker, including ventral and sub- caudal counts that are virtually identical to mine. AMNH 110587 is an adult male road- killed specimen collected by Charles W. Myers on 12 February 1974 approximately 25 km northwest of the type locality. MUSM 2192 is an adult male from “Piura, Peru” (department or city not specified, here assumed to refer to the department) collected by Otavio Ruiz with no addition- al data) MUSM 2192 is soft and in poor condition; its scale counts were deter- mined only with difficulty. (Another spec- imen possibly referable to S. oligozonatus, MUSM 2248, with locality “Piura” but no collector or date, is very soft and deterio- rated. Its color pattern was apparently sim- ilar to MUSM 2192 but its condition pre- cluded accurate scale counts, etc. It will not be considered further here.) Because so few specimens of Sibyno- morphus oligozonatus are known and they exhibit some differences in scutellation and color pattern, I herewith highlight fea- tures suggesting that these are conspecific and note some differences among them. Nonetheless, some characteristics of the SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle specimens make the interpretation of dif- ferences somewhat difficult with so few specimens. For example, the holotype is a subadult male and smaller than the other males, AMNH 110587 and MUSM 2192, whereas BMNH 1935.11.3.108 is a large female (Table 2). The head shape and body proportions of AMNH 110587 were difficult to assess because it is flattened, somewhat desiccated, and has a damaged head. More than likely there would be changes in shape associated with growth as well, rendering comparisons of shape char- acters (e.g., the distinctive head shape of BMNH 1935.11.3.108) between the small holotype and other specimens problemat- ic. Similarly, MUSM 2192 is soft, in poor condition, and has a damaged head. These factors made it difficult to assess the rela- tive contributions of size, sexual dimor- phism, and other sources of variation. Scutellation and Body Proportions. Two of the males, AMNH 110587 and the ho- lotype, are nearly identical in scutellation (Table 2). Especially significant are the very low ventral counts, approached only by a few specimens of Sibynomorphus va- grans among the other species of Ecuador or Peru (male S. vagrans have many more subcaudals than either AMNH 110587 or EPN 3612). On the other hand, MUSM 2192 has considerably more ventrals than either of the other two males. In contrast, the female, BMNH 1935.11.3.108, has considerably more ven- trals than two of the males (EPN 3612, AMNH 110587; Table 2). The difference is probably accounted for by sexual di- morphism and perhaps geographic varia- tion, but the disparity in ventral counts (32) is greater than has been reported for males and females of any other species of Sibynomorphus (see later comparisons herein; Scrocchi et al. 1993). The closest approach to this difference among the northern species of Sibynomorphus is in S. petersi, in which the minimum and maxi- mum ventral counts (male and female, re- spectively) differ by 24. Other differences among the three Ley specimens are found in the configuration of scales in the loreal region. In AMNH 110587 supralabials 2-4 touch the loreal scale (the first supralabial is well separated from the loreal by the posterior nasal scale), whereas supralabials 1-3 border the loreal in BMNH 1935.11.3.108 (Fig. Oy sotTa MUSM 2192 supralabials 2—3 broadly con- tact the loreal but on each side, the first supralabial also touches the loreal at a point. In the holotype, supralabials 2-3 border the loreal. In the AMNH speci- men, supralabials 4 and 5 border the eye, whereas supralabials 3 and 4 border the eye in the other specimens. Scales in the loreal region vary substantially within some other Dipsadini (Cadle, 2005: 71-73; Peters, 1960: 25), and the variations among the referred specimens of Sibyno- morphus oligozonatus do not seem ex- treme. The differences among the speci- mens in the relationships of the suprala- bials, loreal, and eye could reflect slightly different head shapes or proportions (see below). Differences among these speci- mens in other scutellation characters (Ta- ble 2) are typical of those observed in oth- er Dipsadini. BMNH 1935.11.3.108 has one unusual character of the dorsal head plates. Each supraocular is divided by a suture extend- ing from the posterolateral angles of the frontal to the dorsal midpoint of the eye (Fig. 3). These sutures are very regular, well formed, and bilaterally symmetrical (i.e., each supraocular is identically divid- ed). I am not aware of other colubrids with this variation, although Peters (1960: 25) reported fusions among some of the dorsal head plates (e.g., internasals, prefrontals) within Dipsas. BMNH 1935.11.3.108 also has more irregular scales in the temporal region than either AMNH 110587 or EPN 3612, but irregularities in this region are characteristic of many species of Dipsadi- ni. The three specimens differ somewhat in general body form and head shape, but the size differences among specimens and the general conditions of AMNH 110587 and 198 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 3. Head scales of Sibynomorphus oligozonatus (BMNH 1935.11.3.108) in left and right lateral views and in dorsal view. The lateral views indicate the blunt, short appearance of the snout (See also Fig. 4). The peculiar divided supraocular scales (shaded dorsal head plates) are unique to this specimen. MUSM 2192 make these comparisons prob- lematic. Judging from BMNH_ 1935.11.38. 108, adult Sibynomorphus oligozonatus have a peculiarly foreshortened and very blunt snout (Fig. 4). It is not possible to judge precisely the physiognomy of AMNH 110587, especially the head, because it has been flattened. It seems to have a somewhat longer and more acuminate snout than Figure 4. Sibynomorphus oligozonatus. Lateral view of head, BMNH 1935.11.3.108. BMNH 1935.11.3.108, but this could be en- tirely due to artificial deformation. The head shapes of BMNH 1935.11.3.108 and EPN 3612 are similar; the head of MUSM 2192 is damaged but appears to be particularly blunt. Some differences in head size seem ap- parent between the adult specimens of Sibynomorphus oligozonatus, AMNH 110587 (male), BMNH 1935.11.3.108 (fe- male), and MUSM 2192 (male), even ac- counting for distortion brought about by damage to the AMNH specimen. AMNH 110587 has a larger head in relation to SVL (4.0% or 3.7% by HL—mandible and HL-interparietal, respectively) than the BMNH specimen (3.5% and 2.6%, re- spectively) or MUSM 2192 (3.0% by HL— interparietal; lower jaws are damaged, pre- cluding accurate measurement of HL-— mandible). Sexual dimorphism in head length relative to SVL is widespread in SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR ° Cadle snakes—for example, in nearly half of the species and 43% of the genera in one tax- onomically diverse survey (Shine, 1991). However, Shine (1991) found that female snakes commonly had relatively larger heads in most genera (40 of 61 genera, omitting taxa without dimorphism). Thus, if the difference in head proportions be- tween the males and female S. oligozona- tus truly represents sexual dimorphism, it is of a relatively less common _ pattern among, snakes. Color Patterns. The color patterns of specimens referred to Sibynomorphus oli- Zozonan smane masini late n(Eigs slay) 2 510) 5©): Dorsally, all specimens have broad anterior crossbands that begin to fragment at about one-third to one-half the body length. Af- ter the point of fragmentation, a series of middorsal blotches or saddles tends to al- ternate with lateral vertical bars or blotch- es. In BMNH 1935.11.3.108 and MUSM 2192, the middorsal blotches and the lat- eral bars are larger and more regular than in the holotype or in AMNH 110587; the BMNH and MUSM specimens have more complete crossbands than either of the other specimens. The crossbands of all specimens end on the outer one-quarter of the ventral plates. In three specimens, there is a ten- dency, more pronounced in EPN 3612 and AMNH 110587 than in BMNH 1935.11.3.108 or MUSM 2192, for the an- terior crossbands to be offset (i.e., corre- sponding portions of the bands on either side are displaced middorsally). In the BMNH specimen, crossbands 3-5 are off- set, whereas in EPN 3612 and AMNH 110587, all anterior crossbands except the first one or two are offset. None of the complete crossbands in MUSM 2196 are offset. In AMNH 110587, BMNH 1935.11.3.108, and MUSM 2196, the cen- tral parts of the crossbands tend to be somewhat lighter than peripheral parts (i.e., bands appear dark-edged); this char- acter was not recorded for the holotype. The venter of all specimens has narrow irregular rectangles of dark pigment en- 9) croaching from the dorsum onto the edges of the ventral plates. Central parts of the ventral plates are speckled and _ flecked with tiny dark specks, which is much heavier in EPN 3612 and AMNH 110587, less so in BMNH 1935.11.3.108, and least in MUSM 2192. Pigmentation becomes denser on the posterior venter in all spec- imens, but to a lesser extent in BMNH 1935.11.3.108 and MUSM 2192 than in the others. Thus, BMNH 1935.11.3.108 and MUSM 2196 have relatively un- marked venters compared with the holo- type and AMNH 110587. (he top moti Leen ireacie sin sels)NIUN| Tet 1935.11.3.108 is pale yellowish brown with bold black irregular marks all over. Supra- and infralabials in this specimen are pale yellowish with black suture lines (Fig. 4). The dorsal head pattern of the holotype is similar to BMNH 1935.11.3.108, including a pair of irregular bold longitudinal streaks posteriorly and a bold spot on each pre- frontal scale. In AMNH 110587, the top of the head is grayish brown to medium brown without bold markings, except for an irregular spot on the anterior end of each parietal. Its markings are generally smaller and less distinct than in the holo- type or BMNH specimen. Some suture lines on the supralabials and infralabials in AMNH 110587 are grayish brown, and some other grayish brown stippling is pres- ent on the supralabials, but they are not boldly marked as in the holotype or the BMNH specimen. In general, the dark markings on the heads of EPN 3612 and BMNH 1935.11.3.108 are distinctly black- ish, whereas those in AMNH 110587 are grayish brown. MUSM 2192 has irregular spotting on top of the head, including a large spot at the anterior end of each pa- rietal. For several reasons, the identification of MUSM 2192 as Sibynomorphus oligozon- atus is the most problematic of the four specimens. Its ventral count is much high- er (163) than that of either of the other males (145), which brings it closer to the range of ventral counts in male S. williamsi 200 (173-182) (MUSM 2192 was previously identified in the MUSM catalogues as S. williamsi). These two species are other- wise very similar in scutellation (Table 1). The imprecise locality for MUSM 2192, “Piura” [Department], is intermediate be- tween the other nearest localities for S. oli- gozonatus (Catamayo Valley of Loja Prov- ince, Ecuador) and S. williamsi (southern Ancash Department, Peru; see Fig. 43), al- though it is closer to the former. Details of the banding pattern in MUSM 2192 compared with the other re- ferred specimens of S. oligozonatus and S. williamsi convincingly demonstrate that MUSM 2192 should be referred to S. oli- gozonatus. These similarities to S. oligo- zonatus include (contrasting state for S. williamsi in parentheses; see Figs. 40-42) (1) abrupt transition from regular dorsal crossbands anteriorly to fragmented cross- bands posteriorly (transition is gradual); (2) a narrow pale collar just behind the head, with the dark pigment of the first crossband not encroaching onto the head (pigment of first dark crossband extends up onto the head); (3) crossbands with dis- tinctly darkened edges (bands lack dis- tinctly darker edges); (4) crossbands brown (bands usually distinctly blackish in pre- servative, although this could be because of preservation or storage); (5) posterior crossbands usually distinctly narrower than interspaces (posterior crossbands as wide as, or wider, than interspaces); and (6) ven- ter relatively immaculate (venter often, but not universally, dusky or with a ladder-like dark pattern or with dark spots posterior- ly). Although the head shape of MUSM 2192 is difficult to assess because it is dam- aged, it seems to be relatively small and foreshortened, as in S. oligozonatus, com- pared with the longer, more acuminate head in S. williamsi. Considering the highly variable colora- tion and scutellation in some other species of Sibynomorphus, the differences among the four specimens I refer to S. oligozon- atus do not seem extreme, although the disparity in ventral counts (Table 2) Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 between MUSM 2192 and the other males and between the female (BMNH 1935.11.3.108) and two of the males (EPN 3612, AMNH 110587) are remarkable. These specimens should be re-evaluated as additional material of this species ac- cumulates. The following account is based on the collective characteristics of these four specimens. Diagnosis Sibynomorphus oligozonatus is a gray to grayish brown snake with broad dark crossbands anteriorly (narrowing and _ be- coming broken into dorsal and _ lateral blotches posteriorly). Anterior crossbands are much broader than the interspaces and have regular, vertical edges and sometimes are offset. Posterior crossbands are usually much narrower than the interspaces. The venter is unmarked or has irregular dark speckling or spotting concentrated poste- riorly. The head is foreshortened and blunt. The species has a low number of supralabials (6 or 7) and maxillary teeth (12-15). Males of S. oligozonatus have a low number of ventrals (145 in two males, 163 in another) and relatively long tail (24-26% of total length); these characters in the single known female (177 and 22%, respectively), are typical of some other species of Sibynomorphus (see Table 1). Scrocchi et al. (1993) questioned wheth- er Sibynomorphus oligozonatus was diag- nosable from S. vagrans on the basis of the characters given in the original descrip- tion. However, these two species differ in head shape and proportions, as well as sev- eral scale characters. Sibynomorphus oli- gozonatus has a short head and blunt snout (Fig. 4), lacks preoculars in the known specimens (i.e., the preocular is fused with the prefrontal and only a loreal is present between the eye and posterior nasal), and has six or seven supralabials and fewer than 70 subcaudals. Sibynomor- phus vagrans has a more elongate head and more pointed snout (compare Fig. 4 with Figs. 30, 31), has a preocular in ad- dition to a loreal (occasionally 2 preoculars SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle and a loreal), and has eight to 10 supra- labials and more than 70 subcaudals (Table ils): Sibynomorphus oligozonatus differs in color pattern from S. oneilli (see Figs. 8— 10). Both species have dark crossbands that are broader anteriorly than posteriorly (some S. oneilli lack bands; see following species account). In S. oligozonatus, the anterior crossbands are twice or more as broad as the pale interspaces, whereas on the anterior body in S. oneilli, the inter- spaces are about as broad as, or broader than, the dark crossbands. Sibynomorphus oneilli has a bold pattern of dark markings in an irregular checkerboard pattern or paired spots on each ventral plate, whereas the venter of S. oligozonatus is relatively unmarked (AMNH 110587 does have ir- regular dark speckling all over the venter and some larger spots that tend to be con- centrated laterally, but this pattern does not resemble the heavily spotted venter of SONI asec ies), (0), ol?) Sibynomorphus petersi occasionally has relatively broad anterior crossbands remi- niscent of, but less regular than, those of S. oligozonatus. Sibynomorphus petersi has a preocular above the loreal (preocular absent in S. oligozonatus) and usually has eight supralabials (six or seven in S. oli- gozonatus). In addition, male S. petersi have more ventrals (164-183) than S. oli- gozonatus. Sibynomorphus peterst gener- ally has a strongly patterned venter (Fig. 13 and following) compared with the rel- atively less patterned venter of S. oligozon- atus. Sibynomorphus oligozonatus differs from S. vagus in color pattern: broad an- terior crossbands having regular, more or less vertical borders in S. oligozonatus ver- sus either nonbanded (a vague network or fine reticulations) or with narrow, poorly formed, and irregular crossbands in S. va- gus. Sibynomorphus vagus has fewer ven- trals in females, has a preocular scale, usu- ally has a single primary temporal, and has seven or eight supralabials. Sibynomorphus _ oligozonatus differs 201 from S. williamsi in having fewer ventrals (males: 145—163 vs. 173—182: females: 177 vs. 181-188), in head shape (oligozonatus: head slightly broader than neck, short and blunt; williamsi: head broader than neck, longer and acuminate), and in some as- pects of color pattern. In S. williamsi, the first dorsal crossband usually encroaches broadly onto the nuchal region (see Figs. 41, 42), whereas in S. oligozonatus, the first crossband is separated narrowly from the head by a distinct narrow pale collar (Fig. 4). The dorsal bands in S. williamsi are bold and distinct the whole length of the body and tail (even when fragmented into a dorsal and lateral series of blotches), whereas in S. oligozonatus, the anterior bands are much bolder than the posterior ones. In S. oligozonatus the dorsal blotch- es on the posterior body are slightly to much narrower than the pale interspaces; the posterior dorsal blotches in S. william- si are equivalent to or wider than the pale interspaces. The color pattern of the BMNH and AMNH specimens of Sibynomorphus oli- gozonatus is reminiscent of some speci- mens of Dipsas oreas, which perhaps led Parker (1938) to refer the BMNH speci- men to that species (as “Dipsas mikanii or- eas”). However, D. oreas usually has a pair of large oval dark irregular spots on the dorsoposterior portion (primarily parietal scales) of the head (Cadle, 2005: 101—108), a strongly compressed body, and a rela- tively much larger eye (compare Fig. 4 with Cadle, 2005, fig. 21). The relative eye diameter of the BMNH specimen of S. oli- gozonatus, the only adult with an undam- aged head, was 0.51% of SVL, whereas for 12 adult D. oreas, the eye averaged 0.73 + 0.049% of SVL (range: 0.64—0.78%). Males of S. oligozonatus can be distin- guished from D. oreas by having fewer ventrals (144—163 vs. 167-184 in D. oreas) and subcaudals (66—68 vs. 82-91 in D. or- eas). The female referred to S. oligozona- tus is similar to females of D. oreas except for a slightly lower subcaudal count (66 vs. 70-83) and the previously mentioned as- 202 pects of color pattern and body form. See Table 2 and Cadle (2005, table 1). Description Size and Scutellation. Tables 1 and 2 summarize taxonomic data for Sibynomor- phus oligozonatus. Largest male 446 mm total length (330 mm SVL). Largest female 550 mm total length (428 mm SVL). Body somewhat compressed. Tail 24—26% of to- tal length in three males, 22% in one fe- male. Ventrals 145 (two males) to 163 (third known male), 177 in a female; ven- trals preceded by one or two preventrals. Anal single. Subcaudals 66—68 in males, 66 in one female. Loreal and prefrontal bor- dering anterior edge of eye (no separate preocular) (Fig. 3); loreal longer than tall. Postoculars 2 or 3, primary temporals 2, secondary temporals 3 or 4, tertiary tem- porals (when discrete) 3. Supralabials most commonly 6 (3—4 bordering eye); other patterns 7 (3-4) or 7 (4-5). Infralabials 10 (9 on one side in the holotype). Chin shields in 2—3.5 pairs. Maxillary teeth 12— 15 GN = 2). As far as can be discerned with only four specimens, Sibynomorphus oligozonatus shows typical colubrid patterns of sexual dimorphism in size (female larger), ventral counts (female greater), and relative tail length (male longer). Despite the differ- ence in relative tail length, subcaudal counts do not differ between the males and the female. Statistical tests were not performed on these comparisons because only four specimens are known. Color in Life. Orcés and Almendariz (1989) described the color of the holotype as follows (paraphrased): “Diffuse dark brown spots on the top and sides of the head. Dark brown bands on the anterior part of the body four scales wide, narrow- ing to one scale wide on the posterior body. Interspaces anteriorly pale brown, becoming grayish at midbody. Venter whit- ish, finely speckled with brown posterior- ly.” Charles W. Myers annotated his field catalog for AMNH 110587 with the com- ment “venter white,” meaning simply that 9) Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 5. Sibynomorphus oligozonatus. Dorsal and ventral views of AMNH 110587 (Azuay Province, Ecuador). Male, 330 mm SVL. the venter lacked any wash colors such as yellow or orange, which are common on the venters of some colubrids (Charles W. Myers, personal communication); the ven- ter of AMNH 110587 does have some dark spotting. Color in Preservative. The coloration of the four specimens in preservative is very similar (Figs. 1, 2, 5, 6). The dorsal pattern consists of broad anterior crossbands (4— 10 dorsal rows wide) that progressively narrow posteriorly, fragmenting into mid- dorsal and lateral blotches by about mid- body. The crossbands are brown to black- ish brown with a narrow darker border. In- SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR ° Cadle terspaces are gray, grayish white, or light (yellowish) brown. The top of the head is pale grayish or brown with bold dark spots and markings in an irregular pattern. The venter is pale gray without bold markings, but can have dark speckling that increases in density posteriorly. The color tones of the crossbands in MUSM 2192 are some- what lighter brown (with reddish or chest- nut tones) than the other specimens, but this could be preservation artifact (? light- induced fading). The coloration in preser- vative for three specimens is as follows (notes taken 2003 for AMNH_ 110587: 2004 for the others): AMNH. 110587 (Fig. 5): Dorsal ground color on anterior third of body grayish to grayish white, be- coming grayish brown posteriorly. Approximately the anterior third of the body has bold blackish brown crossbands. The anterior five to six crossbands are offset middorsally, have vertical sides (i.e., they ap- pear as squares or vertical rectangles viewed lateral- ly), and tend to have somewhat paler centers than edges (central scales of the crossbands are mottled with dark brown and pale grayish brown, whereas pe- ripheral scales are solid blackish brown). Anterior crossbands are four to five scales wide. Posteriorly the crossbands fragment abruptly into irregular middor- sal and lateral blotches. The anterior dorsal cross- bands encroach onto the outer quarter of the ventral plates. Dorsal scales in the interspaces between cross- bands are finely flecked with minute dark brown specks. Interspaces anteriorly are 2—2.5 scale rows wide, increasing to 3 rows just before the point at which the dorsal crossbands fragment. Venter white, speckled, and irregularly spotted with dark brown, which increases posteriorly. Top of the head brown (grayish brown on parie- tals), heavily flecked and spotted irregularly with dark brown. A single larger irregular spot is present at the anterior edge of each parietal. An indistinct narrow (two scales wide) pale grayish nape collar is behind the mouth commissure, followed by the first dorsal crossband. BMNH 1935.11.3.108 (Figs. 2, 4): Dorsal ground color pale, somewhat yellowish, brown. Crossbands are medium brown with blackish edges <1 dorsal row wide. Under magnification their centers are flecked with minute pale specks, making their centers some- what lighter than the edges. The first neck band is about 6 to 6.5 scale rows wide. The next two cross- bands are 7 to 8 rows wide. Succeeding crossbands narrow to six, five, and four rows before breaking up into a series of dorsal saddles and lateral blotches or bars. All crossbands are complete middorsally and ex- tend ventrally to the outer edges of the ventral plates. On the posterior body, the lateral blotches are very 203 Figure 6. Sibynomorphus oligozonatus. Dorsal and ventral views of MUSM 2192 from an unknown locality in Piura De- partment, Peru. Male, 348 mm SVL. irregular in shape and poorly formed, whereas the dorsal saddles have a more regular shape. The venter is yellowish white with scattered dark markings, which become denser posteriorly and are mostly dis- posed on the lateral portions of the ventral plates. The top of head has bold, crisp (sharp edged) ir- regular blackish reticulations and spots in no distinct pattern. Supra- and infralabials are yellowish with bold black suture lines. Posterior to the region of bold head markings are a pair of scale rows relatively free of markings, forming a narrow pale nape collar; then follows the first crossband. MUSM 2192 (Fig. 6): Dorsal crossbands are pale, somewhat reddish brown, each with a narrow (<1 dorsal scale row wide) dark brown border. Interspac- es grayish. First neck crossband is 11 scale rows wide; then follow 11 to 12 discrete crossbands, progressive- ly narrowing from 7—9 rows wide anteriorly to 4 rows wide posteriorly. Crossbands extend ventrally to the outer edges of the ventral scutes. Pale interspaces are 1.5—2 scale rows wide. The venter is immaculate gray- ish with a few small punctations on the edges of the plates. The top of the head is pale grayish brown with darker irregular spots (large spots on anterior edges of parietals and on other head scales). Supralabials have a few dark lines, but are generally pale. Infra- labials and gular region are pale. Etymology. The specific epithet, oligo- zonatus, comes from the Greek words oli- 204 go- (few) and zona (belt); the suffix -atus indicates possession. It refers to the “few transverse bands, much less numerous than is usual in the genus” (Orcés and Al- mendariz, 1989: 63). Hemipenis The hemipenes of the holotype of Si- bynomorphus oligozonatus are partially everted, and I first report details observ- able on them. Then follows a complete de- scription of the everted organ of AMNH 110587. No differences were observed be- tween the portions of these two organs that could be compared. The hemipenis of the holotype (EPN 3612) is capitate and apparently slightly bi- lobed. The sulcus spermaticus divides within the capitulum, which is spinulate on the visible proximal portion; the spinules apparently correspond to the fringe of spines on the proximal edge of the capit- ulum of the everted organ of AMNH 110587 described below. These spinules in the holotype are nonmineralized or poorly mineralized. Proximal to the capitulum, the distal portion of the hemipenial body bears a battery of enlarged spines, which appear to be mineralized only at their tips. A nude pocket bordered on each side by a fleshy lobe is on the lateral side of the hemipenial body proximal to the above- mentioned battery of spines. The right hemipenis of AMNH 110587 was partially everted upon preservation. Full eversion was effected with the use of methods described by Myers and Cadle (2003). Although the organ is fully everted, it is not likely to be maximally expanded. Hence, the entire surface morphology of the hemipenis can be studied, but it is probably not as fully expanded as a fully everted fresh organ (see Myers and Cadle [2003] for discussion). In particular, the basal portion, which was everted before preservation, appears somewhat dehydrat- ed and not fully expanded; a portion of the asulcate side of the capitulum also appears incompletely expanded. Total length of the everted organ 15 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 mm, bilobed less than 1 mm. The relative proportions of the capitulum to base could not be assessed with accuracy because the base of the hemipenis is rather nonpliant and is probably not extended to its maxi- mal state. The hemipenis is fully capitate and slightly bilobed. The capitulum is en- tirely calyculate and calyces are ornament- ed with fingerlike papillae; some of the more proximal papillae might have min- eralized tips. A fringe of small mineralized spines encircles the proximal edge of the capitulum just distal to the overhang de- marcating the capitulum; these spines are relatively straight and blunt. The calyces cover all aspects of the capitulum, includ- ing the crotch and medial portions of the lobes. Proximal to the capitulum, a battery of enlarged hooked spines about three spines across encircles the entire organ. Proximal to the spinose midsection, the hemipenis bears scattered minute spinules, which ap- pear to be denser toward the sulcate side of the organ. A very long nude pocket is positioned laterally on the organ and ex- tends from the very base of the organ to the battery of spines. The pocket is bor- dered on each side by a prominent lobe. A single large hooked spine approximately twice the size of any other spines in the battery is present at the distal end of the sulcate edge of the nude pocket. At the level of the lobe on the asulcate edge of the pocket (i.e., the lip toward the asulcate side of the organ) are two large spines in a transverse row. Of these, the spine clos- est to the pocket is much larger than the one farther from the pocket, which is po- sitioned nearly in the middle of the asul- cate side (the lack of proper inflation of the base of the organ makes the position of this spine in a fully inflated organ dif- ficult to ascertain). A gap ornamented with minute spinules separates these enlarged spines from the battery of spines around the midsection. The sulcus spermaticus divides just within the capitulum. Its branches extend distally in centrolineal orientation, but ul- SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle timately pass to the outer (lateral) edges of each lobe. It is possible that the orienta- tion of the branches could be somewhat different in a maximally expanded organ because of differential tissue expansion. The branches of the sulcus spermaticus end at the peripheral edge of the tip of each lobe (i.e., not continuing to the cen- ter of the lobe’s apex). Distribution and Natural History Sibynomorphus oligozonatus is known from extreme southwestern Ecuador (Azuay and Loja Provinces) on the western slopes of the Andes and from a specimen without specific locality from Piura De- partment, Peru (Fig. 7). The known ele- vational range is 1,440 to 2,250 m. The holotype of Sibynomorphus oligo- zonatus was found at night on the ground in the process of swallowing a small snail (Orcés and Almendariz, 1989: 63) and in similar ecological conditions as specimens of S. petersi: “arid temperate zone with bushy vegetation that includes algarrobos and cacti; similar to the Brazilian caatinga and corresponding to the regions that Chapman called the arid temperate zone” (Orcés and Almendariz, 1989: 63). Algar- robos refers to trees of the legume family, probably of the genus Prosopis, that are generally characteristic of seasonally dry to arid regions (e.g., mesquite). Chapman (1926: 97-98) described his “Arid Division of the Temperate Zone” thus: The aridity of the treeless portions of the Temper- ate Zone varies from that of valleys with sufficient rainfall to produce pasturage to that of rock-strewn plains where cactus is the prevailing type of vegeta- tion. Generally speaking, the arid Temperate Zone comprises all the unforested areas between 9.000 and 12,000 feet, but where, because of insufficient rain- fall, forests of the Subtropical Zone are lacking, the arid Temperate descends considerably lower and may even meet the Tropical Zone. AMNH 110587 was dead on a road in an area of open grassy slopes in the Rio Jubones valley (Charles W. Myers, person- al communication, and field notes for 12 February 1974). Myers’ notes describe this 205 portion of the valley: “|from about 600 m to above 1,500 m] the Jubones valley has barren, steep, rocky slopes that are mostly covered with short grass and dotted with tree cacti. In places, especially along the river, there is low brush; some patches of sugar cane are grown in small flats near the river. . This section of the valley seems to be in rain shadow.” At the type locality (Orcés and Almen- dariz, 1989) and in the Rio Catamayo val- ley of southwestern Ecuador (BMNH 1935.11.3.108), Sibynomorphus oligozon- atus is broadly sympatric with S. petersi. These species of Sibynomorphus are also broadly sympatric with another dipsadine, Dipsas oreas, but D. oreas may be segre- gated by habitat from the species of Si- bynomorphus (see footnote 7). Sibynomorphus oneilli Rossman and Thomas Figures 8-12 Sibynomorphus oneilli Rossman and Thomas, 1979. Type locality: “NNE Balsas on the road to Abra Chanchillo, Departamento de Amazonas, Peru (ca. 1645 m).” Holotype, LSUMZ 33736. Rossman and Kizirian, 1993. Scrocchi et al., 1993. Carrillo de Espinoza and Icochea, 1995: 17. Sibynomorphus oneilli has been report- ed in the literature from only two males (including the juvenile male holotype) and a female (Rossman and Kizirian, 1993; Rossman and Thomas, 1979). I examined these specimens and refer 14 additional specimens in the MUSM to S. oneilli (see Specimens Examined and Locality Re- cords; another specimen provisionally re- ferred to S. oneilli is excluded from this account because of its uncertain prove- nience: see detailed discussion of MUSM 22.58 at the end of the species account for S. vagrans). The additional specimens sig- nificantly amplify knowledge of intraspe- cific variation and the distribution of S. oneilli. Of the three previously known specimens, Rossman and Thomas (1979: figs. 1, 2) illustrated the holotype and Rossman and Kizirian (1993: fig. 1) illus- trated KU 212600. The third specimen, 206 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 kilometers ON ANE \“E aN aN aN Val ® Sibynomorphus oligozonatus : \ a ~|\ \ Zz 1\ \ \: \ CUSA If aN ee NEN t/a ZA AN ~~, A Sibynomorphus oneilli Nit ~ \ oan bh — 7 Figure 7. Distributions of Sibynomorphus oligozonatus and S. oneilli in southern Ecuador and northwestern Peru. All known localities are plotted (a couple of symbols reflect closely contiguous localities). See Figure 43 for amplification of the southern portion of the distribution of S. oneilli (Ancash Department, Peru). Stippled area is above 1,000 m; hatched area is above 3,000 m. Numbers are locality records and letters are physical features mentioned in the text, as follows: Sibynomorphus oligozon- atus: Ecuador: (1) Santa Isabel; (2) Zhila (type locality); (8) Rio Catamayo valley; Peru: (4) uncertain locality in Piura Department, Peru (symbol arbitrarily placed above the 1,000-m contour in the center of the department). Sibynomorphus oneilli: Peru: (5) between Balsas and Abra Chanchillo (type locality); (6) Abra Gelic; (7) Cajabamba; (8) Hacienda Ticapampa; (9) Huaylas and Hacienda Santa Rosa; (10) Yuracyacu; (11) Huaraz. Physical features: (A) Rio Maranon (the inter-Andean portion is between the Cordilleras Oriental and Occidental; B and C, respectively); (B) Cordillera Oriental; (C) Cordillera Occidental; (D) Rio Santa. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle KU 212599, and others from the MUSM are illustrated herein (Figs. 8—12) Notes on the Holotype and Comparisons with Other Specimens Redescription of the Holotype. The fol- lowing observations on the holotype (LSUMZ 33736) of Sibynomorphus oneilli were made in April 2004 and supplement the original description (Rossman and Thomas, 1979). Notes on the circumstanc- es of its collection are given in the section Natural History. The type is compared with other specimens referred to this spe- cies, and a discussion of these specimens then follows. The holotype is 233 mm in total length, 174 mm SVL (tail 25% of total length). It is a juvenile male, as indicated in the orig- inal description, but it has a distinct um- bilical scar, which suggests that it is a very young juvenile. It is presently in rather poor condition. The specimen seems to have partially cleared since the species was described, perhaps as a result of the qual- ity of its initial preservation and, possibly, exposure to a low concentration of isopro- pyl alcohol at some point during its histo- ry.* The clearing has resulted in loss of de- tails of many aspects of the pattern and made some scale counts (e.g., ventrals) dif- ficult to obtain with accuracy. For exam- ple, the head markings noted by Rossman and Thomas (1979: 5) are no longer dis- cernible without magnification and careful observation, and the dorsal and ventral markings are very indistinct. My scale counts for the holotype are very Close or identical to those reported by * Details concerning the collection and initial pres- ervation of the holotype are given in the section on natural history. Isopropyl alcohol, the standard stor- age medium of the LSUMZ collection, causes clear- ing of museum specimens at concentrations below 45% (Simmons, 2002: 70). The condition of the ho- lotype of Sibynomorphus oneilli was not indicated in the original description, but the photographs provid- ed and the description of the color pattern (Rossman and Thomas, 1979: 5) suggest that the clearing oc- curred subsequent to the original description. bo ~l 0 Rossman and Thomas (1979): Dorsal scales 15-15-13, ventrals 168 (0 preven- trals), subcaudals 77, anal plate single, preoculars 2/2, postoculars 2/2, primary temporals 1/1, secondary temporals 2/2, tertiary temporals 2/2, supralabials 8/8 (4-5 touching eye on each side), infrala- bials 9/8, maxillary teeth 16. The posterior dorsal scale reduction to 13 is unusual and seems to result from the “irregular fusions of the vertebral and paravertebral rows posteriorly” (Rossman and Thomas, 1979: 4); “irregular” in this case refers to the ver- tebral and paravertebral rows that are fused, then separated, several times with no particular regularity. On the posterior body, these fusions result in a “vertebral” row twice as wide as the adjacent rows, whereas anteriorly and at midbody, the vertebral row is scarcely wider than the ad- jacent paravertebral rows. Occasional re- ductions to 13 dorsal rows posteriorly oc- cur in several other species of Sibynomor- phus (e.g., S. mikanii, S. newwiedii, and S. turgidus; Peters, 1960). The top of the head has cleared sub- stantially and, except under magnification, is medium to somewhat darker brown, forming a uniform head cap. Under mag- nification and with good light, the indis- tinct blotches on each parietal and the blotch on the frontal scale mentioned in the original description (Rossman and Thomas, 1979: 5) can be seen. The parietal blotches appear as broad elongate streaks parallel to the interparietal suture and ex- tending from the anterior border of each parietal for most of the length of the scale. The uniform brown head cap is followed by a narrow pale collar and the first cross- band. The body is grayish brown with nar- row darker brown crossbands. The ante- rior neck band is about 2 scales wide. Sub- sequent crossbands are 1.5 scales wide, narrowing to less than 1 scale wide for most of the body. The posterior crossbands are very indistinct. Interspaces anteriorly are about 2 scales wide, broadening to 3 scales for most of body. The venter is largely cleared, but apparently had bold 208 squarish or half-moon-—shaped spots aligned to form irregular lines on the lat- eral edges of the ventral plates. Comparison of the Holotype with Other Specimens. The holotype of Sibynomor- phus oneilli has several unusual character- istics compared with all other specimens referred to this species. These include ventral and subcaudal counts (168 and 77, respectively) at the upper extremities of the character ranges, an unusual reduction to 13 dorsal scales posteriorly (unique), 2/2, preoculars (unique), 9/8 infralabials (nearly unique: no other specimen had 8 infralabials, one other specimen had 9/9), and 1/1 primary temporals (nearly unique: one other individual had 1/1; another had one primary temporal on one side). Some of these characters are unusual not only within Sibynomorphus oneilli, but within the genus as a whole. As a conse- quence of having two preoculars, the lo- real is excluded from the border of the eye. The preocular scales appear to be normally formed and not merely the result of formation of an “extra” suture across a portion of the loreal, as occurs occasionally in Dipsadini (e.g., Cadle, 2005, fig. 1). Elsewhere within the genus, two preocu- lars occur with some frequency in S. pe- tersi and S. vagrans (Table 1), and in an- other specimen of S. oneilli, the loreal on each side was divided to form two lower preoculars in addition to the usual one. The head scales of the holotype are rough- ly the same shape and proportions as in other specimens, although the parietals of the holotype extend farther laterally (cor- related with it having only a single anterior temporal vs. two in most other specimens). However, it does not appear that an upper (second) anterior temporal is simply fused with the parietals in the holotype. Conse- quently, the head of the holotype appears somewhat long and narrow compared with other specimens. The holotype differs in a few details of color pattern from other specimens. al- though color patterns vary considerably within this species. The first dorsal cross- Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 band in the holotype is two dorsal rows wide. Subsequent crossbands are 1.5 rows wide, rapidly narrowing to one scale row wide for most of the body. In most other specimens, the anterior crossbands are three to five scale rows wide, narrowing to two scales (next four to five crossbands) and then to one scale row. In the holotype, bold blotches on the venter are confined primar- ily to the edges of the ventral plates, where- as in virtually all other specimens, the bold markings are distributed across the ventral plates (Figs. 9, 10, 12; Rossman and Kiziri- an, 1993: fig. 1). The holotype has a longi- tudinal dark streak on each parietal that par- allels the interparietal suture, whereas in other specimens, the head is relatively un- marked or has streaks that parallel the pos- terolateral edge of the parietals. New Material of Sibynomorphus oneilli from Ancash and Libertad Departments, Peru After careful consideration of the char- acters and geographical setting of a series of Sibynomorphus in the MUSM from the Rio Marafién and Rio Santa drainages of the Andes in Ancash and Libertad De- partments, Peru, I conclude that they con- form to the concept of S. oneilli developed herein (Fig. 8). As detailed in the section on distribu- tion, Sibynomorphus oneilli inhabits a re- gion of complex topography, and several major rivers and mountain chains separate the referred populations. Primary among these are the Rio Marafién, which sepa- rates the type locality from all other known localities, and the Cordillera Blanca, which separates populations of the Amazonian and Pacific versants. Populations of the Pa- cific versant are known from the Rio Santa drainage, a major river of the western An- des. This geographical setting and the rel- atively few specimens makes the interpre- tation of character variation within this species difficult. One locality (Yuracyacu) is represented by six specimens (four males, two females), but all other localities are represented by three or fewer speci- mens. The holotype of S. oneilli is the only SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle 209 Figure 8. Sibynomorphus oneilli. Representative specimens in dorsal view. Top: left, MUSM 2770 (male, 470 mm SVL) from Hacienda Ticapampa (La Libertad Department); right, MUSM 3103 (female, 469 mm SVL) from Cajabamba (Cajamarca De- partment). Bottom: two specimens from Yuracyacu (Ancash Department): left, MUSM 3211 (female, 413 mm SVL); right, MUSM 3334 (male, 384 mm SVL). specimen from east of the Rio Marafion, and the type locality is the lowest elevation of all known localities (1,646 m). These factors might, in part, explain some of the unusual characters of the holotype. The complexity of the terrain inhabited by these populations enhances the potential for isolation and character differentiation among, populations. The “new” specimens from Ancash De- partment, herein referred to Sibynomor- phus oneilli, were perplexing because they extend the range of variation in scutella- tion and other characters within S. oneilli. In some cases, the extended ranges ap- proach the character variation within S. vagus. Many of these specimens are much larger (nine of 14 specimens >500 mm to- tal length, with the largest 790 mm total length) than had been known for S. oneilli from the three previously known speci- mens (233-374 mm total length). Further complicating their interpretation was the existence, in the Rio Santa valley, of spec- imens lacking discrete crossbands, a char- acteristic of many of the known specimens of S. vagus (see species account). Thus, whether the “new” specimens represented 210 S. oneilli, S. vagus, some mixture of the two, or even a new species, was a real question. All specimens from east of the Cordil- lera Blanca (10 specimens, including the holotype from east of the Rio Marafién) show the banded pattern considered typi- cal of Sibynomorphus oneilli (Rossman and Kizirian, 1993; Rossman and Thomas, 1979) (Fig. 8). Of seven specimens from the Rio Santa drainage (localities Huaylas, Hacienda Santa Rosa, Hacienda Ticapam- pa, and Huaraz), three are typically band- ed (MUSM 2770, 3089, 3395) and one has very narrow and fragmented dorsal cross- bands (MUSM 2662). The other three specimens, each from a different Rio San- ta locality, lack crossbands entirely: MUSM 2687 (Huaylas), 2390 (Hacienda Santa Rosa), and 2660 (Huaraz). Instead, they have a dorsal pattern of fine dark flecks and reticulations reminiscent of the dorsal pattern in S. vagus and some spec- imens of S. petersi. Two lines of reasoning led to the conclusion that all specimens from the Rio Santa valley and from Yura- cyacu in southern Ancash were referable to S. oneilli. First, although the Cordillera Blanca is an impressive mountain range, it is also geologically quite young and probably at- tained heights over 3,000 m only since the late Miocene (Farrar and Noble, 1976; Simpson, 1979). Geographically, it would not be surprising that populations on ei- ther side of the Cordillera Blanca would be closely related to one another. On the other hand, it seems less likely that pop- ulations in the Rio Santa valley would be closely related to populations on the Am- azonian versant far to the north overshad- owed by the Cordillera de Huancabamba (the range of Sibynomorphus vagus). Pop- ulations of the Rio Santa are separated from those of Huancabamba not only by rugged mountains, but also by the rela- tively lower elevations of the Huancabam- ba Deflection region. This area is a bio- geographic discontinuity to many montane elements of flora and fauna that reach Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 their northern or southern distributional limits here (Duellman, 1979; Lynch, 1986; Simpson, 1975, 1979). Second, the character variation within the sample from the Rio Santa valley is more similar to Sibynomorphus oneilli than to S. vagus (Table 3), despite the sim- ilarity of some individuals to S. vagus in color pattern. Banded specimens through- out Ancash and southern Cajamarca/Ama- zonas Departments (e.g., Figs. 8, 9) re- semble the regular and fully crossbanded pattern typical of S. oneilli in scutellation as well as pattern, rather than the unband- ed or anteriorly banded patterns typical of S. vagus. A comparison of selected char- acters of the unbanded specimens of Si- bynomorphus from the Rio Santa valley with both S. oneilli and S. vagus (Table 3) shows that for ventral counts, unbanded males and females from the Rio Santa are within the range of other specimens of S. oneilli, but outside the range for S. vagus. There are strong frequency differences be- tween S. oneilli and S. vagus with respect to temporal scale counts. Temporal scale patterns in the unbanded specimens from the Rio Santa correspond to the predom- inant frequencies in S. oneilli rather than S. vagus. Furthermore, some specimens referred to S. oneilli from east of the Cor- dillera Blanca have fragmented dorsal bands (Fig. 8, lower left), which suggests that there is some lability in the expression of crossbands. I conclude that all available specimens from both sides of the Cordil- lera Blanca in Ancash Department are re- ferable to S. oneilli and that this species is polymorphic in color pattern, as is already well known for some other species of Si- bynomorphus. Diagnosis Sibynomorphus oneilli has a large num- ber (38—63) of dark crossbands on the dor- sum throughout the length of the body when discrete crossbands are present (the most common condition) (Fig. 9). In some specimens, the dorsal crossbands are com- plete only anteriorly; the posterior cross- SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle DAU TABLE 3. COMPARISON OF SELECTED CHARACTERS OF NONCROSSBANDED SPECIMENS REFERRED TO SIBY- NOMORPHUS ONEILLI FROM THE RIO SANTA VALLEY WITH CHARACTER STATES IN S. ONEILLI AND S. VAGUS. VENTRAL COUNTS SUMMARIZED FOR S. ONEILLI AND S. VAGUS ARE RANGES FOLLOWED BY MEAN (IN PAREN- THESES) AND SAMPLE SIZES (N). TEMPORAL COUNTS FOR THESE TWO SPECIES TREAT EACH SIDE OF EACH SPECIMEN AS AN INDEPENDENT OBSERVATION. BILATERAL COUNTS ARE SEPARATED BY A SOLIDUS (LEFT/ RIGHT). THE SAMPLE OF S. ONEILLI INCLUDES ALL SPECIMENS EXCEPT THOSE FROM THE RIO SANTA VALLEY. SEE TABLE 1 FOR SUMMARY STATISTICS OF THESE DATA. Noncrossbanded Specimens from the Rio Santa Valley MUSM 2687 MUSM 2390 MUSM 2660 Female Female Male Sibynomorphus oneilli Sibynomorphus vagus Huaylas Santa Rosa Huaraz (except Rio Santa) (total sample) Ventrals Male = — 159 156-168 (163) N = 8 144-152 (149) N =5 Female 163 168 — NGS 1W/.on GSS) Neamt om @lks4)) Nay Primary temporals YD) DD) 2/1 DNase) CN —sal9)) a (Niels) Secondary temporals 3/3 3/3 3/3 Nee 2)) aN — salts) (iy = 14) On (Nee )) a Nias) Tertiary temporals 4/3 CYB 4/4. Ne) 0 (N = O) ORG NE—aG)) CN eal) 4(N 11) 2a (Nie wal3)) Si (Nee) bands in these specimens are incomplete middorsally and appear as narrow lateral bars. Several specimens from the Rio San- ta valley lack discrete bands entirely; in- stead, the dorsum has an obscure network of dark flecks or reticulations. The anterior crossbands (generally two to three dorsal rows wide) are slightly broader than, or equal in width to, the interspaces, whereas most crossbands are only one dorsal row wide and much narrower than the inter- spaces. The venter is heavily checkered or spotted with large bold markings (round- ed, squarish, or half-moon—shaped) (Fig. 9). Ventrals are 152-168 in males; 163-173 in females. Subcaudals are 62—77 in males; 56-64 in females. Infralabials are 9-13. Sibynomorphus oneilli differs from oth- er species of Sibynomorphus in Ecuador and Peru as follows. In S. oligozonatus, S. vagrans, and S. williamsi, the anterior crossbands are usually twice or more as broad as the interspaces (see illustrations in the species accounts), whereas in S. oneilli, crossbands are usually equivalent to, or narrower than, the interspaces (oc- casionally broader than interspaces but never approaching twice their width). These species also differ from S. oneilli in other characters: S. oligozonatus has fewer supralabials (6-7) and a less boldly pat- terned venter (Figs. 1, 2, 4, 6). Sibyno- morphus williamsi has fewer supralabials (6-7) and a greater number of ventrals (males, 173-182: females, 181—188) than S. oneilli. Sibynomorphus oneilli averages about 10 more ventrals and 15 fewer sub- caudals than S. vagrans when sexes are considered separately (Table 1). Sibynomorphus vagus differs from S. oneilli in temporal scale patterns (Table 1) and by having fewer ventrals (males, 144— 152; females, 151-158). Sibynomorphus petersi has a greater number of ventrals (164—188, sexes combined) and subcaudals Cialessan >—oV-nemalesimO5—147)e tvamis: oneilli and has different color patterns on the head and body. Description Size and Scutellation. Taxonomic data for Sibynomorphus oneilli are summarized in Table 1. The largest specimen is a fe- male 790 mm total length (624 mm SVL). 212 Figure 9. Sibynomorphus oneilli. Dorsal and ventral views of KU 212599 (from near Cajabamba, Cajamarca Department, Peru). Female, 374 mm total length. The largest male is 571 mm total length (440 mm SVL). Body somewhat com- pressed. Tail 22-26% of total length in males, 20-22% of total length in females. Dorsal scales smooth and in 15—15—15 rows in all specimens except the holotype, in which there is a posterior reduction to 13 scale rows (see above Redescription of the Holotype). Vertebral row slightly broader than paravertebral rows. Ventrals 152-168 (averaging 161) in males, 163— 173 (averaging 167) in females; 0-4 pre- ventrals precede the ventral plates. Anal Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 single. Subcaudals 62-77 (averaging 68) in males, 57-64 (averaging 61) in females. Usually, a loreal and a single preocular (superior to the loreal) border the anterior edge of the eye, but the preocular is often fused with the prefrontal (Table 1); loreal squarish or polygonal. The holotype has two preoculars on each side and the loreal is separated from the eye (interposed be- tween the inferior preocular and the pos- terior nasal). MUSM 3376 has a separate preocular but, in addition, the loreal is di- vided vertically and horizontally to form a pair of lower preoculars, thus resulting in three preoculars. Postoculars are virtually always 2 (3 on one side of one specimen). Primary temporals usually 2 (occasionally 1); secondary temporals usually 3 (range 1-4), tertiary temporals usually 3 or 4 (rarely 2). Supralabials 6-10 with the most frequent patterns 8 (4—5 bordering eye) or 7 (3-4 bordering eye). Infralabials 9-13 (most frequently 10 or 11). One pair of infralabials in contact behind mental in all specimens. Chin shields in 2 pairs (N = 3), 3 pairs (N = 10), 3.5 pairs (N = 1), or 4 pairs (N = 4). Maxillary teeth 13-16 (N = 13). The pupil is prolate in preservative. Barring patterns of geographic variation that remain unforeseen because of sam- pling issues,° Sibynomorphus oneilli shows typical colubrid patterns of sexual dimor- phism in size (females attain larger size), ventral counts (female greater; t = 2.545, df = 15, p < 0.05), subcaudal counts (male greater; t = 2.764, df = 12, p < 0.05), and relative tail length (males have longer tails) (Table 1). Color in Life. Unknown. Color in Preservative. The pattern of the holotype is described above, but its poor state of preservation does not warrant a detailed description (see above Rede- ° The female from Santa Rosa is substantially larger than other specimens (790 mm total length). Large size is conceivably characteristic of this population, but that is unverifiable with a single specimen. It could be that the extremely large size of this speci- men is due to vagaries of sampling. The next largest specimen, a male, was 623 mm total length. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle 213 Figure 10. Sibynomorphus oneilli. Representative specimens in ventral view. Top: MUSM 2770 (La Libertad Department; male, 470 mm SVL). Bottom: MUSM 3211 (Ancash Depart- ment; female, 413 mm SVL). scription of the Holotype and Rossman and Thomas, 1979: 5). The coloration of Si- bynomorphus oneilli in preservative, on the basis of the other specimens, follows. Dorsal ground color medium brown or grayish brown, occasionally somewhat red- dish brown; grayish when the stratum cor- neum is removed. Dorsal crossbands and other markings black (Figs. 8, 9). Dorsal crossbands on the body, when present, range from 38 to 63 with no apparent sex- ual dimorphism; crossbands can be frag- mented or even completely absent, as de- scribed below. The venter is dull whitish or with fine dark stippling giving a dusky appearance, upon which are bold black squarish or half-moon—shaped spots in a checkerboard pattern. Ventral spots are usually densely arrayed (Figs. 9, 10), but there is a tendency for specimens lacking dorsal bands also to have less boldly pat- terned venters (see Fig. 12). Gular region Figure 11. Sibynomorphus onelilli. Dorsal and lateral views of head, KU 212599. The skull of this specimen had been dis- sected from the skin, hence the appearance of the eyes and tears in the rostral and mental regions. immaculate, grayish to white. Dorsal and ventral surfaces of the tail are like the cor- responding surfaces of the body, except the dorsal bands are incomplete middor- sally. The top of the head is medium brown, usually with some scattered irregular blackish marks primarily on the parietals and frontal; these marks sometimes in- clude an irregular curved black streak that roughly parallels the posterolateral edge of each parietal (Fig. 11; Rossman and Kizir- ian, 1993: fig. 1). A narrow dark nape bar is frequently present between the first dor- sal crossband and the parietal scales; it ex- tends laterally to just above the posterior supralabials and, when present, can be manifested by dusky shading or a very dis- tinct blackish bar. In a few specimens, the top of the head is unmarked (e.g., MUSM 3211, Fig. 8). Temporal scales are very narrowly edged with black, but without close examination, the impression 1s that the side of the head is unmarked. The su- pralabials are dull whitish with some dark stippling (but no discrete markings) along 214 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 12. Sibynomorphus oneilli. A nonbanded specimen from Huaylas in the Rio Santa valley (Pacific versant, Ancash Department). MUSM 2687 (female, 463 mm SVL). suture lines. Infralabials are dull whitish, unmarked. The dorsal pattern varies by individual. The first crossband is interrupted middor- sally in many specimens, forming a pair of blotches (much higher than wide) on the side of the neck (Figs. 8 [upper and lower right], 9, 11). Other crossbands on the body are likewise sometimes interrupted middorsally. At least occasionally (e.g., MUSM 3211; Fig. 8, lower left), virtually all of the crossbands are interrupted so that the dorsal pattern consists of a series of irregular narrow vertical streaks on the flanks, and in extreme cases (MUSM 2662), the crossbands are extremely frag- mented so as to give the appearance of an irregular pattern. In most specimens, the crossbands on the anterior quarter to third of the body are wider than more posterior crossbands (Figs. 8, 9). Occasionally (MUSM 2770, KU 212600) these anterior crossbands are twice or more the width of the posterior crossbands. Except for these anterior bands, most bands on the body are one scale row or less in width and are much narrower than the interspaces. As described above (New Material of Siby- nomorphus oneilli from Ancash and Lib- ertad Departments, Peru), two females and a male from the Rio Santa valley lack crossbands entirely and have irregular dark flecks over the dorsum (Fig. 12). There seems to be little or no ontoge- netic change in color pattern, as four spec- imens 203-280 mm total length had pat- terns similar to adults; one of these (MUSM 3089; 232 mm total length) had the highest number of dorsal bands re- corded (63), and these were formed by darkened scale edges rather than fully formed bands. Details of the dorsal patterns of KU 212599 and 212600 are as follows: KU 212599 (Fig. 9): Approximately 55 cross- bands are present on the body. On the neck just behind the head is a pair of lateral blackish blotch- es 4 to 5 dorsal rows wide. The following crossband (complete across the dorsum) is 3 scales wide and the following 4 crossbands are two scales wide. Subsequent crossbands are no more than a single dorsal row wide, becoming increasingly fragmented posteriorly, but more or less discrete the length of the body. Anteriorly, the crossbands extend down to the first dorsal row or outer edge of the ventrals; posteriorly the crossbands tend to fragment before reaching the first dorsal row. Interspaces between the crossbands are medium reddish brown, 2 to 3 dorsal rows in width anteriorly and usually 3 rows posteriorly. Many dark flecks are present in the in- terspaces, especially posteriorly. KU 212600: The body has 38 total crossbands (+2 half-bands in between complete bands) (see Rossman and Kizirian, 1993: fig. 1 for dorsal and ventral photographs of this specimen). Crossbands are complete across the dorsum (except the two half-bands) and extend to the outer edge of the ventral plates, where most fuse more or less com- pletely with one or more of the dense squarish spots on the venter. The first crossband behind the head is 4 scales wide, the next five bands are 3 scales wide, the following five are two scales wide, and the rest are one scale wide. The crossbands are wider middorsally than ventrally; they have irreg- ular edges, but are essentially oriented vertically (slightly curved in some cases). Laterally, between SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR ° Cadle many of the crossbands, and in the vertebral/para- vertebral region are scattered irregular black flecks mainly formed by black edging on some of the dor- sal scales. Interspaces anteriorly are two scale rows wide, broadening to three on anterior one-third of body, and then to four scale rows wide posteriorly. Etymology. Sibynomorphus oneilli was named for ornithologist John P. O’Neill, who collected the holotype. The ornitho- logical expeditions of Louisiana State Uni- versity to Peru, many under the guidance of O'Neill, have also made important con- tributions to knowledge of the Peruvian herpetofauna. Distribution Sibynomorphus oneilli is known from the northern end of the Cordillera Ori- ental in extreme southern Amazonas De- partment, Peru, and in the Cordillera Oc- cidental from southern Ancash to southern Cajamarca Department (Fig. 7; see also Fig. 43). Localities are on the Amazonian versant (Rio Marafidn drainage) and Pa- cific versant (Rio Santa and its tributary, the Rio Tablachaca/Tablache) of the An- des. The recorded elevational range is 1,646-3,500 m. In Ancash Department, populations of S. oneilli from the Amazo- nian versant of the Cordillera Occidental are separated from populations of the Pa- cific versant (Rio Santa drainage) by exten- sive mountains of the Cordillera Blanca well over 4,000 m. The type locality is sep- arated from other known localities by the Rio Maranon (but see footnote 15). As here conceived, populations of Si- bynomorphus oneilli inhabit one of the most tectonically active and topographical- ly complex regions of the Andes. The spe- cies occurs on both sides of the Rio Ma- raion, one of the longest and largest rivers descending from the Andes; it separates the Cordillera Oriental from the Cordillera Occidental, the principal chains of the An- des in northern Peru. The Rio Marafoén flows northwesterly until near the Ecua- dorian border, thence turning east and be- coming the major western affluent of the Amazon system. On the Pacific versant, S. 215 oneilli occurs in the valley of the Rio Santa and one of its tributaries, the Rio Tabla- chaca/Tablache (which forms the border between Ancash and La Libertad Depart- ments). The Rio Santa flows northwesterly through the length of Ancash Department, eventually turning westward through the gorge of the Cafion del Pato and emptying into the Pacific Ocean. Lehr et al. (2002: fig. 11) present a photograph of the Rio Santa valley upstream from Huaraz, one of the localities for S. oneilli. The inter-An- dean valleys of the Rio Marafién and Rio Santa are relatively arid and presently form effective barriers for east-west dis- persal of montane flora and fauna (Simp- son, 1979). In addition to large river systems, pop- ulations of Sibynomorphus oneilli are sep- arated by incredibly rugged mountains. Populations in the Rio Maranon drainage are separated from those in the Rio Santa valley by the glacier-laden Cordillera Blan- ca, the highest continuous range in the An- des (see Lehr et al., 2002: fig. 11). This impressive mountain chain contains more than 10 peaks exceeding 6,000 m elevation (the highest being Nevado Huascaran at 6,746 m) and no passes below 4,000 m; most of its area is above 5,000 m (Clap- perton, 1993). Vertical descents on either side of the Cordillera Blanca frequently exceed 4,500 m, and its fault zones are young and very active (Noble et al., 1990). Major uplifts in the late Miocene through Pleistocene brought the mountains to their present height (Farrar and Noble, 1976). The Rio Marafién and the Rio Santa are relatively young rivers that developed in the down-faulted trenches between major uplifts of the Cordillera Oriental and Cor- dillera Occidental (Rio Marafion) or within the Cordillera Occidental (the Rio Santa, flanked on the east and west by the Cor- dillera Blanca and Cordillera Negra, re- spectively). Thus, populations referred to Sibyno- morphus oneilli on either side of major geographic barriers within the species’ range (Rio Marafién, Cordillera Blanca) 216 were probably isolated, geologically speak- ing, relatively recently. Nonetheless, the heavily dissected topography enhances op- portunities for the isolation of these pop- ulations, particularly those of the eastern and western versants, which are separated by inhospitable terrain or unsuitable high- elevation habitats. Sibynomorphus oneilli is one of few reptiles known to have pop- ulations on both the Amazonian and Pa- cific versants of the Andes, and most of these have only become known in con- junction with field work in the area in the last quarter century (Cadle, unpublished data). Natural History The holotype of Sibynomorphus oneilli was collected 8 August 1975 in “semiarid brushland” (Rossman and Thomas, 1979). According to the collector, ornithologist John P. O’Neill (personal communication, 25 April 2006), the holotype was dead in the middle of a gravel highway on an over- cast, cool day; the area had scattered bush- es and was rather overgrazed. It was placed in a jacket pocket for 24 or more hours before preservation, which could in part explain its poor present condition. Two specimens (KU 212599-600) were collected in January and March under rocks in agricultural land (Rossman and Kizirian, 1993). Other specimens with dates of collection have been obtained in January, February, March, April, Septem- ber, October, and December. These months generally correspond to the cooler, wetter part of the year in this portion of the Andes (wet season primarily Novem- ber—April). The three smallest individuals have vis- ible umbilical scars: MUSM 3303 (203 mm total length; 155 mm SVL), MUSM 3089 (232 mm total length; 181 mm SVL), and LSUMZ 33736 (233 mm total length; 174 mm SVL). MUSM 3089 was collected in December 1982; LSUMZ 33736 was collected 8 August 1975. The date of col- lection of MUSM 3303 is unknown. Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Sibynomorphus petersi Orcés and Almendariz Figures 13-27 Dipsas mikanii oreas: Parker, 1934: 271, 1938: 444 (misidentifications). See later discussion of mis- identified literature records. Dipsas variegata variegata: Peters, 1964: 47 (mis- identification). Based on USNM 237040, which was subsequently referred to Sibynomorphus pe- tersi (Cadle and Myers, 2003: 7). Sibynomorphus petersi Orcés and Almendariz (1989). Type locality: “Zhila (2,250 m) 79°17'26"W, 03°39'45"S [see footnote 19], parroquia Ofia, can- ton Girdén, Provincia Azuay [Ecuador].” Holotype, EPN 2659. Almendariz and Orcés (2004). Sibynomorphus vagus: Lehr et al. (2002) (misidenti- fication of SMF 80048, MUSM 20583). Notes on the Type Series of Sibynomorphus petersi The specimens here referred to Siby- nomorphus petersi substantially amplify knowledge of variation in this taxon, which was previously reported only from the ho- lotype (EPN 2659) and two paratypes (EPN 1847, 2660) (Orcés and Almendariz, 1989). I refer other specimens from south- ern Ecuador and eight Peruvian speci- mens to this species (see Specimens Ex- amined and Locality Records). Of the type series, I have examined only a paratype from the type locality, EPN 2660 (Fig. 13). According to Orcés and Almendariz (1989) the holotype is an adult male 763 mm total length, 609 mm SVL, 154 mm tail length (= 20.2% of total length and not the 21.9% reported in the original descrip- tion); 15 [midbody] dorsal scale rows (ver- tebral row slightly enlarged); 176 ventrals, 79 subcaudals, single anal; 9 supralabials (4 + 5 touching the eye), 12 infralabials; four pairs of chin shields; 0/1 preoculars, 2/2, postoculars; 3 + 2 + 3/2 + 3 + 3 tem- porals. These data conform to my own for male Sibynomorphus petersi (Table 1), ex- cept that the tail is shorter than in any male I examined. Diagnosis Sibynomorphus petersi attains a large size (to 771 mm total length in females, 763 mm in males) and has moderately SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle Figure 13. Sibynomorphus petersi. Paratype (EPN 2660, Azuay Province, Ecuador) in dorsal and ventral views. Female, 542 mm SVL. high ventral (164-188) and subcaudal (65— 87) counts. Many features distinguishing this species from others in Peru and Ec- uador are aspects of color pattern. Siby- nomorphus petersi has a dorsal ground col- or of medium brown (brown or grayish in preservative), generally with narrow, dark brown (sometimes dark gray in preserva- tive), irregular zigzag crossbands that ex- tend ventrally to the lowermost dorsal rows or extreme outer edges of the ventral scales (Figs. 13, 14). Three or four anterior crossbands are sometimes broader than the others. Crossbands may be fragmented into a dorsal and lateral series of blotches, and crossbands or blotches are difficult to discern in some individuals. The venter is whitish with bold black or dark brown squarish or half-moon—shaped blotches or streaks; the ventral ground color is some- 217 times heavily invested with brown, a re- flection of a dense peppering of minute flecks and spots. The top of the head is brown or grayish brown with dark brown spots, streaks, or other markings in a highly irregular pat- tern. Nonetheless, one feature of the head pattern seems relatively constant, albeit somewhat variable in precise execution: a large discrete spot is usually present on each prefrontal scale (Fig. 15). The pre- frontal spots are usually symmetrically placed but vary in size, shape, and exact position among specimens (see Color in Preservative and Variation in Pattern). They are usually round but occasionally are irregular in shape. Dark markings on the head have a narrow pale edge (yellow- ish in life), which can be obscure in pre- served specimens. In many specimens the dark head markings also have a pale core (Fig. 15). Sibynomorphus petersi differs from oth- er species of the genus in northern South America as follows: S. vagrans and S. va- gus have fewer ventrals (149-159 and 144-158, respectively) and different color patterns (see descriptions and illustrations herein). Sibynomorphus oneilli usually has narrow, more or less straight-sided (com- pared with zigzag) crossbands and has few- er ventrals and subcaudals than S. petersi (S. oneilli ventrals and subcaudals, male: I5QORIGSPand) 62=7 7 sfemale: 163=173 and 57-64). Sibynomorphus oligozonatus has anterior crossbands that are much broader than the interspaces and, in males, fewer ventrals (145-163) and subcaudals (66— 68): S. oligozonatus has six or seven su- pralabials, whereas S. petersi usually has eight. Sibynomorphus williamsi usually has six supralabials (3-4 touching the eye), whereas S. petersi usually has eight su- pralabials (4-5 touching the eye); S. wil- liamsi also has distinct bold crossbands the entire length of the body and usually a rel- atively unmarked venter (when marked, never bold spots or half-moons). In the collections I surveyed, Sibyno- morphus petersi was commonly misiden- 218 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 14. Sibynomorphus petersi. Dorsal and ventral views of a specimen from southern Ecuador. EPN 4737 (Loja Province, Ecuador). Female, 580 mm SVL. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR ¢ Cadle Figure 15. Sibynomorphus petersi. Top: Lateral view of head (BMNH 1900.3.30.19 from San Pablo, Cajamarca Department, Peru). Bottom: Dorsal view of head (EPN 4737, Loja Province, Ecuador). Both specimens show the large spot on each pre- frontal scale that is characteristic of S. petersi despite some variation (See Figs. 19, 22). In most specimens, the spots are bordered by a narrow pale ring, as seen clearly in the lower figure here. Note also the minute dark spots on individual scales. tified as Dipsas variegata or D. oreas (see discussion of misidentifications in the lit- erature below). Sibynomorphus petersi can be distinguished from all species of Dipsas in western Ecuador and Peru by a few key aspects of color pattern, especially details of the banding pattern on the body and patterns atop the head (see Cadle, 2005; Cadle and Myers, 2003). Scutella- tion characters alone are insufficient for some comparisons because of substantial overlap in scale counts for many species of Dipsadini. Species of Dipsas also have a relatively much larger eye than species of Sibynomorphus; see Cadle (2005) and Ca- dle and Myers (2003) for descriptions and illustrations. The following comparisons show, for subadult and adult S. petersi and D. oreas, the difference in eye size com- 219 pared with SVL (for these comparisons subadults are =351 mm SVL): S. peterst: Adult eye size: 0.552 + 0.081% SVL (N = 4) Subadult eye size: 0.701 + 0.016% SVL UN = 2) D. oreas: Adult eye size: 0.730 + 0.049% SVL CNG — sal) Subadult eye size: 0.840% SVL (N = 1) Misidentifications of Sibynomorphus pe- tersi in the Literature. Sibynomorphus pe- tersi was frequently misidentified in mu- seum collections, and some of these rec- ords have made their way into the litera- ee, IPavikere (Cl 4lo Dyile IGexss aiaiz!) identified specimens of S. petersi from the collections of Clodoveo Carri6n from southern Ecuador as Dipsas mikanii oreas (= D. oreas). Parker cited only six speci- mens from the Carrié6n collection as this species, whereas I located 13 Carrién specimens in the BMNH that were origi- nally identified as “Dipsas mikanii oreas.” All of these except one are here referred to S. petersi (the other is S. oligozonatus; see Specimens Examined and Locality Re- cords). Three specimens can, with reason- able confidence, be associated with BMNH catalogue numbers on the basis of the sex, locality, and segmental counts pro- vided by Parker (1938): a male and two females from “Catamayo Valley” are, re- spectively, BMNH 1935.11.3.110 (= Si- bynomorphus petersi), 1935.11.3.108 (= S. oligozonatus), and 1935.11.3.109 (= S. pe- tersi). I could not confidently associate BMNH numbers with the other three specimens reported by Parker (1934, 1938). Specimens of Sibynomorphus petersi were commonly misidentified as Dipsas variegata, apparently on the basis of ear- lier reports of that species from Ecuador and Peru (e.g., Peters, 1960, 1964). How- ever, Cadle and Myers (2003) showed that some previously published records of 220 “Dipsas variegata variegata” and “Dipsas variegata nicholsi” from Ecuador and Peru were based on misidentifications of other species, including D. andiana, D. oreas, D. peruana, and S. petersi.!° Peters’ (1964: 47) record of “Dipsas variegata variegata” from western Ecuador is based on USNM 237040, which was subsequently referred to Sibynomorphus petersi (Cadle and My- ers, 2003: 7). Lehr et al. (2002) referred two specimens of S. petersi, SMF 80048 and MUSM 20583 from Ancash Depart- ment, Periytons og2use lheyacleanlyranc not that species and one of them is here referred to S. petersi (see footnote 22). Cadle and Myers (2003) were specifically con- cerned with Peruvian and Ecuadorian records of Dip- sas variegata discussed by Peters (1960, 1964) and Steindachner (1902). We failed to note more recent reports of “D. variegata” from southeastern Peru (Doan and Arriaga, 2002; Duellman and Salas, 1991). In 2001, I had examined one of two specimens re- ferred to D. variegata from Cusco Amazonico (Duell- man and Salas, 1991), KU 214858, and concluded that it is not the same taxon as D. variegata Duméril, Bibron, and Duméril from Venezuela and the Guay- anan region. On this basis, Duellman (2005: 371) list- ed the Cusco Amaz6énico material as “Dipsas sp.” Doan and Arriaga (2002) apparently collected the same taxon from a site near Cusco Amazoénico and referred all the material to D. variegata. KU 214858 (Duellman, 2005: 371 and pl. 183), a female, has more ventrals and a different color pat- tern than specimens of Dipsas variegata from Vene- zuela and the Guayanan region examined by Cadle and Myers (2003: table 1), although ventral counts recorded in the literature, including those for a syn- type reported in the original description, are greater (Cadle and Myers, 2003: footnote 5; Duméril et al., 1854: 478). KU 214858 is similar in color pattern and high ventral counts to specimens tentatively referred to D. variegata from the mouth of the Amazon (Cadle and Myers, 2003: footnote 5; Cunha and Nascimento, 1993). This taxon has not turned up in some com- prehensive collections from central Amazonia (Mar- tins and Oliveira, 1998). Clearly, the status of these specimens from southeastern Peru referred to D. var- iegata (Doan and Arriaga, 2002; Duellman and Salas, 1991) needs clarification, and Dipsas variegata itself needs a thorough range-wide study. These issues do not affect Cadle and Myers’s (2003) conclusions on the basis of Peters’ (1960, 1964) and Steindachner’s (1902) records of “Dipsas variegata” from Peru and Ecuador, none of which are that species. Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Description Size and Scutellation. Table 1 summa- rizes basic characteristics of Sibynomor- phus petersi. The largest specimen is a fe- male 771 mm total length, 595 mm SVL. The largest male is the holotype, 763 mm total length, 609 mm SVL (Orcés and Al- mendariz, 1989). Tail 22-28% of total length in males (with the holotype an out- lier at 20% on the basis of measurements in the original description), 20-24% of to- tal length in females. Body somewhat compressed. Dorsal scales smooth, in 15— 15-15 rows. The vertebral row is slightly broader than, to approximately twice the width of, the adjacent paravertebral rows; the vertebral row seems relatively broader in Ecuadorian compared with Peruvian specimens. Ventrals 164-183 (averaging 173) in males, 173-188 (averaging 179) in females. Ventrals preceded by one to three preventrals. Anal single. Subcaudals 75-87 (averaging 81) in males, 65—77 (averaging 73) in females. Usually a loreal and a single preocular border the eye (preocular su- perior to the loreal); occasionally two or three preoculars were present, but the preocular was never fused with the pre- frontal (Table 1). The loreal varies in shape from squarish (e.g., Fig. 15, top) to more elongate (Fig. 23, bottom). Postoculars 2 (rarely 3). Primary temporals usually 2 (oc- casionally 1). Secondary temporals usually 3 (range 2-4). Tertiary temporals 2-4. Su- pralabials usually 8 (range 7-9) with 4-5 bordering the eye. Infralabials usually 10 or 1] (range 9-12). One or two pairs of infralabials in contact behind the mental, with the following frequencies in the sam- ple: first pair of infralabials in contact (11), first two pairs in contact (6), one infralabial contacts two on the opposite side (4). Three pairs (N = 15), 3.5 pairs (N = 1), or four pairs (N = 3) of squarish or polyg- onal chin shields. Maxillary teeth 14—17 (N = 1:0), Sexual Dimorphism. Compared with fe- males, male Sibynomorphus petersi have significantly fewer ventrals (¢ = 2.408, df SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle = 22, p < 0.05), significantly more sub- caudals (t¢ = 4.760, df = 18, p < 0.001), and greater relative tail lengths. Females attain a greater body size than males. The five largest females were 681-771 mm to- tal length (541-595 mm SVL), and four of these were >700 mm total length. The five largest males were 568-657+ mm total length (422-520 mm SVL) and three of these were >600 mm total length. Large females appear to have somewhat different body proportions than smaller individuals of either sex, but larger samples of well- preserved specimens would be needed to test this observation rigorously. Addition- ally, because localities for S. petersi are represented by few individuals (most fre- quently one or two specimens per locality), it is difficult to separate divergence in characters because of sex from geographic or other sources of variation. Nonetheless, large females tend to have rather narrow, elongate heads and relatively small eyes; smaller individuals have shorter, more rounded, bulbous heads and larger eyes. Large females have rather stout bodies, es- pecially posteriorly, compared with males (this comparison ignores the two females that were clearly carrying eggs). Sex differences in head size and shape and body size and shape, such as those ap- parent in Sibynomorphus petersi, are rel- atively common in snakes (reviews in Bon- net et al., 1997: Shine, 1989, 1993). The head shape differences most often reflect intersexual dietary differences (e.g., Cam- illeri and Shine, 1990), whereas body shape differences, in which females have a stouter, robust body, most likely reflect the reproductive needs of female snakes to accommodate eggs or fat body stores (Lourdais et al., 2006). Corresponding to the relatively longer SVLs of females compared with males, there is sexual dimorphism in the number of crossbands on the body. Females aver- aged 38 + 8.01 bands on the body (range: 31-59, N = 12; ignoring a clear outlier at 59, the range is 31-44 with a mean of 36.0 + 4.74). Males averaged 31.2 + 1.69 bands 220 (range: 29-35; N = 10). The overall means are significantly different (unequal vari- ACES ig ROA Oat gigs = 22208.) = 0105)) as well as the comparison ignoring the fe- male outlier (unequal variances, t’, = 3.049, too = 2.77, 0.01 < p < 0.02). Superimposed on this pattern of sexual dimorphism is geographic variation in the number of crossbands (Peruvian speci- mens tend to have greater numbers of crossbands; see below), but even consid- ering only Ecuadorian specimens, females average about four more bands on the body than males. Geographic Variation. Notable geo- graphic trends in scutellation or body pro- portions are few in Sibynomorphus peter- si, as shown for ventrals, subcaudals, and relative tail lengths in Table 4. None of these characters show statistically signifi- cant differences between Ecuadorian and Peruvian samples when sexes are com- pared separately. Three of six Peruvian specimens have a single anterior temporal on each side (the others have two), where- as all Ecuadorian specimens have either two or three anterior temporals. No other scutellation differences among Ecuadorian and Peruvian specimens were noted. Per- haps significantly, the only specimen of S. petersi from the Amazonian versant, MUSM-JEC 12884, has the lowest ventral count (164) of all specimens (range in the rest of the sample: 170-188; N = 23). No other characters of this specimen were un- usual (unfortunately, it is a badly damaged road kill). In addition to the sexual differences in- dicated above, the number of body cross- bands varies geographically, but sample sizes are too small to thoroughly analyze the trend. The only Peruvian male with a full complement of crossbands has 35 bands on the body, whereas the range among Ecuadorian males was 29-32. Three Peruvian females had 41, 44, and 99 crossbands, whereas the range among Ecuadorian females was 31-38 and one outlier at 43. Thus, it seems likely that for bo bo TABLE 4. DY) Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 (GEOGRAPHIC COMPARISON OF SELECTED CHARACTERS IN SIBYNOMORPHUS PETERSI. THE TWO MALES FROM PERU HAD BROKEN TAILS, SO NO SUMMARY STATISTICS ARE AVAILABLE FOR SUBCAUDALS AND RELATIVE TAIL LENGTHS FOR MALES FROM PERU. NONE OF THESE CHARACTERS SHOW STATISTICALLY SIGNIFICANT DIFFERENCES BETWEEN ECUADORIAN AND PERUVIAN SAMPLES IN SEPARATE COMPARISONS BY SEX.. Ecuador Ventrals Male 170-183 (ASS) == AES N 79) Female 173-188 LSeSh= =o O6NUNE—sS) Subcaudals Male (537 81.0 = 3.66 (N = 8) Female 65=1) (oro ONE—eo) Tail length/total length Male 0222003 OD == OLO25 (NE —s)) Female 0.20—0.24 0.22 = 0.01 (N = 7) both sexes, northern specimens tend to have fewer crossbands than southern ones. Color in Life. The color pattern of Si- bynomorphus petersi in life consists of var- ious shades of brown (yellowish brown to dark brown) or gray. The dorsum is usually distinctly crossbanded, but three individ- uals from Peru have very indistinct cross- bands. Bands are characteristically narrow and undulating or zigzag. The venter is heavily marked with bold spots, and the top of the head has complex dark mark- ings, including the consistent presence of a large spot on each prefrontal (details be- low). The following color notes are avail- able for individual specimens: FMNH 232569 (Fig. 16 female, Rio Zana Study Site, 494 mm SVL): Dorsum medium rich brown with dark brown markings: a middorsal series of irregular spots and a lateral series forming oblique incomplete crossbands. Dorsal ground color lighter anteriorly than posteriorly. Head medium rich brown with dark irregular spots and markings on most scales. Gular region cream with dark brown peppering on lower labials (upper labials also have this peppering but the ground color is darker). Ven- ter whitish anteriorly with large dark spots forming irregular, roughly parallel, series of broken lines on each side. Entire venter peppered with dark brown, which becomes denser posteriorly. Ventral Peru Total Sample 16276 164-183 I7OL0-=SSn( Nea) 73:0) == > SON CN —siae) 174-182 173-188 S22 92S oon NG — ae) 178.5 + 4.33 (N = 13) 65+ to 74+ 79-87 (N = 2) 81.0 + 3.66 (N = 8) 70-74 65-77 WZ. = J9ll (N = 4) 73.0 + 3.69 (N = 12) O.21+ to 0:26 0.22-0.28 (N = 2) 0.25 + 0.02 (N = 8) 0.20—0.24 0.20—-0.24 0.22 = 0.02 (N = 4) 0.22 = 0:01 (N= 1) surface of tail also peppered with dark brown. (Field notes of J. E. Cadle) KU 121309 (Fig. 17. male, southern Ecuador, 351 mm SVL): Ground color medium brown ex- tending to bottom of lower scale row; dorsum spot- ted and barred with darker brown, often edged with black; top of head medium brown with black spots; upper labials gray brown; chin white and dark brown; iris gray at top and bottom with broad brown band through it; tip of tongue black, re- mainder red. (Field notes of John D. Lynch) KU 142804 (Fig. 18. male, southern Ecuador, 377 mm SVL): Dorsum brown with darker brown crossbars faintly outlined with creamy tan. Venter creamy white with black spots. Iris tan. (Field notes of William E. Duellman) LSUMZ 27374 (male, northern Peru, 520 mm SVL): Dorsal ground color gray, markings black or very dark gray (gray-brown). Light flecking and iso- lated scales on edges of bands white. Venter paler gray with black spots and white spots, more exten- sive anteriorly. (Field notes of Richard Thomas for RT 3059) Color in Preservative and Variation in Pattern. In preservative, the pattern ele- ments (crossbands, head markings, ventral markings) remain distinct. In preserved specimens that have not lost the stratum corneum, the ground colors, crossbands, and other markings retain their brown col- or tones, although they are duller than in SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle Figure 16. Sibynomorphus petersi. Dorsal and ventral views of FMNH 232569 (Rio Zana Study Site, Cajamarca Depart- ment, Peru). Female, 494 mm SVL. The crossbands in this specimen are fragmented into a series of irregular vertebral and lateral spots. life. With loss of the stratum corneum, the brown tones present in life are gray (in some older specimens, the ground color of such specimens is ash white). There is both sexual dimorphism and geographic variation in the number of crossbands on the body, as covered in previous sections. Most specimens of Sibynomorphus pe- tersi have a series of narrow ZIGZag OY wavy crossbands that are complete across the dorsum (i.e., not alternating side to side). These crossbands usually have a very nar- row light border and jagged edges (Fig. 14). A band on the nape and the following two to four crossbands are usually, but not 223 Figure 17. Sibynomorphus petersi. Dorsal and ventral views of KU 121309 (Loja Province, Ecuador). Male, 351 mm SVL. always, broader than the others (Fig. 14). These anterior crossbands are sometimes incomplete across the vertebral line. Crossbands on the neck are occasionally absent and replaced by a broad longitudi- nal streak on each side (Fig. 18). In some individuals, the crossbands are fragmented laterally into a series of spots that can maintain the appearance of crossbands or are so fragmented that individual bands 22 Figure 18. Sibynomorphus petersi. KU 142804 (Loja Prov- ince, Ecuador). Male, 377 mm SVL. The dark streak on the side of the neck is characteristic of many specimens of S. petersi. 4 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 are difficult to discern, as described for three Peruvian specimens below. The top of the head in Sibynomorphus petersi is brown or grayish brown with dark brown markings. A large discrete spot is present on each prefrontal (Fig. 19); al- though it varies in size, position, and shape, it is a constant feature of the pat- tern. Despite the variation in form, the prefrontal spots are sufficiently diagnostic of this species that specimens that other- wise differ greatly in pattern retain this character (compare Figs. 15, 19, 22, and the photos of corresponding dorsal pat- terns). Other large spots or reticulations are present on the frontal, parietals, and supraoculars; these can fuse with one an- other to form larger patterns (Fig. 19, low- er right). In virtually all cases, the dark head markings have a narrow pale edge. The venter has a whitish ground color with bold black or dark brown squarish or half-moon—shaped spots. These spots vary Figure 19. Sibynomorphus petersi. Dorsal views of heads. Top: left, KU 142804; right, KU 121309; both from Loja Province, Ecuador. Bottom: left, FMNH 232569; right, BMNH 1900.3.30.19; both from Cajamarca Department, Peru. These specimens show the characteristic prefrontal spots in S. petersi and some of the variation in the size, shape, and position of the spots. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle 225 in density, from series arrayed at the lat- eral edges of each ventral plate (Fig. 14) to denser arrays all across the venter and occasionally arrayed into larger patterns of streaks (e.g., Figs. 13, 16). The venter is often heavily peppered with fine dark brown. The dorsal bands do not encroach onto the ventral plates or encroach only to their extreme lateral edges. Individual scales in Sibynomorphus pe- tersi are typically finely flecked with mi- nute dark brown spots, which are best seen under magnification. These spots seemingly appear on any scales of the body (dorsal and ventral) or head, with some variation in their density among _ speci- mens. Figure 15 (top) shows the pattern on head scales particularly well; for some of the variation, see Figures 17 (bottom), IG), DH. avavel DR, Three Peruvian specimens, SMF 80048 (Fig. 20), FMNH 232569 (Fig. 16), and stomach contents associated with a Micru- rus mertensi, ANSP 31806 (see Fig. 26), lack discrete crossbands over most or all of the body. In FMNH 232569 and ANSP 31806, a series of vertebral spots marks the position of the crossbands, and indistinct lateral bars are visible when the specimens are submerged in alcohol in good light. These unbanded specimens otherwise re- tain characteristic pattern elements of this species, as elaborated below. Other Peru- vian specimens are banded with some var- iation: bold bands present the whole length of the body and tail (e.g., Fig. 21) to specimens with narrow, somewhat frag- mented bands (e.g., Fig. 27). The color pattern in preservative of SMF 80048 (Figs. 20, 22), one of two southernmost specimens of Sibynomor- phus petersi known and one of three un- banded specimens, is as follows: Dorsal pattern indistinct, with no discrete bands. Ground color (dorsally, ventrally, and on head scales) brown, each scale on all of these areas finely speckled with minute dark brown specks. A large round dark brown spot on each prefrontal positioned toward the midline (Fig. 22). A similar, but more smudged spot on the anterior end of each parietal, more smudged on the right than on the left side. Dark suture lines to many dorsal scales give the appearance of irregular flecking on the dorsum. Posteri- orly these dark edges tend to align into diagonal lines, but anteriorly they are more irregular. Nowhere are there regular lines or bands. The gular region is yellow- ish and unpatterned except for a very few minute flecks. The venter is heavily marked with irregular to half-moon— shaped dark brown spots, more or less aligned into irregular rows. In addition, very fine dark flecks cover the venter. The ground color of the venter is paler and more yellowish brown than the dorsum (medium brown). The smallest specimen examined (USNM 237040 from southern Ecuador, 184 mm SVL) still has an unfused umbil- ical scar. It has irregular crossbands over the dorsum similar to some adult patterns and otherwise does not differ substantially from the color pattern of adults. Thus, there is little or no ontogenetic change in color pattern. To summarize, several features of the color pattern of Sibynomorphus petersi are relatively constant and are helpful in iden- tifying this species: (1) Narrow zigzag crossbands with irregular edges, and often a longitudinal lateral neck bar. (2) A spot, usually rounded and of varying size and position, on each prefrontal scale. These spots are sometimes positioned toward the midline near the suture between the pre- frontals and sometimes more laterally; they are usually on the posterior half of the prefrontals (Figs. 15, 19, 22, 23). Like oth- er markings on top of the head, these spots are often surrounded by a narrow pale border. Even in specimens that otherwise differ greatly in pattern, including those lacking definitive crossbands on the dor- sum, these spots are invariably present (Fig. 22). (3) Minute dark brown speckling on individual dorsal and ventral scales (e.g., Fig. 15, top). In specimens with less distinct dorsal bands, the bands are usually 226 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 20. Sibynomorphus petersi. Dorsal and ventral views of SMF 80048 (Ancash Department, Peru). Female, 492 mm SVL. This is one of two southernmost specimens of S. petersi known. The specimen entirely lacks discrete crossbands, instead having a dorsal pattern of fine streaks and reticulations. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR *¢ Cadle DAT Figure 21. of MCZ 178047 (Bosque Cachil, Cajamarca Department, Peru). Female, 543 mm SVL. This specimen is more com- pletely banded than most specimens of S. petersi. Sibynomorphus petersi. Dorsal and ventral views Figure 22. Sibynomorphus petersi from Ancash Department, Peru. Dorsal view of head (SMF 80048). The head is relatively unpatterned compared with most S. petersi (compare Figs. 19, 23) but still retains the characteristic prefrontal spots. Figure 23. Sibynomorphus petersi. Dorsal and lateral views of head (MCZ 178047). The prefrontal spots in this specimen are exceptionally large and show the surrounding pale ring particularly well. Also visible are the minute dark specks that pepper individual scales on most specimens of S. petersi (es- pecially visible on the posterior dorsal head scales and lateral scales around the jaw angle). indicated as a discrete increase in the den- sity of this characteristic dark peppering. (4) The venter is heavily marked with bold half-moons or squarish spots (Fig. 21), of- ten in longitudinal arrays. In addition, the venter is often heavily invested with fine dark brown minute spots. Etymology. Sibynomorphus petersi was named for James A. Peters, former curator in the Division of Amphibians and Rep- tiles at the National Museum of Natural History. Hemipenis Field-everted hemipenes were available for two specimens of Sibynomorphus pe- tersi: KU 142804 (Loja Province, Ecuador) and LSUMZ 27374 (Piura Department, Peru). These organs are virtually identical and the following detailed description is based on KU 142804. The hemipenis of KU 142804 is nonbi- lobed (single) and fully capitate (Fig. 24). i) i) Figure 24. Hemipenis of Sibynomorphus petersi in sulcate view (KU 142804). Total length of the organ is approximately 13 mm. Length of the capitulum on the sulcate side approximately 7 mm. The sul- cus spermaticus divides within the capit- ulum and has more or less centrolineal branches, but each branch ultimately pass- es somewhat laterally to end at the outer (lateral) periphery of the tip of the organ. Proximally, the hemipenis has minute spines all over, although these are sparse, especially on the asulcate side. A battery comprising about three rows of enlarged hooked spines completely encircles the midsection of the organ just proximal to the capitulum. These spines are larger ad- jacent to the sulcus spermaticus than on the asulcate side. The capitulum is completely ornament- ed with papillate calyces. Calyces are well developed, but proximally, the longitudinal connections between them tend to break down so as to give the appearance of weakly developed, but densely arrayed, flounces. The calyces are surmounted by large blunt papillae. Because the proximal calyces tend to be poorly formed, there is the appearance of about three rows of pa- pillae (or papillate flounces) encircling the base of the capitulum. Probing with a fine needle suggests that these proximal papil- lae might be somewhat mineralized. A very large nude pocket (3.5 mm long) is present on the lateral side of the organ 8 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 (i.e., midway between the sulcate and asul- cate surfaces); it extends from the very base of the organ to the battery of spines on the midsection. The pocket has thick- ened borders, which are especially prom- inent at the middle of the pocket. A small, but distinct, fleshy lobe is present on the asulcate side of the pocket slightly more distal than its midpoint. Distally, the pock- et is bordered by two large hooked spines on one side and a similar spine on the oth- er. These spines are larger than any of those in the midsection battery and are po- sitioned at the proximal border of the bat- tery. In LSUMZ 27374, the spines at the distal edge of the nude pocket are not as closely associated with the pocket as in KU 142804. In particular, the large pair of spines on the asulcate edge of the nude pocket could be viewed in either specimen as having no special association with the nude pocket. Distribution and Natural History Sibynomorphus petersi is known from southwestern Ecuador (Azuay and Loja Provinces) and northern Peru south to southern Ancash Department (10°09’S) (Fig. 25; see also Fig. 43). The Ecuadorian localities and all Peruvian localities except one are on the Pacific versant of the An- des. MUSM-JEC 12884 documents that the species also occurs east of the conti- nental divide in the Rio Chotano drainage, which flows northward and joins the Rio Huancabamba to form the Rio Chamaya, a western affluent of the Rio Marafion. The documented elevational range is 1,457—2,250 m for Ecuadorian localities and 1,250—3,120 m for Peruvian localities. Two specimens of Sibynomorphus pe- tersi were obtained at the Rio Zafa Study Site (Cajamarca Department, Peru). FMNH 232569 was found at midmorning recently killed on a trail through secondary growth (brushy hillsides, agricultural land) on 2 May 1987 (early dry season). The freshness of the specimen on a hot day suggests that it might have been active during the day when killed. Another spec- kilometers ® Sibynomorphus petersi aah A Sibynomorphus vagus S XN aN = NN ~ \ ~ SS \ NY Nal m@ Sibynomorphus vagrans N=7\ =F 2 iS Ne Zin ~\ Figure 25. Distributions of Sibynomorphus petersi, S. vagrans, and S. vagus in southern Ecuador and northwestern Peru. All known localities are plotted (locality 3 represents multiple contiguous localities). See Figure 43 for amplification of the southern portion of the distribution of S. petersi (Ancash Department, Peru). Stippled area is above 1,000 m; hatched area is above 3,000 m. Numbers are locality records, as follows: Sibynomorphus petersi: Ecuador. (1) Zhila (type locality, north) and El Tablon (south); (2) Rio Catamayo valley; (3) Loja city and vicinity (multiple localities); Peru: (4) Canchaque; (5) Paraguay, Rio Chotano (Amazonian versant); (6) Rio Zana Study Site; (7) San Pablo; (8) Bosque Cachil; (9) Malvas; (10) Cajacay. Sibynomorphus vagus: Peru: (11) Huancabamba. Sibynomorphus vagrans: Peru: (12) Bellavista. 230 imen from the Rio Zafia Study Site had been swallowed head first by a Micrurus mertensi, ANSP 31806 (Fig. 26). The Mi- crurus is 573 mm SVL (total length 652 mm) and the Sibynomorphus (not sepa- rately catalogued) is approximately 465 mm SVL (total length 613 mm); the spec- imen was collected by a local resident in early November 1988 (late dry season/ transition to rainy season).''’ MCZ 178047 was collected 12 October 1992 at Bosque Cachil at the edge of the humid forest; Ca- dle and Chuna (1995) and Sagastegui et al. (“2003” [2004]) describe this forest. MUSM-JEC 12884, a road kill in poor condition, was collected 10 August 1994 near Paraguay in the Rio Chotano valley (Cajamarca Department, Peru) and is the only specimen from the Amazonian ver- sant (Fig. 25). This area once harbored ex- tensive montane humid forests, but by 1994, these forests had been reduced to scattered patches, mainly on steep hillsides and in riparian strips. Otherwise, pasture and agricultural land were extensive. The area was dissected by rocky streams with occasional waterfalls. My field notes de- scribed the forest as “rich tropical mon- tane humid forest with abundant tree ferns and large bromeliads. Canopy 20—30 m high, with some palms. Abundant epi- phytes with a diversity of orchids. Second- ary growth forest on some steep hillsides in areas of former landslides.” MUSM- JEC 12884 was collected in a deforested area. Ancillary field data accompany a few other specimens of Sibynomorphus peter- si. KU 142804 was collected 23 July 1971 at the base of an agave plant growing on a mud fence by day. KU 121309 was col- '' The Sibynomorphus is intact except that scales covering the head and approximately the anterior 25— 30 mm of the body are digested away. The specimen has very indistinct crossbands and the diagnostic as- pects of head pattern could not be evaluated. Ventral counts and body measurements were determinable within narrow limits because the exposed vertebral column and skull were still intact; however, no head scales remained. Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 26. Prey and predator: Sibynomorphus petersi (top) from the stomach of Micrurus mertensi (bottom; ANSP 31806). Both photos are to the same scale. Approximate total length of the Sibynomorphus is 613 mm. Total length of the Micrurus is 644 mm. The dorsal crossbands in the Sibynomorphus were visible only under alcohol and in good light. lected 12 June 1968 underneath a rock buried in soil along the edge of a stream. LSUMZ 27374 was found 1 December 1974 by day under a log in a partly over- grown pasture (Richard Thomas, field notes). SMF 80048 was collected 2 Feb- ruary 1997 under a rock exposed to the sun (Lehr et al., 2002: 383). The first specimen of Sibynomorphus petersi to make its way into collections from Peru is BMNH 1900.3.30.19 (Figs. 15, 19, 27), collected at San Pablo, Caja- SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle * EM “ te & zy? Pou! aug? Y 2 6 il @ q ob% Bat tee ? ly © Figure 27. Sibynomorphus petersi. Dorsal and ventral views of BMNH 1900.3.30.19 (San Pablo, Cajamarca Department, Peru). Female, 595 mm SVL. marca Department by Perry O. Simons on 5 November 1899 (the date comes from Simons’ itinerary published by Chubb, 1919; see Cadle, 1998: footnote 6). Si- mons’ South American herpetological col- lections were described by Boulenger (1899, 1900, 1901, 1902a,b), but he men- tions no snakes other than Philodryas si- monsii, Homalocranium marcapatae (= Oxyrhopus marcapatae), and two species of Elaps: E. simonsii (= Micrurus pyrrho- cryptus) and E. regularis (= M. annella- tus). I visited San Pablo and vicinity 16— 17 August 1994, and my field notes indi- cate that “all natural habitat in the broader area has been destroyed—now only agri- cultural land, secondary forest, and some brushy vegetation along fence rows . . . few streams. Vegetation included much Eucalyptus and Opuntia as well as culti- vated agaves. Terrain [was] very steep and at this time very dry.” Two females from southern Ecuador 231 (BMNH 1924.6.24.110, 550 mm SVL, 720 mm total length; BMNH 1933.6.24.110, 541 mm SVL, 711 mm total length), dates of collection unknown, each contained five large eggs (determined by palpation). The smallest specimen examined (USNM 237040, 184 mm SVL) still has an unfused umbilical scar and was collected 2 August 1954. The region of southern Ecuador and northern Peru from which Sibynomorphus petersi is known has a complex topography and, consequently, a diversity of environ- ments, including relatively arid inter-An- dean valleys (e.g., Rio Catamayo) and hu- mid forests on the Andean slopes (Chap- man, 1926). Few specific data are available for the ecological distribution of S. petersi within this region, although some localities (e.g., the Rio Catamayo valley of southern Ecuador and the slopes of the Andes in southern Ancash Department, Peru) are relatively more arid than others (e.g., Rio Zaha Study Site and the single known lo- cality on the Amazonian versant). Orcés and Almendariz (1989: 59) reported the following concerning the Ecuadorian lo- calities for S. petersi (see also the species account for S. oligozonatus): The places ... ecologically correspond to the life zone of Lower Montane Dry Forest . . . that Chap- man called Arid Temperate. Annually, rainfall reaches 500 to 1000 mm. Geographically, to this zone belong the dry plains and canyons of the inter- Andean valleys (toward the interior basins), at al- titudes between 2000 and 3000 m above sea level, in the edges of the Lower Montane spiny steppes and Lower Montane humid forest. The natural veg- etation ... has been replaced by cultivated land and there remain few areas covered by algarrobos. In Peru, several localities for Sibyno- morphus petersi are associated with rem- nants of humid forests occupying the west- ern Andean slopes or (one specimen) the more humid eastern slopes. Three locali- ties (Canchaque, Rio Zafia Study Site [= Bosque Monteseco], and Cachil) are among the best studied and most diverse humid forests on the western slopes of the Andes in Peru (Dillon et al., 1995; Sagés- 232 tegui et al., “2003” [2004]). Although these are humid forests (with abundant epi- phytes and other indicators), the region ex- periences a pronounced dry season as well. Cadle and McDiarmid (1990: 765—766) and Cadle (2005: 70-71) gave general de- scriptions and climatic regimes of the Rio Zafia Study Site, and Cadle and Chuna (1995: 21-24) described Bosque Cachil. The two specimens of Sibynomorphus petersi from the Rio Zafia Study Site are from an area of transition (1,250—1,400 m) between the humid forests on the slopes (above about 1,500 m) and the dry forest characteristic of lower elevations (<1,000 m). Whether S. petersi occurs at lower el- evations within the dry forest zone is un- known. Despite intensive sampling within humid forests between 1,500 and 2,500 m at the Rio Zafia Study Site,” the species was not obtained at these elevations, sug- gesting that it might be more characteristic of the transition zone or the dry forest it- self. The three specimens from the Rio Zaha Study Site and Bosque Cachil were collected in the early and late dry season (May, October, November). The only specimen from the Amazonian versant, MUSM-JEC 12884, was collected 10 Au- gust 1994; the Amazonian versant is more continually humid than the Pacific versant. A specimen collected by Richard Thom- as (LSUMZ 27374) was obtained as part of faunal studies undertaken by the LSUMZ beginning in 1974, when Thomas and Theodore A. Parker III established a camp near the summit of the Cordillera de Huancabamba (3,050 m) and worked on the western slope of the cordillera in late November and early December (approxi- mately the transition between the dry and rainy seasons). LSUMZ 27374 was collect- ed at 2,073 m. Parker et al. (1985: 170), ' Field work at the study site covered 55 days dur- ing the early dry season (May—June 1989) and 18 days during the rainy season (January 1991). A basecamp was established at 1,800 m within the humid forest and collections were made from about 1,000—3,000 m, with greater emphasis between 1,500 and 2,500 m. Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 under the name “Cruz Blanca,” de- scribed the general environment: The west slope was covered by a mixed evergreen forest (including Clusia, Oreopanax, Podocarpus, and Polylepis). ... Human pressure on this habitat is increasing ... and forest is being cleared from above and below. Only scattered patches of forest exist below 2150 m. A mule trail descends about 1000 m from Cruz Blanca through mature forest, continuing downward through cleared lands to Canchaque. We worked the forest along this trail almost daily during our visits. Treeline forest is dominated by Polylepis, with a canopy 4 to 6 m high. Taller trees cover the lower slopes and stream valleys. Arboreal bromeliads are conspicuous, but tree ferns are absent (or very scarce), and Chus- quea bamboo is uncommon. ... Foggy, cloudy weather occurred regularly on all three visits. Lehr et al. (2002) characterized Malvas and Cajacay (Ancash Department; SMF 80048, MUSM 20583), two localities from which they obtained specimens, as “mon- tane steppe.” Their photograph (Lehr et al., 2002: fig. 10) of the environment at Malvas shows open, mountainous terrain with shrubs, agaves, and eucalyptus trees. Lehr et al. (2002) described montane steppe as having “an arid climate with in- creases in precipitation and decrease in temperature during the [austral] summer. .. The vegetation is sparse and forest negligible in extent.” Sibynomorphus petersi is sympatric with S. oligozonatus in southwestern Ecuador (see account for S. oligozonatus). Sibyno- morphus petersi is also broadly sympatric with another dipsadine, Dipsas oreas, an the two species’ geographic ranges are very similar (Fig. 25; Cadle, 2005: fig. 23). Sibynomorphus petersi and D. oreas are known to be sympatric at two localities on the western slope of the Andes in northern Peru: the Rio Zafia Study Site (Cajamarca Department) and between Canchaque and the summit of the Cordillera de Huanca- bamba (Piura Department). However, the '3 These forests are referred to under the name “Canchaque” or “Bosque Canchaque” in much of the floral and faunal literature on this area (e.g., Cadle, 1991; Dillon et al., 1995; Sagdstegui et al., “2003” [2004]). SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle two species might be segregated by habi- tat, with S. petersi occurring in more mesic to arid habitats than D. oreas (see footnote On Sibynomorphus vagrans (Dunn) Figures 28-32 Pseudopareas vagrans Dunn, 1923: 187. Type locali- ty: “Bellavista [Cajamarca Department], Peru.” Holotype, MCZ 17284. Pseudopareas vagus vagrans: Amaral, “1929"a [1930]: 33; “1929"b [1930]: 201; “1929’c [1930]: 75. Sibynomorphus vagrans: Peters, 1960: 161, pl. IVe; 1965: 13. Orcés and Almendariz, 1989. Scrocchi et al., 1993. Carrillo de Espinoza and Icochea, 1995. Sibynomorphus vagrans is known only from the type series collected by G. K. No- ble, all from Bellavista and originally cat- alogued in the Museum of Comparative Zoology. Noble’s series originally com- prised the holotype and 26 paratypes, of which some were exchanged to other col- lections (see footnote 25). I am aware of one other specimen from a “Bellavista” whose reference to S. vagrans is highly questionable (MUSM 2258); it is discussed separately at the end of this species ac- count and is not included in the diagnosis and description of S. vagrans presented here. Notes on the Holotype of Sibynomorphus vagrans The holotype of Sibynomorphus. va- grans. (MCZ 17284) is an adult female in good condition, 557 mm total length, 425 mm SVL. Tail (132 mm) 23% of total length. Dorsal scales in 15-15-15 rows, 156 ventrals, anal scale single, and 73 sub- caudals. Preoculars 1/2, postoculars 2/2, temporals 1 + 2 + 3 on each side. Su- pralabials 10 with 4—6 touching the eye on the left, 8 with 4—5 touching the eye on the right. Infralabials 12/12. One pair of infralabials is in contact behind the mental and there are three pairs of polygonal chin shields. Diagnosis Sibynomorphus vagrans is a brownish or grayish snake with a variable number LSS (5-10) of broad dark crossbands anteriorly, much wider than the interspaces, and a posterior pattern consisting of an obscure network-like pattern, irregular flecks, or narrow, somewhat indistinct, irregular crossbands (much narrower than inter- spaces) (Fig. 28). The venter is unmarked (most specimens) or has a few dark spots that never form dense arrays. This species has a relatively low number of ventrals (149-155 in males, 154—159 in females) and relatively high number of subcaudals (SO-87 in males, 72—79 in females). The top of the head is ornately patterned and the nape has a narrow pale collar with a characteristic obtuse angular shape. Sibynomorphus vagrans is distinguished from all other species of the genus in northern South America by its combina- tion of relatively low ventral counts and high subcaudal counts (Table 1). Sibyno- morphus vagus 1s similar to S. vagrans in ventral counts but lacks well-defined cross- bands anteriorly (and they are narrower than interspaces when present), lacks a pale nape collar, and has fewer subcaudals (60—65 in males, 56—60 in females). In ad- dition, S. vagrans has a longer tail on av- erage (25-29% of total length in males, 23-26% of total length in females) than S. vagus (23-24% in males, 20-21% in fe- males). Distinct broad anterior crossbands sim- ilar to those of S. vagrans are found only in S. oligozonatus and S. williamsi. Siby- nomorphus oligozonatus differs from S. vagrans in having a foreshortened snout (compare Fig. 4 and Figs. 30, 31), fewer subcaudals, and fewer supralabials (6-7 vs. 8-9 in S. vagrans). Sibynomorphus wil- liamsi differs from S. vagrans in having 6 supralabials (3-4 touching the eye) com- pared with 8 (usually) with 4-5 touching the eye in S. vagrans, higher ventral counts (173-188), lower subcaudal counts (56-73), and in S. williamsi, the dorsal crossbands are distinct virtually the entire length of the body (sometimes dorsal sad- dles and alternating lateral blotches pos- teriorly). Sibynomorphus oneilli differs 234 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 28. Sibynomorphus vagrans Paratype (MCZ 17439). Male, 387 mm SVL. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle from S. vagrans in having more ventrals (=152 in males, 2160 in females), fewer subcaudals (62—77 in males, 57—64 in fe- males), a shorter tail (22-26% of total length in males, 20-22% in females), a less ornately patterned head, narrower anterior dorsal crossbands, and a venter heavily marked with bold black spots. Sibynomor- phus petersi differs from S. vagrans in hav- ing more ventrals (>160), a primarily brown dorsum with narrower crossbands (when present), interspaces brown (com- pared with whitish in S. vagrans), and a venter densely patterned with dark brown markings. Description Size and Scutellation. Descriptive data for Sibynomorphus vagrans are presented in Table 1. The largest specimen is the fe- male holotype, 557 mm total length, 425 mm SVL. The largest male is 543 mm total length, 387 mm SVL. Tail 25-29% of total length in males, 23-26% of total length in females. Body moderately compressed. Dorsal scales smooth and in 15—15-—15 rows. The vertebral row is not enlarged or only very subtly enlarged compared with paravertebral rows. Ventrals 149-155 (av- eraging 151) in males, 154-159 (averaging 157) in females (but see Table 1, footnote 2). Anal single. Subcaudals 80-87 (aver- aging 83) in males, 72-79 (averaging 74) in females. The loreal and a single pre- ocular border the eye in most specimens (preocular superior to the loreal); the lo- real is longer than tall. In three specimens, the loreal is vertically divided, resulting in a small triangular or trapezoidal preocular in addition to the usual one between the eye and the prefrontal scale. Postoculars 2 or, rarely, 3. Primary temporals usually 2 (occasionally 1 or 3). Secondary temporals 2 or 3. Tertiary temporals usually 3 (oc- casionally 4). Supralabials usually 8 (rarely 9 or 10) with 4—5 bordering the eye (rarely 4—6 or 3-5). Infralabials usually 11 or 12 (range 9-13). One pair of infralabials in contact behind the mental in all specimens (N = 18). Generally 3 pairs of chin shields 235 (16 specimens); occasionally 2 pairs (2 specimens) or 2.5 pairs (1 specimen). Max- illary teeth 15-17 (N = 9). The pupil is prolate (Myers, 1984) in preserved speci- mens. Peters (1960: 162—163), who examined five specimens not seen in this study (see footnote 25), noted individual variations in scutellation within Sibynomorphus va- grans that were not observed in my sam- ple. The only consequential difference be- tween Peters’ (1960) description and the one here is that Peters indicated the range of subcaudals for male S. vagrans as 74— 87 (my data: 80-87). There are a few dis- crepancies between my description and Peters’ (1960) on one hand, and the orig- inal description (Dunn, 1923: 187-188). Dunn reported a range in ventral counts (sexes combined) of 142—155, whereas Pe- ters’ count (149-160) and mine (149-159) are nearly identical. Dunn reported the maxillary tooth count as “about 12 equal teeth,” whereas my counts for nine speci- mens were 15—17 and Peters’ for 11 spec- imens were 14-15. Dunn reported that one specimen had a divided anal scale; this specimen is probably MCZ 17444, which has a partially divided anal plate. Sibynomorphus vagrans exhibits typical patterns of sexual dimorphism for colu- brids. In comparison to females, males have significantly fewer ventrals (t = 6.692, di = 17, p < 0.0001), significantly more subcaudals (¢ = 7.892, df = 17, p < 0.0001), and greater relative tail lengths (Table 1). Color in Life. Unknown. Dunn (1923: 188) used the expression “belly yellow” in describing the holotype, which might re- flect the ventral color still apparent when he described the species. The venter of the type series as of 2003 was grayish white. Color in Preservative. The color pattern of Sibynomorphus vagrans is very consis- tent among the specimens examined. All specimens have a series of five to 10 broad, bold dark brown crossbands anteriorly that become narrower and increasingly frag- 236 Figure 29. Sibynomorphus vagrans Paratype (MCZ 17442). Male, 230 mm SVL. mented posteriorly (Figs. 28, 29). The an- terior crossbands are widest middorsally (generally spanning 5-9 middorsal scale rows) and narrower laterally (spanning 3— 6 dorsal rows). The bands in some speci- mens have a distinctly darker brown nar- row border (Figs. 28-30). Bands extend ventrally to the outer edge of the ventral scales. The first dorsal crossband is sepa- rated from the dark spotting on the top of the head by a distinct narrow (2 scales or less) whitish collar just posterior to the mouth commissure; the collar connects to the pale gular region on each side and forms a characteristic obtuse angle mid- dorsally with the vertex directed posteri- orly (Fig. 30). In MCZ 17447, the first Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 crossband behind the head is interrupted middorsally and thus forms an elongated blotch on each side of the neck; the cross- bands in other specimens are complete across the dorsum. The dorsal ground col- or is dull grayish brown. The venter of most specimens is immaculate (dull gray- ish or whitish), but some specimens have a variable amount of dark brown flecks or spots, especially concentrated toward the outer edges of the ventral scales. The top of the head is ornately marked with irregular dark brown spots and blotches (Fig. 30). A consistent feature of the head pattern is a broad, irregular band extending across the posterior edge of the prefrontals, curving over the canthus, and extending ventrally to the top half of the loreal scale (Figs. 30, 31). This band is in- terrupted at the suture line between the prefrontals in some specimens (Fig. 30, lower left) and occasionally on the lateral portion of the prefrontals on one or both sides (Fig. 30, upper right). Nonetheless, it is a consistent feature of head pattern in all the specimens examined. The suprala- bial sutures are outlined with dark brown in all specimens and the same is true for the infralabial sutures in most specimens; in a few specimens, the sutures of the chin shields and other gular scales are also marked. Three small juveniles (MCZ 17446, 17451-52; SVLs of 181, 163, and 147 mm, respectively) do not differ in color pattern from adults. MCZ 17451-52 still have ob- vious umbilical scars, so the pattern prob- ably does not change ontogenetically. Etymology. Dunn (1923: 187) did not elaborate on the meaning of his species epithet, saying only that Sibynomorphus vagrans was “a species closely allied to [S. vagus| but quite distinct.” To reflect his supposition that the two species were closely related, Dunn might have used the name vagrans (originally of Anglo-French and Germanic origin) because it is a lat- inized cognate associated with the Latin word vagus (meaning wandering or roam- ing or indefinite; from vagor, to wander). SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle DT Figure 30. Sibynomorphus vagrans. Dorsal views of heads. Top: MCZ 17436 (left), MCZ 17440 (right). Bottom: MCZ 17442 (left), MCZ 17439 (right). These illustrate the ornate patterning atop the head of this species and two characteristic features of the pattern: an angular pale nape collar and a narrow dark band across the posterior portions of the prefrontal scales. The prefrontal band is complete in MCZ 17436 (top left) and shows varying degrees of interruption in the other three (see also Fig. 31). Figure 31. MCZ 17436 (top), MCZ 17442 (bottom). The ventrolateral por- tion of the dark prefrontal band is visible immediately in front of the preocular and extending to the upper edge of the loreal (see Fig. 30, left upper and lower). Sibynomorphus vagrans. Lateral views of heads. However, the similarity between the two species ends there if my supposition that the name of S. vagus refers to its color pattern (see Description in its species ac- count), because the color pattern of S. va- grans bears little resemblance to that of S. vagus. Hemipenis Inverted Hemipenis. The following de- scription of the hemipenis of Sibynomor- phus vagrans is based on the inverted or- gan of MCZ 17438, an adult male, 381 mm SVL. The organ extends to the suture between subcaudals 6 and 7. It is slightly bilobed, dividing at the level of the proxi- mal edge of subcaudal 6. Total length of organ 12.5 mm, bilobed distally for ap- proximately 1 mm. The major retractor muscle is divided proximally for 1-1.5 mm. The hemipenis is capitate, and the ca- pitulum bears well-developed papillate ca- lyces. The sulcus spermaticus is on the 238 dorsal surface of the inverted organ and divides just within the capitulum; its tips end just short of the distal tip of the organ. The undivided portion of sulcus sperma- ticus is 5—5.5 mm in length; the divided portion is 6—6.5 mm in length. Proximal to the capitulum is a broad battery of robust spines encircling the entire organ; the bat- tery is about three spines in width adjacent to the sulcus, broadening to about four or five spines in width on the asulcate side. The battery is separated from the capitu- lum by a narrow gap on the asulcate side, whereas the battery approaches the capit- ulum adjacent to the sulcus spermaticus. Proximal to the battery of enlarged spines, the hemipenis is ornamented with minute spinules, including the borders and lobes adjacent to a basal nude pocket. The basal nude pocket is 4 mm long and extends from the base of the organ to the battery of spines. The pocket is located in the dorsomedial wall of the inverted organ and thus is probably somewhat lateral in the everted organ. On its asulcate edge, the pocket is bordered by a prominent lobe on its proximal half and a somewhat smaller lobe on the sulcate edge. Everted Hemipenis. The left hemipenis of MCZ 17439 (adult male, 387 mm SVL) was everted from a preserved specimen according to techniques described by My- ers and Cadle (2003) (Fig. 32). Before eversion, the hemipenis extended to the distal half of subcaudal 8. It was slightly bilobed, dividing at the suture line be- tween subcaudals 7 and 8. Total length of the uneverted organ was 15.5 mm, divided for approximately 1 mm. The retractor muscle was divided for about 1 mm, each tip inserting onto a lobe of the hemipenis. After eversion, the hemipenis measured 16 mm total length. The divided portion of the sulcus spermaticus was 7.5 mm in length. The everted hemipenis is nonbilobed or only very slightly bilobed, fully capitate, and with the sulcus spermaticus dividing within the capitulum. The lobes appear asymmetrical in the preparation (Fig. 32), Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 32. Hemipenis of Sibynomorphus vagrans in sulcate (left) and asulcate (right) views (MCZ 17439). This hemipenis was manually everted with the use of methods detailed by My- ers and Cadle (2003). but this very likely reflects a puncture in one of the lobes and its consequent lack of full inflation. The capitulum is orna- mented with well-developed calyces sur- mounted by spinulate papillae. Proximal spinules are longer and apparently more mineralized than distal ones. A fringe of spinules borders the proximal overhang of the capitulum. The sulcus spermaticus is more or less centrolineal but the tip of each branch passes somewhat laterally to lie nearly on the lateral sides of the lobes. The tips of the branches end on the outer periphery of the lobes, short of the center of the apex of the organ. Proximally, the hemipenis is ornamented with minute spinules. The midsection of the hemipenial body is or- namented with a battery of large, relatively straight spines encircling the organ; this battery is about three spines across adja- cent to the undivided portion of the sulcus spermaticus and about four spines across on the asulcate side. Individual spines are relatively long, narrow, straight, and SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle hooked only at their tips (in contrast to the robust, strongly hooked spines typical of many colubrid hemipenes, including other Sibynomorphus). Proximal to the battery of spines on the asulcate side, but not clearly separated from the battery, are two large, hooked spines that are larger than those in the battery. A similar large spine is present near the undivided portion of the sulcus spermaticus. A large nude pocket (total length 4 mm) is present basally on the lateral surface of the hemipenis; its distal end is bordered by the large hooked spines just mentioned. The pocket is also bordered on each side by a well-developed lobe, the asulcate lobe somewhat larger than the sulcate lobe. The entire basal portion of the organ be- low the encircling battery of spines (in- cluding the lobes on either side of the nude pocket, but not the pocket itself) is ornamented with minute spinules. Distribution and Natural History Sibynomorphus vagrans is known only from the type locality, Bellavista, Cajamar- ca Department, Peru, which lies at 421 m elevation in the subhumid inter-Andean valley of the Rio Marafién (Amazonian versant) (Fig. 25). Dixon and Huey (1970: 38) described the general environment around Bellavista as “thorn scrub ... [with] standing dead Cereus [columnar cactus]”, although Dixon (personal com- munication, March 2007) stated that the area was wet enough to support coffee plantations; Dixon did not obtain S. va- grans at Bellavista 21-28 November 1968. Noble (1921) included a photographic ap- pendix which shows “The [Rio] Marafién at Bellavista”; his caption states that “the valley of the Marafién above Tutumberos [Amazonas Department on the Rfo Ma- ranon a few kilometers downstream from Bellavista] is decidedly barren except for the cane and the dense thickets that crowd the river banks” (Noble, 1921: 493). All known specimens of S. vagrans were ob- tained by G. K. Noble, who collected around Bellavista in September 1916. Two 239 specimens (MCZ 17451—52; SVLs 163 and 147 mm, respectively) have obvious um- bilical scars and probably represent young of the year. Sibynomorphus vagrans comes from a region frequently traveled by field biolo- gists in northern Peru (e.g., Dixon and Huey, 1970; Dixon and Wright, 1975) be- cause it lies along the main route from the coast to the upper Rio Marafion, the step- ping stone to the northern Peruvian Am- azon. That the species is known only from specimens collected at the type locality nearly a century ago suggests that it might have a very localized distribution. Sup- porting this suggestion is that Noble did not obtain S. vagrans at another locality close to Bellavista—Perico—only about 35 km from Bellavista on the same side of the Rio Maranon. The MCZ catalogues record more than 750 reptiles obtained by Noble at Bellavista and more than 300 at Perico. According to his field notes and Noble (1921), Noble used local resident collec- tors (“frog hounds”) to obtain specimens on the 1916 expedition to Peru. He ob- tained 27 specimens of Sibynomorphus vagrans at Bellavista (it was the most abundant snake in the collection), which suggests that the species was relatively common at that time. These specimens ac- count for more than twice as many as the next most abundant snake represented in the Bellavista collection (12 specimens of Leptodeira septentrionalis; see Duellman, 1958: 80), and 41% of the total number of snakes from that locality (66 specimens representing seven species of snakes ac- cording to the online catalogue of the MCZ collection). An Enigmatic Specimen from “Bellavista” Referred to Sibynomorphus oneilli Apart from the type series, MUSM 2258 (Fig. 33) is the only specimen known to me that has been referred to Sibynomor- phus vagrans (as reflected in the MUSM catalogues). According to the MUSM cat- alogues, MUSM 2258 was taken from the stomach of a “Micrurus mertensi” (MUSM 240 Figure 33. Sibynomorphus cf. oneilli from “Bellavista” (Peru) of unknown location (MUSM 2258). Male, 328 mm SVL. 2256) collected 25 July 1958 from “Bella- vista”; the catalogue entry for the depart- ment “Cajamarca” might have been added at a later time. The specimen is problem- atic because many of its characteristics are unlike specimens of S. vagrans of the type series (Table 1) and, in fact, are more like S. oneilli. However, no species of Micrurus is recorded from the range and elevations known for S. oneilli, as judged from dis- tribution maps and elevational data in Campbell and Lamar (2004). Only two known localities for S. oneilli, the type lo- cality at 1,646 m and Hacienda Santa Rosa at 1,800 m, are at elevations that seem Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 likely at all to harbor Micrurus species, al- though it is certainly possible that S. oneilli occurs at lower elevations as well. Resolv- ing the identity of the Sibynomorphus should go hand in hand with study of the Micrurus and the locality. I have been un- able to undertake a thorough study of MUSM 2256 in comparison with other Pe- ruvian Micrurus, and, like the Sibynomor- phus, it has some unusual characteristics. I herewith present my assessment of these specimens but leave the ultimate resolu- tion of this problem to others. The locality for the two specimens, “Bellavista,” could refer to the well-known herpetological locality in the Rio Marafién valley of Cajamarca Department, the type locality of Sibynomorphus vagrans. How- ever, a search of the GeoNet Names Serv- er for “Bellavista” in Peru recovered 70 entries, of which several were within the conceivable ranges and elevations of S. va- grans and S. oneilli. Moreover, the identity of MUSM 2256 as “Micrurus mertensi” seems questionable if the locality is the “Bellavista” of the type locality of Siby- nomorphus vagrans. Micrurus mertensi is a snake of the Pacific coastal region of northern Peru (Campbell and Lamar, 2004), whereas the very similar species, M. peruvianus, is found in the upper Rio Ma- rafon valley and, indeed, is known from the “Bellavista” of Noble’s collections (specimens in MCZ). Nonetheless, MUSM 2256 has some characteristics that are more similar to M. mertensi than M. pe- ruvianus, and the relationship (or conspe- cificity) of these two very similar species is unclear (Campbell and Lamar, 2004). MUSM 2256 is a male tricolored coral snake (without triads), 655 mm total length, 567 mm SVL, 225 ventrals, 46 sub- caudals, 1 + 1 temporals, 7 supralabials (3-4 touching the eye), 7 infralabials, 1 preocular, and 2 postoculars. The size is more than 100 mm longer than the maxi- mum reported for M. peruvianus but is typical for M. mertensi. The ventral count is 23 more than the maximum for male M. peruvianus and 4 more than the maximum SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle reported for male M. mertensi (data from Campbell and Lamar, 2004). Without fur- ther comparative study of Micrurus from northern Peru, I do not feel confident in making a call on the identity of MUSM 2256. The Sibynomorphus, MUSM 2258 (Fig. 33), is a male, 425 mm total length, 107 mm tail length (25% of total), with 15—15— 15 dorsal rows, 157 ventrals (+2 preven- trals), 72 subcaudals, anal single, 8 (4—5)/7 (3-4) supralabials, 11/10 infralabials (the first pair in contact behind the mental), a small preocular on each side superior to loreal, loreal and preocular in contact with eye, 2/2 postoculars, 2/2 primary tempo- rals, 2/3 secondary temporals, and 2 pairs of chin shields. MUSM 2258 is distinctly banded the whole length of the body with 39 black crossbands (Fig. 33). The anterior cross- bands are broader than the posterior ones; a series of crossbands at midbody is inter- rupted middorsally and tend to alternate. The first (neck) band is 4 scales wide; then follow two bands that are 3 scales wide, whereas most of the crossbands on the body are 1—1.5 scales wide. Interspaces are 1-2 scales wide anteriorly; most of the in- terspaces are 3 scales wide, occasionally 4. The top of the head is gray with an irreg- ular smudged spot on each parietal; the remainder of head is immaculate grayish. The gular region is grayish and unmarked. The venter is heavily checkerboarded with black squarish and trapezoidal markings on a grayish white ground color (Fig. 33). MUSM 2258 differs from the type se- ries of Sibynomorphus vagrans in several scale and pattern characteristics. Com- pared with male S. vagrans, MUSM 2258 has 2 more ventrals and 8 fewer subcau- dals than the next closest respective counts among male specimens of S. vagrans (Ta- ble 1). The supralabial pattern on one side, 7 (3-4 touching the eye), was observed in no other specimen of S. vagrans, although these patterns are highly variable within species. Perhaps the most striking differ- ences between MUSM 2258 and _speci- 241 mens of S. vagrans are notable differences in the dorsal and ventral patterns. For these comparisons, the contrasting char- acters of S. vagrans are indicated in pa- rentheses: in MUSM 2258 the body is dis- tinctly crossbanded for its entire length (distinctly banded only anteriorly; compare Figs. 28, 29, 33); the anterior broad cross- bands are 3-4 scales wide (generally 5 or more but occasionally 3 or 4); the top of the head is mostly uniform grayish (top of the head ornately patterned with dark ir- regular markings; Fig. 30); the venter is densely patterned with bold black mark- ings on a pale ground color (venter im- maculate or with scattered obscure dark spots). The band across the prefrontal scales typical of S. vagrans (see above de- scription) is indistinct in MUSM 2258 and could only be discerned with close inspec- tion; it would certainly not have been no- ticed had I not been looking carefully for it, whereas in typical S. vagrans, this band is very obvious (as noted also by Peters, 1960: 162). MUSM 2258 also lacks the pale angular nape collar characteristic of S. vagrans; although MUSM 2258 has a pale nape collar, it is more rounded than is characteristic of S. vagrans and is sep- arated middorsally from the parietal scales only by slightly more than 1 nuchal scale, compared with about 3 scales in S. va- grans. On the other hand, all of the scutella- tion characteristics of MUSM 2258 are typical of Sibynomorphus oneilli (Table 1), including ventral and subcaudal counts that are at the approximate midrange of the corresponding counts of male S. oneil- li. Moreover, the color pattern of MUSM 22.58, including details of the dorsal cross- bands, the heavily patterned venter, the uncomplicated pattern atop the head, and details of the pale nape collar, are much more similar to other specimens of S. oneilli (see above species account) than to S. vagrans. In conclusion, were it not for uncertainties concerning the locality and its association with a Micrurus of uncertain identity, I would unhesitatingly refer ho bo 4 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 34. Sibynomorphus vagus. Illustration of the holotype from Jan and Sordelli (1860-1881: 37¢ Livraison, pl. VI, fig. 2). MUSM 2258 to Sibynomorphus oneilli rather than to S. vagrans. Further research on the identity of the Micrurus and pos- sible identities of the “Bellavista” where the specimens were collected is needed to confidently resolve the present ambigui- ties. Sibynomorphus vagus (Jan) Figures 34—39 Leptognathus vagus Jan, 1863: 100-101. Holotype (Fig. 34), a specimen in the Museo Civico di Storia Naturale, Milan (not examined; probably destroyed during the bombing of 1943), Jan and Sordelli, 1860-1881: 37° Livraison, pl. VI, fig. 2. Type local- ity: “Hong Kong.” Cope (1868: 136) examined the holotype and stated that it is “probably South American.” Leptognathus vaga: Cope, 1868: 136. [brief descrip- tion of the holotype; see below] Pareas vagus: Boettger, 1888: 145. Pseudopareas vagus: Boulenger, 1896: 462. Werner, 1922: 200. Dunn, 1923: 187. Pseudopareas vagus vagus: Amaral, “1929"a [1930]: 33; “1929”b [1930]: 201; “1929”c [1930]: 75. Sibynomorphus vagus: Peters, 1960: 164, 1965: 14. Rossman and Thomas, 1979. Rossman and Kiziri- an, 1993. Orecés and Almendariz, 1989. Scrocchi et al., 1993. Carrillo de Espinoza and Icochea, 1995. Notes on the Holotype of Sibynomorphus vagus The holotype of Sibynomorphus vagus, illustrated by Jan and Sordelli (1860-1881) (Fig. 34), was a specimen in Milan’s Museo Civico di Storia Naturale. Jan’s (1863: 100— 101) diagnosis of Leptognathus vagus ap- peared in a key to the genus, in which L. vagus was diagnosed by the following characteristics: [15] scale rows, loreal in contact with the eye, and 8 supralabials. Jan and Sordelli’s (1860-1881) illustration of the holotype shows the following char- acteristics: body slightly compressed; smooth dorsal scales in 15 rows anteriorly and at midbody; supralabials 8 with 4—5 touching the eye on each side, 2-4 bor- dering the loreal; infralabials 10/10, the first pair in contact behind the mental; three pairs of polygonal chin shields (the first pair somewhat elongate) followed by three preventrals (the first with a partial suture); loreal and preocular bordering the anterior edge of the eye (preocular supe- rior to loreal); loreal longer than tall; post- oculars 2/2; primary temporals I/1, sec- ondary temporals 2/2, tertiary temporals 2/2 (with the upper secondary temporal fused with the upper tertiary temporal on the left side); anal single; subcaudals 50. The anterior body has a series of irregular lateral vertical markings one scale row or less in width and separated by 2—3 rows. Middorsally is a series of smaller irregular SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle flecks that are aligned with or slightly off- set from the lateral series. Top of the head without distinct pattern (stippled uniform- ly); a slight concentration of dark pigment is on suture lines of the posterior supra- labials. The ventrals and subcaudals are heavily patterned with bold half-moon markings; on the posterior body, these tend to align into longitudinal rows on the outer portion of the ventrals. The characters of the holotype as de- rived from Jan and Sordelli’s illustration are consistent with the variation within the sample of Sibynomorphus vagus I exam- ined (Table 1) except for the low number of subcaudals. On this basis, Dunn (1923) inferred that the holotype must be a fe- male, which seems a reasonable assump- tion. Cope’s (1868: 136) brief description of the holotype contains two remarks at variance with the illustration by Jan and Sordelli. Cope stated that “of the preocu- lars nothing can be said” (the illustration shows the preocular and loreal contacting the eye) and “four pairs of genials” (the illustration shows three pairs of genials, but the first preventral has a partial suture and may have been the fourth pair of gen- ials recognized by Cope). Cope (1868) first suggested that the ho- lotype of Sibynomorphus vagus came from South America, but no other specimens were reported until Dunn (1923) de- scribed the series obtained by G. K. Noble at Huancabamba, Peru (Piura Depart- ment), in 1916. Dunn (1923: 187) merely stated that Noble’s specimens “apparently represent the type of the genus [Pseudo- pareas|.” Only half a dozen specimens of S. vagus, all from the vicinity of Huanca- bamba, have been obtained since Noble collected his specimens (see Specimens Examined and Locality Records). Dunn (1923) undoubtedly associated Noble’s series with the illustration by Jan and Sordelli because of the close match to details of scutellation and pattern. How- ever, those details also largely match spec- imens of the later-described S. oneilli with fragmented dorsal crossbands, as de- 243 scribed herein (see species account and Table 1). The subcaudal count in Jan and Sordelli’s illustration (Fig. 34) is equally low for both S. oneilli and S. vagus. Only the details of the temporal scale patterns IMENeEMOlOLyperON oS. DAZUS (Ly a2) more closely match specimens from Huan- cabamba than they do specimens of S. oneilli, as indicated in the discussion of nonbanded specimens of S. oneilli from the Rio Santa valley and comparisons of them to S. vagus (see Tables 1 and 3 and the species account for S. oneilli). Thus, even though the differences in temporal scales between these two species are not absolute, the preponderance of characters discernible in the illustration by Jan and Sordelli (1860-1881) matches S. vagus sensu Dunn (1923) more closely than S. oneilli. Confirmation of the ventral count and sex of the holotype would provide cor- roboration (see Table 1), but that might be impossible if the specimen was destroyed during the bombing of 1943, as was the case with most or all of the older speci- mens in the Museo Civico di Storia Na- turale. Diagnosis Sibynomorphus vagus is characterized by a relatively low number of ventrals (144-152 in males, 151—158 in females) and subcaudals (60-65 in males, 56—60 in females), a dorsal pattern lacking well-de- fined crossbands or with narrow irregular crossbands (much narrower than_ inter- spaces anteriorly), and a venter heavily marked with large squarish spots, often concentrated laterally on the ventral plates (Figs. 34, 35). The top of the head is rel- atively unmarked or has an obscure pat- tern. Dorsal crossbands, when present, are usually not well defined, are often present only anteriorly, and usually have irregular borders. Sibynomorphus vagus differs from other species of the genus in northern South America except S. vagrans and S. oligo- zonatus in having a low number of ventral scales. Sibynomorphus vagrans is similar 244 Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 35. to S. vagus in ventral counts (Table 1) but has bold, well-defined anterior crossbands that are much broader than the interspac- es (Figs. 28, 29), has more subcaudals (80— 87 in males, 72—79 in females), and has a longer tail than S. vagus (Table 1). Siby- nomorphus oligozonatus differs from S. vagus in having well-defined bold anterior crossbands that are much broader than the interspaces (Figs. 1, 2, 4, 5) and a relative- ly unpatterned venter (heavily marked in S. vagus), and in lacking a preocular scale (present in S. vagus). Sibynomorphus pe- tersi and S. williamsi have >160 and >170 ventrals, respectively (sexes combined), and different color patterns. Male and fe- male S. oneilli have 152-168 and 163-173 ventrals, respectively. Description Size and Scutellation. Descriptive data for Sibynomorphus vagus are presented in Table 1. The largest specimen is a male, 483 mm total length, 368 mm SVL; the largest female is 408 mm total length, 326 mm SVL. Tail 23-24% of total length in males, 20-21% of total length in females. Body only slightly compressed. Dorsal scales in 15—15—15 rows, smooth. The ver- tebral row is very slightly broader to about Sibynomorphus vagus from Huancabamba, Cajamarca Department, Peru (MCZ 17422). Male, 291 mm SVL. 1.5 times as broad as the paravertebral rows. Ventrals 144-152 (averaging 149) in males, 151-158 (averaging 154) in females. Anal single. Subcaudals 60-65 (averaging 63) in males, 56-60 (averaging 57) in fe- males. Loreal and preocular bordering eye (preocular superior to the loreal). Loreal longer than tall. Postoculars 2 (rarely 1). Primary temporals usually 1 (rarely 2). Secondary temporals 2 (rarely 3). Tertiary temporals usually 2 (range 0-3). Suprala- bials 7 or 8, usually with 4-5 bordering the eye (occasionally 3-4). Infralabials usually 9 or 10 (range 8-11). One pair of infrala- bials in contact behind the mental (8 spec- imens), or a single infralabial contacts two on the opposite side (2 specimens). Chin shields in 2 pairs (N = 1), 3 pairs (N = 7), or 4 pairs (N = 2). Maxillary teeth 14-16 (N = 5). The pupil is prolate (Myers, 1984) in preserved specimens. Sexual Dimorphism. The largest speci- men is a male (Table 1), but with only two adult females and three adult males, it seems likely that this is a sampling artifact rather than an uncommon pattern of sex- ual dimorphism (none of the other species of Sibynomorphus show this unusual pat- tern, although S. petersi comes. close). Other sexually dimorphic characters in Si- SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle bynomorphus vagus follow the typical col- ubrid patterns. Compared with females, male S. vagus have significantly fewer ven- trals ({ = 2.680, df = 8, p < 0.05), signif- icantly more subcaudals (t¢ = 7.200, df = 6, p < 0.001), and greater relative tail lengths (Table 1). Dunn (1923: 187) com- mented that females have less ventral spotting than males, but that difference is not apparent considering the new material (see also Rossman and Kizirian, 1993, fig. Diy Color in Life. Unknown. Cope (1868: 136) might have seen the relatively freshly preserved holotype, which he described as “above wood-brown, with indistinct cross series of spots. Below yellowish, tessellated with brown.” Color in Preservative. The dorsal ground color is gray to grayish brown. An- teriorly, there may be a series of ill-de- fined, irregular, narrow (one scale row in width) crossbands; poorly defined reticu- lations; or a vague dark network (Figs. 35, 36). Posteriorly, the dorsum can be uni- color grayish, ‘with a poorly defined ob- scure network, or poorly defined cross- bands can extend for a variable length along the body (see Rossman and Kizirian, 1993: fig. 2). The venter is usually boldly marked with dark squarish or half-moon— shaped markings or spots on a gray to grayish brown ground color; in some spec- imens (e.g., MCZ 17420), these markings are reduced to small dark spots mainly concentrated on the anterior venter. The top of the head is gray to grayish brown, usually with a few scattered darker spots, especially on the posterior head plates. The lateral surface of head is generally un- marked and lacks extensively darkened su- ture lines on the labial scales (Figs. 37, 38). There seemingly is no substantive on- togenetic change in color pattern, as sev- eral small individuals, including one with a visible umbilical scar (MCZ 17426; 137 mm SVL), do not consistently differ from larger ones in pattern. Etymology. The common meaning of Figure 36. Sibynomorphus vagus from Huancabamba, Ca- jamarca Department, Peru (MCZ 17421). Male, 368 mm SVL. the specific epithet, vagus, is wandering or roaming. However, ane word can also mean diffuse or indefinite (as in vague). I suspect this is a reference to the diffuse dorsal pattern of Sibynomorphus vagus compared with other species of Leptog- nathus known at the time of its descrip- tion—as seen especially by comparing the illustration of S. vagus with the other spe- cies illustrated on the same plate by Jan and Sordelli (1860-1881): Dipsas incerta, S. mikanii, and the Asian colubrid Pareas laevis (all in the genus Leptognathus at the 246 Figure 37. Sibynomorphus vagus. Dorsal views of heads. Top to bottom: KU 219121, MCZ 17422, MCZ 17421. The dark spots or smudges on the parietal scales are characteristic of most specimens of S. vagus. time); these are much more boldly marked than S. vagus. Hemipenis Inverted Hemipenis. The following de- scription of the inverted organ of Sibyno- morphus vagus is based on MCZ 17422 (adult male, 291 mm SVL). The hemipenis extends to the level of the suture between subcaudals 10 and 11. It is slightly bilobed, dividing at the level of the suture between subcaudals 9 and 10. Total length of the organ 15 mm, divided distally for 2 mm. The major retractor muscle is divided for approximately 1—1.5 mm at its insertion. The organ is ornamented with minute Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 38. Sibynomorphus vagus. Lateral views of heads. KU 219121 (top), MCZ 17422 (bottom). spines basally and distally is fully capitate. The capitulum is ornamented with papil- late calyces. The sulcus spermaticus (in the dorsal wall of the organ) divides within the capitulum at the level of the distal portion of subcaudal 4; its branches end at the dis- tal tips of the lobes. The undivided portion of the sulcus spermaticus is 6 mm long; its divided portion is 7.5 mm long. Proximal to the capitulum, a battery of spines completely encircles the hemipenial body. The battery is three spines in width adjacent to the sulcus spermaticus and broadens to four or five spines wide on the asulcate side. A very long basal nude pock- et, 4.5 mm in length, extends in the dor- somedial wall of the organ (lateral surface in the everted condition) from the base of the hemipenis to the battery of spines. The distal end of the pocket is nested within the battery of spines. The asulcate edge of the pocket is bordered by a prominent lobe extending from the base to the mid- point of the pocket. The lobe is a narrow flange of tissue proximally, and is most prominent at about the midpoint of the SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle Figure 39. Hemipenis of Sibynomorphus vagus in sulcate view (MCZ 17421). This hemipenis was manually everted with methods detailed by Myers and Cadle (2003). pocket. The hemipenis proximal to the en- circling battery of spines is ornamented with minute spines everywhere except within the nude pocket. Everted Hemipenis. The left hemipenis of MCZ 17421 (adult male, 368 mm SVL) was everted from the preserved specimen with the use of techniques described by Myers and Cadle (2003) (Fig. 39). Before eversion, the hemipenis extended to the suture between subcaudals 9 and 10. It was slightly bilobed, dividing at the suture between subcaudals 7 and 8. Total length of the organ was 15.5 mm, divided for approx- imately 2 mm. The retractor muscle was di- vided for approximately 1.5 mm, each tip inserting onto a lobe of the hemipenis. At the distal end of the inverted hemipenis, a fold of the tissue that would form the apex of the everted organ extends between the inverted lobes. Consequently, the distal end of the inverted organ was rather trilobed, with a fold of the capitular tissue extending posteriorly between the branches of the di- vided retractor muscle, and larger than ei- ther lateral lobe (i.e., tissue that would be- 247 come the lobes of the everted organ). The folding of the apical tissue is perhaps only a convenient way of accommodating tissue that is incorporated into the expanding api- cal region of the everted organ; it has no apparent counterpart in the everted organ. This peculiar structure was not noted in the above-described inverted hemipenis (MCZ AZ, The everted organ was 15 mm in length and the sulcus spermaticus was divided for 7 mm. The hemipenis is fully capitate, with only a very slight indication of bilo- bation (perhaps nonbilobed if fully ex- panded and inflated, which was not at- tempted because of the delicacy of the preparation). The sulcus spermaticus di- vides within the capitulum and its tips are more or less centrolineal in orientation, but they pass somewhat laterally and end at the lateral edges of the apex of the or- gan. The capitulum is ornamented with well-developed papillate calyces. Proximal to the capitulum is a battery of large hooked spines encircling the organ, three to four spines across all around. A large basal nude pocket (3.5 mm in length) is on the lateral side of the organ. It is bordered on its asulcate side by a large lobe and to- ward its distal end by a pair of large spines, one on each edge. These spines are larger than any in the battery encircling the or- gan. The base of the hemipenis below the battery of spines, including the lobes bor- dering the nude pocket (but not the tissue within the pocket) is ornamented with mi- nute spines. The hemipenes of Sibynomorphus va- gus and S. vagrans are very similar to one another but differ in the shape of their en- larged spines. The spines on the organ of S. vagus are comparatively shorter and more robust, and the entire spine forms a curved hook, whereas the enlarged spines on the hemipenis of S. vagrans are rather narrow, somewhat straight, and hooked only at their distal tip. Distribution and Natural History Sibynomorphus vagus is known only from the vicinity of Huancabamba, Piura 248 Department, Peru (Fig. 25). The docu- mented elevational range is 1,810 to 1,920 m. Huancabamba lies in the steep valley of the Rio Huancabamba, a tributary of the Rio Marafon and thus is on the Am- azonian versant (and not the Pacific slopes as occasionally stated; e.g., Rossman and Thomas, 1979: 6). Noble (1921) described the general en- vironment surrounding Huancabamba during August 1916 (when the series in the MCZ was collected) as follows: Huancabamba lies on a slope overlooking a mountain torrent. The country on either side is steep and barren. Still, there are pockets of less dry areas and in these we found tropical animals and plants. Huancabamba seemed to be the mingling point of several diverse biotas. Bananas and cacti grew side by side ... the dominant facies of the fauna seemed tropical, although much out of place in such a barren region. Parker et al. (1985: 171) described the area in June/July 1980: Huancabamba lies in a narrow rain-shadow val- ley. ... Although this arid valley is densely settled and intensely cultivated, a few small remnants of desert scrub remain within walking distance of the city. The vegetation |of this desert scrub] is com- posed of dense shrubbery and scattered Acacia trees and clumps of columnar cacti. The farmlands above this dry terrain eventually give way to humid forest at about 2150 m. Noble’s field notes (AMNH Herpetolo- gy Department archives) indicate that he worked in and around Huancabamba from 7 to 27 August 1916. There are references in his notes during this period to collecting birds (most references), small mammals, and frogs (including Gastrotheca), but no mention of snakes. His entry for 27 August includes the comment “There are so few birds or beasts of interest about Huanca- bamba that I have decided to move on to some place else.” “4 The AMNH Herpetology Archives contain only Noble’s notes for July-August of the 1916 Harvard Peruvian Expedition, with coverage to just past Huancabamba. The whereabouts of the remainder of the notes (through September, covering the impor- tant Noble localities Bellavista and Perico) is un- Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Some literature records for “Sibynomor- phus vagus” are questionable or errone- ous. Lehr et al. (2002: 134) referred two specimens from the Pacific slopes of southern Ancash Department, Peru (SMF 80048, MUSM 20583) to S. vagus; these specimens are S. petersi (see above species account). Specimens from Ancash, Caja- marca, La Libertad, and San Martin De- partments were also identified as S. vagus (Carrillo de Espinoza and Icochea, 1995). These records were presumably based on the specimens I refer to S. oneilli (see spe- cies account in New Material of Sibyno- morphus oneilli from Ancash and Libertad Departments, Peru), but I did not see specimens from San Martin." Very little is known of the natural his- tory of Sibynomorphus vagus. Six speci- mens were taken from under rocks in a semiurban setting (“outside a tourist ho- tel”) or in a cleared field in January 1991 (Rossman and Kizirian, 1993). KU 135180 was collected 10 May 1970, and the MCZ series was collected in August 1916. The smallest specimen examined, MCZ 17426 (SVL 137 mm), has the umbilical scar be- ginning to fuse. Sibynomorphus williamsi Carrillo de Espinoza Figures 40-42 Sibynomorphus williamsi Carrillo de Espinoza, 1974: 3. Holotype, MUSM 2170 (original number “Of. 0492” given in the original description)'® (Fig. 40). Type locality: “Jicamarca. Radio-observatorio. Dis- known. A possible repository is the archives of Har- vard University, but their existence was not pursued during this study. The portion of the notes at AMNH contain almost no information on the collections, es- sentially being a diary of the journey rather than ac- tual field notes. '° If the specimen(s) from San Martin are found and also prove to be Sibynomorphus oneilli, it would be only the second record, in addition to the holo- type, from east of the Rio Maranon. '® The herpetological collections of the MUSM originally used separate numbering systems for dif- ferent taxonomic groups; hence “Of.,” referring to “Ofidios.” With reorganization of the collection com- mencing in the 1980s, all specimens were renum- bered in a continuous series. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle &, “aQy% 249 Figure 40. Sibynomorphus williamsi. Holotype (MUSM 2170) from Jicamarca (vicinity of Lima). Male, 404 mm SVL. trito Lurigancho, Provincia Lima, Departamento Lima” [Peru]. Sibynomorphus williamsi has heretofore been reported only from coastal “lomas” formations in the Department of Lima (Carrillo de Espinoza, 1974); see Distri- bution and Natural History below. Speci- mens in the MUSM show that the species has a broader distribution in the central coastal region of Peru, including Andean foothills and slopes. Notes on the Holotype of Sibynomorphus williamsi The type series of Sibynomorphus wil- liamsi consisted of a male holotype (MUSM 2170) and six female paratypes. I studied the holotype and four of the six paratypes. Three of the paratypes were from the type locality (MUSM 2173-74; the other not seen), one from the Lomas de Lurin (MUSM 2171), one from the Lo- mas de Pachacamac (not seen), and one from the Lomas de Lachay (MUSM 3400). These localities, except the Lomas de La- chay, are in the immediate vicinity of the city of Lima; the Lomas de Lachay are on the coast approximately 80-90 km NW of Lima (see Fig. 43). The holotype of Sibynomorphus wil- liamsi (MUSM 2170; Fig. 40) is an adult male in good condition, 530 mm in total length, 126 mm tail length (tail 24% of to- tal length). It has 15-15-15 dorsals, 174 ventrals (+2 preventrals), single anal, 67 subcaudals, 0 preoculars (loreal and pre- frontal border the anterior edge of the eye), 3/2 postoculars, 6/6 supralabials with 3-4 bordering the eye on each side (the third supralabial on the right side is divid- ed horizontally to form a “subocular”), 10/ 11 infralabials, 3 + 4 + 4 temporals on each side, 3 pairs of polygonal chin shields, 1 pair of infralabials in contact behind the mental, and 13 maxillary teeth. It has 35 dorsal crossbands or blotches. The right hemipenis is everted. I am unable to reconcile my data for the holotype with some scale counts and mea- surements given by Carrillo de Espinoza (1974: 4, table 1), who reported 186 ven- trals, 63 subcaudals, and 10/10 infralabials in the holotype, and body measurements of 545 mm (“cuerpo”), 118 mm (“cola”), and 20 mm (“cabeza”). The high number of ventrals and low number of subcaudals (see Description) suggest that data for one of the female paratypes were mixed up 250 with those of the holotype, but data pre- sented for the paratypes (Carrillo de Es- pinoza, 1974: table 1) do not closely match my data either. Nonetheless, our observa- tions of the holotype are of the same spec- imen, as indicated by the sex and locality of the specimen and other details, such as the divided right third supralabial (shown in Carrillo de Espinoza, 1974: fig. 4), the everted right hemipenis, and details of banding pattern (Fig. 40; Carrillo de Es- pinoza, 1974: fig. 1). Diagnosis Sibynomorphus williamsi is character- ized by a high number of ventral scales (173=182 in males, 1SI=188) in’ females): usually 6 supralabials with 3—4 bordering the eye, a relatively short tail (21-24% of total length in males, 19-22% in females), and a head distinctly wider than the neck, with the first dorsal crossband usually ex- tending up onto the wide posterior part of the head. The dorsum is strongly cross- banded from head to tail (835-48 bands: posterior bands might be broken into dor- sal and lateral blotches, offset to varying degrees, but they retain their boldness). Anterior crossbands can be twice the width of posterior crossbands, but they are always wider than the pale interspaces, which are 1—2 scale rows wide the length of the body. Sibynomorphus williamsi differs from other species of the genus in western South America as follows (see also previ- ous species accounts). Sibynomorphus va- grans and S. vagus have fewer ventral scales (<160 in each case) and different color patterns. Sibynomorphus oligozona- tus has a short, blunt snout; a head that is not distinctly wider than the neck; and fewer ventrals in males (145—163 in three specimens). In S. williamsi, the dorsal crossbands (including posterior blotches) are bold the entire length of the body (Figs. 40, 41), whereas in S. oligozonatus, the anterior crossbands are much more well defined than the posterior blotches. Sibynomorphus petersi and S. oneilli differ Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Figure 41. Sibynomorphus williamsi. Representative speci- mens in dorsal view. Top: MUSM 3025 from Ancash Depart- ment (female, 427 mm SVL). Bottom: MUSM 3137 from Lima Department (male, 435 mm SVL). from S. williamsi in color patterns (see their species accounts) and in having 8 su- pralabials with 4-5 touching the eye. Description Size and Scutellation. A summary of character data for Sibynomorphus william- si is presented in Table 1. The largest specimen is a female 718 mm total length, 561 mm SVL; largest male is 563 mm total length, 435 mm SVL. Tail 21-24% in males, 19-22% of total length in females. Body somewhat compressed; the head is distinctly wider than the neck. Dorsal scales in 15-15-15 rows. Vertebral row not, or only scarcely, enlarged. Ventrals 173-182 (averaging 177) in males, 181— 188 (averaging 184) in females. One to three preventrals precedes the ventral SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle plates. Anal single. Subcaudals 67—73 (av- eraging 69) in males, 56—66 (averaging 62) in females. Loreal Anil prefrontal border- ing anterior edge of eye (no separate pre- ocular); loreal longer than tall. The first su- pralabial usually makes broad contact with the loreal (occasionally only narrow con- tact); supralabials 2 and 3 also border the loreal. Postoculars 2 (occasionally 3). Pri- mary temporals QO" oO: secondary tempo- rals usually 3 or 4 (rarely 2), tertiary tem- porals usually 3 or 4 (rarely 5). Supralabi- als 6 with 3-4 touching the eye (rarely 7 with 3-4 touching the eye). Infralabials usually 10 (range 8-11). One pair (N = 17 specimens) or two pairs (N = 1) of infra- labials in contact behind the mental. Chin shields in three pairs (N = 11) or two pairs (N = 3). Maxillary teeth 12-14 (N = 6). The consistency of the fused preocular— prefrontal is remarkable in this species com- pared with some other Dipsadini (e.g., Si- bynomorphus oneilli, S. petersi, S. vagrans, and various Dipsas spp.; Table 1; Cadle, 2005; Cadle and Myers, 2003). No separate preocular is present in any specimen. Three specimens have unusual configurations of the loreal and surrounding scales: in MUSM 19236, the loreal is expanded posteriorly so that the prefrontal contacts the eye very nar- rowly, MUSM 3137 and MUSM 2173 show the same condition on one side only; addi- tionally, on the contralateral side of MUSM 2173, the loreal contacts the supraocular, oc- cluding contact between the prefrontal and eye. A few other variations were observed: one specimen (MUSM 19236) has an irreg- ular row of suboculars on the left side sep- arating the supralabials from the eye, and a single subocular on the right side of the ho- lotype is formed by horizontal division of the third supralabial (see Carrillo de Espinoza, 1974, fig. 4). No consistent or substantive differences in either scutellation or color pattern were noted between the Ancash and Lima Department series. Sibynomorphus williamsi shows typical patterns of sexual di- morphism for colubrids. In comparison to females, male S. williamsi have significantly fewer ventrals (¢ = 4.560, df = 13, p < 251 : ie 3 53 ai A, os i A. sg fr ? m A ‘'. : .. = & it + \ \y i ie m+ Eom }- 7) mae 1. r ae : , a = \aaeA, Se e Bee 8 eee Ke 5 Me y e os< , a Kee, » & At . teers Sees, Fi a : w Rh. Lae oo st SS = Co 24 Figure 42. Sibynomorphus williamsi. Dorsal and ventral views of a specimen from Lima Department, MUSM 10034. Female, 531 mm SVL. 0.001), significantly more subcaudals (t = 4.365, df = 12, p < 0.01), and greater rel- ative tail lengths (Table 1). Color in Life. Unknown. In a relatively recently preserved specimen (MUSM 10034; Fig. 42) the dorsal crossbands are blackish and the interspaces are medium brown with darker brown spots and stip- ling. The venter is grayish brown, invest- ed with dark grayish irregular markings across each ventral plate. Color in Preservative. Sibynomorphus williamsi exhibits a consistent color pat- tern. The dorsum is strongly crossbanded anteriorly, although the bands tend to 252 break up into dorsal and lateral blotches, sometimes offset, on the posterior half to two-thirds of the body. The first dorsal crossband extends well up onto the wide part of the head in most specimens (Figs. 4], 42). At their widest point (dorsal half), individual crossbands are 6 to 9 dorsal scale rows wide on the anterior part of the body, reducing to 3 or 4 rows wide pos- teriorly. Ventrally, individual crossbands are 2 to 4 rows wide, and they extend onto the lateral edges (up to approximately one- third) of the ventral plates. The interspac- es are very narrow (1 or 2 scale rows wide) the length of the body. The total number of crossbands is 35-48. The dorsal cross- bands and interspaces are highly contrast- ing in most specimens, apparently very dark brown or black in life with much pal- er interspaces. Ventral patterns of the specimens examined were of three forms: (1) pale whitish with fine dark peppering; (2) rather dusky, especially posteriorly; or (3) whitish or dusky with a posterior series of large rounded or squarish spots on the lateral edges of the ventral scales. Occa- sionally dark pigment was concentrated in narrow lines across the belly, creating a ladder-like appearance. The head of Sibynomorphus williamsi is brown to grayish with blackish markings. Usually some of the labial scale sutures are lined with black and often irregular dark spots are present on the top and sides of the head, especially the parietal scales. The color pattern of the two smallest in- dividuals, MUSM 3400 (243 mm total length, 198 mm SVL) and MUSM 2867 (199 mm total length, 159 mm SVL), is similar to that of adults, which indicates that little, if any, ontogenetic change in Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 color pattern occurs. The venter of these specimens is very dusky and with a dark ladder-like pattern formed by a dark irreg- ular line across each ventral. The tops of their heads are also very dark. Etymology. Sibynomorphus williamsi was named for Ernest E. Williams, former curator of herpetology and professor of bi- ology at the Museum of Comparative Zo- ology, Harvard University. Hemipenis The following description of the hemi- penis of Sibynomorphus williamsi is based on the fully everted right organ of the ho- lotype (MUSM 2170); a sketch of the or- gan was given in the original description (Carrillo de Espinoza, 1974). Total length of the organ 12 mm, bilobed distally for about 1.5 mm. The hemipenis is slightly bilobed, fully capitate, and with the sulcus spermaticus dividing within the capitulum. The capit- ulum is set off from the proximal portion of the organ by a well-developed capitular groove. Calyces bearing fleshy papillae or- nament the capitulum, but the papillae on the more proximal calyces are spinulate (i.e., have mineralized tips). The sulcus spermaticus divides within the proximal edge of the capitulum and its branches are centrolineal, but distally they pass to the lateral edges of the hemipenial lobes and end at their peripheral distal tips. Proximal to the capitulum is a central section or- namented all around the organ with three to four rows of stout spines. On the asul- cate side of the hemipenis is a pair of en- larged spines just proximal to the midsec- tion battery. The basal area below the cen- tral spinose section is ornamented with mi- => Figure 43. Distributions of Sibynomorphus williamsi, S. petersi, and S. oneilli in Lima and Ancash Departments, Peru, and adjacent La Libertad Department (locality 12 only). Inset shows the area covered by the larger map (outlined department bound- aries are Lima to the south, Ancash to the north). Numbers refer to specimen localities and letters to physical features as follows: Localities: S. williamsi: (1) Lima and vicinity: Canto Grande, Lomas de Atocongo, Lomas de Lurin, Jicamarca (type locality), Lomas de Pachacamac; (2) Santa Eulalia; (3) Lomas de Lachay; (4) Huanchay; (5) Yumpe (west) and Huayllacayan (east). Sibynomorphus petersi: (6) Cajacay; (7) Malvas. Sibynomorphus oneilli: (8) Yuracyacu; (9) Huaraz; (10) Huaylas; (11) Hacienda Santa Rosa; (12) Hacienda Ticapampa. Physical features: (A) Rio Maranon; (B) Cordillera Blanca; (C) Rio Santa; (D) Cordillera Negra. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle 2000 m contour above 4000 m 50 100 km @ Sibynomorphus williamsi A Sibynomorphus petersi m = Sibynomorphus oneilli 254 nute spines. A large basal nude pocket is present on the lateral surface of the organ, but is positioned almost on the asulcate side (i.e., it seems displaced far more to- ward the asulcate side of the hemipenis than in other species of Sibynomorphus). The pocket is 5.5 mm long and has a lobe at its distal end on the asulcate edge and a large spine at the distal end on the sul- cate edge. Distribution and Natural History Sibynomorphus williamsi is known from coastal “lomas formations” from the vicin- ity of Lima north to the Lomas de Lachay, from the Andean foothills east of Lima, and from the Andean slopes of southern Ancash Department, Peru (Fig. 43). The known localities span 10 m to at least 2,990 m, but possibly as high as 3,626 m elevation—an extraordinary elevational range matched in few other snakes of which I am aware. The uncertainty in the upper elevational extent is based on spec- imens from Huayllacayan, Ancash Depart- ment, which could have come from lower elevations than the town itself (see foot- note 27). However, there is a dichotomy in the elevational range of localities in Lima Department (10—1,036 m) compared with Ancash Department (1,857—?3,626 m). This might reflect either collection biases or environmental differences, but the ab- sence of information on microhabitats or capture circumstances for Sibynomorphus williamsi precludes adequate evaluation. The new records of Sibynomorphus wil- liamsi in the Andean foothills and slopes are significant in showing that the species is not restricted to the coastal lomas for- mations but could have a broader distri- bution in the Peruvian coastal region and western Andean slopes. Given the extreme environmental conditions that pertain to much of the Peruvian coastal region, a snake such as S. williamsi might be infre- quently encountered on the surface and could be overlooked through much of the region. Perhaps significantly, many speci- mens have come from the heavily popu- Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 lated region around Lima where chance encounters are more likely to record the species’ presence. No detailed observations seem to have been recorded for the circumstances of cap- ture for any specimen of Sibynomorphus williamsi. The lomas formations, from which some specimens have come (e.g., the lomas of Lurin, Jicamarca, Pachacamac, and La- chay), are plant communities with a highly endemic flora characteristic of the Peruvian and Chilean coastal region (Rundel et al., 1991). These communities develop where there is sufficient topographic relief to in- tercept the low, dense clouds characteristic of the winter months, thus providing local mesic environments in this otherwise arid region. The formations from the vicinity of Lima and north are floristically among the most diverse lomas because of the increased moisture in this part of the coast compared to farther south. Fog drip at the Lomas de Lachay, from which S. williamsi is known, can reach nearly 700 mm per year (Rundel et al., 1991), which is far more than the amount of actual precipitation. Plant com- munities at Lachay are a diverse assemblage of herbaceous and semiwoody vascular plants, lichens, terrestrial and arboreal bro- meliads, and dense stands of small trees with abundant epiphytes (mosses, lichens, ferns) (Rundel et al., 1991: 32). The topog- raphy interacts with the prevailing climate to produce a wide array of microenviron- mental conditions, including moist woody ravines, lichen-covered rocky terrain, and sandy shrub habitats. Whether S. williamsi is restricted to particular microhabitats with- in this region is unknown. Also, the Lomas de Lachay are topographically and environ- mentally more diverse than some other lo- mas localities for S. williamsi (at Lachay the lomas reach nearly 1,000 m elevation). For example, the Lomas de Lurin and Lomas de Pachacamac are of much lower relief and with much simpler and more xeric vegeta- tion communities. Referring to localities in the vicinity of Lima from which the type series was ob- tained, Carrillo de Espinoza (1974: 5) stat- SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle ed that Sibynomorphus williamsi was “a species adapted to a xerophytic environ- ment where Tillandsias and other decid- uous plants grow; of nocturnal habits, pos- sibly semiarboreal because of the com- pressed nature of its body.” She did not elaborate on the observations suggesting nocturnal habits. One locality for S. wil- liamsi, Santa Eulalia (Lima Department, 1,036 m), has been characterized as a semidesert (Koepcke, 1961: 35). Because they are above the level of in- terception of the winter clouds and fogs characteristic of the Peruvian coast, the lo- calities for Sibynomorphus williamsi in An- cash Department (Fig. 43) are notably more arid than the localities at lower elevations in Lima Department. The region has been characterized as montane or bush steppe (Koepcke, 1961; Lehr et al., 2002). Three specimens from Lima Depart- ment with dates of collection were taken in January, February, and October. Six specimens from Ancash Department were collected in February, March, or April. These months, particularly January—April, correspond to the warmer, drier months along the central coastal region of Peru. Dense clouds form along the Peruvian coast from May to October, often with a ceiling of only 150-300 m (Rundel et al., 1991: 9), and cool weather prevails. One female (MUSM 10034, month of collection not recorded; 531 mm SVL; Fig. 42), contained six eggs, as determined by palpation through the body wall. The smallest individual, MUSM 2867 (157 mm SVL) was collected 9 January 1978. Leptognathus atypicus Cope, 1874: Incertae Sedis Leptognathus atypicus Cope, 1874a: 65-66. Holotype unknown. Type locality: “some portion of the Pe- ruvian Andes, from an elevation of twelve thousand feet.” Pseudopareas atypicus: Boulenger, 1896: 463. Wer- ner, 1922: 200. Sibynomorphus turgidus (part): Amaral, “1929"a [1930]: 33. Peters, 1960: 158, 1965: 13. Peters and Orejas-Miranda, 1970: 278. The identity of Leptognathus atypicus 9) ay ey Cope (1874a) has not been conclusively resolved. Neither Peters (1960: 160) nor I located its holotype in the Academy of Natural Sciences of Philadelphia, and it is not among the Cope types acquired by the American Museum of Natural History (Charles W. Myers, personal communica- tion, 1998; see Myers 1982: footnote 7) or the National Museum of Natural History (Cochran, 1961). Cope’s description of Leptognathus atypicus in its entirety follows (Cope, 1874a: 65-66): Belonging to sec[tion] vi of my monograph of this genus [Cope, 1868: 107-108], that is, with smooth equal scales, and short geneials separated by but one pair of labials from the symphyseals. The body is rather stout, and, unlike most of the genus, nearly cylindric; the neck is not contracted, but the head is wide and flat, and the muzzle short. Scales rather wide; in fifteen rows. Rostral plate subtriangular; a subquadrate loreal; no preocular; postoculars 2; temporals 2 X 3. Frontal and pari- etals broad and short, the latter truncate. Superior labials six, eye over third and fourth, fifth and sixth elongate. Inferior labials ten, mostly transverse; four pairs of geneials, all except the first broader than long. Tail short, m. .047 in length, from a total Oley 243) Color pale, with black transverse spots, which are wide anteriorly (the second covers seven trans- verse rows of scales) and become gradually narrow- er, having a width of only two cross-rows on the hinder part of the body. Posteriorly their lateral ends are broken off, and alternate with the dorsal portion. A few small blotches on the ends of the gastrosteges. The description is cursory, even for Cope. For example, he reported neither ventral nor subcaudal counts. Several characters (15 [midbody] scale rows, a rather stout, cylin- drical body, a short muzzle, a loreal but no preocular, and a pale dorsum with wide black blotches anteriorly, narrower posteri- orly) suggest a species of Sibynomorphus, but it is conceivable that it could also be a species of Dipsas. Cope’s (1868) concept of Leptognathus included species now appor- tioned among Dipsas, Sibon, Sibynomor- phus, and Tropidodipsas, and his “sections” of the genus were not based on characters now considered to be systematically infor- mative, that is, “smooth equal scales, and 256 short geneials separated by but one pair of labials from the symphyseals [= mental scale].” Amaral (“1929’a [1930]: 33) considered Leptognathus atypicus identical to Siby- nomorphus turgidus, and Peters (1960: 158) formally placed it in the synonymy of S. turgidus, where it has remained since (e.g., Peters and Orejas-Miranda, 1970). Both authors were perhaps guided by Cope’s referral of the species to his “sec- tion vi” of Leptognathus, which, at the time L. atypicus was described, included only S. turgidus (Cope, 1868: 108). Peters (1960: 160) noted that the differences be- tween S. turgidus and the type of L. atyp- icus as described by Cope, particularly la- bial counts, appeared as variants within his sample of S. turgidus. Peters (1960: 160) also questioned Cope’s locality because of the high elevation (3,660 m), which was perhaps understandable because the only montane species of Sibynomorphus known at the time, S. vagus, was known only from <2,000 m elevation. Nonetheless, because scutellation and color patterns within this group of snakes are highly variable, Cope’s brief descrip- tion precludes a rigorous determination of the identity of Leptognathus atypicus. For example, with little amplification, the char- acters given by Cope also describe Siby- normorphus oneilli, S. oligozonatus, and S. williamsi, which were unknown to Peters, and these are more logical on geographic grounds than S. turgidus. The known up- per elevational limits for these species are, respectively, about 3,500, 2,250, and (pos- sibly) 3,600 m (see previous species ac- counts). One character in Cope’s descrip- tion, six supralabials with the third and fourth touching the eye, is uniquely shared by S. oligozonatus and S. williamsi among the northern species of Sibynomorphus (Table 1). Given the lack of details in Cope’s description, the identity of L. atyp- icus will remain questionable unless the type is rediscovered. I see no reason to question the type lo- cality of Leptognathus atypicus as Peters Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 did, although the elevation (3,660 m) is ex- ceptionally high for a species of Sibynomor- phus or the related genus Dipsas. Nonethe- less, we still have much to learn about snake distributions in the Andes. For example, the discovery of a new population of S. oneilli extended the known elevational range of that species from 1,645 m to over 3,000 m (Rossman and Kizirian, 1993), and S. wil- liamsi might occur well over 3,000 m (but see footnote 27). Ancillary information in Cope’s descrip- tion lend credibility to the locality data. Cope (1874a) stated that the type of L. atypicus was presented to the ANSP “along with two fishes.” Cope (1874a) de- scribed one of the fishes as Protistius se- motilus (now Basilichthys semotilus) and stated that the other was Trichomycterus dispar (Tschudi). The holotype of Protis- tius semotilus is still extant (ANSP fish col- lection 14404), and both species of fish oc- cur in the Andes of Peru. Thus, unless there was a mislabeling or mix-up before these specimens were presented to Cope, it seems likely that the type of L. atypicus is also from Peru. Other circumstantial details suggest, but cannot prove, that Peru might be the ori- gin of Leptognathus atypicus. Cope’s work on the South American herpetofauna in the 1870s was dominated by the collec- tions of the naturalist James Orton (1830- 1877), primarily from Peru. Orton’s second expedition in 1873 traversed the Peruvian Andes, including a “side excursion” to Lake Titicaca (Orton, 1875: xiv). Indeed, Cope (1874a: 70-72) included an appendix describing two reptiles denoted explicitly as sent by Orton, and a paper later the same year (Cope, 1874b) described the bulk of Orton’s 1873 collections. In addi- tion to the holotype of Protistius semotilus, the ANSP fish collection has two speci- mens of Trichomycterus dispar collected by Orton in 1873 at Tinta (Cuzco Depart- ment; ANSP 21251, 21355), which could well be the specimens of that species to which Cope referred (Cope, 1874a: 66) (one reading of the expression “two fishes” SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle is “two kinds, or species” of fishes, rather than two specimens). Trichomycterus dis- par is not mentioned in the later paper covering most of the fish collection (Cope, 1874b). It is plausible, and perhaps likely, that the holotype of L. atypicus was sent to Cope by Orton and that it was obtained on his 1873 expedition to Peru. HEMIPENIAL MORPHOLOGY IN SIBYNOMORPHUS AND OTHER DIPSADINI As outlined in the following section, I consider the Dipsadini to comprise species in four nominal genera of Neotropical molluscivores: Dipsas, Sibon, Sibynomor- phus, and Tropidodipsas. These are part of a larger clade, the Dipsadinae (sensu Zah- er, 1999), essentially the Central American xenodontine clade of Cadle (1984a,b, 1985) with slightly different composition and many taxa listed “incertae sedis” by Zaher (1999). This is not equivalent to the Dipsadinae of Peters (1960), a much smaller clade whose composition is dis- cussed in the next section. As part of Dips sadinae, Sibynomorphus, along with spe- cies of several other genera (e.g., Dipsas, Atractus, Leptodeira, and Imantodes), is a South American representative of a clade with a great diversity of species throughout tropical Mesoamerica. The Dipsadinae has been characterized on the basis of hemipenial synapomor- phies. I here summarize the morphology of the hemipenes of Sibynomorphus, as gleaned from the previous species ac- counts, put their variation in the context of other Dipsadinae and Dipsadini, and characterize interspecific variation in hemipenial morphology among species of Sibynomorphus. Myers and Cadle (1994) suggested three hemipenial characters as synapomorphies of Dipsadinae, despite some variation within this group and con- vergent occurrences in other clades of col- ubrids (Zaher, 1999): (1) unicapitation, (2) either unilobed or slightly bilobed, and (3) sulcus spermaticus dividing distally within the capitulum. 257 Hemipenes of at least 27 species of Dip- sadini, including species of all four genera, have been described (Cadle, 2005; Cadle and Myers, 2003, and herein), although the descriptions vary in the degree of de- tail presented. Nonetheless, as inferred from these published descriptions and il- lustrations, hemipenes of the Dipsadini are very similar in ornamentation. Cadle and Myers (2003: 14-15) briefly reviewed these descriptions, described the organs of Dipsas nicholsi and D. andiana, and cor- rected a few observations in previously published descriptions (see also Cadle, 2005). Hemipenes of Dipsadini include the following characters in addition to those considered synapomorphies of Dip- sadinae: (1) calyces with well-developed papillae ornamenting the capitulum (the apillae on the proximal calyces are spi- nulate); (2) a well-developed capitular groove; (3) a battery of enlarged spines of varying width (generally three to four rows of spines across but varying from one to five) encircling the hemipenial body prox- imal to the capitulum; (4) an exceptionally long nude pocket extending from the base of the hemipenis to, or nearly to, the bat- tery of spines around the midsection; and (5) usually two or more basal spines that are larger than, and separated from, spines in the midsection battery. Proximal to the battery of enlarged spines on the midsec- tion, hemipenes of Dipsadini are orna- mented with minute spines overall. A brief survey of published descriptions and un- published data indicates that these char- acters commonly appear elsewhere among Dipsadinae (e.g., see illustrations of Rhad- inaea in Myers, 1974), although compara- tive studies could reveal differences in their manifestation. In the following sec- tion, I briefly compare hemipenes of the northern species of Sibynomorphus and compare variation in these and other char- acters in Sibynomorphus vis-a-vis other Dipsadini and Dipsadinae. Comparison of the Hemipenes of Sibynomorphus Hemipenes of the five species of north- ern Sibynomorphus for which the organs 258 are known (all except S. oneilli) are very similar to one another, but they seem to offer few characters that link two or more of the northern species of Sibynomorphus to the exclusion of others. References to Sibynomorphus in the following discussion refers explicitly to these five northern spe- cies only. No other hemipenes of Sibyno- morphus have been described in sufficient detail to make meaningful comparisons. Cei (1993) and Scrocchi et al. (1993) brief- ly described organs of several southern species. (1) Bilobation and Proportions of the Hemipenial Body. The organs are nonbi- lobed (S. petersi) or slightly bilobed. The bilobation is greatest in S. williamsi and S. vagus (approximately 12-13% bilobed compared with <7% in S. vagrans and S. oligozonatus). The degree of bilobation varies somewhat among species of Dipsa- dini, as seen from published illustrations of various species of Dipsas, Sibon, and Tropidodipsas (reviewed by Cadle and Myers, 2003). However, no species of Dip- sadini seems to have more than slight bi- lobation, and that exhibited by S. williamsi and S. vagus is about as much as has been observed in any species; Porto and Fer- nandes (1996) illustrate the hemipenis of D. neivai, which shows a similar degree of bilobation. The capitulum composes a large pro- portion of the hemipenial body in Siby- nomorphus. In the species examined here- in, the capitulum is approximately 50% or more of the body on the sulcate surface (see Figs. 32, 39). A review of notes and hemipenial illustrations in the literature suggests that this could be proportionally large compared with many other Dipsa- dinae. Further comparative studies are needed to determine whether useful sys- tematic variation in this character exists. (2) Orientation of the Sulcus Spermati- cus. In hemipenes of all species of Siby- nomorphus described herein, the orienta- tion of branches of the sulcus spermaticus is centrolineal, but their distal tips pass somewhat to the outer edges of the lobes Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 (Figs. 32, 39). In typical centrolineal sulci, the branches “diverge moderately and ex- tend in relatively straight lines, to lie on the same side ... as the forking point of the sulcus” (Myers and Campbell, 1981: 16). Thus, the condition in Sibynomorphus might be perceived as somewhat inter- mediate between centrolineal and centrif- ugal orientation (in which the branches di- verge greatly, to lie ultimately on opposite, lateral sides of the organ; Myers and Campbell, 1981: 16). This somewhat in- termediate condition has been noted in other Dipsadini, including Dipsas andiana and D. nicholsi (Cadle and Myers, 2003). However, Zaher et al. (2005: 22, fig. 3) de- scribed a seemingly very similar orienta- tion of the sulcus spermaticus of Atractus albuquerquei as “centrifugal.” Published illustrations of hemipenes of other Dipsadini generally show more con- sistently centrolineal orientations of the sulcus (Kofron, 1982, 1985a,b), although the branches in D. neivai appear to pass somewhat lateral as well (Porto and Fer- nandes, 1996). The tips of the sulcus sper- maticus in the five species of Sibynomor- phus surveyed end at the lateral edges of the lobe(s) rather than passing to the api- cal surface of the organ, i.e., to the center of the lobe(s). This condition is similar to that in D. andiana and D. nicholsi (see Ca- dle and Myers, 2003: fig. 6, bottom). (3) Spines Encircling the Hemipenial Body. A regular array of spines completely encircles the hemipenial body just proxi- mal to the capitulum in Sibynomorphus. The array is consistently about three rows across, although in S. vagrans and S. vagus the array broadened to about four rows on the asulcate surface. This battery of spines appears to be a constant feature of hemi- penes of Dipsadini, but its width varies from one to five rows in published descrip- tions (Cadle and Myers, 2003; Kofron, 1987). Observations of Dipsas oreas (Ca- dle, 2005: 108) suggest that this battery can vary in width intraspecifically so that too much should probably not be made of interspecific differences on the basis of ob- SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR ° Cadle servations of few organs. The size of the spines in the midsection battery varies within Sibynomorphus and in other Dip- sadini (e.g., Sibon; Kofron, 1982, 1985b, 1987, 1988), but these are usually distinct- ly smaller than the enlarged basal spines characteristics of Dipsadini (character 7 below). (4) Presence and Size of the Basal Nude Pocket. One unusual feature of hemipenes of Dipsadini seems to be the exceptional size of the basal nude pocket, which is commonly 30% or more of the overall length of everted organs. It extends from the base of the organ to, or almost to, the encircling battery of spines. The extremes as currently known are in species of Si- bynomorphus, in which the pockets in everted organs of S. vagus and S. vagrans are 23% and 26% of the hemipenial length, respectively, whereas in S. william- si, the pocket is 46% of the everted organ length. Too much should not be made of these quantitative comparisons until some potential sources of variation in relative pocket size are investigated more thor- oughly, including the effects of preserva- tion, overall size, and degree of inflation. Current data suggest that the size of the nude pocket relative to hemipenial size is greater in inverted organs than in everted ones. For example, the relative pocket size in inverted versus everted hemipenes of S. vagrans and S. vagus, as described herein, are 32% versus 26% and 30% versus 23%, respectively. Some published descriptions of hemi- penes of Dipsadini fail to mention a basal nude pocket but Cadle and Myers (2003: 15) pointed out that the pocket had prob- ably been overlooked in several of these, as examination of additional specimens showed the pocket to be present. Peters (1960) did not mention the pocket at all in his brief characterizations of the hemipe- nes of Dipsas, Sibon, and Sibynomorphus. I now suspect that the pocket has been overlooked in all species of Dipsadini in which the pocket was not reported in pub- lished descriptions (e.g., MacCulloch and 259 Lathrop, 2004; McCranie, 2006; Passos et al., 2005; Scrocchi et al., 1993; additional citations and observations in Cadle and Myers, 2003: 15) and that this character will prove universal in Dipsadini. Cadle and Myers (2003: footnote 7) pointed out that longitudinal folds on the base of the hemipenial body might need to be sepa- rated with fine forceps or needles to vi- sualize the pocket and that the pocket is especially difficult to demonstrate if the organ is tied off with thread. I encourage workers to make efforts to verify the exis- tence of the pocket during study of hemi- penes. The exceptional size of the pocket in Dipsadini might prove to be another synapomorphy of the clade, although more extensive quantitative comparisons need to be made, as well as a broader survey to verify the presence or absence of a nude pocket in other Dipsadinae. A basal nude pocket is present in some other Dipsadinae (and many other colu- brids), including Geophis (Downs, 1967; Myers, 2003); Imantodes (Myers, 1982); the Rhadinaea flavilata, R. godmani, and R. vermiculaticeps groups (Myers, 1974); and Atractus (Hoogmoed, 1980; Myers and Schargel, 2006; Savage, 1960; Zaher et al., 2005). Moreover, judging from my ex- perience with dipsadines, the failure to mention a nude pocket in published de- scriptions cannot be taken as definitive ev- idence of its absence. For example, pho- tographs of hemipenes of three species of Leptodeira appear to show a basal nude pocket (Duellman, 1958: pls. 26, 27), even though pockets were not mentioned in any descriptions (the presence of pockets is in- dicated by what appears to be a deep fur- row bordered by lobes such as commonly ornament the nude pockets in other snakes, as seen especially in Duellman’s pl. 26, no. 1). Thus, the taxonomic distribu- tion of basal nude pockets among Dipsa- dinae might be broader than indicated by a literature survey of hemipenial descrip- tions. The exceptional length of the nude pocket in Dipsadini is seemingly ap- proached in some species of the genus 260 Atractus, in which the pocket in inverted organs of some species extends distally to the level of the division of the sulcus sper- maticus (Hoogmoed, 1980; Savage, 1960). On the other hand, Zaher et al. (2005) de- scribed the nude pocket on the hemipenis of A. albuquerquei as “small.” (5) Position of the Basal Nude Pocket. The pocket is consistently on the lateral surface of the hemipenial body in Siby- nomorphus (i.e., about midway between the sulcate and asulcate surfaces, as dis- cussed in Methods), although in S. wil- liamsi it seemed farther displaced toward the asulcate surface than in the other spe- cies. Cadle and Myers (2003) reported var- iation in the position of the basal nude pocket in different species of Dipsadini, with all species examined except two hav- ing the pocket positioned laterally as in Si- bynomorphus. Cadle and Myers (2003) re- ported that in one specimen each of Dip- sas variegata and Sibon nebulatus, the nude pocket was on the asulcate surface of the hemipenis. A review of my original notes suggests that this might have been overstated because the notes indicate that the pocket in these two specimens “is off- set from the midpoint of the asulcate side”—a loose description that might be no different from the laterally positioned pockets observed in other Dipsadini. Ca- dle (2005) and Cadle and Myers (2003) re- ported a laterally positioned nude pocket in five other species of Dipsas and Sibon. Nonetheless, Kofron (1982, 1985b, 1987, 1988) described the nude pocket in several species of Sibon and Dipsas as an “asulcate pocket” (species for which he used this descriptor include S. annuliferus, S. carri, S. fasciatus fasciatus, S. fischeri fischeri, S. fischeri kidderi, S. sartorii, D. brevifacies, and D. gaigeae). However, dif- ferent observers could apply terminology differently, so this apparent variation should be verified before acceptance at face value. In addition, the manner of preparation or differential inflation of this pliant tissue could affect the position of Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 the nude pocket, resulting in apparent in- tra- or interspecific variation. (6) Lobes Associated with the Nude Pocket. The nude pockets in Sibynomor- phus oligozonatus and S. vagrans are bor- dered on each side by a prominent lobe. The lobe on the asulcate edge of the pock- et (i.e., toward the asulcate side of the or-— gan) is larger than the lobe toward the sul- cate side. The nude pockets in S. petersi, S. vagus, and S. williamsi are bordered | only by a single lobe on the asulcate edge of the pocket. The lobes are usually near _ the midpoint of the pocket, but at least in S. williamsi, the single lobe was near the distal end. Other Dipsadini show variation in this feature as well: Dipsas nicholsi and D. elegans have a single lobe on the asul- . cate edge of the pocket; D. andiana, D. oreas, and D. ellipsifera have a lobe on each side of the pocket (Cadle, 2005; Ca- dle and Myers, 2003). In D. ellipsifera, the lobes are asymmetrical, with the asulcate lobe being larger than the lobe on the sul- cate edge. In dipsadine species with two lobes, the lobe on the asulcate edge is of- ten (P? usually) larger than the sulcate lobe, and in species in which only one lobe is present, it consistently seems to be the lobe on the asulcate edge. One or more lobes are commonly associated with hemi- - penial nude pockets in diverse colubrids having a pocket, and the variation could merit further study. (7) Enlarged Spines on the Hemipenial Body. In everted hemipenes of the species of Sibynomorphus described herein, usu- ally two enlarged spines are present on the asulcate side of the organ below, or at the lower edge of, the encircling battery of spines (sometimes more closely associated with the nude pocket, as in S. petersi). In addition, another enlarged spine is present on the distal sulcate edge of the nude pocket. In everted organs, these enlarged spines are often, but not universally, clear- ly separated by a gap from spines in the midsection battery (in inverted hemipenes the enlarged basal spines are generally not clearly separated from spines in the mid- SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle section battery). Basal spines are generally notably enlarged compared with spines in the midsection battery, but because spines in the battery vary in size (see character 3 above), enlarged basal spines might not be clearly differentiated if there is not a dis- tinct gap separating the basal spines from the battery (I recorded only a single asul- cate spine in S. vagus, but that could re- flect lesser size distinction in the specimen studied between the enlarged basal spines and those of the midsection). Enlarged basal asulcate spines are fre- quently present in other Dipsadini judging from figures and descriptions in the liter- ature (e.g., Kofron, 1982, 1985b, 1987; Porto and Fernandes, 1996), but their number varies. For example, Cadle and Myers (2003) reported 2—4 basal spines in Dipsas nicholsi (the number varied among specimens), and in a single organ of D. an- diana, two enlarged basal spines were present in addition to two enlarged spines associated with the distal asulcate edge of the nude pocket (Cadle and Myers, 2003). The position of the enlarged basal spines varies somewhat, sometimes being more closely associated with the asulcate edge of the nude pocket and in other cases deployed more clearly on the asulcate side of the organ. This variation could reflect _ differences in hemipenial preparations (e.g., degree of inflation) or their interpre- tation, but side-by-side comparisons are needed to verify the consistency of these placements. Moreover, Kofron (1985b) re- ported that the largest spines in the hemi- penis of Sibon carri were on the sulcate side of the organ. (8) Miscellaneous Variation. A few unique or unusual hemipenial characters seemingly pertain to individual species of Sibynomorphus. The proximal calyces on the hemipenis of S. petersi have weakly developed longitudinal walls, giving the _ appearance of weakly developed flounces on the proximal edge of the capitulum. This morphology is observed occasionally in colubrids, but I am unaware of other species of Dipsadini with incipient flounc- 261 es. Species of Sibynomorphus (and other Dipsadini; see Cadle and Myers, 2003) vary in a few characters such as spine form (e.g., narrow, relatively straight, and hooked only at their tips in S. vagrans, compared with relatively robust and strongly hooked in the other species). The calycular papillae vary in length (e.g., lon- ger im 9. vagrans than in S. vagus ), but I have been unable to compare organs of most species simultaneously to objectify these differences. Similarly, my notes in- dicate a distinct fringe of mineralized spi- nules or small spines on the proximal edge of the capitulum in S. oligozonatus and S. vagrans. SIBYNOMORPHUS AND THE DIPSADINI: DIRECTIONS FOR FURTHER RESEARCH In this paper, I lay some foundation for future work on Sibynomorphus by outlin- ing patterns of variation in the Ecuadorian and Peruvian species, thus providing a framework for interpreting new specimens or data from northwestern South America. The systematic conclusions herein should facilitate a better understanding of the re- lationship (if any) between the Sibyno- morphus species of Ecuador and Peru, and those of southern South America. De- tailed comparisons of the two sets of spe- cies remain for future research. Recent studies on the systematics and biology of Neotropical molluscivores (e.g., Cadle, 2005; Cadle and Myers, 2003; Fer- nandes, 1995; Fernandes et al., 2002; this paper) have already refined knowledge of the species-level taxonomy and distribu- tions since the group was last reviewed (Peters, 1960). Nevertheless, much re- mains to be done. With over 60 species (including nominal taxa in the genera Dip- sas, Sibon, Sibynomorphus, and Tropido- dipsas, as outlined by Peters and Orejas- Miranda, 1970, and subsequently de- scribed species), the Neotropical mollus- civores comprise a highly diverse and tightly knit clade of Neotropical snakes (see ensuing discussion). More changes in 262 the understanding of species limits can be expected. Moreover, the phylogenetic re- lationships among the species have been controversial (Fernandes, 1995; Kofron, 1985a; Wallach, 1995). Although this study does not attempt to address broader rela- tionships among species of molluscivores (beyond comparisons among the species covered herein), the summaries of char- acter variation contained herein should ul- timately be useful to that end. In the fol- lowing sections, I outline some of the problems as I see them and pose some questions for future research on this group : The Systematics of Dipsadini—An Outline of Some Problems The Content of Dipsadini. Within Dip- sadinae, Sibynomorphus has long been considered closely related to the genera Dipsas, Sibon, and Tropidodipsas (e.g., Dunn, 1935, 1951), with the exception that Peters (1960) did not consider Tro- pidodipsas in his review of “Dipsadinae.” Peters did not elaborate on his exclusion of Tropidodipsas, although he may have been following Boulenger (1894, 1896), who included Tropidodipsas as then known in the Colubrinae and recognized the Amblycephalidae for Asian mollusci- vorous colubrids and most species of Dip- sas, Sibynomorphus, and Sibon then known. Zaher (1999) referred to the pu- tative clade comprising Dipsas, Sibon, Si- bynomorphus, and Tropidodipsas as Dip- sadini and I follow that usage here. Underwood (1967) observed that Dip- sas, Sibon, and Sibynomorphus have an ex- tensive tracheal lung, which is lacking in Tropidodipsas. Wallach (1995) interpreted the presence of a tracheal lung as a de- rived character of Dipsas, Sibon, and Si- bynomorphus, and explicitly excluded Tro- pidodipsas from the Dipsadini because it lacks this derived character.'’ However, '7 Wallach (1995: 479) also noted three characters of visceral topography that “separate the dipsadines from Tropidodipsas,” but it was not clear whether Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 that evidence could equally well be taken as a synapomorphy of Dipsas, Sibon, and Sibynomorphus within Dipsadini (includ- ing Tropidodipsas), rather than a reason for excluding Tropidodipsas from Dipsa- dini. At the least, Tropidodipsas shares with Dipsas, Sibon, and Sibynomorphus a behavioral propensity to consume gastro- pods as nearly the sole constituent of its diet (Cadle and Greene, 1993). Wallach (1995) viewed the dietary specialization as a derived feature linking Tropidodipsas with the other dipsadine genera but felt that, by extension, “a case could be made for the gastropod diet as a synapomorphy of the Dipsadini plus Pareatini |Aplopel- tura and Pareas| (Wallach, 1995: 479). However, the Pareatini lack the hemipen- ial and other synapomorphies of Dipsadi- nae (see Zaher, 1999), the broader clade to which the Dipsadini belongs, and more- over, some specializations for molluscivory in the two groups differ in details of anat- omy (Brongersma, 1956, 1958; Zaher, 1999: 20). Those differences indicate that the dietary specializations of the Pareatini and Dipsadini are convergent in these groups. Thus, there is no reason not to consider “molluscivory” a synapomorphy of Dipsas, Sibon, Sibynomorphus, and Tropidodipsas within the Dipsadinae. The behavioral trait for dietary specialization can also be viewed as a proxy for an array of other traits associated with the physical ability to subdue these prey (head musculoskeletal system, head glands, dentition, head sca- lation; Peters, 1960) and, very likely, bio- chemical traits associated with the sensory apparatus involved in prey detection and recognition. For these reasons, I suggest that the Dipsadini be defined to include Tropidodipsas along with Dipsas, Sibon, these were derived characters. One of these, the po- sition of the posterior tip of the lung, does not seem to clearly differentiate Tropidodipsas from some spe- cies of each of the other three genera of Dipsadini (Wallach, 1995: table 1), contrary to the text discus- sion (Wallach, 1995: 479). SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle and Sibynomorphus, as already advocated by Zaher (1999). Although the morpholog- ical traits associated with molluscivory have been repeatedly described in general terms for the Dipsadini or portions thereof Coe Dunn lool wreters, 9605 519; Scott, 1967), a modern analytical study of their distribution across species is needed. When incorporated into a formal phylo- genetic analysis of Dipsadini, this ap- proach should yield valuable insights into the evolution of feeding specializations within a speciose clade. Sibynomorphus in the Context of Other Dipsadini: Is Sibynomorphus Monophylet- ic? Species of Dipsadini have had an es- pecially unsettled generic taxonomy, as ex- amination of the synonymy of virtually any species will readily show (e.g., Cadle, 2005; Cadle and Myers, 2003). Useful re- views of the taxonomy are Peters (1960: 12-15 and, with special reference to Si- bynomorphus, 142-144), Smith and Taylor (1945: 49), and Kofron (1985a: 85-86). The current generic taxonomy derives from Peters (1960), notwithstanding some disagreement about the recognition or composition of the included genera (e.g., Fernandes, 1995; Kofron, 1985a; Wallach, 1995). The generic taxonomy should be re- vised as understanding of relationships among species of Dipsadini improves, but that understanding is, as yet, elusive. De- spite many suggested revisions of the ge- neric taxonomy of Dipsadini, methodolog- ical or sampling problems pertain to many previous considerations of the issue. Here I outline some of the issues and point to- ward ways of possible resolution. Kofron (1985a) synonymized Tropido- dipsas with Sibon on the basis that species of both genera shared a “unique” postero- lateral process on the maxilla (compared with its absence in three species of Dipsas he examined); the process was preceded by a notch on the dorsal surface of the bone. However, Scott (1967) had reported presence of the maxillary process in D. gracilis, and Boulenger’s (1896: 461) illus- tration of the skull of Dipsas indica shows 263 a maxillary notch similar to that portrayed by Kofron (1985a: figs. 1, 2). As discussed above, Wallach (1995) resurrected Tropi- dodipsas on the basis of tracheal lung and other visceral characteristics. These stud- ies have contributed substantially to our knowledge of morphological characteris- tics in species of Dipsadini. However, the species and character sampling for a di- verse clade such as Dipsadini has generally been less than desirable in previous broad taxonomic studies; they have essentially taken an “exemplar” approach, wherein a few species are taken to represent many others. For example, Kofron (1985a) used three species to represent Dipsas in his comparisons despite conflicting character data already reported for another (Scott, 1967). Despite the rarity of some dipsa- dines in collections, my impression is that there is much more material available than has yet been incorporated into broader phylogenetic studies. I encourage more comprehensive surveys of species and characters to flesh out the character dis- tributions already noted in previous stud- ies (Fernandes, 1995; Kofron, 1985a; Wal- lach, 1995). Since Peters’ (1960) review, Sibynomor- phus has been understood to comprise more “generalized” or terrestrial members of the Dipsadini. Species of Sibynomor- phus lack highly compressed bodies, en- larged heads and eyes, elongate bodies and tails, and enlarged vertebral scale rows (characters generally associated with ar- boreality in snakes). However, these char- acters also apply to most species of the re- lated genus Tropidodipsas (sensu Wallach, 1995: see also Cadle, 1984a; Kofron, 1985a), a diverse assemblage of Mexican and Central American species that Peters (1960) did not consider in his revision of the Dipsadinae (= Dipsadini of present understanding, i.e., the genera Dipsas, Si- bon, Sibynomorphus, and Tropidodipsas). Although Sibynomorphus are primarily terrestrial snakes, observations suggest that at least some species occasionally climb: S. ventrimaculatus climbs a meter 264 or more (Cechin and Oliveira, 2003) and S. neuwiedii and S. mikanii have been de- scribed as “semi-arboreal” (Sazima, 1992) or “terrestrial/arboreal” (Rodrigues, 2003). Furthermore, Peters (1960: 19) had pointed out that “one of the greatest ob- structions to a satisfactory analysis of snakes of [the tribe Dipsadini] is that spe- cies with nearly every combination of .. . characters [associated with mollusc-feed- ing and arboreality] can be found.” In view of this, Peters’ (1960: 12—19, 141-145) dis- cussion of generic delimitations within this group seems arbitrary. Cadle and Myers (2003) noted that most species of Dipsa- dini for which any data are available share some unusual characters of the hemipenes (e.g., very large basal nude pockets) and defensive behaviors, but none of these characters appeared to be restricted to particular genera of this group. The pos- sibility should be addressed whether the species of Sibynomorphus are a phenotyp- ically similar, but not monophyletic, set of species whose peculiar distribution is purely artifactual because of an erroneous understanding of relationships. Only a broader study of phylogeny at the species level among snakes of the tribe Dipsadini can resolve this issue. Compared with Sibon and Tropidodipsas, Sibynomorphus shares with Dipsas some derived characters of the gular region: no- tably, the shape of the chin shields (squarish or polygonal rather than elongate and _nar- row), often more than two pairs of chin shields, and absence of the mental groove. Sibon and Tropidodipsas have correspond- ing character states that might be consid- ered plesiomorphic because they are wide- spread among Dipsadinae and other colu- brids: one or (usually) two pairs of narrow, elongate chin shields similar to those typi- cally found in colubrids, and a mental groove (in Sibon often only the anterior chin shields are conspicuously elongate). Sibynomorphus and Dipsas also are usu- ally said to share a suite of musculoskeletal features of the head associated with mollus- civory that are more derived than those of Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 Sibon and Tropidodipsas (Kofron, 1985a; Peters, 1960; Savage, 2002), including an in- ward-turned maxilla with medially directed teeth, pterygoids that are parallel (as op- posed to diverging), pterygoids with a flat ventral surface (as opposed to having a cen- tral fossa), mandibles bowed in a vertical plane, and a pivot joint between the dentary and compound bone of the lower jaw (Kof- ron, 1985a: 88; Peters, 1960). In reality, these characters have not been investigated in most species of either genus. For exam- ple, with the exception of a few skull char- acters described for S. lavillai (Scrocchi et al., 1993), the nature of these characters is unknown in the five species of Sibynomor- phus described since 1970, and at least some of the characters have been reported in species of Sibon and Tropidodipsas (e.g., Fernandes, 1995; Scott, 1967). Newly de- scribed species of Sibynomorphus have been referred to the genus by a combination of “key” external and dentition characters. . usually used in generic recognition (e.g., Pe- ters and Orejas-Miranda, 1970). Over the years, several authors have rec- ommended synonymizing Sibynomorphus — with Dipsas (e.g., Brongersma, 1958; Kof- ron, 1990: 220; Parker, 1926: 206; Smith and Taylor, 1945: 49), a name that Boulen- ger (1896), among others, applied to a sin- gle species, D. indica. The two genera were traditionally distinguished by the presence (Sibynomorphus) or absence (Dipsas) of pterygoid teeth until Parker (1926) indicated that interspecific variation in this character was nearly continuous. Parker also reported intraspecific variation in the number of pterygoid teeth within “Sibynomorphus oreas” (= Dipsas oreas). That evidence is a red herring, however, inasmuch as the BMNH specimens Parker cited belong to two different species (Ca- dle, 2005: 114). Nonetheless, Peters’ (1960) data show that absence of pterygoid teeth is approached in very few species of Dipsadini (e.g., D. indica, D. variegata) and that many species of Dipsas (sensu Pe- ters, 1960) have as many, or more, ptery- goid teeth as species of Sibynomorphus. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle Peters’ (1960: 144) justification for maintaining Sibynomorphus distinct from Dipsas seems particularly arbitrary: “I agree in part with Parker [1926]. I feel that the generic distinction between the many species formerly included in the genus Si- bynomorphus and ... the genus Dipsas is not valid, but I use the name Sibynomor- phus in an extremely restricted sense, con- fining it to the stem forms found in south- erm Brazil and neighboring countries.” It is unclear how the two species of Sibyno- morphus from northern Peru known at the time, S. vagrans and S. vagus, fit into this concept. Peters (1960) provided little in the way of differential diagnoses for these two genera other than a few external char- acteristics in his “Key to the Genera” (Pe- ters, 1960: 19), e.g., body form, head and eye size, relative enlargement of the ver- tebral scale row. Evidence for the mono- phyly of Sibynomorphus is thus particular- ly weak. In an as yet unpublished doctoral dis- sertation dealing with a broad sampling of Dipsadinae, Fernandes (1995) argued that Sibynomorphus should be synonymized with Dipsas because his phylogenetic anal- ysis recovered one character purportedly shared by two species of Sibynomorphus (S. neuwiedii and S. ventrimaculatus) and two species of the Dipsas catesbyi group (D. catesbyi and D. pavonina) to the ex- clusion of other Dipsas (seven additional species included in the analysis). It is be- yond the scope of this report to present an extended critique of Fernandes’ (1995) study, but taxon and character sampling is- sues alone suggest that synonymizing the two genera is premature. Only two of 12 species of Sibynomorphus (S. neuwiedii and S. ventrimaculatus) and nine of 30+ species of Dipsas were included in the tax- on sampling for formal phylogenetic anal- ysis (Fernandes, 1995: figs. 7, 8). None of the northern species of Sibynomorphus were investigated. The derived character linking the two species of Sibynomorphus and two members of the D. catesbyi group (D. catesbyi and D. pavonina) was an en- 265 larged L-shaped Harderian gland. Al- though this character state was coded as present in S. neuwiedii (Fernandes, 1995: table 3), the list of material studied does not indicate that the head glands of this species were investigated (Fernandes, 1995: appendix B), and the character was scored as unknown in two of the nine spe- cies of Dipsas examined. Furthermore, the type species of Sibynomorphus is S. mi- kanii Schlegel, a species that Fernandes (1995) did not include in the formal phy- logenetic analysis, so from a purely no- menclatural viewpoint, synonymizing Si- bynomorphus with Dipsas on that basis entails some unwarranted assumptions about the phylogenetic unity of Sibyno- morphus. In addition to these issues, some analyt- ical aspects of Fernandes’ (1995) study need more careful consideration before making further changes in the generic tax- onomy of these snakes. These include more thorough evaluation of intraspecific character variation, character scorings, and more robust tree searching and evaluation methods. For example, intraspecific vari- ability (or error in character scoring) in the purported character linking Sibynomor- phus with Dipsas catesbyi and D. pavoni- na is suggested by Brongersma’s (1958: 11) observation that the Harderian gland is en- larged in D. catesbyi but “small” in D. indica and D. pavonina, contrary to Fer- nandes’ (1995) finding. Even as some authors have called for synonymizing Sibynomorphus with Dip- sas, no one has specifically addressed whether Sibynomorphus itself (sensu Pe- ters, 1960, and including subsequently de- scribed species) is monophyletic. Although quantification and comprehensive compar- isons at the species level are needed, for present purposes, Sibynomorphus is dis- tinguished from Dipsas by lacking physical characteristics associated with arboreality (highly compressed and attenuate body, prehensile tail, and different head, eye, and tail proportions; Peters, 1960). Many more characters should be investigated in 266 a broad spectrum of species of Sibyno- morphus and Dipsas from throughout their geographic ranges. A search should also be made for new characters that could potentially inform phylogenetic studies of Dipsadini. Among these are perhaps patterns of sexual di- morphism. Cadle (2005: 77) noted that several species of Dipsas had unusual pat- terns of sexual dimorphism, including one species (D. elegans) in which males had greater numbers of ventral plates than fe- males (i.e., the reverse of the common pat- tern in colubrids); two other species, D. ellipsifera and D. oreas, lacked sexual di- morphism in ventral plate numbers, which is also an uncommon pattern. A subse- quent review of Peters’ (1956: fig. 12) data for D. catesbyi suggested that the unusual pattern observed in D. elegans pertained to D. catesbyi as well, and perusal of sev- eral papers on other Dipsadini hinted at possible unusual patterns of sexual dimor- phism in other species. These include con- siderable overlap in the ventral and sub- caudal counts of male and female Sibon annulifera, S. sartorii sartorii, S. dimidia- ta, and S. sanniola (Kofron, 1988, 1990). Unfortunately, Kofron presented only ranges for most comparisons, and he com- bined male and female counts in other pa- pers so that it is impossible to discern any trends (a practice followed also by Savage and McDiarmid, 1992). However, Kofron (1990) reported mean values for male and female ventrals in S. s. sanniola as 155 and 153, respectively. This suggests little or no sexual dimorphism in ventral counts for this species (the character ranges of males and females were nearly identical as well). In contrast to these unusual patterns, the Ecuadorian and Peruvian species of Si- bynomorphus are not unusual with respect to patterns of sexual dimorphism: they show the patterns most common in colu- brids generally. Whether the southern spe- cies of Sibynomorphus show typical or un- usual patterns of sexual dimorphism is not clear (e.g., data for S. lavillai from Scroc- chi et al. [1993] showed little difference Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 between males and females for ventral counts). Nonetheless, this character might provide an additional distinction between some species of Sibynomorphus and some Dipsas. Broader and more quantitative surveys could prove fruitful. Sibynomorphus (or a part thereof) could thus be plesiomorphic relative to Dipsas in terms of ecomorphological characters as- sociated with arboreality and with respect to patterns of sexual dimorphism. These plesiomorphic characters in Sibynomor- phus do not support the monophyly of Si- bynomorphus per se, but they do provide a differential diagnosis from Dipsas until more detailed studies are conducted. Fur- ther changes to the taxonomy of these snakes should be commensurate with an increase in understanding of a suite of characters and their variation across a broad sampling of nominal taxa of Dipsa- dini—something that has been lacking in previous examinations of the generic tax- onomy. The Peculiar Distribution of Sibynomor- phus. Certainly one outstanding feature of Sibynomorphus as currently understood is its odd distribution, which is unlike most other Neotropical reptiles and amphibians (Fig. 44). Lest this peculiarity be con- strued as “evidence” for the nonmonophy- ly of Sibynomorphus, it is worth briefly comparing the distribution to other South American. species. The six species of Si- bynomorphus that form the focus of this report (the “northern species” in the fol- lowing discussion) are distributed in south- ern Ecuador and the northern half of Peru, and these are species of the Pacific versant (coast, Andean slopes, and western inter-Andean valleys) or of the eastern in- ter-Andean valleys on the Amazonian ver- sant (Rio Maranoén and its affluents, Rio Huancabamba and Rio Chotano). Then follows a broad geographic gap before the genus appears again in the dry woodlands of southern Bolivia (Fugler and Cabot, 1995; Fugler and De la Riva, 1990) and the Brazilian state of Mato Grosso, whence it is broadly distributed south of the main SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR ° Cadle 267 Figure 44. The widely disjunct distribution of species of Si- bynomorphus in western Peru and Ecuador, and south of the Amazon basin. The distribution of the “northern” species is compiled from records herein. The distribution of the “south- ern” species is compiled from Peters (1960), Cei (1993), Ley- naud and Bucher (1999), and various other faunal works. Question marks indicate my inability to find reliable records through a large part of the Brazilian cerrado. Similarly, the dis- tribution might be more continuous in the borderlands between Bolivia, Paraguay, and Argentina. course of the Amazon. About 1,500 km (straight line distance) separates the clos- est points in the two parts of the distri- bution, not to mention the rugged terrain of the Andes and parts of forested lowland Amazonia, where no species of Sibyno- morphus seems to occur. South of the Amazon, species of Siby- nomorphus (the “southern species”) occur in an extraordinary range of major habi- tats, including cerrado, caatinga, Atlantic forest, chaco, pantanal, and dry deciduous woodlands (Brites and Bauab, 1988: Cei, 1993; Leynaud and Bucher, 1999; Marques et al., 2000; Norman, 1994; Pe- ters, 1960; Rodrigues, 2003; Vanzolini, 1948). There seem to be no definite rec- ords from within the Amazonian forest. The collective distributions of the south- ern species of Sibynomorphus are remark- ably similar to that of the colubrid snake genus Psomophis, except that Psomophis is conspicuously absent from the Atlantic Forest and caatinga (Myers and Cadle, 1994) (Rodrigues [2003] described the presence of S. mikanii in caatinga as relic- tual). Apart from S. vagrans and coastal populations of S. williamsi, the northern species of Sibynomorphus are montane species and occur at higher elevations than are known for any of the southern species. With the exception of Sibynomorphus neuwiedii, which is apparently restricted to forested regions of the Atlantic Forest biome) (Hoge ‘et ale, 1978/1979" 1981; 177), southern species of Sibynomorphus primarily occur in relatively open habitats, often with reduced rainfall. The ecological (particularly microhabitat) distribution of the northern species of Sibynomorphus is not well known, but subhumid to arid en- vironments prevail in this region. Locali- ties for most northern species, as docu- mented in the natural history sections of the species accounts, are referred to fre- quently by descriptors such as arid habitats with cacti, semiarid brushland, dry decid- uous forests, thorn scrub, arid rain shadow valleys, and montane steppe. Nonetheless, a few records for S. petersi (e.g., Rio Zaiia Study Site, Bosque Cachil, Canchaque, Rio Chotano valley) suggest that this spe- cies, at least, occurs in the dry/humid for- est ecotone, if not actually within humid montane forests (see species account), and other species might occur in more mesic areas, such as riparian habitats, than their general habitat associations might imply. Species of Sibynomorphus are not unique among Dipsadini in their occur- rence in subhumid to xeric habitats. A few species of the other genera occur in hab- itats such as dry deciduous forests, thorn forests, or deserts (e.g., Dipsas brevifacies and D. gaigeae, Sibon sanniola, Tropido- 268 dipsas annulifera). Nevertheless, species of Dipsas, Sibon, and Tropidodipsas over- whelmingly inhabit more mesic to humid environments than species of Sibynomor- phus. In the case of broad macrosympatry between D. oreas and S. petersi (see its species account) in southern Ecuador and northern Peru, these two species might be segregated by habitat, with S. petersi oc- curring in more arid environments than D. oreas (see footnote 7). The disjunct distribution of Sibynomor- phus is unusual but not without parallels. The Pacific coastal and western Andean region of northern Peru and Ecuador (the generalized distribution of the northern species of Sibynomorphus) harbors a dis- tinctive and largely endemic herpetofauna. Few components of this fauna have been subjected to phylogenetic studies, but three examples illustrate parallels with the distribution of Sibynomorphus (Fig. 45). In the frog genus Ceratophrys (“Lep- todactylidae” or Ceratophryidae), C. stolz- manni (southwestern Ecuador/northwest- ern Peru) is the sister taxon to C. calcar- ata—C. cornuta (Caribbean Colombia/Ve- nezuela and the Amazon basin), and these three species, composing the subgenus Stombus, are the sister group to the sub- genus Ceratophrys (Lynch, 1982), which occurs in forested and nonforested habi- tats from southern Bolivia and northern Argentina east throughout southern Brazil Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 (Espinoza et al., 2004; Etheridge, 1995). Thus, an area cladogram for Liolaemidae is similar to that of Sibynomorphus (Fig. 45A), with the additional geographic com- ponents of the southern Andes and coast. Finally, among tropidurid lizards, Mi- crolophus (Fig. 45D), from western Ec- uador and Peru (including inter-Andean Maranon valley of northern Peru and the Galapagos Islands), is the sister group to other tropidurid genera (Tropidurus, Plica, Uracentron, Eurolophosaurus, and Stro- bilurus) (Frost et al., 2001), whose geo- graphic distribution encompasses Amazon- ia, the Guayanan region, and virtually all of austral South America (i.e., south of the Amazon basin). Again, Amazonian and Guayanan taxa represent geographic com- ponents not found in Sibynomorphus. Al- though the examples in Figure 45 seem most similar to the distribution of Siby- nomorphus, other possible examples might be found in genera such as Stenocercus (Tropiduridae) and Philodryas (Colubri- dae), which have coastal/Andean compo- nents in Peru, Ecuador, or both and which are broadly distributed east of the Andes and in austral South America. Each of these examples includes geo- graphic components (Amazonia, southern Andes and southern Pacific coast, northern South America) not known in Sibynomor- phus. Nonetheless, a complete phyloge- netic analysis of Dipsadini at the species (Fig. 45B). Thus, apart from the existence of a widespread Amazonian species (C. cornuta) and a northern South American species (C. calcarata), the distributional relationships of the western South Amer- ican species, C. stolzmanni, are similar to distribution patterns portrayed by species of Sibynomorphus as that genus is pres- ently understood. The lizard Ctenoblepharys (Liolaemi- dae; Fig. 45C) is endemic to loose sand habitats of coastal central Peru. It is the sister group to Phymaturus + Liolaemus of the southern Andes and Pacific coast (Argentina—Chile), and cis-Andean South America from Argentina to southern Brazil level might eventually show that these “missing” geographic components are rep- resented by species whose relationship to Sibynomorphus is not yet demonstrated (i.e., species of Dipsas). This is especially true given that monophyly of Sibynomor- phus is not yet demonstrated, and its re- lationship to other Dipsadini (particularly the diverse array of Amazonian and north- ern South American Dipsas) is presently murky. Moreover, the cladograms in Figure 45 illustrate two alternative area relationships among species of western Ecuador/Peru, austral South America, and Amazonia. In Ceratophrys (Fig. 45B) the western Ec- SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle 269 A Sibynomorphus B Ceratophrys Amazonia/Northern Western Ecuador/Peru NactralSouthvAgenics Shih Acierica Austral South America yd 7 C Ctenoblepharys Austral South America Southern Andes and Coast Austral South America Ctenoblepharys Phymaturus D Microlophus Western Ecuador/Peru/ z : Austral South America Chile Austral South America Northern South America Microlophus Eurolophosaurus Strobilurus Uracentron/Plica @ 7 ? Z Z y 7 7 Figure 45. Area cladograms illustrating geographic components (areas of endemism) shared between Sibynomorphus and other taxa of western South America. “Austral South America” here refers to all of cis-Andean South America south of Amazonia; the exact area delimited varies somewhat in each case (see discussion and references in the text). Solid lines in the cladograms indicate shared geographic components. Dashed lines indicate one or more geographic components not present in Sibynomor- phus. (A) Geographic components in Sibynomorphus. Neither the monophyly of the genus itself nor its geographic components (“northern” and “southern” species groups) is strongly supported (see text). (B) Geographic components in the frog Ceratophrys (Leptodactylidae or Ceratophryidae). (C) Geographic components in the lizard Ctenoblepharys (Liolaemidae). (D) Geographic components in the lizard Microlophus and related lizard genera (Tropiduridae). uador/Peru taxon is most closely related to American taxa. For Ctenoblepharys (Fig. the Amazonian species, whereas in Micro- 45C), the area cladogram is identical with lophus (Fig. 45D), the Amazonian taxa are _ that of Sibynomorphus (i.e., no Amazonian more closely related to the austral South component is present) except that species 270 of Liolaemus also occur in the southern Andes and the southern Chilean coast, geographic components also unknown in Sibynomorphus. Depending on how the phylogenetic re- lationships among the northern and south- em species of Sibynomorphus and the spe- cies of Dipsas are ultimately resolved, the area cladogram for Sibynomorphus could resemble any of the examples in Figure 45 or more complex geographic relationships. For example, three possible resolutions of the area cladogram for Sibynomorphus are shown in Figure 46, in which dashed lines to the geographic component “Amazonia” represent as yet undemonstrated relation- ships of the northern (western Ecuador/ Peru; Fig. 46A) or southern (austral South America) species of Sibynomorphus (Fig. 46B). The former resolution is compatible with the area cladogram for Ceratophrys (Fig. 45B), and the latter is compatible with the area cladogram for Microlophus (Fig. 45D). Many more complex resolutions are possible, of which the example in Figure 46C is but one example (ie., the northern and southern species of Sibynomorphus are each independently related to different spe- cies groups of Amazonian Dipsas). The ex- ample in Figure 46C would imply that Si- bynomorphus as it is currently known is polyphyletic. The possibilities outlined in Figure 46 are speculative, but they indicate ways in which some outstanding phylogenetic and biogeographic enigmas concerning Siby- nomorphus might ultimately be resolved. Figure 46 also provides guidelines for con- structing any rigorous test for relationships between Sibynomorphus or any of its geo- graphic components and other Dipsadini. At the least, northern and southern species of Sibynomorphus must be included in analyses, preferably with a broad taxonom- ic and geographic representation of South American species of Dipsas as well. Unless tests are constructed such that these alter- natives can potentially be discerned, we are left with more ambiguity than clarity. Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 A Amazoni Western Ecuador/Peru Austral South America 8 (Atlantic Forest) Amazonia z Western Ecuador/Peru Austral South America Cc Western Ecuador/Peru Austral South America Figure 46. Examples of possible resolutions of “missing” geographic components in the area cladogram of Sibynomor- phus (Fig. 45). These area cladograms assume a complex re- lationship between geographic components of Sibynomorphus and species of Dipsas, which currently is speculative (see text). (A) Resolution corresponding to the area cladogram of Ceratophrys (Fig. 45B). (B) Resolution corresponding to the area cladogram of Microlophus (Fig. 45D); a possible alter- native is indicated by the additional geographic component “Atlantic Forest,’ which is a component of “Austral South America” used in Figures 45 and 46. (C) A more complex scenario in which the northern and southern species of Siby- nomorphus are each related to different species of Dipsas in Amazonia, which implies polyphyly of Sibynomorphus as cur- rently understood. Many more resolutions of these area clad- ograms are possible, but all depend on the precise relationship among the northern and southern species of Sibynomorphus and other species of South American Dipsadini. These rela- tionships are largely unexplored. Research Needs on the Northern Species of Sibynomorphus More Field Work and Collections are Needed. Several topics addressed in this study need additional research based on SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle more samples than currently exist. System- atic studies of some tropical snakes are hampered because few specimens from widely scattered localities are available, and species of Sibynomorphus in Ecuador and Peru are no exceptions. In extreme cases, species can be represented by unique or nearly unique specimens as, for example, in the species of “rare snakes” of the genera Geophis and Atractus studied by Myers (2003). None of the species of Sibynomorphus considered herein is as ab- solutely rare as the species considered by Myers (2003). However, interpreting pat- terns of variation and inferring species lim- its was difficult in several cases because of inferred intrapopulational or geographic variation combined with few specimens sampled across the complex terrain of the Andes. Even with the new material re- ported herein, one of the species is known from only four specimens, and three spe- cies are known from three or fewer local- ities. Great variability in systematic char- acters has been documented within spe- cies of Sibynomorphus (Rossman and Ki- zirian, 1993; this study) and related genera (e.g., Dipsas; Cadle, 2005; Cadle and My- ers, 2003). The variation occurs at several levels (individual variation within popula- tions, sexual dimorphism, and geographic variation among populations) and with re- spect to scutellation and color patterns. Teasing apart the sources of variation in taxa represented by few specimens is par- ticularly difficult. Because of the variability of informative systematic characters within and between population samples and few specimens spread over a topographically and environ- mentally diverse region of South America, the available material of Ecuadorian and Peruvian Sibynomorphus is insufficient to fully clarify their systematics. Thus, the identity of some populations proved diffi- cult to resolve. Because of the extensive variation in characteristics such as color patterns and scutellation, the species limits recognized herein should be reviewed as 271 more material and denser geographic sam- pling become available. In particular, the concepts of Sibynomorphus oligozonatus, S. petersi, and S. oneilli need reassess- ment. Full characterization of S. oligozon- atus is hampered by few specimens and rather large disparity among specimens in a few characters, such as ventral counts within males and between males and fe- males. For Sibynomorphus petersi and S. oneil- li, the problems of species limits have more to do with character variation among localities. The Peruvian samples of S. pe- tersi are heterogeneous in color patterns among themselves and compared with Ec- uadorian samples. Ecuadorian specimens tend to be more homogeneous in color patterns, but they also form a more co- hesive geographical unit than Peruvian samples, which are spread in a linear array along the western Andean front (Fig. 25). For S. oneilli, the problem is that the type specimen differs in several characters from virtually all other specimens referred to this species, and it is the only specimen currently known from the Cordillera Ori- ental. Efforts should be made to obtain ad- ditional specimens (particularly adults) from the vicinity of the type locality to ver- ify whether the unusual characters of the holotype pertain to this individual or re- flect population or regional characteristics. Additional field work is needed to flesh out the distributions of Sibynomorphus in Peru in more detail. In particular, field work at the northern end of the Cordillera Oriental in Peru (east of the Rio Maranon in La Libertad, San Martin, and Amazonas Departments) should be undertaken to understand more fully the distribution of S. oneilli in this region (currently known only from a single specimen, the holotype, from east of the Rio Marafidn). Similarly, S. petersi is known from a single specimen on the Amazonian versant, and the extent of its distribution east of the continental divide should be established. There is also a broad geographic gap in the distribution 272, of S. petersi on the Pacific versant between southern Ancash Department and the next locality to the north, San Pablo (Cajamarca Department) (Fig. 25). Sibynomorphus vagus and S. vagrans are known only from single locations, and the latter species has apparently not been collected since Noble collected the type series in 1916, even though it seemed to be common at that time (see species account). As currently known, the distribution of Sibynomorphus williamsi along the Peru- vian coast is peculiarly circumscribed. Specimens have been recovered from sev- eral localities in the immediate vicinity of Lima and from the relatively well-studied Lomas de Lachay just north of Lima (Fig. 43). The lomas formations extend from northern Chile to northern Peru and, al- though their general aspect becomes drier from the vicinity of Lima southward, the climate in general for this region is rela- tively homogeneous (Rundel et al., 1991). Comprehensive field surveys of the lomas formations should reveal whether S. wil- liamsi is more broadly distributed in this habitat along the Peruvian coast. These environments have rarely been given at- tention by zoologists, and new discoveries are likely. (Pearson and Ralph [1978] stud- ied bird, mammal, and lizard communities of a loma in southern Peru, but their sam- pling protocol was unlikely to reliably sam- ple secretive or highly seasonal snakes.) A further paradox concerning the distri- bution of S. williamsi concerns its eleva- tional distribution. All initially reported lo- calities were from coastal lomas formations less than 1,000 m in elevation (and all, ex- cept the Lomas de Lachay, <300 m) (Car- rillo de Espinoza, 1974). The localities in Ancash Department differ in elevation (>1,800 m) and environment (more xeric) from these coastal formations. The breadth of the elevational distribution of S. williamsi (10 to perhaps 3,600 m) is ex- traordinary for a species of snake and in- vites more detailed distributional studies and population comparisons to determine Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 whether all are, in fact, the same species." In particular, field studies of coastal areas in southern Ancash Department could de- termine whether the upland populations are linked to any at lower elevations in this region. Similarly, additional study of the Andean foothills east of Lima should de- termine whether S. williamsi occurs more generally in this area (the species is cur- rently known only from one inland locality in Lima Department, Santa Eulalia). Currently, the only known case of sym- patry among any of the northern species of Sibynomorphus is the presence of S. pe- tersi and S. oligozonatus at their type lo- cality in Azuay Province, Ecuador. It seems likely that these two species are broadly sympatric throughout the highland areas of southern Ecuador and (possibly) northern Peru (Figs. 7, 25). Potential areas of sympatry for species of Sibynomorphus in Peru include southern Ancash Depart- ment, where three species (S. oneilli, S. petersi, and S. williamsi) occur in close proximity (Fig. 43). Further field work is needed to determine whether there is mi- crosympatry among these species and, if so, to what extent their resource use pat- terns are similar. Systematic and Phylogenetic Issues. Much work remains to be done on the sys- tematics of Sibynomorphus in the broad sense and achieving a well-supported phy- logenetic hypothesis for its relationships within Dipsadini. Chief among the out- standing questions is whether Sibynomor- phus, as it has been understood since Pe- ters (1960) (i.e., the northern + southern species), is monophyletic. Fernandes (1995) formalized the suggestion raised many times previously that Sibynomor- phus stemmed from within Dipsas, but the possibility also exists that Sibynomorphus '8 Two other snakes of the Pacific versant also have huge elevational ranges. Mastigodryas heathii and Philodryas simonsii are known from near sea level. The former ranges to at least 2,500 m and the latter to over 3,000 m in the Andes of northern Peru (Ca- dle, unpublished data). SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle is polyphyletic with respect to Dipsas. For example, the two geographic units (north- ern and southern) could be independently related to different clades within Dipsas or there could be an even more complex re- lationship among species of the two genera (see above discussion and Figs. 45 and 46). These are unexplored questions. This study yielded few clues as to rela- tionships among any of the species re- viewed. Three species, Sibynomorphus oli- gozonatus, S. williamsi, and S. vagrans, re- semble one another in several characters. Their color patterns are less variable intra- specifically, and they are more similar to one another than any of the other three northern species, which are quite variable in color pattern. Sibynomorphus oligozon- atus, S. williamsi, and S. vagrans all have highly contrasting dorsal patterns (bold crossbands on relatively pale ground col- ors) and relatively unmarked venters com- pared with S. oneilli, S. petersi, and S. va- gus. Sibynomorphus oligozonatus and S. williamsi share some characters that are rare or absent in other species of the ge- nus: (1) They share the uncommon char- acteristic of having six supralabials with the third and fourth bordering the eye—a characteristic seen elsewhere in some specimens of S. lavillai and S. ventrima- culatus (Serocchi et al., 1993), but not among other northern species of Sibyno- morphus (Table 1). (2) They consistently lack separate preoculars (i.e., the preocular is always fused with the prefrontals in the specimens examined) and have elongate loreal scales. Among the northern species of Sibynomorphus, lack of a separate pre- ocular was observed elsewhere only in some specimens of S. oneilli but was re- ported as a consistent feature of S. mikanii and the usual condition in S. ventrimacu- latus (Cei, 1993). (3) They tend to have fewer maxillary teeth than the other north- ern species. The color patterns of S. oli- gozonatus and S. williamsi are similar ex- cept for the less distinct bands/blotches on the posterior body of S. oligozonatus and 273 its broader anterior crossbands compared with S. williamsi. On the other hand, Sibynomorphus oli- gozonatus shares with S. vagrans the pres- ence of two lobes associated with the nude pocket on the hemipenial body (one asul- cate lobe in the other northern species). The color patterns of these two species are also very similar, including broad anterior crossbands, less distinct posterior cross- bands, and relatively unmarked venters (Figs. 1, 3-6, 28, 29). It is unclear whether any of these similarities among S. oligo- zonatus, S. vagrans, and S. williamsi are indicative of close phylogenetic relation- ships in the absence of broader compari- sons. Also, as perceived by Dunn (1923), S. vagrans is similar to S. vagus in having low ventral counts compared with the oth- er species. Further comparative studies of all spe- cies referred to Sibynomorphus are need- ed, particularly in the context of other Dipsadini. Three of the northern species, Sibynomorphus oneilli, S. petersi, and S. williamsi attain larger sizes (males, 563— 763 mm total length; females, 718—790 mm total length; Table 1) than any of the southern species (generally <500 mm total length in males and females; Cei, 1993; Scrocchi et al., 1993). Intraspecific varia- tion in scutellation is generally much less in Sibynomorphus than in Dipsas (Peters, 1960; this study; cf. Cadle, 2005; Cadle and Myers, 2003). Such comparative stud- ies should yield insights not only into the systematics of this group, but into aspects of their natural history. Natural History Studies. The paucity of data on the natural history of all northern species of Sibynomorphus is evident from the species accounts contained herein. Mi- crohabitat and behavioral data are almost completely lacking for all species, and only a single dietary record (a gastropod) is available (see S. oligozonatus). Two rec- ords document predation on S. petersi and S. cf. oneilli by coral snakes (Micrurus). Three records document clutch sizes for S. petersi (two clutches of five eggs each) 274 and S. williamsi (six eggs), but dates of col- lection are not available for these speci- mens, and nothing concerning reproduc- tion is known for the other northern spe- cies of Sibynomorphus. Most specimens with recorded dates of collection have been obtained in the austral summer (i.e., generally between October and April), al- though S. petersi has also been obtained during the austral winter (June—August) in southern Ecuador. Seasonal activity and reproductive patterns for these species need to be documented with greater pre- cision. The proximity of two species, S. williamsi and S. oneilli, to major popula- tion centers (Lima vs. Huardz and Caja- marca, respectively), offers significant op- portunities for detailed local natural his- tory studies. Finally, two aspects of variation within Sibynomorphus and the Dipsadini deserve further study relative to the natural history of these snakes: color and scales. A puz- zling aspect of variation in several species of northern Sibynomorphus (S. oneilli, S. petersi, S. vagus) is considerable intra- and interpopulational variation in color pat- tern. Each of these species varies in pat- tern from relatively unicolor (or finely re- ticulated or flecked) to strongly banded (see species accounts and illustrations herein). This variation poses a systematic question: Are species limits correctly in- ferred, and do the color patterns thus truly represent intraspecific variation? Assum- ing that the answer to the first question is “yes,” how is the color pattern variation re- lated to the natural history of the snakes? Species of Dipsadini are widely sus- pected to mimic pit vipers in color pat- terns and behaviors (Greene and Mc- Diarmid, 2005). Such mimicry has been specifically suggested for Sibynomorphus neuwiedii (Sazima, 1992), and defensive behaviors similar to that of S. neuwwiedii have been reported for S. mikanii (Cadle and Myers, 2003). Perhaps the extreme pattern polymorphism in several northern species of Sibynomorphus is associated Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 with specific and, as yet unknown, defen- sive behaviors or mimicry systems. Some northern species of Sibynomor- phus are known to be regionally sympatric with species of Bothrops: S. oligozonatus and S. petersi with B. lojanus and B. os- bornei in southern Ecuador (Campbell and Lamar, 2004; Parker, 1930, 1932, 1938); S. oneilli and S. williamsi with B. pictus, B. roedingeri, or both in northern and central Peru (inferred from distribu- tions in Campbell and Lamar, 2004). In addition, some of these species of Both- rops are highly polymorphic in color and pattern (e.g., B. pictus; Campbell and La- mar, 2004). Parallel studies of geographic color pattern variation could shed light on potential mimicry systems in these snakes, as could a study of populations at localities in which high intrapopulational pattern variation has been documented (e.g., S. oneilli in the Rio Santa valley of Ancash Department, Peru; S. vagus from the vi- cinity of Huancabamba, Piura Depart- ment, Peru). Although current evidence suggests that little or no ontogenetic change in color pattern occurs in Sibyno- morphus, more detailed investigations of this phenomenon should be conducted. Do potential models (Bothrops) and mim- ics (Sibynomorphus) undergo parallel on- togenetic color changes, as is known in some other snake mimicry systems (Greene and McDiarmid, 2005)? A second intriguing aspect of the biol- ogy of Sibynomorphus and other Dipsa- dini (particularly Dipsas) is the extreme variation in external scalation (Cadle, 2005; Cadle and Myers, 2003; Peters, 1960). The variation in standard scutella- tion characters, particularly of the head, seems greater than occurs in many other colubrid groups. It calls for study in mech- anistic terms (What developmental pro- cesses result in the variation?); in function- al biology terms (What is the functional significance of the variation?); and in evo- lutionary terms (What is the phylogenetic distribution and origin of the variation?). It is tempting to link variability in scu- SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle 2 tellation to the molluscivorous diet of Dip- sadini (and perhaps some correlated changes such as the musculature underly- ing the temporal scales; see Dunn, 1951). Indeed, the absence of a mental groove and the peculiar form of the chin shields have been linked specifically with the abil- ity to consume gastropods (Gans, 1952: footnote 5; Peters, 1960: 17-18). But that can be only part of the story because Si- bynomorphus and Dipsas are more de- rived in these two characters than are Si- bon and Tropidodipsas (see above discus- sion). Sibynomorphus itself is less variable in some head scale characters than are at least some Dipsas (e.g., scales in the loreal and temporal regions). The variation among species could be related to differ- ent functional requirements for different prey, and different species of Dipsadini are already known to have different prey pref- erences and correlated morphology (e.g., in Sibynomorphus; LaPorta-Ferreira et al., 1986: Sazima, 1989). Functional and com- parative studies of head scale variation in snakes could shed light on this phenome- non in Dipsadini. A fruitful avenue of re- search also includes a variational study of the Asian molluscivores (Pareatini). Do their patterns of variation entail the same scales and to the same degree of variation as in Dipsadini? This could suggest in- sights into how the variation might be re- lated, if at all, to molluscivory in colubrids. Species of Sibynomorphus in Ecuador and Peru offer many opportunities for deepening our understanding of the biol- ogy of the South American radiation of Dipsadini. Hopefully, the analyses pre- sented here will stimulate further studies of these snakes to resolve outstanding sys- tematic and phylogenetic questions and will serve as a point of departure for im- proving knowledge of their natural history. ACKNOWLEDGMENTS The following collection personnel of- fered indispensable and generous loans and other assistance during the course of this study: Ted Daeschler and Ned Gil- ~l Ol more (ANSP); Charles J. Cole, linda Ford, and Charles W. Myers (AMNH); E. Nicholas Arnold, Colin J. McCarthy, and Mark Wilkinson (BMNH); Ana Almendar- z (EPN); Robert F. Inger, Maureen Kear- ney, Alan Resetar, and Harold K. Voris (FMNH); Hugo Alamillo, Juan M. Guay- asamin, John Simmons, Omar Torres-Car- vajal, ard Linda Trueb (KU); James Han- ken and José P. Rosado (MCZ): César Aguilar, Nelly Carrillo de Espinoza, and Jests Cordova (MUSM); Barbara Stein and David B. Wake (MVZ): and W. Ronald Heyer, Roy W. McDiarmid, Robert Wil- son, and George R. Zug (USNM). I es- pecially thank Ana Almendariz (EPN) and Jestis Cordova and Cesar Aguilar (MUSM) for kindnesses on visits to their institu- tions. Ana Almendariz arranged for the loan of the holotype of S. oligozonatus and critical specimens of S. petersi; these were hand carried to the U.S. by John Simmons (KU), and for those efforts I am extremely grateful. Personnel of the FMNH provid- ed work space and hospitality during much of the work on this project. I am additionally grateful to many col- leagues for specific and critical assistance during this study. Charles W. Myers called my attention to the specimen of Sibyno- morphus oligozonatus he collected and permitted NE WO jOREOLIKS ies hemipenis; provided copies of the field notes of G. K. Noble in the AMNH Department of Her- petology archives; and provided copies of two crucial references, including a scan of a plate from Jan and Sordelli. Grace Tilger of AMNH assisted in ferreting out AMNH archival sources. Roy W. McDiarmid also provided copies of Jan (1863) and plates from Jan and Sordelli. John P. O'Neill gave me details on the circumstances of collec- tion of the holotype of Sibynomorphus oneilli. Charles Q. Forester took photo- graphs of several specimens in the MUSM. Richard Thomas provided copies of field notes on specimens he collected. Karsten Hartel advised me about the fish identities and distributions associated with the description of Leptognathus atypicus. 276 William Alverson, Robin Foster, and es- pecially Michael O. Dillon helped with un- derstanding plant communities. Field work was supported by the Field Museum of Natural History, the American Philosophical Society, the Putnam Fund of the Museum of Comparative Zoology, The David Rockefeller Center for Latin Amer- ican Studies (Harvard University), and a faculty grant from School of Arts and Sci- ences of Harvard University. Additional support from Conservation International permitted the examination of specimens in Quito. My field work would have been im- possible without the efforts and camara- derie of my field companions: Pablo Chu- na Mogollon, Camilo Diaz, Michael O. Dillon, the late Alwyn H. Gentry, José Guevarra Barreto, Segundo Leiva, Pedro Lezama, Rosa Ortiz de Gentry, Raul Qui- roz, Abtindio Sagastegui Alva, José Santis- teban, and Helena Siesniegas. I am grate- ful to the Direccién General Forestal y de Fauna del Perti and the Museo de la Univ- ersidad Nacional de San Marcos in Lima for their many years’ support of my efforts to understand the biodiversity of Peru. I am grateful to Charles W. Myers and an anonymous reviewer for helpful, detailed comments on the manuscript, and for sav- ing me from a few errors (any that remain are mine). SPECIMENS EXAMINED AND LOCALITY RECORDS AMNH American Museum of Natural History, New York ANSP Academy of Natural Sciences of Philadelphia BMNH The Natural History Museum, London EPN Escuela Politécnica Nacional, Quito, Ecuador FMNH Field Museum of Natural His- tory, Chicago KU Museum of Natural History, University of Kansas, Lawrence MCZ Museum of Comparative Zoolo- gy, Harvard University, Cam- bridge Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 MUSM Museo de Historia Natural de San Marcos, Lima, Peru National Museum of Natural History, Smithsonian Institution, Washington, D.C. USNM Sibynomorphus oligozonatus ECUADOR: Azuay: Zhila, Parroquia Ona: ‘Canton? Giron, 2.250mime(Os2o04s: 79°12’W],!°9 EPN 3612 (holotype). About 6 km airline W Santa Isabel, 1,440 m, Rio Jubones drainage [03°16'S, 79°19’W], AMNH 110587. Loja: Catamayo Valley, 30 km W Loja [approximately 04°01'S, 79°20’W], BMNH 1935.11.3.108. PERU: PruRA: No specific locality, MUSM 2192, 2248. Sibynomorphus oneilli PERU: ANCASH: Provincia Antonio Raimondi: Yuracyacu, 3,500 m [09°15’S, TOL W298) MECISIVINS2 TI 3296m So0ar 3334, 3392, 3448. Huardz [3,060 m:; 09°32’S, 77°32'W], MUSM 3395, 2660. Provincia Huaylas: Huaylas [2,721 m; 08°52'S, 77°54'W], MUSM 2662, 2687, 3089. Provincia Huaylas: Hacienda Santa Rosa, 1,800 m [now Provincia Santa: ap- proximately 08°46’S, 78°08'W ?!], MUSM 2390. AMAZONAS: 5,400 ft. (1,646 m) on road between Balsas and Abra Chanchillo [06°49'S, 77°59’'W], LSUMZ 33736 (ho- '8 Coordinates for this locality given by the GeoNet Names Server (03°30’S, 79°12’W) differ from those given in the original description (03°39'45’S, 79°17'26"W). Consultation with several maps sug- gests that the former are more accurate. The Global Gazetteer gives the elevation of Zhila as 2,818 m. > Instituto Geografico Militar, Departamento de Ancash, 1:400,000, 1979. "1 Stiglich (1922) indexes two haciendas in Ancash Department with the name “Santa Rosa”: one in the District of Macate, the other in the District of Pueblo Libre. All of the territory of Pueblo Libre is 22,000 m, whereas Macate has significant area <2,000 m, and I infer this is the Santa Rosa referred to. Macate has subsequently been incorporated into Santa Prov- ince (IGM Departamento de Ancash, 1:400,000, 1979). The coordinates are an approximation on the basis of the closest territory to Macate that is <2,000 m. MUSM 2390 was collected by Roberto Donoso- Barros, but no date was recorded. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle lotype). CAJAMARCA: Abra Gelic, 13 km E Celendin, 3,080 m [06°51'S, 78°08’W], KU 212599. Provincia Cajabamba, 10 km SSE Cajabamba, 2, 900 m [approximately 07°37'S, 78°03'’W]|, KU 212600. Provincia Cajabamba: Cajabamba [2,654 m; 07°37’S, 78°03'W]|, MUSM 3103. LA LIBERTAB: Provincia Santiago de Chuco: Hacienda Ticapampa |= Hacienda Ticabamba, Dis- trito de Mollepata; Stiglich, 1922; approx- imately 08°11’S, 77°57'W, the coordinates of Mollepata], MUSM 2770. Sibynomorphus cf. oneilli PERU: “Bellavista,” MUSM 2258 (see text discussion at end of species account for S. vagrans). Sibynomorphus petersi ECUADOR: Azuay: Zhila, Parroquia Ona, Canton Girén, 2,250 m [03°30’S, 79°12'W; see footnote 19], EPN 2659 (ho- lotype; not seen), 2660 (paratype). Loja: Loja City, 2,200 m [04°00’S, 79°13'W], BMINED 1924624711 0-111 1930:1.30:7, 1930.10.12.24 25, 1931.2.12.30-31, BMNH 1933.6.24.110-111. Vicinity of Loja City, 1,280 ft: [2.219 m; 04°00’S, 79°13’W], BMNH 1935.11.3.111. 0.5 km E Loja City on road to Zamora, 6,765 feet [2,063 m:; 04°00'S, 79°13’W]|, USNM 237040. 2 km FE, Loja, 2,200 m [04°00’S, 79°13’W], KU 121309. 10 km E Loja City [04°00’S, 79°11'W], EPN 4737. Catamayo Valley, 30 km W Loja, 4,780 ft. [1,457 m; 04°01’S, 79°20’W], BMNH 1935.11.3.109-10. 7.3 km N Loja, 2,080 m [03°58'S, 79°13’W], KU 142804. Cant6n Saraguro, El Tablén, 03°28'42"S, 79°10’30"W, EPN 1847 (not seen; Orcés and Almendariz, 1989). PERU: ANCASH: Malvas, 3,120 m [09°55'S, 77°39'W], SMF 80048. Cajacay, 2,599 m [10°09'S, 77°26’W], MUSM 20583 (not seen; Lehr et al., 2002).22 Ca- * T assume that the identity of MUSM 20583 is the same as SMF 80048, which I examined. Both speci- mens are from the same collection and from geo- graphically close localities (see Lehr et al., 2002). PATCH JAMARCA: 3-4 km N Paraguay (along road from Querocoto to Hacienda La Granja), PAZ ON aie |O62210S 79205 W|-22 MU SM= JEC 12884. Approximately 1 km NE (air- line) of Monte Seco on trail between Mon- te Seco and Chorro Blanco, Rio Zafia, 1,250 m [06°51'S, 79°06’W], FMNH 932569. El Chorro, 1 km N (airline) Mon- te Seco, 1,400 m, Rio Zana, ANSP 31806 (= stomach contents, not separately cata- logued, associated with a Micrurus mer- tensi (see Fig. 26). San Pablo [2,400 m; see Cadle, 1998: footnote 6; 07°07'S, 78°50'W], BMNH 1900.3.30.19. Bosque de Cachil, 3 km (airline) SW Contumaza, 2,200—2,400 mt LO7223' Savor wb MCZ 178047, Pr URA: 33 km SW Huancabamba, below (west) crest of the Cordillera Huancabam- ba [“below camp, ca. 6800 ft” (2,073 m) fide field notes of Richard Thomas; ap- proximately 05°21'S, 79°34'W],* LSUMZ DIAL Sibynomorphus vagrans PERU: CAJAMARCA: Bellavista [421 m; Oar 74S WOOO NY | ea MiGZa k7284) = (holo: type), 17428, 17436—52 (paratypes). Sibynomorphus vagus PERU: [PrurA]: Huancabamba [1,929 OMA Sem De Save INGZ2 17420=23) 23 Coordinates derived from 1: 100,000 Carta Na- cional Incahuasi, Departamento Cajamarca (IGM, 1965, reprinted 1973). *4 This locality is roughly equivalent to “Can- chaque” or “Cruz Blanca” in some literature on this locality. See footnote 13. » Paratypes of Sibynomorphus vagrans comprised 26 specimens (Dunn, 1923), but Barbour and Loy- eridge (1929) indicated only 23 [sic] at that time: MCZ 17430-53 [= 24 specimens]. Several paratypes were exchanged to other collections: Instituto Butan- tan (17429-30, 17435), University of Illinois Museum of Natural History (17431), Hebrew University of Je- rusalem (17432), AMNH (17433, 17453), and the Naturhistorischen Museums in Wien, Austria (17434). These exchanged specimens were not ex- amined for this study. 278 17425-2627"; Huancabamba (1,900 m), KU 135180; Huancabamba (1,820 m), KU 219121-22. 5.7 km N Huancabamba on road “to Salalay W920 im KUr 219123: MUSM 15464—65 (not seen; Rossman and Kizirian, 1993). Sibynomorphus williamsi PERU: ANCASH: Huayllacayan [3,626 m (?)?*; 10°15’S, 77°26'W; also spelled Huayacayan], MUSM 2665. Provincia Bol- ognesi: Huanchay [1,857 m; 10°31’S, 77°25'W ; on the Quebrada Huanchay, a tributary of the Rio Pativilca], MUSM 3025. Provincia Bolognesi, Distrito Huayl- lacayanw umpes 22930 r mnie O alta. sp 77°29’W], MUSM 3378, 3412-14, 3475. LIMA: Distrito San Juan de Lurigancho: Canto Grande [311 m; 11°59’S, 77°01’W], MUSM 10034. Lomas de Atocongo [to 540 m; 12°08'S, 76°54’'W], MUSM 19236. Lomas de Lachay [300-900 m; 11°21'S, 77°23'W], MUSM 2668 (paratype), 3400. Lomas de Lurin [10 m; 12°17'S, 76°52’W], MUSM 2171 (paratype). Radio Observa- torio de Jicamarca [approximately 300-400 mae lay {Se Ova 2eVV eNOS Mie On Che= lotype), 2173-74, 2867 (paratypes). Lomas de Pachacamac [75 m; 12°14’S, 76°53’W] °° MCZ 17424 and 17427 were exchanged to the Instituto Butantan. 27 According to the IGM 1:400,000 Ancash De- partment map, Huayllacayan is between 3,000 and 4,000 m. According to the Global Gazetteer, its ele- vation is 11,899 ft. (= 3,626 m), which is much higher than the next lower recorded elevation for S. william- si (2,989 m). Another specimen, MUSM 3376, is list- ed from the same locality in the MUSM catalogues but explicitly notes the elevation as “2,700 m.” The latter specimen is a juvenile with no discernible dor- sal pattern, but all scutellation features are similar to S. williamsi except for 8 supralabials (4-5 touching the eye); in addition, the venter appeared to have large bold spots, which is uncharacteristic of most specimens of S. williamsi. For these reasons, I ex- cluded MUSM 3376 from the study, but it indicates that specimens from this locality could come from lower elevations near Huayallcayaén, in which case the upper elevational extent of S. williamsi is not known with precision. Another interpretation is that the cat- alogue entry “Huayllacayén” refers to the political unit district (Distrito) and not to the town of the same name. Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 (Carrillo de Espinoza, 1974). Provincia Huarochiri: Santa Eulalia [1,036 m: 11°51'S, 76°40’W], MUSM 3137. LITERATURE CITED ALMENDARIZ, A., AND G. ORCES V. 2004. Distribu- cién de algunas especies de la herpetofauna de los pisos: altoandino, temperado y subtropical. Politécnica (Revista de Informaci6én Técnico— Cientifica, Quito), 25(1): 97-149 + 2 pls. and 1 foldout map. AMARAL, A. DO. “1929”a [1930]. Estudos sobre ophi- dios neotropicos. XVII—valor systemdatico de varias formas de ophidios neotropicos. Memorias do Instituto Butantan, 4: 3-68. . -1929”b [1930]. Estudos sobre ophidios neo- tropicos XVIII. Lista remissiva dos ophidios da Regiao Neotropica. Memorias do Instituto Bu- tantanw4 7S ale . “1929’c [1930]. Contribuigéo ao conheci- mento dos ophidios do Brasil. IV. Lista remissiva dos ophidios do Brasil. Memorias do Instituto Butantan, 4: 71-125. BARBOUR, T., AND A. LOVERIDGE. 1929. Typical rep- tiles and amphibians in the Museum of Compar- ative Zoology. Bulletin of the Museum of Com- parative Zoology, 69(10): 205-360. BOETTGER, O. 1888. Materialien zur herpetologisch- en Fauna von China II. (2. Erneute Aufzahlung der Reptilien u. Batrachier des chinesischen Reiches). Berichte iiber die Thitigkeit des Of fenbacher Vereins fiir Naturkunde, 26—28: 53— 191+: 2 plst BONNET, X., R. SHINE, e& NAULLEAU, AND M. VACH- ER-VALLAS. 1997. Sexual dimorphism in snakes: different reproductive roles favour different body plans. Proceedings of the Royal Society of Lon- don B, 265: 179-183. BOULENGER, G. A. 1894. Catalogue of the snakes in the British Museum (Natural History). Volume II. London: British Museum (Natural History). x O82) 20%pls. . 1896. Catalogue of the snakes in the British Museum (Natural History). Vol. 3. London: Brit- ish Museum (Natural History). xiv + 727 pp. + 25 pls. . 1899. Descriptions of new reptiles and ba- trachians collected by Mr. P. O. Simons in the Andes of Ecuador. Annals and Magazine of Nat- ural History, series 7, 4(24): 454-457. . 1900. Descriptions of new batrachians and reptiles collected by Mr. P. O. Simons in Peru. Annals and Magazine of Natural History, series 7, 6(32): 181-186. . 1901. Further descriptions of new reptiles collected by Mr. P. O. Simons in Peru and Bo- livia. Annals and Magazine of Natural History, series 7, 7(42): 546-549. . 1902a. List of the fishes, batrachians, and reptiles collected by the late Mr. P. O. Simons in SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle the Provinces of Mendoza and Cordova, Argen- tina. Annals and Magazine of Natural History, series 7, 9(53): 336-339. . 1902b. Descriptions of new batrachians and reptiles from the Andes of Peru and Bolivia. An- nals and Magazine of Natural History, series 7, 10(59): 394—402. : BRITES, V. L. D. C., AND F. A. BAUAB. 1988. Fauna ofidiana do municipio de Uberlandia, Minas Ger- ais—Brasil. I. Ocorrencia na area Urbana. Re- portes da Centro Cientifico Biomedico da Univ- ersidade Federal Uberlandia, 4(1): 3-8. BRONGERSMA, L. D. 1956. The palato-maxillary arch in some Asiatic Dipsadinae (Serpentes). Pro- ceedings of the Koninklijk Nederlandse Akade- mie van Wetenschappen, Amsterdam, series C, Biological and Medical Sciences, 59(4): 439-446. . 1958. Some features of the Dipsadinae and Pareinae (Serpentes, Colubridae). Proceedings of the Koninklijk Nederlandse Akademie van Wetenschappen, Amsterdam, series C, Biological and Medical Sciences, 61(1): 7—12. CADLE, J. E. 1984a. Molecular systematics of Neo- tropical xenodontine snakes. II. Central Ameri- can xenodontines. Herpetologica, 40(1): 21—30. . 1984b. Molecular systematics of Neotropical xenodontine snakes. III. Overview of xenodonti- ne phylogeny and the history of New World snakes. Copeia, 1984(4): 641-652. . 1985. The Neotropical colubrid snake fauna: lineage components and biogeography. System- atic Zoology, 34(1): 1-20. . 1989. A new species of Coniophanes (Ser- pentes: Colubridae) from northwestern Peru. Herpetologica, 45(4): 411-424. . 1991. Systematics of lizards of the genus Stenocercus (Iguania: Tropiduridae) from north- ern Peru: new species and comments on rela- tionships and distribution patterns. Proceedings of the Academy of Natural Sciences of Philadel- phia, 143: 1-96. . 1996. Systematics of snakes of the genus Geodipsas (Colubridae) from Madagascar, with descriptions of new species and observations on natural history. Bulletin of the Museum of Com- parative Zoology, 155(2): 33-87. . 1998. New species of lizards, genus Steno- cercus (Iguania: Tropiduridae), from western Ec- uador and Peru. Bulletin of the Museum of Comparative Zoology, 155(6): 257-297. . 2005. Systematics of snakes of the Dipsas oreas complex (Colubridae: Dipsadinae) in west- em Ecuador and Peru, with revalidation of D. elegans (Boulenger) and D. ellipsifera (Boulen- ger). Bulletin of the Museum of Comparative Zo- ology, 158(3): 67-136. CADLE, J. E., AND P. CHUNA M. 1995. A new lizard of the genus Macropholidus (Teiidae) from a re- lictual humid forest of northwestern Peru, and notes on Macropholidus ruthveni Noble. Bre- viora, 501: 1—39. 279 CADLE, J. E., AND H. W. GREENE. 1993. Phyloge- netic patterns, biogeography, and the composi- tion of Neotropical snake assemblages, pp. 281— 293. In R. E. Ricklefs and D. Schluter (eds.), Species Diversity in Ecological Communities: Historical and Geographical Perspectives. Chi- cago: University of Chicago Press. CADLE, J. E.. AND R. W. MCDIARMID. 1990. Two new species of Centrolenella (Anura, Centrolen- idae) from the western slope of the Andes in northern Peru. Proceedings of the Biological So- ciety of Washington, 103(3): 746-768. CADLE, J. E., AND C. W. MYERS. 2003. Systematics of snakes referred to Dipsas variegata in Panama and western South America, with revalidation of two species and notes on defensive behaviors in the Dipsadini (Colubridae). American Museum Novitates, 3409: 1-47. CAMILLERI, C., AND R. SHINE. 1990. Sexual dimor- phism and dietary divergence: differences in tro- phic morphology between male and female snakes. Copeia, 1990(3): 649-687. CAMPBELL, J. A., AND W. W. LAMAR. 2004. The Ven- omous Reptiles of the New World. Vols. 1 and 2. Ithaca, New York: Comstock Publishing Associates. xviii + 1-476 + 28 pp. (Vol. 1); xiv + 477-870 + 28 pp. (Vol. 2); 1500 pls. on 127 pp.; 8 maps. CARRILLO DE ESPINOZA, N. 1974. Sibynomorphus williamsi nov. sp. (Serpentes: Colubridae). Pub- licaciones del Museo de Historia Natural “Javier Prado,” serie A (Zoologia), 24: 1-16. CARRILLO DE ESPINOZA, N., AND J. ICOCHEA. 1995. Lista taxonodmica preliminar de los reptiles vi- vientes del Perti. Publicaciones del Museo de Historia Natural de la Universidad Nacional de San Marcos, serie A (Zoologia), 49: 1-27. CECHIN, S. Z., AND J. L. OLIVEIRA. 2003. Sibyno- morphus ventrimaculatus (southern snail-eater): mating. Herpetological Review, 34(1): 72. CEI, J. M. 1993. Reptiles del Noroeste, Nordeste y Este de la Argentina. Torino: Museo Regionale di Scienze Naturali. 949 pp. CHAPMAN, F. M. 1926. The distribution of bird-life in Ecuador; a contribution to a study of the ori- gin of Andean bird-life. Bulletin of the American Museum of Natural History, 55: 1-784. CHUBB, C. 1919. Notes on collections of birds in the British Museum, from Ecuador, Peru, Bolivia, and Argentina. Ibis, 1919: 1-55, 256-290. CLAPPERTON, C. 1993. Quaternary Geology and Geo- morphology of South America. Amsterdam: El- Sevier xvi 4-779) pp. COCHRAN, D. M. 1961. Type specimens of reptiles and amphibians in the U.S. National Museum. Bulletin of the United States National Museum, 220: 1-291. Cope, E. D. 1868. An examination of the reptilia and batrachia obtained by the Orton expedition to Equador and the upper Amazon, with notes on other species. Proceedings of the Academy of Natural Sciences of Philadelphia, 20: 96-140. 280 . 1874a. Description of some species of rep- tiles obtained by Dr. John F. Bransford, assistant surgeon United States Navy, while attached to the Nicaraguan Surveying Expedition in 1873. Proceedings of the Academy of Natural Sciences of Philadelphia, 26: 64-72. . 1874b. On some Batrachia and Nematogna- thi brought from the Upper Amazon by Prof. Or- ton. Proceedings of the Academy of Natural Sci- ences of Philadelphia, 26: 120-137. 1895. The classification of the Ophidia. Transactions of the American Philosophical So- ciety, 18: 186-219 + pls. xiv—xxxiii. CUNHA, O. R. DA, AND F. P. DO NASCIMENTO. 1993. Ofidios da Amaz6nia. As cobras da regiao leste do Para [2nd ed.]. Boletim do Museu Paraense Emilio Goeldi, série Zoologia, 9(1): 1-191. DILLON, M. O., A. SAGASTEGUI ALVA, I. SANCHEZ VEGA, S. LLATAS QUIROZ, AND N. HENSOLD. 1995. Floristic inventory and biogeographic anal- ysis of montane forests in northwestern Peru, pp: 251-269. In S. P. Churchill, H. Balslev, E. For- ero, and J. L. Luteyn (eds.), Biodiversity and Conservation of Neotropical Montane Forests. Bronx: The New York Botanical Garden. DINGERKUS, G., AND L. D. UHLER. 1977. Enzyme clearing of alcian blue stained whole small ver- tebrates for demonstration of cartilage. Stain Technology, 52: 229-232. DIXON, J. R., AND R. B. HUEY. 1970. Systematics of the lizards of the gekkonid genus Phyllodactylus of mainland South America. Natural History Mu- seum of Los Angeles County, Contributions in Science, 192:1—78. DIXON, J. R., AND J. W. WRIGHT. 1975. A review of the lizards of the iguanid genus Tropidurus in Peru. Natural History Museum of Los Angeles County, Contributions in Science, 271: 1-39. DOAN, T. M., AND W. A. ARRIAGA. 2002. Microgeo- graphic variation in species composition of the herpetofaunal communities of Tambopata re- gion, Peru. Biotropica, 34(1): 101-117. DOWLING, H. G. 1951. A proposed standard system of counting ventrals in snakes. British Journal of Herpetology, 1: 97-99. DOWLING, H. G., AND J. M. SAVAGE. 1960. A guide to the snake hemipenis: a survey of basic struc- ture and systematic characteristics. Zoologica, AS; 17-31. Downs, F. L. 1967. Intrageneric relationships among colubrid snakes of the genus Geophis Wagler. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 131: 1-193. DUELLMAN, W. E. 1958. A monographic study of the colubrid snake genus Leptodeira. Bulletin of the American Museum of Natural History, 114(1): 1-152 + 31 pls. . 1979. The herpetofauna of the Andes: pat- terns of distribution, origin, differentiation, and present communities, pp. 371-459. In W. E. Duellman (ed.), The South American Herpeto- Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 fauna: Its Origin, Evolution, and Dispersal, Mon- ogr. 7. Lawrence: The Museum of Natural His- tory, University of Kansas. . 2005. Cusco Amazé6nico: The Lives of Am- phibians and Reptiles in an Amazonian Rainfo- rest. Ithaca, New York: Cornell University Press. xv + 433 pp. + 236 pls. on 32 pp. DUELLMAN, W. E., AND A. W. SALAS. 1991. Annotat- ed checklist of the amphibians and reptiles of Cuzco Amazonico, Peru. Occasional Papers of the Museum of Natural History, University of Kansas, 143: 1-13. DUMERIL, A. M. C., G. BIBRON, AND A. DUMERIL. 1854. Erpétologie Générale ou Histoire Natu- relle Compléte des Reptiles, Vol. 7. Paris: Li- brairie Encyclopédique de Roret, premiére par- tie, vii + 4 + xvi + 780 pp.; deuxiéme partie, xii + 781-1536 pp. DUNN, E. R. 1923. Some snakes from northwestern Peru. Proceedings of the Biological Society of Washington, 36: 185-188. . 1935. The snakes of the genus Ninia. Pro- ceedings of the National Academy of Sciences, USA, 21(1): 9-12. . 1951. The status of the snake genera Dipsas and Sibon, a problem for “quantum evolution.” Evolution, 5(4): 355-358. ESPINOZA, R. E., J. J. WIENS, AND C, R. TRACY. 2004. Recurrent evolution of herbivory in small, cold- climate lizards: breaking the ecophysiological rules of reptilian herbivory. Proceedings of the National Academy of Sciences, USA, 101(48): 16819-16824. ETHERIDGE, R. 1995. Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Lio- laeminae (Reptilia: Squamata: Tropiduridae). American Museum Novitates, 3142: 1—34. FARRAR, E., AND D. C. NOBLE. 1976. Timing of late Tertiary deformation in the Andes of Peru. Geo- logical Society of America Bulletin, 87: 1247-1250. FERNANDES, R. 1995. Phylogeny of the Dipsadine Snakes. Ph.D. Dissertation, University of Texas at Arlington, 115 pp. FERNANDES, R., D. S. FERNANDES, AND P. PASSOS. 2002. Leptognathus latifasciatus Boulenger, 1913, a junior synonym of Dipsas polylepis (Bou- lenger, 1912) (Serpentes, Colubridae). Boletim do Museu Nacional (Rio de Janeiro), nova série, Zoologia, 493: 1-7. FRANCO, F. L. 1994. O género Sibynomorphus Fit- zinger, 1843, no Brasil (Colubridae; Xenodonti- nae; Dipsadini). Dissertagaéo de Mestrado, Pon- tificia Universidade Catélica do Rio Grande do Sul, Rio Grande do Sul, Brasil, 148 pp. FRANCO, F. L., P. SANCHES, AND C. A. FALCETTI. 2002. Variagio morfolégica e distribuigao geo- grafica das serpentes dormideiras do complexo de espécies associadas a Sibynomorphus neuwie- di (Ihering, 1911) (Serpentes, Colubridae, Dip- sadinae). Resumos do III Simpésio do Programa Biota/FAPESP (Fundagao de Amparo a4 Pesquisa SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR * Cadle do Estado de Sao Paulo, FAPESP. Centro de Re- feréncia em Informagéo Ambiental, CRIA). Ab- stract online at: www.biota.org.br/publi/banco/ indexPshow+91144183. Accessed 10 July 2006. FRANCO, F. L., P. SANCHES, C. A. FALCETTI, AND R. FERNANDES. 2003. Designagao de lect6tipo para Sibynomorphus neuwiedi (Ihering, 1911) e re- descrigao do holétipo de Leptognathus garmani Cope, 1887 (Serpentes, Colubridae; Dipsadinae). IV Simposio de Avaliagaéo Biota (Fundagao de Amparo a Pesquisa do Estado de Sao Paulo, FA- PESP. Centro de Referéncia em Informagao Am- biental, CRIA). Abstract online at: www.biota. org. br/publi/banco/indexPshow+ 90869164. Ac- cessed 10 July 2006. FROST, D. R., M. T. RODRIGUES, T. GRANT, AND T. A. Titus. 2001. Phylogenetics of the lizard genus Tropidurus (Squamata: Tropiduridae: Tropiduri- nae): direct optimization, descriptive efficiency, and sensitivity analysis of congruence between molecular data and morphology. Molecular Phy- logenetics and Evolution, 21(3): 352-371. FUGLER, C. M., AND J. CABOT. 1995. Herpetologica boliviana: una lista comentada de las serpientes de Bolivia con datos sobre su distribucién. Ecol- ogia en Bolivia, 24: 40-87. FUGLER, C. M., AND I. DE LA RIvA. 1990. Herpe- tologia Boliviana: lista provisional de las serpien- tes conocidas en el pais. Museo Nacional de His- toria Natural (Bolivia) Comunicacion, 9: 22—53. GANS, C. 1952. The functional morphology of the egg-eating adaptations in the snake genus Da- sypeltis. Zoologica, 37(4): 209-244 + pls. I-IV. GREENE, H. W., AND R. W. MCDIARMID. 2005. Wal- lace and Savage: heroes, theories, and venomous snake mimicry, pp. 190-208. In M. A. Donnelly, B. I. Crother, C. Guyer, M. H. Wake, and M. E. White (eds.), Ecology and Evolution in the Trop- ics, a Herpetological Perspective. Chicago: Uni- versity of Chicago Press. HOoOGE, A. R., I. L. LAPORTA, AND S. A. ROMANO HoGE. “1978/1979” [1981]. Notes on Sibyno- morphus mikanii Schlegel 1837. Memorias do Instituto Butantan, 42/43: 175-178. HOOGMOED, M. S. 1980. Revision of the genus Atractus in Surinam, with the resurrection of two species (Colubridae, Reptilia). Notes on the her- petofauna of Surinam VII. Zoologische Verhan- delingen, Leiden, 175: 1-47. HOPKINS, D. J. (ed.) 1977. Merriam-Webster’s Geo- graphical Dictionary, Third Edition. Springfield, Massachusetts: Merriam-Webster, Inc. 26 + 1361 pp. JAN, G. 1863. Elenco sistematico degli Ofidi descritti e disegnati per ’Iconographia Generale. Milan: A. Lombardi. vii + 143 pp. JAN, G., AND F. SORDELLI. 1860-1881. Iconographie générale des ophidiens. Vols. 1-3, Livrs. 1—50. Milan: Jan and Sordelli. 300 pls. KOEPCKE, H. W. 1961. Synékologische studien an 281 der Westseite der peruanischen Anden. Bonner Geographische Abhandlungen, 29: 1-320. KOFRON, C. P. 1982. A review of the Mexican snail- eating snakes, Dipsas brevifacies and Dipsas gai- geae. Journal of Herpetology, 16(3): 270-286. . 1985a. Systematics of the Neotropical gastro- pod-eating snake genera, Tropidodipsas and Si- bon. Journal of Herpetology, 19(1): 84-92. . 1985b. Review of the Central American col- ubrid snakes, Sibon fischeri and S. carri. Copeia, 1985(1): 164-174. . 1987. Systematics of Neotropical gastropod- eating snakes: the fasciata group of the genus Sibon. Journal of Herpetology, 21(3): 210-225. . 1988. Systematics of neotropical gastropod- eating snakes: the sartorii group of the genus Si- bon. Amphibia Reptilia, 9: 145-168. . 1990. Systematics of neotropical gastropod- eating snakes: the dimidiata group of the genus Sibon, with comments on the nebulata group. Amphibia Reptilia, 11: 207-223. LAPORTA-FERREIRA, I. L., M. DA GRACA SALOMAO, AND P. SAWAYA. 1986. Biologia de Sibynomor- phus (Colubridae—Dipsadinae )—reprodugao e habitos alimentares. Revista Brasileira de Biolo- gia, 46(4): 793-799. LEHR, E., G. KOHLER, AND B. STREIT. 2002. Die Herpetofauna von Mittelperu entlang eines Transektes von der pazifischen Kiiste bis in die Hochanden (Amphibia et Reptilia). Faunistische Abhandlungen: Staatliches Museum fiir Tierkun- de Dresden, 22(24): 361-392. LEYNAUD, G. C., AND E. H. BUCHER. 1999. La fauna de serpientes del Chaco sudamericano: diversi- dad, distribucién geografica y estado de conser- vacion. Miscelanea de la Academia Nacional de Ciencias, Cordoba, Argentina, 98: 1-46. LOURDAIS, O., R. SHINE, X. BONNET, AND F. BRIS- CHOUX. 2006. Sex differences in body composi- tion, performance and behaviour in the Colom- bian rainbow boa (Epicrates cenchria maurus, Boidae). Journal of Zoology, 269: 175-182. LYNCH, J. D. 1982. Relationships of the frogs of the genus Ceratophrys (Leptodactylidae) and their bearing on hypotheses of Pleistocene forest re- fugia in South America and punctuated equilib- ria. Systematic Zoology, 31: 166-179. . 1986. Origins of the high Andean herpeto- logical fauna, pp. 478-499. In F. Vuilleumier and M. Monasterio (eds.), High Altitude Tropical Biogeography. Oxford: Oxford University Press. MACCULLOCH, R. D., AND A. LATHROP. 2004. A new species of Dipsas (Squamata: Colubridae) from Guyana. Revista de Biologia Tropical, 52(1): 239-247. MARQUES, O. A. V., A. ETEROVIC, AND W. ENDO. 2000. Seasonal activity of snakes in the Atlantic Forest in southeastern Brazil. Amphibia Reptilia, 4pap AOS SLI MARTINS, M., AND M. E. OLIVEIRA. 1998. Natural history of snakes in forests of the Manaus region, 282 Central Amazonia, Brazil. Herpetological Natu- ral History, 6(2): 78-150. MCCRANIE, J. R. 2006. New species of Sibon (Squa- mata: Colubridae) from northeastern Honduras. Journal of Herpetology, 40(1): 16-21. Myers, C. W. 1974. The systematics of Rhadinaea (Colubridae), a genus of New World snakes. Bul- letin of the American Museum of Natural His- tory, 153(1): 1-262. . 1982. Blunt-headed vine snakes (Imantodes) in Panama, including a new species and other revisionary notes. American Museum Novitates, 2738: 1-50. . 1984. Subcircular pupil shape in the snake Tantalophis (Colubridae). Copeia, 1984(1): 215— 216. . 2003. Rare snakes—five new species from eastern Panama: reviews of northern Atractus and southern Geophis (Colubridae: Dipsadinae). American Museum Novitates, 3391: 1—47. MYERS, C. W., AND J. E. CADLE. 1994. A new genus for South American snakes related to Rhadinaea obtusa Cope (Colubridae) and resurrection of Taeniophallus Cope for the “Rhadinaea” brevi- rostris group. American Museum Novitates, 3102: 1-33. . 2003. On the snake hemipenis, with notes on Psomophis and techniques of eversion: a re- sponse to Dowling. Herpetological Review, 34(4): 295-302. MYERS, C. W., AND J. A. CAMPBELL. 1981. A new genus and species of colubrid snake from the Si- erra Madre del Sur of Guerrero, Mexico. Amer- ican Museum Novitates, 2708: 1—20. MYERS, C. W., AND W. E. SCHARGEL. 2006. Morpho- logical extremes—two new snakes of the genus Atractus from northwestern South America (Co- lubridae: Dipsadinae). American Museum Novi- tates, 3532: 1-13. MYERS, C. W., AND L. TRUEB. 1967. The hemipenis of an anomalepidid snake. Herpetologica, 23(3): 935-238. NOBLE, D. C., E. H. MCKEE, T. MOURIER, AND F. MEGARD. 1990. Cenozoic stratigraphy, magmatic activity, compressive deformation, and uplift in northern Peru. Geological Society of America Bulletin, 102: 1105-1113. NOBLE, G. K. 1921. A search for the marsupial frog. Natural History, 21: 474493. NorMaN, D. R. 1994. Anfibios y Reptiles del Chaco Paraguayo, tomo 1. San José, Costa Rica: David Norman. vi + 281 pp. + 32 pls. ORCES V. G., AND A. ALMENDARIZ. 1989. Presencia en el Ecuador de los colubridos del género Si- bynomorphus. Politécnica (Revista de Informa- ci6n Técnico—Cientifica, Quito), 14(3): 57-67. ORTON, J. 1875. The Andes and the Amazon; or, Across the Continent of South America. 3rd ed. New York: Harper & Brothers. 645 pp. + 2 fold- out maps. PARKER, H. W. 1926. Description of a new snake Bulletin Museum of Comparative Zoology, Vol. 158, No. 5 from Trinidad. Annals and Magazine of Natural History, series 9, 18: 205-207. . 1930a. A new colubrine snake from Ecuador. Annals and Magazine of Natural History, series 10, 5: 207-209. . 1930b. Two new reptiles from southern Ec- uador. Annals and Magazine of Natural History, series 10, 5: 568-571. . 1932. Some new or rare reptiles and am- phibians from southern Ecuador. Annals and Magazine of Natural History, series 10, 9: 21-26. . 1934. Reptiles and amphibians from south- ern Ecuador. Annals and Magazine of Natural History, series 10, 14: 264-273. . 1938. The vertical distribution of some rep- tiles and amphibians in southern Ecuador. An- nals and Magazine of Natural History, series 11, 2: 438—450. PARKER 2 Al. TE as) ss CHULENBE RG Gye GRAVES, AND M. J. BRAUN. 1985. The avifauna of the Huancabamba region, northern Peru, pp. 169-188. In P. A. Buckley, M. S. Foster, E. S. Morton, R. S. Ridgely, and F. G. Buckley (eds.), Neotropical Ornithology. Washington, D.C.:: American Ornithologists’ Union. Passos, P., R. FERNANDES, AND M. PORTO. 2005. Geo- graphical variation and taxonomy of the snail-eating snake Dipsas albifrons (Sauvage, 1884), with com- ments on the systematic status of Dipsas albifrons cavalheiroi Hoge, 1950 (Serpentes: Colubridae: Dipsadinae). Zootaxa, 1013: 19-34. PAYNTER, R. A., JR. 1993. Ornithological Gazetteer of Ecuador. 2nd ed. Cambridge, Massachusetts: Museum of Comparative Zoology. xi + 247 pp. PEARSON, O. P., AND C. P. RALPH. 1978. The diversity and abundance of vertebrates along an altitudinal gradient in Peru. Memorias del Museo de His- toria Natural “Javier Prado,” 18: 1—97. PETERS, J. A. 1956. An analysis of variation in a South American snake, Catesby’s Snail-sucker (Dipsas catesbyi Sentzen). American Museum Novitates, 1783: 1-41. . 1960. The snakes of the subfamily Dipsadi- nae. Miscellaneous Publications of the Museum of Zoology, University of Michigan, 114: 1-224. . 1964. Supplemental notes on snakes of the subfamily Dipsadinae (Reptilia: Colubridae). Beitrage zur Neotropischen Fauna, 4: 45—50. . 1965. Liste der rezenten Amphibien und Reptilien: Colubridae (Dipsadinae). Das Tier- reich, 81: i-viii + 1-18. PETERS, J. A., AND B. OREJAS—MIRANDA. 1970. Cat- alogue of the Neotropical Squamata: Part I. Snakes. Bulletin of the United States National Museum, 297: 1—347. PoRTO, M., AND R. FERNANDES. 1996. Variation and natural history of the snail-eating snake Dipsas neivai (Colubridae: Xenodontinae). Journal of Herpetology, 30(2): 269-271. RODRIGUES, M. T. 2003. Herpetofauna da caatinga, pp. 181-236. In I. R. Leal, M. Tabarelli, and J. SIBYNOMORPHUS (COLUBRIDAE) IN PERU AND ECUADOR ° Cadle M. Cardoso da Silva (eds.), Ecologia e Conser- vacao da Caatinga. Recife, Brasil: Universidade Federal de Pernambuco. ROSSMAN, D. A., AND D. A. KIZIRIAN. 1993. Variation in the Peruvian dipsadine snakes Sibynomorphus oneilli and S. vagus. Journal of Herpetology, 271): 87-90. ROSSMAN, D. A., AND R. THOMAS. 1979. A new dip- sadine snake of the genus Sibynomorphus from Peru. Occasional Papers of the Museum of Zo- ology, Louisiana State University, 54: 1-6. RUNDEL, P. W., M. O. DILLON, B. PALMA, H. A. MOONEY, S. L. SULMON, AND J. R. EHLERINGER. 1991. The phytogeography and ecology of the coastal Atacama and Peruvian deserts. Aliso, 13(1): 1-50. SAGASTEGUI ALVA, A., I. SANCHEZ VEGA, M. ZAPATA CRUZ, AND M. O. DILLON. “2003” [2004]. Div- ersidad Floristica del Norte de Pert, Tomo II: Bosques Montanos. Trujillo, Peru: Universidad Antenor Orrego de Trujillo. 305 pp. SAVAGE, J. M. 1960. A revision of the Ecuadorian snakes of the colubrid genus Atractus. Miscella- neous Publications of the Museum of Zoology, University of Michigan, 112: 1—86. . 2002. The Amphibians and Reptiles of Costa Rica, a Herpetofauna between Two Continents, between Two Seas. Chicago: The University of Chicago Press. xx + 934 pp. SAVAGE, J. M., AND R. W. MCDIARMID. 1992. Redis- covery of the Central American colubrid snake, Sibon argus, with comments on related species from the region. Copeia, 1992(2): 421-432. SAZIMA, I. 1989. Feeding behavior of the snail-eating snake, Dipsas indica. Journal of Herpetology, 23(4): 464-468. . 1992. Natural history of the jararaca pitviper, Bothrops jararaca, in southeastern Brazil, pp. 199- 216. In J. A. Campbell and E. D. Brodie, Jr. (eds.), Biology of the Pitvipers. Tyler, Texas: Selva. ScoTT, N. J., JR. 1967. The colubrid snake, Tropi- dodipsas annulifera, with reference to the status of Geatractus, Exelencophis, Chersodromus an- nulatus, and Tropidodipsas malacodryas. Copeia, 1967(2): 280-287. SCROCCHI, G., M. PORTO, AND L. REy. 1993. Des- cripcion de una especie nueva y situacion del gé- nero Sibynomorphus (Serpentes: Colubridae) en la Argentina. Revista Brasileira de Biologia, 53(2): 197-208. SHINE, R. 1989. Ecological causes for the evolution of sexual dimorphism: a review of the evidence. Quarterly Review of Biology, 64(4): 419-461. . 1991. Intersexual dietary divergence and the evolution of sexual dimorphism in snakes. Amer- ican Naturalist, 138(1): 103-122. . 1993. Sexual dimorphism in snakes, pp. 49— S658 the Ae seigel and™ |p a Collisii(eds:), Snakes: Ecology and Behavior. New York: Mc- Graw Hill, Inc. SIMMONS, J. E. 2002. Herpetological Collecting and 283 Collections Management, revised ed. (Herpeto- logical Circular no. 31). Ithaca, New York: Soci- ety for the Study of Amphibians and Reptiles. vi + 153 pp. SIMPSON, B. B. 1975. Pleistocene changes in the flora of the high tropical Andes. Paleobiology, 1(3): 273-294. . 1979. Quaternary biogeography of the high montane regions of South America, pp. 157-188. In W. E. Duellman (ed.), The South American Herpetofauna: Its Origin, Evolution, and Dis- persal, Monogr. 7. Lawrence: Museum of Natu- ral History, University of Kansas. SMITH, H. M., AND E. H. TAYLOR. 1945. An anno- tated checklist and key to the snakes of Mexico. Bulletin of the United States National Museum, 187: iv + 1-239. SOKAL, R. R., AND F. J. ROHLF. 1981. Biometry, the Principles and Practice of Statistics in Biological Research. 2nd ed. New York: W. H. Freeman. xviii + 859 pp. STEINDACHNER, F. 1902. Herpetologische und ich- thyologische Ergebnisse einer Reise nach Siida- merika mit einer Einleitung von Therese Prin- zessin von Bayern. Denkschriften der Kaiserlich- en Akademie der Wissenschaften in Wien (Math- ematische-naturwissenschaftliche Klasse), 72: 89-148 + pls. 1-5. STEPHENS, L., AND M. A. TRAYLOR, JR. 1983. Orni- thological Gazetteer of Peru. Cambridge, Mas- sachusetts: Museum of Comparative Zoology. vi a= AMAL jo). STIGLICH, G. 1922. Diccionario Geografico del Pert. Vols. 1 and 2. Lima: Torres Aguirre. 1193 pp. UNDERWOOD, G. 1967. A Contribution to the Clas- sification of Snakes. London: British Museum (Natural History). x + 179 pp. VANZOLINI, P. E. 1948. Notas sObre os ofidios e la- gartos da Cachoeira de Emas, no municipio de Pirassununga, Estado de Sao Paulo. Revista Brasileira de Biologia, 8(3): 377-400. WALLACH, V. 1995. Revalidation of the genus Tro- pidodipsas Giinther, with notes on the Dipsadini and Nothopsini (Serpentes: Colubridae). Journal of Herpetology, 29(3): 476-481. WERNER, F. 1922. Synopsis der Schlangenfamilie der Amblycephaliden und Viperiden nebst Gbersicht Uber die kleineren Familien und die Colubriden der Acrochordinengruppe. Auf Grund des Bou- lengerschen Schlangenkatalogs (1893-1896). Ar- chiv fur Naturgeschichte [Berlin], 8: 185-244. ZAHER, H. 1999. Hemipenial morphology of the South American xenodontine snakes, with a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemipenes. Bulletin of the American Museum of Natural History, 240: 1-168. ZAHER, H., I. SOUZA, D. J. GOWER, E. HINGST-ZAH- ER, AND N. J. DA SILVA, JR. 2005. Resdescription of Atractus albuquerquei (Serpentes: Colubridae: Dipsadinae), with comments on geographical distribution and intraspecific variation. Papéis Avulsos de Zoologia, 45(2): 19-32. Li. enee Atos, ahs dd the ay Atte “ban by ae , . hae 4 nies : At Wt ee Ni ay 4 on a Mg iyi A ne ing Lit, allan ane Wats Fah Bs) AM bie po nee ae ; hee 43 te ( et oe i eS wv ie Rulletin OF THE Museum of Comparative Loology Anatomy of Eocaecilia micropodia, a-Limbed Caecilian of the Early Jurassic FARISH A. JENKINS, JR., DENIS M. WALSH, AND ROBERT L. CARROLL a ST SD Se ay a ea Ee RTs RP HARVARD UNIVERSITY VOLUME 158, NUMBER 6 CAMBRIDGE, MASSACHUSETTS, U.S.A. 20 August 2007 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY - BreEviora 1952— BULLETIN 1863— Memoirs 1865-1938 JounsoniA, Department of Mollusks, 1941-1974 OCCASIONAL PAPERS ON MOLLUusks, 1945— SPECIAL PUBLICATIONS. 1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and Evolution of Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere- dinidea (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. | 4. Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology and Evolutionary Biology: Essays in Honor of Emest E. Williams. 725 pp. 6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the M. C. Z., Vol. 108.) Reprinted 1971. ) Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. ‘ Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) | | Orinthological Gazetteers of the Neotropics (1975-). Peter’s Check-list of Birds of the World, vols. 1-16. : Oeste e of the New England Zoological Club 1899-1947. (Complete sets only.) rg | Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. eT ee This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2007. ANATOMY OF EOCAECILIA MICROPODIA, A LIMBED CAECILIAN OF THE EARLY JURASSIC FARISH A. JENKINS, JR.,1 DENIS M. WALSH, AND ROBERT L. CARROLL? CONTENTS I NGS EVAYG tg ee! LO ck Ree | dnt ah ek, Se LT ee 285 lantomisicin@mess eset pee es cae Ty Frat 286 Matenals andevicthodses.-- Fe ee 288 DD OST ELON Ses hee sot ah OO hae 290 Skid ae wee tee SE ee 290 Sit ROMs set ee pone Pete ks Aen uh 290 Sculpturing of the Skull Roof 299 PAL ate thee ra" BG se oon 05x Fy Thy A, 299 OseBasnler ee sia: eee see LAL: Scie 304 Plemnospneneid sen 2k sk Aes 310 SiOLECESOVEN EEN GTC eae Aes aaa ate leek py et 310 Stapes-quadnate sys ries eee 312 Wpencriliuua feel See tase es 316 Ibeweng law 7. 27 ven 8 a) veel See ot 318 Comparative Dental Morphology and OOLM MC OMMES, Ao. 2 8: LE 32m osteranial skeleton 22 ae th ee ees 330 QYWELE 1S ee Se en OE Se ee ee ee 330 Rosteranial Axial Skeleton .... 2. Bom AMS pel iee ne SRC Le a 331 Werepral Mmegiouss 2 2 ee ea! Ba5 ostarantalmineeion ka eee) ee tan Ba5 AMtenor Dorsal ecion es) 2) see ee 336 Middoxsal depron y..t.i sata ea 338 Posterior Dorsal, Sacral, and Caudal METH TAC rien es wn ae hee De ah 338 Comparative Anatomy of the Vertebral CP co earin a eee Oe HO eee Re? BR oe i OE Bt Ss 342 MppendieulariSkelotony:2 1/26 e Few ay 343 Anatomical Location of the Forelimb —__ 343 SeAzpMlOCOIACOldye sf es8 18 1s Py 9 343 PARAM T DIG Frese wae awl one ee 2h eel Ee 343 ' Department of Organismic and Evolutionary Biology, and Museum of Comparative Zoology, Har- vard University, Cambridge, Massachusetts 02138. E-mail: fjenkins@oeb.harvard.edu. * Department of Philosophy and Institute for the History and Philosophy of Science and Technology, University of Toronto, Toronto, Ontario M5S 1J5, Canada. E-mail: denis.walsh@utoronto.ca. ° Redpath Museum, McGill University, Montreal, Quebec H3A 2K6, Canada. E-mail: Robert.carroll@ megill.ca. Bull. Mus. Comp. Zool., 158(6): 285-366, August, 2007 TS ACIS eRe eee ee ee See 344 | CET on ie 0 Se ees oa abet 26 a 347 See ee ree at ee eee. ee are 347 else eae ere. ee Pee ee eee ty 347 ELTA ee ee ee ee ee A 349 A IIGYAL Sb tae tail Nha Wi Solin eaten: tty es. le aah 349 "Er, CVD LE ashe laa Sal re Ree cea acs = Net ia ali 350 TS Se ee i oe Re SA Woe ee 302 AD IS CHISSIOM et: sean eR ee ne ee de BE Vase Mechamicsi sen Rete Bae The Habitus of Eocaecilia micropodia 355 Zygokrotaphy and Stegokrotaphy —__ 356 Eocaecilia as a Stage in Caecilian EL CICS i pee eee’ SMR Da eee 358 RCM OW le OPONETIS ew een Te 361 Appendix: A List of Eocaecilia micropodia DOGO ee ee 361 bares eic tar ey Gri ree OR de OL aie en ee 362 ABSTRACT. Eocaecilia micropodia, an Early Jurassic caecilian from the Kayenta Formation of northeast- ern Arizona, is structurally comparable to Recent gymnophionans in numerous aspects but also pos- sesses characters that are primitive or appear to be uniquely derived. The skull of Eocaecilia exhibits such distinctively caecilian features as (1) a sulcus along the orbital rim indicating the presence of a ten- taculum; (2) an os basale representing consolidation of the supraoccipital, exoccipital, basisphenoid, basi- occipital, pleurosphenoid, and parasphenoid ele- ments; (3) an internal naris posterior to the premax- illary—maxillary suture and medial to the tooth rows on the vomer and palatine; (4) enlarged nasal cap- sules; and (5) an olfactory eminence on the vomer. As in Recent caecilians, the lower jaw comprises a pseudodentary and pseudoangular that are joined along an elongate, oblique suture. The pseudoangular bears a robust internal process and an elongate re- troarticular process. The teeth are bicuspid and ped- icellate, but are minute in size and are more numer- ous than in most living caecilians. Several features of the skull and lower jaw of Eocaecilia are unexpectedly divergent from the pat- tern known in Recent gymnophionans. The apparent 285 286 fusion of the stapes and quadrate is unique. The obliquely oriented, more or less planar jaw joint would appear to provide little stability, and is thus structurally and functionally unlike that known in any other caecilian. The internal process of the lower jaw is very robust, and projects into the adductor cham- ber. Eocaecilia also presents primitive and/or transition- al features that might be expected in forms repre- senting an intermediate stage in the development of a specialized life style. The skull retains separate ju- gal, quadratojugal, postparietal, and Ptabular (or Psu- pratemporal) bones, elements that in living forms are either co-ossified with adjacent bones or lost. The limb girdles and limbs of Eocaecilia are a primitive retention, but the reduction in their relative size would appear to be transitional toward the limbless, gymnophionan condition. Elongation of the body in Eocaecilia is estimated to be comparable to that in primitive extant gymno- phionans, but a precise comparison cannot be made because of uncertainty over the number of vertebrae. In general, the postcranial axial skeleton is relatively primitive. Intercentra are present. The parapophyses are not protracted as elongate processes, as in living forms, nor is there a pronounced longitudinal keel on the ventral aspect of centra. In contrast to the atlases of Recent caecilians, an interglenoid tubercle is pre- sent. Processes projecting from the internal walls of the neural canal of the atlas and at least the next four postatlantal vertebrae represent attachment points for a suspensory ligament of the spinal cord. Such pro- cesses have not been previously reported in living caecilians but are now known to be present in rep- resentatives of various families (ichthyophiids, typhlo- nectids, and caeciliids). Neural spines are absent in the postatlantal and dorsal regions, as in living cae- cilians. Haemal arches are present in the tail, and distal caudal vertebrae bear posterodorsally recurved neural processes, as in rhinatrematids. Although our knowledge of caecilian evolution and diversity now extends into the Early Jurassic, Eocae- cilia micropodia does not provide sufficient evidence to securely recognize the origin of gymnophionans among known Paleozoic amphibians. The discovery of an operculum in Eocaecilia micropodia is novel confirmatory evidence that the opercular apparatus is a character shared with other lissamphibians. The ab- sence of a separate operculum in extant gymnophion- ans may be hypothesized to relate to the loss of the shoulder girdle and the muscular link between the girdle and operculum. Nonetheless, a substantial morphological and temporal gap still intervenes in the identification of caecilian origins. Lissamphibia still remains at best a crown group concept, without a securely rooted stem in the Paleozoic. INTRODUCTION Fossil caecilians are sparsely represent- ed in the geological record. Initially known Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 only from an isolated vertebra from the Paleocene of Brazil (Estes and Wake, 1972), subsequent finds—vertebrae from the Paleocene of Bolivia (Rage, 1986) and the Cretaceous of Sudan (Evans et al., 1996; Werner, 1994) and the Miocene of Columbia (Hecht and Laduke, 1997)—of- fered scant evidence with which to probe the evolutionary history of the group. More recently, Evans and Sigogneau-Rus- sell (2001) described the fragmentary re- mains of a primitive caecilian, Rubricacae- cilia monbaroni, from the Lower Creta- ceous of Morocco, adding further perspec- tive on the Mesozoic record of caecilians. With fossil caecilians so rare, the discovery of Eocaecilia micropodia from the Early Jurassic Kayenta Formation of northeast- ern Arizona (Jenkins and Walsh, 1993) was particularly significant. This earliest known caecilian, abundantly represented by nu- merous cranial and postcranial specimens, possesses a number of features, including intercentra and limbs, which are primitive for the order but unknown in living rep- resentatives. Our descriptive account doc- uments Early Mesozoic caecilian diversity in terms of characters that are clearly ple- siomorphic, as well as features that are au- tapomorphic for gymnophionans. In this study, we provide detailed doc- umentation of the skull and postcranial skeleton of Eocaecilia micropodia, making anatomical comparisons with major fea- tures of Recent caecilians as well as with those few fossil forms that are known. We explore several specializations of E. micro- podia in a functional context and finally consider the phylogenetic significance of this taxon in relation to current interpre- tations—and uncertainties—regarding the evolutionary history and relationships of caecilians. The taxonomic terms “Apoda” and “apodans,” used in reference to extant taxa as a group (cf. Trueb and Cloutier, 1991), is preoccupied (Dubois, 2004; Duellman and Trueb, 1986) and has been abandoned in this account. We follow the suggestion of Frost et al. (2006) that “Gymnophiona” ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 287 Figure 1. Reconstruction of the skull of Eocaecilia micropodia in (A) dorsal, (B) ventral, (C, D) lateral, and (E) occipital views. Major features, including size and general proportions, are based on the type (MNA V8066), but some details have been drawn from other specimens. (MCZ 9169, 9015; MNA V8059, 8062). Postmortem compaction of the skulls renders the lateral and occipital views less reliable than the dorsal and ventral reconstructions. In D, the posterior end of the jugal (hatched area) has been truncated to reveal the dorsal margin of the stapes-quadrate. 288 be restricted to extant, limbless caecilians but do not adopt their proposed systematic reordering in face of incomplete results and ongoing analyses referred to in their work. We employ “caecilian” as an appro- priate term for gymnophionans and their fossil relatives. MATERIALS AND METHODS Eocaecilia micropodia is represented by 40 specimens. Only two specimens are of more or less complete skulls with articu- lated lower jaws, but both have been sub- jected to postmortem compaction and fracturing. Nine specimens represent par- tial skulls (with or without lower jaws) in various stages of disarticulation, and six specimens are of isolated mandibles. Ten specimens are primarily postcranial: ver- tebrae, both in isolation and in short, ar- ticulated series; disarticulated appendicu- lar elements; or both. The remainder of the collection consists of variably complete associations of cranial and postcranial bones. All of the material derives from a single locality, a quarry at Gold Spring (35°45'35"N, 111°04'51"W), approximately 300 m WSW of Gold Spring, Adeii Eechii Cliffs, Coconino County, Arizona, USA, in the silty facies of the Lower Jurassic Kay- enta Formation. The Kayenta fauna, which also includes the anuran Prosalirus bitis, is probably at least Pleinsbachian in age if not slightly older (Shubin and Jenkins, 1995; Jenkins and Shubin, 1998). A com- plete list of the specimens is provided in the Appendix. The jaw musculature of Ichthyophis glutinosus was studied by R.L.C. by frontal and sagittal serial sections and through dis- sections of preserved specimens prepared by David Dilkes. Bemis et al. (1983) dis- cuss the diverse terminology of caecilian jaw muscles in the literature, reflecting perspectives that varied from morpholog- ical description to determination of ho- mology. The present account follows the nomenclature of Lakjer (1926; also em- ployed by Siive-Séderbergh, 1945), who based his identifications on the relation- Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 ships of muscles to the branches of cranial nerve V, which are readily established in all living amphibian groups. Other descrip- tions of caecilian jaw muscles have com- monly used the terminology of Edgeworth (1935), which differs significantly from that now broadly applied to all groups of terrestrial vertebrates, e.g., Carroll and Holmes (1980) for frogs and salamanders, Rieppel (1980) for squamates, and Bram- ble (1978) for mammals. Institutional Abbreviations FMNH_ Field Museum of Natural His- tory, Chicago, Illinois Louisiana State University Mu- seum of Zoology, Baton Rouge, Louisiana LSUMZ MCZ Museum of Comparative Zool- ogy, Harvard University, Cam- bridge, Massachusetts MNA Museum of Northern Arizona, Flagstaff, Arizona Abbreviations Used in Figures Id dorsal branch of the olfactory tract Iv ventral branch of the olfactory tract II ‘optic nerve V trigeminal nerve V, ophthalmic division of V Ve maxillary division of V V3 mandibular division of V VII facial nerve AME adductor mandibulae externus AMI adductor mandibulae internus AMI(pr) deep head of the adductor mandibulae internus (profun- dus) AMI(su) superficial head of the adduc- tor mandibulae internus (su- perficialis) AMP adductor mandibulae posterior AMP(lg) adductor mandibulae posterior longus art articular facet art pa articular facet for pseudoan- cular articular facet for pterygoid art pt pal art qu pt art St=q ANATOMY OF EOCAECILIA MICROPODIA *¢ Jenkins, Walsh, and Carroll articulation between pterygoid and palatine articulating surface for quad- rate ramus of pterygoid articular facet for stapes-quad- rate atlas basin on the dorsal surface of the cultriform process basipterygoid process foramen for the carotid artery coracoid foramen cultriform process depressor mandibulae ectopterygoid external naris frontal fibula femur intramandibular foramen fenestra ovalis groove for pterygoideus muscle glenoid groove for articulation with the sphenethmoid groove for pterygoideus muscle humerus haemal arch intercentrum interhyoideus interhyoideus posterior intermandibularis internal naris internal process jugal jugular foramen lateral (labial) tooth row levator quadrati maxilla medial (lingual) tooth row mandibular (adductor) fossa maxillopalatine metatarsal nasal nasolacrimal duct os basale otic capsule occipital condyle operculum OY ver cd ver d ver patl WE VS 289 otic ridge for contact with skull roof orbit parietal pseudoangular palatine processus conchoides pseudodentary possible pelvic element postfrontal phalanges internal process of pedicle pleurosphenoid premaxilla postparietal parietal ridge prefrontal pterygoid pterygoideus muscle quadrate quadratojugal quadrate ramus of the ptery- goid rib radius retroarticular process scapulocoracoid sphenethmoid septomaxilla possible septomaxilla squamosal stapes stapedial foramen stapes-quadrate Ptabular tibia tentacle tentacular opening tentacular sulcus ulna vomer vertebra caudal vertebra(e) doral vertebra(e) postatlantal vertebra vascular foramen vomerine sulcus 290 DESCRIPTION Skull The skull of Eocaecilia micropodia was recognized by Jenkins and Walsh (1993) as sharing a suite of features that are distinc- tively caecilian, including a well ossified, compact skull roof, a tentacular sulcus, and a large retroarticular process on the lower jaw. The skull appears to have been rela- tively low, as in most modern caecilians, but all specimens have been dorsoventrally crushed to a variable degree. In dorsal view (Fig. 1A), the most conspicuous fea- ture is the extension of the braincase be- hind the posterior margin of the skull roof; as a consequence, the jaw articulation is situated in a relatively anterior position (Fig. 1D). The orbits are large compared with those in most Recent caecilians but are small relative to those in Paleozoic tet- rapods of similar body size (e.g., gymnar- thrids, goniorhynchids, and amphibamids). The mouth, which Jenkins and Walsh (1993) reconstructed as only slightly sub- terminal, is here reconstructed with essen- tially no premaxillary overhang, in contrast to the subterminal condition typical of gymnophionans. The skull table and cheek are a continuous bony shield, as in some Recent caecilians. Evidence of variation in skull size is modest. The length and width of the type (MNA V8066, Figs. 2-4), which is the most complete and least distorted speci- men, are 12.7 and 8 mm, respectively. Length was determined in the midline from the tip of the rostrum to a line trans- versely tangential to the occipital condyles and is equivalent to Lessa and Wake’s (1992) measurement 5 in their morpho- metric study of Dermophis mexicanus. Width was determined from the most broadly separated points on the sides of the skull; this measurement approximates but is not exactly equivalent to “skull width at jaw articulation,” measurement 3 of Lessa and Wake (1992, appendix 1; incor- rectly labeled as “40” on fig. 1). Estimates of other skull lengths and widths are 13.5 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 and 8.2 mm (MCZ 9169, Fig. 5) and 9.8 and 7.2 mm (MCZ 9015), respectively. The skull of MNA V9346 (Fig. DS ! mm in length but does not permit a reli- able measure of width. MNA V8062 (Fig. 8) is too incomplete to provide a direct linear estimate of length, but on the basis of the dimensions of the frontal bone, the skull would appear to have been compa- rable in size to that of the holotype. No single specimen shows all elements, but nearly all of the dermal skull can be reconstructed by comparing and compiling structural data from the following speci- mens: the type specimen, MNA V8066 (Figs. 2-4), a nearly complete skull pre- pared in dorsal and ventral views; MCZ 9169 (Fig. 5), most of a skull exposed in dorsal view; MCZ 9015 (Fig. 13A, B), a nearly complete skull exposed in dorsal and ventral aspects; MNA V8059 (Fig. 7), exhibiting palate, jaws, both stapes-quad- rates, and the underside of the skull roof; MNA V8062 (Fig. 8), a right antorbital re- gion; and MNA V9346 (Fig. 11, a palate and lower jaws in ventral view. Further study of the material since Jen- kins and Walsh’s (1993) initial description has resulted in some modifications to the previously interpreted pattern of dermal bones. In some cases, a coating of Glyptal applied to the external surface of the skull roof illuminated narrow, pigmented bands on either side of barely visible sutures. The new findings are the presence of distinct postparietals and Ptabulars/Psupratempor- als; concrete evidence for the presence of both a postorbital and a postfrontal in the circumorbital series is lacking. Skull Roof. Almost all of the bones that ancestrally compose the amphibian skull roof are present. In contrast to all Recent caecilians, the skull in Eocaecilia micro- podia retains the postparietal, jugal, quad- ratojugal, and Ptabular/Psupratemporal; the last three are evident in the type. The sculpturing of the bones indicates that the dermis was firmly adherent to the skull roof, as in Recent taxa. Postparietals are not readily apparent in ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll the type specimen and were not recog- nized as distinct elements by Jenkins and Walsh (1993). However, separate postpar- ietals are evident in MCZ 9169 (Fig. 5), in which they are suturally delineated from surrounding bones. Once observed in this specimen, the presence of postparietals in the holotype was detected through an ap- plication of the Glyptal technique de- scribed above, which revealed the sutural lines separating parietals from postparie- tals. Similar bands became evident along both sides of the sutures separating pari- etals and frontals and at the anterior mar- gin of the frontal in the type, but not in other specimens, in which the bones are more coarsely sculptured. In most modern caecilians (e.g., in Ichthyophis glutinosus, Fig. 6A), the parietals are longer than the frontals, as would be expected if they in- corporated the area that was originally formed by the postparietals. The postpar- ietals exhibit a slightly recessed area of smooth bone surface posteriorly, which was presumably covered by an anterior ex- tension of the dorsal trunk musculature, as is the case in living caecilians. An irregular, inconsistently shaped bone lies in the position of a tabular or supra- temporal in Paleozoic tetrapods. This ele- ment, which we tentatively interpret as a Ptabular (Figs. 1, 2, 5, 7), appears to have been relatively loosely articulated with ad- jacent dermal bones, overlying ventral lap- pets of the surrounding bones without be- ing suturally interdigitated. In the type (Fig. 2), the right tabular is close to a nor- mal position, but the left has slipped over the left side of the os basale. The left tab- ular is also displaced posteriorly in MCZ 9169 (Fig. 5). The posterior margin of the tabular is not complete in any specimen, obviating the possibility of determining whether this bone reached the posterior margin of the skull, as is the case in most Paleozoic tetrapods. The large squamosal, best preserved in MNA V8059 (Fig. 7), forms much of the cheek region. The great width of the squa- mosal suggests that it probably reached 291 the postparietal, posterior to the tabular. The smooth, thickened medial margin would appear to have abutted the parietal and tabular by a squamous suture in which the adjacent bony margins are reciprocally beveled (rather than by serrate interdigi- tation). The articular relationship between the squamosal and dermal skull roof in Eocaecilia micropodia thus is comparable to that in many stegokrotaphic gymno- phionans in which the squamosal is at- tached to the skull roof by a sutural liga- ment; the ligament in some taxa is so broad that an apparent “gap” appears on dried skulls. In none of the specimens of E. micropodia is the posterolateral portion of the squamosal sufficiently well pre- served in dorsal view to determine the configuration of the occipital margin or show the area from which the depressor mandibulae originates in Recent caeci- lians. The anterior extremity of the squa- mosal narrows to an acute terminus be- tween the frontal and postfrontal. In Eocaecilia micropodia, the ventral part of the cheek is formed by a large ju- gal, which extends rostrad to form the pos- teroventral margin of the orbit, and a long, slender quadratojugal (Figs. 1C, D, 2). In Recent caecilians, a separate quadratojugal is not known to be retained in the adults of any species; thus, the squamosal extends to the inferior margin of the skull. Peter (1898) proposed that the quadratojugal of Ichthyophis glutinosus has been incorpo- rated into the quadrate as an anteriorly projecting, laminar process, an interpreta- tion that was adopted by de Villiers (1936) and Visser (1963). In a developmental study of two species of Epicrionops, Reiss (1996) identified the “quadratojugal pro- cess” as a feature appearing late in ontog- eny but presented no evidence for its der- ivation from a quadratojugal. Studies of Dermophis mexicanus by Wake and Han- ken (1982) and Gegeneophis ramaswamii by Miiller et al. (2005) concluded that the quadratojugal is absent. The interpretation of Marcus et al. (1935: 411) that a quad- ratojugal (“quadratomaxillare”) anlagen is 292 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 1mm Figure 2. Skull of the type specimen of Eocaecilia micropodia (MNA V8066) in dorsal view. present during development is rendered moot by the observations of Wake and Hanken (1982: 211), who noted several sources of possible error, in addition to the inclusion of different genera in a “single” growth series. Thus, at best, the evidence for the retention of a quadratojugal vestige in living caecilians is equivocal; yet, there is no doubt that a distinct quadratojugal persisted as the posterolateral marginal el- ement of the dermal skull roof until at least the Jurassic. The large, rectangular nasal, most clear- ly preserved on the left side of MCZ 9169 (Fig. 5), is incised rostrally at the premax- illary articulation. The nasals are missing ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 293 1mm jf Figure 3. Skull of the type specimen of Eocaecilia micropodia (MNA V8066) in ventral view. Both pterygoids (pt) have been broken transversely by postmortem crushing against the adjacent internal process (int p) of the pseudoangular; in life, each internal process projected into the adductor chamber above the level of the pterygoids. in the type (Fig. 2) and on the right side to the narial margin. A prefrontal is pre- of MCZ 9169 (Fig. 5), suggestive evidence that they might have been only loosely at- tached to the frontals. The long, rectangular prefrontal, best preserved in MNA V8062 (Fig. 8), extends from the midpoint of the dorsal orbital rim sent in the Ichthyophiidae, Uraeotyphli- dae, and Scolecomorphidae, but the bone has been lost or become incorporated into the maxillopalatine in other taxa (Nuss- baum and Wilkinson, 1989). The configuration of the posterior or- 294 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Figure 4. The skull of the type specimen of Eocaecilia micropodia (MNA V8066) in (A) dorsal and (B) ventral views (stereo- photographs). bital margin in Eocaecilia micropodia is somewhat equivocal because all specimens are to some degree damaged in this area. In most Paleozoic tetrapods two bones—a postfrontal and a postorbital—contribute to the posterior orbital margin. Jenkins and Walsh’s reconstruction (1993, fig. 2b) of E. micropodia depicted both bones, an interpretation based primarily on the right side of the holotype (Fig. 2) where there ANATOMY OF EOCAECILIA MICROPODIA ¢ Jenkins, Walsh, and Carroll 295 1mm Figure 5. Eocaecilia micropodia, dorsal view of the skull of MCZ 9169. Left stapes-quadrate has been removed (see Fig..20). On the right side, the stapes-quadrate is intact and nearly in its natural position, exposing much of the dorsal and lateral surface. The posterior portion of the bone was removed from the left side so that.it can be viewed in all aspects, but the anterior extremity was not complete. appears to be two elements. Both have been displaced ventrally by crushing, and the adjoining margins are sufficiently ob- scured that it is not possible to determine definitively whether there are two bones separated suturally or a single element fractured longitudinally. Several other skulls, however, provide evidence that fa- vors the latter interpretation. The left side of MCZ 9169 (Fig. 5) shows only a single bone composing the posterodorsal margin of the orbit. Similarly, but a single element forms the posterodorsal orbital margin in MNA V8059 (Fig. 7); on the left side are 296 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 1mm imm cf D mp ees Figure 6. Skull roof and palate of gymnophionans. (A) Skull roof of /chthyophis glutinosis. (B) Palate of Ichthyophis glutinosis. (C) Palate of Gymnopis multiplicata (FMNH 189131). (D) Palate of Epicrionops petersi. Plates A and B drawn are rendered from photographs in Taylor, 1969; plate D is reproduced from Nussbaum (1977, fig. 1). part of the jugal and a maxillary fragment, and on the right, a sutural contact with the prefrontal is preserved. Cognizant of the limited evidence and the lack of a com- plete orbital margin, we nonetheless favor the interpretation that E. micropodia pos- sessed only a single postfrontal/postorbital element. No living caecilian possesses two elements in this region. Representatives of only two families (Ichthyophiidae, Uraeo- ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 297 Figure 7. Eocaecilia micropodia, MNA V8059, ventral view of the skull roof and palate. typhlidae) possess a putative postfrontal, which could be distinct or partially or en- tirely fused to the maxillopalatine (Nuss- baum and Wilkinson, 1989). The “post- frontal” of living caecilians is thus a con- ventional designation for the single ossifi- cation that nearly encircles the eye (cf. Ichthyophis glutinosus, Fig. 6A; Uraeo- typhlus narayani, Nussbaum, 1979, fig. 1), occupying the circumorbital margin that in E. micropodia is formed by the prefrontal, postfrontal, and jugal bones. 298 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 1mm Figure 8. Eocaecilia micropodia, MNA V8062, dorsolateral view of a partial skull of which only the anterior part is preserved. The lacrimal bone presents the most equivocal case for interpretation of any of the skull roof elements. In the only spec- imen for which the lateral surface of the antorbital region is articulated (MNA V8062, Fig. 8), there appears to be a nar- row triangular space between the posterior portion of the prefrontal and maxilla that might have been occupied by a lacrimal. Although a lacrimal is not known in the adult of any Recent caecilian, a separate element in the nasal region of embryonic Ichthyophis glutinosus was identified by Peter (1898, fig. 20) as a turbinale (= lac- rimal). Similarly, Marcus et al. (1935, fig. 5) identified a lacrimal in their reconstruc- tion of a 68-mm embryo of Grandisonia alterans (cf. Wake and Hanken, 1982, ta- ble 1). Wake and Hanken (1982), however, found no evidence of a separate lacrimal ossification center in their study of Der- mophis mexicanus; in view of their cau- tionary assessment of the evidence pre- sented by Marcus et al. (1935) for multiple ossification centers (including that for the quadratojugal, postparietal, periorbital, ec- topterygoid, and interparietal), the fate of the lacrimal in caecilian history must be regarded as unresolved. The premaxilla, which does not appear to overhang the lower jaw as it does in many Recent caecilians, has a long poster- odorsal process that fits into a slot in the nasal and a very narrow extension that oc- cupies a groove along the anterodorsal margin of the maxilla. As is the case for almost all specimens of upper and lower jaws, the crowns of the teeth have disar- ticulated postmortem from the pedicels; only in the premaxilla of MNA V9346 do a few crowns appear to be preserved in situ. The premaxilla in MNA V8059 (Fig. 7) bears approximately 10—12 tooth posi- tions (10 as actual pedicels and 2 as spaces wide enough to accommodate a pedicel). The pedicels are formed as elongate, nar- row tubes that are fused to the inside of the jaw margin. The diameter of premax- illary pedicels was measured with an op- tical micrometer on specimens in which individual pedicels are well preserved. Four pedicels on MNA V8059 have di- ameters of 0.10, 0.12, 0.13, and 0.13 mm. One premaxillary pedicel on MNA V9346 ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll has a diameter of 0.14 mm, and an adja- cent (disarticulated) tooth crown measures 0.12 mm in basal diameter and 0.22 mm in apicobasal height. Our present interpretation of the sep- tomaxilla in Eocaecilia micropodia differs from that originally proposed (Jenkins and Walsh, 1993, fig. 2b) and is based on a dis- placed bone in a single specimen (Fig. 8). The bone might have contributed to the posterior margin of the narial opening, as in the case in some extant caecilians. A septomaxilla is present in ichthyophiids, rhinatrematids, uraeotyphlids, and scole- comorphids but is absent as a separate el- ement in typhlonectids and caeciliids (Nussbaum, 1977, 1979; Taylor, 1969; Wake, 2003). The maxilla is a long, narrow bone that contributes to the anteroventral margin of the orbit for a short distance. The most completely preserved maxilla is on the right side of the type (Fig. 2), but the in- ferior margin and maxillary teeth are ob- scured by the overlying, displaced lower jaw. No specimen shows the entire maxil- lary tooth row. The number of maxillary teeth is probably in the range of 32-39, an estimate based on the length of the maxilla in the type (4.9 mm) and the number of pedicels per millimeter on maxillary frag- ments of other specimens. Pedicels per millimeter varied from 6.5 on a left ante- rior maxillary fragment of MNA V9346 to 8 in a right anterior maxillary fragment of MNA V8059 (seven well-preserved maxil- lary pedicels on the latter specimen range from 0.1 to 0.13 mm in diameter, with a mean of 0.11 mm). Intermediate values of 7.9 were measured from other maxilla fragments (MCZ 1956, MNA V8059, and MNA V8062). In contrast to Recent caecilians, the maxilla is not fused to the palatine; a su- ture delineating the adjacent borders of these bones is clearly preserved in MNA V8062 (Fig. 8). The margin of the maxilla that borders the anteroventral rim of the orbit is incised by a shallow, broad sulcus in both the type of Eocaecilia micropodia 299 (Fig. 2) and in MCZ 9156 (Figs. 9, 10C). The sulcus is confluent with a comparably smooth depression in the adjacent pala- tine. We interpret this feature as marking the course of the tentacle as originally pro- posed by Jenkins and Walsh (1993). Early ontogenetic development of the tentacle in Dermophis mexicanus is in close associa- tion with the eye, from which various com- ponents of the tentacular apparatus are co- opted (e.g., the Harderian gland, extraocu- lar muscles and nerves, and lacrimal ducts, among others; Billo and Wake, 1987; Wake, 1992). Although the tentacle in most living caecilians passes through an aperture that is separate from the orbit (Fig. 10A), in rhinatrematids the tentacu- lar opening is not separate (Fig. 10B) but is merely “a small longitudinal slit” along the anterior margin of the orbit (Nuss- baum, 1977: 7). In fossil caecilian material, the presence of a tentacular apparatus could only be inferred with certainty if a separate tentacular foramen were present, but the slight depression of the orbital margin in E. micropodia (Fig. 10C) is at least consistent with the expected primi- tive position of this organ. Sculpturing of the Skull Roof. Most cra- nial specimens of Eocaecilia micropodia possess a shallow rugosity on the superfi- cial surface of the dermal bones compa- rable to that developed in some larger, ex- tant caecilians (e.g., Gymnopis spp.). The skull of the type specimen also exhibits small, scattered foramina (Fig. 2). These foramina, which distribute nerves and nu- trient vessels, are most numerous on the bones surrounding the orbit and adjacent to the jaw margins, as in living species (Taylor, 1969), but also occur sporadically on the bones that border the midline of the skull roof. Palate. The palate of Eocaecilia micro- podia demonstrates that basic features re- tained among extant caecilians were ac- quired by the Jurassic, in contrast to the pattern of dermal bones of the skull, which is only modestly altered from that common among Paleozoic amphibians. A principal 300 feature is a continuous row of teeth that spans each vomer and palatine, paralleling the row on the premaxilla and maxilla (Figs. 1B, 3, 7). The most complete series of pedicels is preserved on the right side of the type (MNA V8066, Fig. 3) where the vomer and palatine each bear approx- imately 17 teeth. In this series, nine well- preserved palatal pedicels selected for measurement range in diameter from 0.1 to 0.14 mm (mean 0.12 mm) and are com- parable in size to those on the maxilla and premaxilla measured from other speci- mens. On MNA V8059, five pedicels that could be measured range from 0.11 to 0.15 mm in diameter (mean 0.13 mm). A series of 13 pedicels on the vomer of MCZ 9242 (Fig. 12) is the most perfectly pre- served palatal row available with which to assess detailed geometry. All the pedicels are slightly oval in cross section, with the labiolingual axis slightly longer than the mesiodistal axis. Labiolingual axes range from 0.14 to 0.18 mm (mean 0.17 mm), whereas mesiodistal axes range from 0.12 to 0.14 mm (mean 0.135 mm). The height of the pedicels ranges from 0.35 to 0.37 mm. Lingual to and between the 11th and 12th pedicel in this series is an inverted tooth crown (circular in cross section, with a diameter of 0.1 mm). The distance between the premaxillary— maxillary and vomerine—palatine tooth rows varies among extant caecilians, being relatively narrowly separated in species of Ichthyophis and well separated in species of Caecilia (Taylor, 1969). Direct assess- ment of this feature in E. micropodia is precluded by the fact that, in all specimens with an intact palate, the lower jaws re- main in position, concealing the most lat- eral part of the palate and the premaxil- lary—maxillary tooth row. However, in view of the close juxtaposition of the tooth rows of the lower jaw, the separation of the pal- atal and marginal rows was probably only slightly greater (for further analysis, see Lower jaw). The vomers meet anteriorly along a me- dian suture for about half their length. Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Posteriorly, the medial margins of the vo- mers are separated by the relatively broad rostral end of the cultriform process of the parasphenoid component of the os basale, which intervenes to complete the bony palate in this area (Fig. 1B). The degree to which the cultriform process is exposed between the recessed medial margins of the vomers in Eocaecilia micropodia would appear to be intermediate between the extremes observed in extant caecilians (cf. Trueb, 1993: 275, table 1; Wake, 2003). In the rhinatrematid Epicrionops petersi (Fig. 6D), the vomers meet ros- trally only along a very short suture; the cultriform process is extensively exposed between them. In ichthyophiids (Fig. 6B) and many other caecilians, by contrast, the vomers are united for nearly their entire length along the median suture. Intraspe- cific variations can also be substantial, as in Gymnopis; in G. multiplicata (Taylor, 1969, fig. 34), the vomers are separated for almost their entire length, whereas in G. multiplicata proxima (Fig. 6C; see also Taylor, 1969, fig. 35), the vomers are unit- ed for more than half their length. In most Recent caecilians, the vomer is pierced by a large foramen (in some cases two) anteromedial to the internal naris (Taylor, 1969). In Ichthyophis glutinosus (Fig. 6B), this opening provides passage for the palatine branch of the facial nerve (VIL) and a vein from the snout and dorsal surface of the palate; a rostrocaudally aligned neurovascular sulcus leads to the foramen’s palatal opening. In I. glutinosus, the vomerine foramen and sulcus on the palatal surface correspond in position to a ridge on the dorsal surface of the vomer; this ridge, the olfactory eminence (Fig. 14A), divides the large nasal sacs into me- dial and lateral cavities (Badenhorst, 1978, fig. 9; Schmidt and Wake, 1990). Although there are no large vomerine foramina in Eocaecilia micropodia, numerous small fo- ramina are variably developed on the bone’s palatal surface. A consistent feature is a sulcus on the palatal surface of the vomer, anteromedial to the internal naris. ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 301 Figure 9. Eocaecilia micropodia, MCZ 9156, the disarticulated elements of a skull. For details of the maxilla (m) and tentacular sulcus (ten s), see Figure 10C. The depression is evident on both vomers of MNA V9346 (v s, Fig. 11) and on the single vomer present in MCZ 9242 (Fig. 12). The corresponding elevation of the dorsal surface of the vomer might possibly represent an olfactory eminence (cf. Fig. 14A). The internal nares of Eocaecilia micro- podia open medial to the internal row of teeth, rather than adjacent to the suture between the maxilla and premaxilla as in most Paleozoic tetrapods. Both the vomer and palatine contribute to the margin of the internal naris, a condition that is com- 302 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Figure 10. tens (A) A skull and lower jaw of /chthyophis glutinosus (skull drawn from Taylor, 1969, fig. 2; lower jaw drawn from Sarasin and Sarasin, 1887-1890, pl. 15, fig. 3). (B) Lateral view of the skull of Epicrionops petersi (reproduced from Nussbaum, 1977, fig. 1). (C) A right maxilla of Eocaecilia micropodia (MCZ 9156) exhibiting a tentacular sulcus (ten s). mon to representatives of all extant fami- lies: rhinatrematids (Epicrionops, Nuss- baum, 1977, fig. 1), ichthyophiids (Ichth- yophis, Taylor, 1969, figs. 2-7; Wake, 2003), uraeotyphlids (Uraeotyphlus, Nuss- baum, 1979, fig. 1; Uraeotyphlus narayani, Wake, 2003, fig. 6B), scolecomorphids (Crotaphatrema, Scolecomorphus, Nuss- baum, 1985, figs. 1, 4; Scolecomorphus uluguruensis, Wake, 2003, fig. 6E), caeci- liids (Caecilia, Oscaecila, Herpele, Taylor, 1969, figs. 17-26, 42; Oscaecilia ochroce- phala, Wake, 2003, fig. 9H; Herpele, Wake, 2003), and typhlonectids (Typhlo- nectes, Potamotyphlus, Taylor, 1969, figs. 12-14; Typhlonectes natans, Wilkinson and Nussbaum, 1997, fig. 9C). A few cae- ciliids exhibit an apparently derived con- dition in which the internal narial opening is completely surrounded by the palatine (maxillopalatine) (e.g., Dermophis parvi- ceps, Geotrypetes seraphini, Taylor, 1969, ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 303 Figure 11. figs. 30, 40, 41; Dermophis mexicanus, Wake and Hanken, 1982, fig. 2b; Geotry- petes seraphini, Wake, 2003, fig. 8E). In other species of Dermophis, however, both the vomer and palatine contribute to the narial margin (D. glandulosus, D. occiden- talis, Taylor, 1969, figs. 31, 32). In various species of Siphonops, the internal naris is either variably bordered by the vomer and palatine or completely enclosed by the pal- atine (Taylor, 1969, figs. 36-39; Wake, 2003). Eocaecilia micropodia, MNA V9346, ventral view of the palate, both stapes-quadrates and lower jaws. The pterygoid of Eocaecilia micropodia (Figs. 1, 3), together with the palatine and vomer, contributes to the relatively primi- tive configuration of the palate as a broad plate that laterally borders large interpter- ygoid vacuities. The articulation between the pterygoid and palatine, which is evi- dent in the holotype where the two bones have partially separated from one another (Fig. 3), appears to have been composed of simple, overlapping surfaces. Both the quadrate ramus and the ascending process 304 of the pterygoid are preserved in MNA V9346 (Fig. 11). The elongate quadrate ra- mus has an extensive articulation with a sulcus on the inferior surface of the stapes- quadrate. In contrast to the condition in Recent caecilians, the quadrate ramus ap- pears to have extended well posterior to the jaw articulation (cf. Figs. 1B, 3). Both the palatal ramus of the pterygoid and the parasphenoid component of the os basale are covered with pedicellate denticles. As reconstructed, the rostrocaudal length of the pterygoid in Eocaecilia micropodia is approximately half of the length of the skull, which is longer than in most (if not all) extant caecilians (see Taylor, 1969). Al- though a separate pterygoid persists in rhinatrematids and ichthyophiids, in ty- phlonectids and caeciliids the bone is or- dinarily fused either to the quadrate or maxillopalatine (Nussbaum, 1977) or, in the case of scolecomorphids, possibly pre- sent only as a vestige (Scolecomorphus vit- tatus, Nussbaum, 1985, fig. 4) or lost al- together (Crotaphatrema bornmuelleri, Nussbaum, 1985, fig. 1; cf. Brand, 1956, for Scolecomorphus uluguruensis). An ectopterygoid could not be identified in Eocaecilia. Nussbaum and Wilkinson (1989: 32) noted that a small “ectoptery- goid” [sic] is present in some extant cae- ciliids. In Gymnopis multiplicata (Fig. 6C) and Grandisonia alternans (Taylor, 1969, figs. 34, 47), for example, this small bone forms the anteromedial palatal margin of the adductor chamber. Slight disarticula- tion of most skulls of E. micropodia has disrupted the palatal margins of the ad- ductor chamber; in a few more intact spec- imens, the associated lower jaw is com- pacted against the region in which an ec- topterygoid might be expected. The coro- noid eminence, although low, extends anteriorly to approximately the midpoint of the jaw’ length; if an ectopterygoid were present, the bone would have prob- ably been much reduced in size so as not to impinge upon the coronoid eminence during jaw closure. Os Basale. The braincase of Eocaecilia, Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 as in all Recent caecilians, is constructed from two components: the os basale pos- teriorly and the sphenethmoid anteriorly. As in other caecilians, the os basale of Eocaecilia is a single bone that has been interpreted to represent a consolidation of ossification across a large area primitively occupied by separate bones (supraoccipi- tal, exoccipitals, basioccipital, basisphe- noid, pleurosphenoid, and parasphenoid). This massive element, which incorporates the otic capsules (presumably formed from the fusion of the prootic and opisthotic) as well as posterior elements of the endo- chondral braincase, is represented by nu- merous specimens (Figs. 2-5, 7, 11, 12, 15). In the holotype, the maximum width of the os basale, measured across the otic capsules, is estimated to be 3.5 mm; the estimated maximum length of the bone, which includes the cultriform process, is 10.4 mm. Dorsally, the os basale has a sutural junction with the postparietals and forms a roof over the most posterior portion of the braincase and otic capsules; as in Re- cent caecilians, a median suture divides the two halves of the braincase above the foramen magnum (cf. Figs. 1, 2; Taylor, 1969). The dorsal surface of each half of the os basale in this region is slightly con- cave and bears fine, longitudinal striae, features that represent the attachment of epaxial muscles. A low ridge delineates the boundary between the skull’s dorsal roof and occipital surface. A complete occiput is represented only in the holotype, but certain details (such as the size and shape of the foramen mag- num and the relative position of the occip- ital condyles) are distorted or obscured by postmortem compaction. Protuberant oc- cipital condyles are preserved in two spec- imens (Figs. 13A, B, 15A, B, 16). The fac- ets for the atlas are oval, slightly convex, face medially and slightly ventrally, and ap- pear to have been situated ventral to the foramen magnum. A jugular foramen is present at the base of the occipital condyle (Fig. 3), closely approximating the ar- ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll st-q Cul pr 305 Figure 12. Eocaecilia micropodia, MCZ 9242, disarticulated elements of skull. rangement in most modern forms. Slightly anteroventral to the jugular foramen is a small carotid foramen. The ventral portion of the os basale in- corporates the parasphenoid and is config- ured accordingly: posteriorly broad in the region of the otic capsules and anteriorly narrow in the region representing the cul- triform process (Figs. 1B, 3). At the an- terior border of the otic capsule, where the os basale narrows abruptly, one might ex- pect to find a basicranial articulation with the pterygoid (as is common in Recent caecilians, Fig. 18E); no such articulation is discernible in Eocaecilia (Figs. 15A, 16, 17). The dorsal surface of the area repre- senting the cultriform process bears a groove adjacent to the lateral margins for the articulation of the sphenethmoid (gr art sph, Fig. 12). The dorsal surface of the os basale is also marked by three depres- sions. The largest of these extends from the posterior margin of the braincase ros- trally to the posterior terminus of the grooves for the sphenethmoid. At the ros- tral end of the depression is an elevated, hemicircular margin set transversely across the cultriform process that defines a very shallow basin (b cul pr, Fig. 12). Anterior to this depression are two additional shal- low fossae, one on each side of the mid- line. 306 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 ANATOMY OF EOCAECILIA MICROPODIA ¢ Jenkins, Walsh, and Carroll 307 A B | pm external naris nin olfactory Bs skin pm eminence area af nasal capsule tae \- Nasal sac Figure 14. olfactory eminence en sphenethmoid naso-lacrimal duct _7naso-lacrimal duct 2 int na cartilaginous connection between sphenethmoid ~eye and cheek ON %, 6, jaw mm. cartilage (A) The head of Ichthyophis glutinosis (FMNH 121528) in dorsal view, with skin and bones removed to reveal the extent of the nasal apparatus. The openly stippled area on the right side indicates the extent of the entire nasal sac. The finely stippled area indicates the forebrain and dorsal olfactory tract. On the left side, the nasal sac has been removed to expose the dorsal surface of bones underlying the nasal apparatus. (B) The head of Epicrionops petersi (LSUMZ 27324) in horizontal section (right side) to show the elaboration of nasal sac contents. The cartilages surrounding the area adjacent to the external nares are comparable in position and relative size to the entire nasal capsule in frogs, salamanders and Paleozoic amphibians. The lateral aspect of the os basale, and specifically the otic capsule, is appressed to the massive stapes-quadrate; therefore, the structure of the capsule is best re- vealed by specimens in which the stapes- quadrate is missing or was removed (Figs. 15A-—C, 16). An anterolaterally descending ridge marks the line of sutural contact with the skull roof (o r, Fig. 17A). The otic cap- sule of MNA V8063 is estimated to be ap- proximately 2.0 mm in length and 1.6 mm in height. The lateral face of the otic cap- sule is dominated by a large fenestra ovalis < which, in MNA V8063, is 1.4 mm in length and 0.6 mm in height. In MCZ 9169, the fenestra is irregularly oval in shape (length 1.7 mm; height 0.8 mm). The inferior mar- gin is a gently convex, slightly thickened flange demarcating a narrow boundary be- tween the external and internal surfaces of the capsule. The superior and posterior margins of the fenestra are broadly round- ed, and the transition from the external to internal surface is more gradual. The os basale bears a number of prom- inent foramina, the largest of which is lo- Figure 13. The skull of Eocaecilia micropodia (MCZ 9015) in (A) dorsal and (B) ventral views (stereophotographs). The jaw articulation between the stapes-quadrate and pseudoangular (arrows) is shown in the dorsal view. The ventral view reveals the extensive field of pedicels on the cultriform process. (C) A partial skull of Eocaecilia micropodia (MCZ 9237) in ventral view (stereophotographs). 308 cated at the anterior margin of the otic capsule, where the otic and pleurosphen- oid regions join. On the basis of the com- parable arrangement in modern caecilians (in particular, Dermophis mexicanus), this foramen is presumed to convey cranial nerve V (Fig. 15B). A second foramen lies at the anterolateral margin of the otic cap- sule, slightly ventral and lateral to that for V; elevated on a slight eminence, this fo- ramen is interpreted as the exit of cranial nerve VII (Fig. 15B, C). A third, small fo- ramen opens onto the posterolateral Mar- gin of the parasphenoid portion of the os basale and is confluent with a long sulcus on the lateral surface of the parasphenoid; both features are interpreted as represent- ing the passage of the carotid artery (Fig. 15B). Although the overall configuration of the os basale of Eocaecilia micropodia is comparable to that in living caecilians, dif- ferences are apparent in a number of de- tails. First, the shape of the cultriform pro- cess is distinctive (cf. Figs. 1, 3, 6). Illus- trations of the palates of modern caecilians by Taylor (1969) and Wake (2003) reveal a diversity of cultriform process outlines, none of which is especially similar to that of E. micropodia, which is elongate (rela- tive to skull length) and narrow. The con- stricted posterior end of the cultriform process is distinctly demarcated from the broad capsular part of the os basale. Fi- nally, the lateral margins of the process, rather than converging at an acute angle as in many living taxa, remain subparallel for much of their length and converge dis- tinctly only near the apex. Second, the palatal surface of the cul- triform process bears an irregular array of pedicellate teeth (although no specimen preserves a tooth crown in place). The di- ameters of the pedicel bases on the cultri- form process are more variable and, on av- erage, smaller than those found on the jaws. On the type specimen (MNA V8066, Fig. 3), a sample of 14 well-preserved ped- icels range from 0.07 to 0.13 mm in di- ameter; the mean of 0.09 mm is less than Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 the means of 0.11—0.12 mm observed from tooth rows of the upper and lower jaws. On MNA V8071, a sample of eight pedi- cels on the cultriform process has a mean diameter of 0.1 mm, whereas the estimat- ed diameters of several poorly preserved pedicels on the associated mandible is 0.13 mm. Third, the condyles and the facets in Eocaecilia micropodia are relatively small- er than those of gymnophionans. An index of the relative size of the atlanto-occipital articulation is the bilateral breadth of the condyles (the distance between the lateral margins of the two condylar facets) ex- pressed as a percentage of skull width. The specimen of E. micropodia in which the condyles appear undistorted (MCZ 9015, Fig. 13B) reveals their bilateral breadth to be 25% of the skull width. Indices taken from MCZ gymnophionan specimens or the literature reveal a range of values be- tween 33 and 45%: the rhinatrematid Ep- icrionops petersi, 39% (Nussbaum, 1977, fig. 1); the uraeotyplid Uraeotyphlus na- rayani, 41% (Nussbaum, 1979, fig. 1); the typhlonectid Typhlonectes compressicau- da, 33% (MCZ 24524); the ichthyophiid Ichthyophis glandulosus, 34% (MCZ 14003); the scolecomorphid Scolecomor- phus uluguruensis, 39% (MCZ 12284); and various caeciliids: Afrocaecilia taitana, 45% (MCZ 20021); Dermophis mexicanus, 38% (MCZ 126357); Gegeneophis ramas- wamii, 38% (MCZ 29456); Geotrypetes seraphini, 42% (MCZ 3424); Gymnopis multiplicata, 36% (MCZ 29265); Oscaeci- lia ochrocephala, 40% (MCZ 9591); Schis- tometopum gregorii, 39% (MCZ 20070); and Siphonops annulatus, 36% (MCZ 19402). The articular facets of E. micropodia are relatively small as well. In MCZ 9015, the greatest width of a facet (measured along the long axis of the oval) is 0.7 mm, or 10% of the estimated skull width of 7.2 mm compared with values of 15% in Siphon- ops annulatus (MCZ 19405), and 17% in Gegeneophis ramaswamii (MCZ 29452) ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 309 o con 1mm ver patl Figure 15. The os basale of Eocaecilia micropodia (MNA V8063) in (A) dorsal and (B) ventral views. For stereophotographs of this specimen in ventral view, see Figure 16. (C) An otic capsule (MCZ 9169) in lateral view exhibiting the fenestra ovalis and associated operculum. (D) A partial braincase and the otic capsules in ventral view (MCZ 9167). 310 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Figure 16. The braincase of Eocaecilia micropodia (MNA V8063) in ventral view (stereophotographs). For labeled drawings of this specimen, see Figure 15A, B. and Typhlonectes compressicauda (MCZ QA524). A fourth difference is that the ventral aspect of the os basale of Eocaecilia mi- cropodia lacks any indication on its ventral surface of the attachment of subvertebral cranial depressor musculature, specifically the longus capitis. In many Recent caeci- lians, a pair of facetlike depressions ante- rior to the foramen magnum, separated in the median plane by a low ridge, mark this attachment site (cf. Figs. 3, 6B—D). The muscle flexes the skull ventrally, a move- ment that Bemis et al. (1983) suggested would be employed in both feeding and burrowing. Pleurosphenoid. Anteromedial to the otic capsule a large pleurosphenoid forms the lateral wall of the braincase, extending anteriorly along the lateral margin of the parasphenoid portion of the os basale to- ward the sphenethmoid (Fig. 16), as in modern caecilians. The pleurosphenoid does appear to have been fully fused to the parasphenoid (Figs. 15, 16). In MCZ 9167, the pleurosphenoid is approximately 3.4 mm long and 1.9 mm high. The anterior margin of the pleurosphenoid is recessed, and a small gap, estimated to be 1.4 mm in width, separates the anterior margin of this bone from the posterior margin of the sphenethmoid (Fig. 17A). This gap is likely to have accommodated the egress of cra- nial nerves II, III, and IV. Sphenethmoid. The sphenethmoid of Eocaecilia micropodia, which is not well preserved or visible in its entirety in any one specimen, is reconstructed (Fig. 17) on the basis of the holotype, MNA V8066, and MNA V8059 (Fig. 7). The spheneth- moid appears to conform generally to the pattern in modern caecilians (Fig. 18). In the holotype, the sphenethmoid is esti- mated to be 3.8 mm in length, or approx- imately half of the length of the para- sphenoid portion of the os basale. This well-ossified, complex element represents the rostral end of the braincase. The pos- terior half of the sphenethmoid forms the thin lateral walls of the anterior part of the braincase (the walls of the posterior part of the braincase are contributed by the pleurosphenoids, Fig. 17A). The anterior end of the sphenethmoid is a robust, ex- panded structure that forms the transverse posterior wall of the nasal chambers. As in ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll dll Figure 17. (A) Reconstruction of the braincase in Eocaecilia micropodia in \ateral view. The reconstruction is based on the holotype (MNA V8066; Figs. 2, 3) and MNA V8059 (Fig. 7). The sphenethmoid (sph) component of the braincase in (B) dorsal and (C) ventral views. modern taxa, the bone serves a supporting role in the anterior portion of the skull by extending dorsoventrally from the cultri- form process of the os basale to the ventral surface of the skull roof. The spheneth- moid is not exposed dorsally as a compo- nent of the skull roof, nor is it in most Recent caecilians: rhinatrematids (e.g., Epicrionops, Nussbaum, 1977, fig. 1; Wake, 2003, fig. 5A), ichthyophiids (Ichth- yophis sp., Wake, 2003, fig. 5G), uraeo- typhlids (Uraeotyphlus narayani, Wake, 2003, fig. 6A), scolecomorphids (Scoleco- morphus uluguruensis, Wake, 2003, fig. 6D), and typhlonectids (Typhlonectes com- pressicauda and T. natans, Wake, 2003, fig. 6G, J; Potomotyphlus kaupii, Wilkinson and Nussbaum, 1997, fig. 8B; see also Tay- lor, 1969; Wiedersheim, 1879). Dorsal ex- posure of the sphenethmoid does occur in various extant caeciliids in which the bone appears as a median element between the frontals (Taylor, 1969; Wake, 2003). The anterior portion of the sphenethmoid of E. micropodia also extends toward the lateral margin of the skull. In Epicrionops, the sphenethmoid continues as a cartilaginous posterior wall of the narial passage (Fig. 14B), possibly adding an additional but- tress against the compressive forces of burrowing. The same process occurs in Ichthyophis (the lamina orbitonasalis of Visser, 1963). The sphenethmoid in Eocaecilia micro- podia is pierced by several foramina. One, located on the posterior face of the lateral 312 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Ts NMA HEL, ATs ie Figure 18. Braincase of Dermophis mexicanus based on a specimen described by Wake and Hanken (1982) in the collection of Marvalee H. Wake, Department of Integrative Biology, University of California, Berkeley. Sohenethmoid in (A) left lateral, (B) dorsal, (C) anterior, and (D) posterior views. (E) Os basale in left lateral view. wing (Figs. 1B, 3; V,, Fig. 17A, C) is com- parable in position to a foramen in Recent taxa (e.g., Ichthyophis) that conveys the deep ophthalmic branch of cranial nerve V. Another foramen, visible in both lateral and ventral aspects (v f, Fig. 17A, C), en- ters the sphenethmoid at a level slightly above the opening for deep ophthalmic branch and is interpreted (by comparison with Ichthyophis) to represent a vascular foramen. Stapes-quadrate. Although the skull and lower jaw of Eocaecilia micropodia resem- ble those of Recent caecilians in many as- pects, the structure of the stapes and jaw suspension is unlike that in any other known gymnophionan and, furthermore, appears to have no parallel among other terrestrial vertebrates. There is a single el- ement situated in the stapedial and quad- rate region that has characteristics of both bones. As would be expected of a quad- ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll rate, the bone is firmly attached to the skull in three areas: posteroventrally to the quadrate ramus of the pterygoid (Fig. 11), dorsally to the back of the skull table (Figs. 1A, C, 2), and anteriorly to the underside of the squamosal (Figs. 5, 19B). Further- more, the bone bears a distinct facet for articulation with the lower jaw (Figs. 13A, 19B, 20D, 21C). As would be expected of a stapes, the bone is closely associated with the otic capsule and is traversed by a rel- atively large foramen; the size of the fo- ramen is consistent with a vascular passage (i.e., a stapedial artery, Figs. 2, 11, 19-21) rather than a narrow conduit expected for a nerve (such as the chorda tympani, which traverses the quadrate). The nature of the attachment to the otic capsule, how- ever, is less well defined than in Recent caecilians, in which the thickened edges of the footplate (e.g., in Dermophis mexican- us, Fig. 22) articulate with the margins of the fenestra ovalis. This large and com- plexly shaped single element can be termed a stapes-quadrate, although there is no evidence of sutural fusion in any specimen. Nearly all aspects of the stapes- quadrate can be seen in one or more spec- imens, yet in no single specimen is the en- tire bone both complete and fully exposed; integrating a reconstruction (Fig. 21) proved difficult and required certain sim- plifications to be consistent in all views. The stapes-quadrate is a large bone (ap- proximately one-third of the skull length) that extends, as a posterior process, behind the dermal skull roof and between the re- troarticular process of the lower jaw and the otic region of the os basale (Fig. 2). Although the posterior process of the sta- pes-quadrate resembles the stem or shaft of a stapes (by analogy with those in Pa- leozoic tetrapods; cf. Lombard and Bolt, 1988, figs. 3-9), its medial surface lies lat- eral to the fenestra ovalis and thus occu- pies a position that would be expected of a footplate. Rather than fitting into the fe- nestra ovalis, as in modern caecilians (Fig. 10B), the medial surface is large enough to have completely overlapped the margins 313 of the fenestra. However, in the type, which is the least distorted specimen, nei- ther stapes-quadrate lies in intimate con- tact with the fenestra ovalis (Fig. 3). The presence of an ossicle in this region (see below, Operculum) is additional evidence favoring the interpretation that the stapes actually had withdrawn from contact with the otic region (and assumed principally a jaw suspensory function and only second- arily, by bone conduction, an auditory function). The caudal end of the posterior process of the bone is free (i.e., lacks any bony contact). The lateral surface of the distal end of the process is rugose; the ru- gosity is slightly raised along the dorsolat- eral margin of the process and could rep- resent evidence of tendinous attachment. The adjacent medial surface of the lower jaws retroarticular process is also rugose. A deep, triangular fossa on the anterior surface of the stapes-quadrate (Figs. 20C, 21E), medial to the area of articulation with the lower jaw and dorsolateral to the articulation with the quadrate ramus of the pterygoid, represents an extension of the adductor chamber. The anterodorsal por- tion of the stapes-quadrate extends rostrad as a narrow lamina (Fig. 21A—D) that con- tributes to the lateral margin of the ad- ductor chamber. We interpret the rugosity on the dorsal surface of the lamina as an area of attachment to the undersurface of the lower margin of the cheek. A large stapedial foramen traverses the bone obliquely from dorsolateral to medial surfaces in a ventromedial and somewhat posterior direction (Figs. 2, 11, 19-21). Laterally, the foramen is distinctly oval (the long and short diameters of which measure 0.4 and 0.2 mm in MNA V8066; in MCZ 9169, the diameters are 0.5 and 0.25 mm, respectively); medially, the fo- ramen is nearly circular. The foramen opens onto the concave medial surface of the posterior process in a position slightly rostral to the fenestra ovalis. Rostral to the foramen’s dorsolateral opening, an anter- odorsal extension of the stapes-quadrate apparently attaches to the underside of the 314 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Figure 19. Stapes-quadrates of Eocaecilia micropodia. (A) Left side of the type specimen (MNA V8066) in oblique posterolateral view, showing articulation with pseudoangular. (B) Lateral view of right stapes-quadrate of MCZ 9169. skull roof, near or at the occipital margin (a relationship preserved only in the ho- lotype, Fig. 2). An approximately oval articular facet is situated at the middle of the lateral surface of the stapes-quadrate (Figs. 19B, 21C). In MCZ 9235, the dimensions of the facet in the anteroposterior and dorsoventral planes are, respectively, 1.1 and 0.8 mm; comparable dimensions are seen in MCZ DCO Ge 0.9 conn) pee Wine away Onna formed by the apposition of this facet with another on the medial surface of the pseu- doangular (Figs. 19A, 24C, 26B; see also Jenkins and Walsh, 1993, fig. le). The sur- faces of both facets are of “unfinished” bone, evidence of a cartilaginous covering and a synovial joint. The facet on the sta- pes- -quadrate faces principally laterally but is inclined slightly ventrally (Fig. 29A): the relatively flat surface, which exhibits a very shallow anteroposterior concavity, con- trasts markedly with the complexly config- ured quadrate and pseudoangular facets in Recent gymnophionans (Fig. 29B, G).An- terior to this facet is a rounded, transverse bar that forms the posterior margin of the adductor chamber; relative to the lower jaw, the bar is positioned at a point just posterior to the insertion of the adductor muscles. The finished surface of the bone lacks any indication of articular cartilage; ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll aU UI art qu pt 1mm ; st f Figure 20. Left stapes-quadrate of Eocaecilia micropodia, MCZ 9169. (A) dorsal, (B) ventral, (C) anterior, (D) lateral, (E) medial, (F) posterior views. therefore, the bar appears not to be a com- ponent of the jaw joint. In most gymnophionans the quadrate and stapes retain separate identities. The apparent fusion of the stapes-quadrate in Eocaecilia micropodia represents a more derived condition than that of some Re- cent caecilians in which the quadrate and stapes are closely abutted and even artic- ulate by means of a synovial joint. In a ju- venile Ichthyophis glutinosus, for example, de Jager (1939b) described a synovial joint between the stapes and quadrate. Serial sections of adult Ichthyophis glutinosus (Fig. 23) and a specimen of Epicrionops petersi examined by R.L.C. confirm an ex- tensive contact between the quadrate and stapes Yet 1 micropodia is not unique among known caecilians for having com- pletely departed from the ancestral con- dition of a distinct stapes and quadrate. Scolecomorphids lack stapes as well as fo- ramina ovales (Nussbaum, 1985). Speci- mens of Boulengerula boulengeri were re- ported by de Villiers (1936, 1938) to vari- ably exhibit a synovial articulation between the stapes and quadrate or partial fusion of the two bones. de Jager (1939a) de- scribed an incipient fusion of the stapes and quadrate in Dermophis mexicanus, but this condition was not confirmed by Wake and Hanken (1982: 214), who reported that “the stapes is always free of the quad- rate....” In D. mexicanus, however, the quadrate fuses with the pterygoid (Wake and Hanken, 1982), a condition that Law- son (1963) believed to be present in Hy- pogeophis rostratus. The massive stapes-quadrate, intimately articulated with the quadrate ramus of the pterygoid and functioning in the jaw artic- ulation, presumably could have transmit- ted vibrations by bone conductance. Sta- pes in Recent caecilians serve both audi- tory and structural roles, linking the brain- case (otic capsule) with the cheek (quadrate). Although the modern caecilian stapes is not a component of an impedance matching system, as in amniotes and most frogs, the apparatus does respond to low- 316 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Figure 21. Reconstruction of the right stapes-quadrate of Eocaecilia micropodia based on MCZ 9169 and MNA V8066, V9346. (A) dorsal, (B) ventral, (C) lateral, (D) medial, (E) anterior views (E drawn from MNA V8059). frequency vibrations (Wever, 1975; Wever and Gans, 1976). Operculum. An ossicle is preserved with several specimens, either in proximity to the fenestra ovalis or stapes-quadrate, or in isolation. In MCZ 9242 (op, Fig. 12) and MNA V8054 (Fig. 25G) the ossicle is a thin, oval plate resting against the stapes- quadrate. The medial surface facing the fenestra ovalis is concave. In both MCZ 9242 (Fig. 12) and the holotype, the edges are thickened, as in the case of gymno- phionan stapedial footplates that articulate with the margins of a fenestra ovalis (Fig. 22). In MCZ 9156 (Fig. 9), the ossicle lies against the retroarticular process, close to the jaw articulation. In the holotype (Fig. 3), the ossicle appears to have been turned outward and lies immediately adjacent to the fenestra ovalis. The variable positions in which the ossicle is preserved are evi- dence that this element was not rigidly at- tached but presumably was anchored by soft tissue. In MCZ 9169 (Fig. 15C), how- ever, the ossicle was exposed by removal of the stapes and, although fragmented, was found to be closely associated with the fenestra ovalis; the reconstructed shape of the ossicle is an elongate oval that would have occupied most or all of the fenestra ovalis, the margins of which are well pre- served. ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 1mm Figure 22. Stapes of the Dermophis mexicanus in (A) medial, (B) anterior, (C) ventral, (D) dorsal, and (E) lateral views. Drawn from a specimen described by Wake and Hanken (1982) in the collection of Marvalee H. Wake, Department of Integrative Biology, University of California, Berkeley. The oval shape and otic association of the ossicle of Eocaecilia micropodia are comparable to that found in the opercula of frogs (Wever, 1973) and salamanders (Monath, 1965), and on this evidence, the ossicle can be proposed as a homologous opercular element. Unlike Recent caeci- lians, E. micropodia retained a shoulder girdle to which an opercularis muscle could have attached. No extant caecilian is known to have an operculum. Duellman and Trueb (1986: 387) state that the oper- culum in caecilians is either absent or fused with the footplate of the stapes and elsewhere (1986: 306) cite Marcus (1935) as suggesting that the operculum in Hy- pogeophis “... has been incorporated into the stapedial footplate, as ... in pletho- dontid salamanders.” However, Marcus ac- tually made no so such claim, having been unable to identify any vestige of an oper- culum in his developmental study: “Von ei- nem operculum finde ich auch in der En- twicklung keine Spur” (Marcus, 1935: 145). Although Marcus (1935) did suggest 317 that part of the otic capsule is incorporated into the stapedial footplate in Hypogeo- phis, Wake and Hanken’s (1982) study of Dermophis mexicanus revealed only a sin- gle ossification center for the stapes, with no evidence of a separate origin for the footplate. On the basis of the limited evi- dence available, we conclude that the operculum in Recent gymnophionans has been lost. An assessment of the possibility that the ear ossicle in Eocaecilia micropodia is ac- tually a reduced stapes, rather than an operculum as here interpreted, can only be made with reference to structures in putatively basal gymnophionans (i.e., the rhinatrematids Epicrionops and Rhinatre- ma; Nussbaum, 1977, 1979; Wilkinson, 1992, 1996; Wilkinson and Nussbaum, 1996). In these forms, the stapes is a rel- atively massive bone that fits tightly into the margins of the fenestra ovalis; a ros- trally directed stem articulates via a syno- vial joint with the posterior surface of the quadrate, and a stapedial foramen travers- es the base of the stem. Although the sta- pes-quadrate of E. micropodia possesses a stapedial foramen, the bone does not “fit” within the fenestra ovalis. Postmortem dis- tortion obviates the possibility of deter- mining whether the stapes-quadrate con- tacted or simply lay in close apposition to the otic capsule. Certainly the concave medial surface of the posterior process of the stapes-quadrate exhibits no structural feature that would conform to the fenes- tral aperture. Iwo interpretations are therefore possible. First, the ear ossicle of E. micropodia is a reduced stapes, and the correspondingly enlarged quadrate has captured the course of the stapedial artery. Alternatively, the ossicle is a true opercu- lum, and the stapes retained the primitive relation to the stapedial artery but lost the primitive seating of a footplate within the fenestra ovalis. We favor the latter inter- pretation on several grounds. Given the in- timate association of stapes and quadrate in numerous extant taxa, the fusion of these two elements is not an altogether 318 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Figure 23. Horizontal section of the quadrate region of Ichthyophis glutinosus exhibiting an extensive cartilaginous contact between the stapes and quadrate, with both bones contributing to the facet for the pseudoangular. Based on serial sections in the collection of Marvalee H. Wake, Department of Integrative Biology, University of California, Berkeley. unexpected specialization. Second, postu- lating the loss of an arterial foramen from one element and its reappearance in an- other would require at least some corrob- orative evidence, of which there is none. Lower Jaw. Among the most durable of skeletal elements, the lower jaws of Eocae- cilia micropodia are represented in artic- ulation with several skulls (Figs. 3, 11, 13), as well as by numerous isolated specimens (Figs. 24-26). In contrast, no bones could be recognized as belonging to the hyo- branchial apparatus, a not unexpected finding because the hyobranchial appara- tus does not ossify in the adults of Recent caecilians. Eocaecilia micropodia shares with most living caecilians the following unique com- bination of mandibular features: The jaw comprises two distinct bones— the pseudodentary and the pseudoangu- lar—which extensively overlap along their medial and lateral surfaces, respectively, and are thus joined along a wide suture that transects the jaw obliquely (Fig. 25C, D). There is no trace of any other separate ossification. The pseudoangular, which in- corporates the articular, forms the mandib- ular articulation with the skull (Figs. 24C, G, 25F, 26B). A lingual (or splenial) row of pedicellate teeth, which parallels the primary labial ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll row, extends posteriorly from the symphy- sis to about the midpoint of the labial row (Figs. 24E, 26A). Data presented by Taylor (1968) demonstrate that the lingual row is a highly variable feature among gymno- phionans. A row containing 2—20 teeth is present in representatives of three families (rhinatrematids, typhlonectids, and uraeo- typhlids) but is altogether absent in sco- lecomorphids (Scolecomorphus spp., Cro- taphatrema bormuelleri). Of the two gen- era of ichthyophiids, a lingual row is pre- sent in Ichthyophis but not developed in Caudacaecilia. Among caeciliids, many genera lack lingual teeth (Boulengerula, Brasilotyphlus, Dermophis, Leutkenotyph- lus, Microcaecilia, Mimosiphonops, Parvi- caecilia, Pseudosiphonops, Siphonops) but even more possess them (Caecilia, Gege- neophis, Geotrypetes, Grandisonia, Gym- nopis, Herpele, Hypogeophis, Idiocranium, Indotyphlus, Oscaecilia, Praslinia, Schis- tometopum, Sylvacaecilia; for the last ge- nus, see Wake, 1987a). A well-defined mandibular fossa is de- veloped on the dorsal surface of the pseu- doangular for insertion of the adductor musculature (Figs. 24C, 27C). In the depths of the fossa is a foramen (Fig. 27C) whereby V, and accompanying vasculature are conveyed into an intramandibular ca- nal. There is no adductor fenestra, as in anurans and salamanders. The pseudoangular bears a robust inter- nal process that projects dorsomedially (Figs. 24D, 25C, H). The apicobasal height and anteroposterior width (mea- sured across its base) of the internal pro- cess is relatively greater, and in some cases substantially greater, than that found in a sample of extant gymnophionans (Table 1). The pseudoangular also bears an elongate, posteriorly projecting retroarticular pro- cess that in life likely extended just caudal to the level of the occiput (Figs. 1A, 25D, 26B, 27C). The robustly constructed re- troarticular process is slightly less than 20% of the lower jaw length, and thus shorter than those measured from selected extant taxa, which range from 20 to 32% 319 (Table 1). The posterior terminus in a number of specimens is obliquely truncat- ed (Fig. 27D) so as to face posterolaterally, although this geometry is not seen in every specimen (e.g., Fig. 24C). A slightly raised area developed on the terminus extends onto the dorsal aspect of the process (Fig. 24A, C); the area appears to represent a muscle insertion, probably for the inter- hyoideus posterior. A rugose depression, on some specimens linear (Fig. 25B) but on others irregular (Fig. 24C, G), is situ- ated on the dorsal aspect of the process and might represent the insertion of the depressor mandibulae. The arrangement of neurovascular fo- ramina is also similar to those in Recent taxa. In Hypogeophis rostratus (Lawson, 1963) and other living caecilians, the man- dibular branch of cranial nerve V and the mandibular artery and vein pass into the lower jaw via a foramen in the floor of the adductor fossa. A comparable arrangement occurs in Eocaecilia micropodia, although two foramina, rather than one, are present (Fig. 24C, G). The neurovascular bundle is distributed to two regions. First, the in- tramandibular branches of the mandibular vasculature and the ramus intramandibu- laris of V exit about midway along the me- dial surface of the jaw through a conspic- uous intramandibular foramen. A compa- rable foramen is present in Eocaecilia mi- cropodia (Figs. 24D, 25C, 27B). Second, the external and alveolar branches of both the vasculature and nerve V_ continue through the ramus toward the symphysis. Small foramina on the medial and lateral surfaces of the symphysial region provide access to surrounding soft tissue; these are evident in E. micropodia (Figs. 24A, 27A, B). As in living forms, E. micropodia pos- sesses a number of vascular foramina along the trough between the two tooth rows that served the spongy tissue surrounding the tooth rows. In Dermophis and Gymnopis, the ramus alveolaris of cranial nerve VII enters the lower jaw through a foramen that lies pos- terior to the base of the internal process, 320 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 art st-q mf Pe Figure 24. The lower jaws of Eocaecilia micropodia. MNA V8070 in (A) lateral, (B) medial, (C) dorsal (see also Fig. 26B), and (D) ventromedial views. In D the jaw is oriented in the plane of the internal process to show the length of the process. (E) A pseudodentary (MCZ 9152) in dorsal view; see also Fig. 26A. (F) A pseudoangular (MNA V8058) in medial view. (G) MNA V8058, a pseudoangular and partial pseudodentary (rostral end missing) in oblique dorsomedial view. approximately in the transverse plane of the jaw joint (for an illustration of Hypo- geophis rostratus, see Lawson, 1963, fig. 9a, b). In Eocaecilia micropodia, the fo- ramen is comparably positioned relative to the jaw joint but is separated from the in- ternal process which is relatively larger and located in a more rostral position (Figs. 3, 24F, 27A—-C). The foramen vari- ably occurs either on the ventral margin of the bone or slightly displaced onto either the medial or lateral surface. In modern ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 321 oe pe teS El aay yu DUD Y - the teeth are more numerous than in any oth- er known genus of Caeciliids . . . [with] 40 to 48 teeth on each side of the upper jaw, and about as many in the outer mandib- ular series on each side; about 45 inner mandibular teeth altogether.” Close ex- amination of Boulenger’s illustration (1909, fig. 1b) shows a unilateral count of 47 premaxilla—maxillary teeth, 40 vomero- palatine teeth, and 47 or 48 and 25 or 26 in the labial and lingual pseudodentary rows, respectively. Subsequent authors re- port comparably high counts; Parker (1941) cited 36-48 premaxilla—maxillary teeth, and Nussbaum and Wilkinson’s (1989: 37) diagnosis of this monotypic ge- nus includes “teeth small, uniform in size, more than 50 per row, except for the splenials.” No data on the diameter and height are available in the literature; mea- surements made with an ocular microme- A composite reconstruction of Eocaecilia micropodia depicting the relative size of skull, vertebrae, limb girdles and ter of the teeth illustrated by Boulenger (1909, fig. 1b) yield a tenuous estimate of an apicobasal height of 0.17 mm, which is in the range observed for Eocaecilia mi- cropodia teeth. Thus P. cooperi is the only known living caecilian with teeth in equiv- alent numbers and size as those in E. mi- cropodia, and it is therefore regrettable that little is known of the dietary habits of this species; Nussbaum (1984) was unsuc- cessful in his earlier attempts to collect this apparently rare caecilian but has since reported (personal communication) col- lecting several specimens. The Cretaceous caecilian Rubricacaeci- lia monbaroni appears to have had fewer teeth in the lower jaw than Eocaecilia mi- cropodia—with 28 in the labial pseudo- dentary row and only two in the lingual (splenial) row (Evans and Sigogneau-Rus- sell, 2001). Yet, like E. micropodia, the teeth of R. monbaroni are very small, with a basal diameter of about 0.3 mm (esti- mated from Evans and Sigogneau-Russell, 2001, fig. 4B). Although the base of the crowns are thus about twice that of E. mi- cropodia, both fossil taxa have, by com- parison to those of most living caecilians, diminutive dentitions. Postcranial Skeleton Overview. Postcranial bones are abun- dant among the available materials of Eocaecilia micropodia; most elements, with the exception of the pelvis, manus, and pes, are represented by multiple spec- imens. The relative completeness of this assemblage provides a sound basis for a reconstruction that depicts the relative size of the skull, vertebrae, and limbs (Fig. 31). ANATOMY OF EOCAECILIA MICROPODIA ¢ Jenkins, Walsh, and Carroll Although the number of vertebrae cannot be precisely determined on the basis of available material, there is sufficient evi- dence from several specimens, reviewed below, that the presacral region of E. mi- cropodia was elongate, as in modern cae- cilians. A distinct tail was present, com- parable to the postcloacal region of rhin- atrematids, ichthyophiids, and uraeotyph- lids (Nussbaum, 1977; Nussbaum and Wilkinson, 1989). Relative to the size of the vertebrae, the limb bones and girdles are diminutive. Postcranial Axial Skeleton In Recent caecilians, the vertebral col- umn can be most simply described as be- ing composed of an atlas, a large number of trunk vertebrae, and, in taxa in which postcloacal vertebrae occur, caudal verte- brae. The subtly gradational nature of the axial column poses a challenge to distin- guishing vertebral regions. In a study of Hypogeophis rostratus, Lawson (1963: 271) concluded that “with the exception of the first or atlas ..., [the vertebrae] are essentially of a uniform structure without regional variation.” In contrast, Wake (1980a) identified regional differences in both structure and growth allometry in Dermophis mexicanus, Ichthyophis gluti- nosus, and Typhlonectes compressicauda that characterize “cervical,” midbody, and posterior vertebrae. Although Taylor (1977) initially suggested that modifica- tions of the anterior four or five vertebrae were sufficiently distinctive to designate them cervicals, Wake (1980a) demonstrat- ed on the basis of the taxa included in her study that the number of distinctive ante- rior vertebrae is much larger. The first 20 vertebrae typically possess a longitudinal nuchal keel; short, widespread parapoph- yses; and broad, relatively flat pre- and postzygapophyses; Wake suggested that these “cervical” features are related to sta- bilizing and elevating the head during bur- rowing. Comparison of vertebral regions in Eocaecilia micropodia with those of mod- ool ern caecilians cannot be undertaken with certainty throughout the axial skeleton be- cause no specimen preserves a complete vertebral column. Nonetheless, the atlas is well known, and the identity of the second vertebra (axis) and other vertebrae associ- ated closely with the atlas is secure; these are referred to as postatlantal (PA) verte- brae, rather than cervical (C), for they largely lack those features that character- ize the “cervicals” of living caecilians (fide Wake, 1980a). If the precise location of the shoulder girdle along the postatlantal ver- tebral column were known, a cervical re- gion might be securely identified. The shoulder girdle and forelimb elements in MCZ 9169 (Fig. 35) are disarticulated and not certainly in place with respect to the associated vertebral column. Lacking evi- dence in any specimen of enlarged ribs on postatlantal vertebrae, which might also indicate the placement of the shoulder gir- dle, the length of the “cervical” region re- mains moot. A sacral region is known, as is the caudal series. The fact that the anal- ysis and description of the vertebral col- umn is based on several incomplete spec- imens removes the possibility of using within-column variations in size as a useful descriptor. Wake (1970: 33) expressed the opinion that “without fossil evidence, identification of the centrum of modern amphibians [frogs, salamanders, and caecilians] with the pleurocentrum of fossil forms is not warranted.” Eocaecilia micropodia pro- vides the desired evidence, for caecilians at least, by possessing small, crescentic in- tercentra. The gymnophione centrum thus appears to represent a pleurocentrum. In- tercentra occur in specimens in which the atlas, axis, and other vertebrae of the post- atlantal region are preserved in articula- tion. Intercentra also occur in a specimen interpreted as representing the anterior dorsal region. The absence of intercentra in other series of vertebrae could possibly reflect regional variation, but postmortem loss cannot be ruled out. Atlas. The atlas, represented in several 332 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 1mm Figure 32. The atlas of Eocaecilia micropodia (MCZ 9231) in (A) lateral, (B) anterior, and (C) posterior views (stereophoto- graphs). specimens (MCZ 9167, 1969, 9171; MNA V8059, V8066), is best preserved in MCZ 9231 (Fig. 32), which serves as the primary basis for the following description. The length of the atlas centrum, excluding the pyramidal interglenoid tubercle that pro- longs the floor of the neural canal rostrad (Fig. 32A), is 1.5 mm. An interglenoid tu- bercle, a feature found in salamanders (Francis, 1934), certain microsaurs (Car- roll and Gaskill, 1978, figs. 115, 116), and albanerpetontids (McGowan, 1998), is not known to be present in any Recent gym- nophionan (Wake, 1970). The two condy- lar facets (or cotyles) extend from the ven- trolateral aspects of the interglenoid tu- bercle to the lateral margin of the cen- trum. The dorsomedial part of each condylar facet thus faces ventrolaterally and the lateral part rostrad (Fig. 32B). The anterior width of the centrum measured across the facets is 1.8 mm. On the ventral surface of the centrum is a distinct notch between the condylar facets; a rounded, median keel occupies the posterior ventral half of the centrum. A small, presumably vascular foramen occurs on the lateral side of the centrum. The posterior end of the centrum (Fig. 32C), excavated by a deep notochordal fossa, is 0.8 mm wide and 0.85 mm high. No evidence exists of a diapophysis. MCZ 9169 (Fig. 33A) reveals that the head of the first rib articulated with the Figure 33. Vertebrae of Eocaecilia micropodia (MCZ 9169). (A) An articulated atlas and axis in lateral view. (B) Articulated vertebrae interpreted to be the sixth and seventh vertebrae of the vertebral column, lateral view. ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll atlantoaxial intercentrum and that the tu- berculum articulated with the axial di- apophysis. A foramen on the lateral sur- face of the base of each pedicle is a feature that is also present in other postatlantal vertebrae of Eocaecilia micropodia. A comparably situated intravertebral fora- men, which communicates with the neural canal, occurs in the atlas in at least some Recent taxa: Gymnopis multiplicata (MCZ 29265) and Dermophis mexicanus, Ichth- yophis glutinosus, and Typhlonectes com- pressicauda (Wake, 1980a). A foramen on the medial wall of the pedicle of the atlas and the following four postatlantal verte- brae in the holotype of E. micropodia (MNA V8066, Fig. 34) is evidence of a complete passage through the pedicle; therefore, the canal is most likely an intra- vertebral foramen for neuronal egress (rather than a vascular channel into the bone). Anteriorly the laminae (i.e., the dorsal roof of the neural arch) extend as an archlike prolongation over the intergle- noid tubercle and probably closely ap- proached the margin of the foramen mag- num. The anteroposterior length of the arch from the anterior margin of the lam- inae to the tips of the postzygapophyses is 2.3 mm, or 0.8 mm longer than the length of the centrum. The postzygapophyses, rather than being distinct processes, are united across the midline by a thin lamina of bone. The orientation of the postzyga- pophysial facets is nearly horizontal; they face only slightly laterally. The central floor of the neural canal is relatively flat, but the laminae composing the roof are distinctly arched; the neural canal is thus hemitu- bular, in contrast to the condition typical of Recent caecilians in which the neural canal is tubular in cross section (for addi- tional details of the atlantal neural canal, see the description of MNA V8066 below). The atlantal spinous process is represented only by a low tubercle; the spinous process appears to bifurcate posteriorly into a pair of faint ridges that extend posterolaterally onto the postzygapophyses. The atlas of Eocaecilia micropodia re- 333 sembles the incomplete atlas of Rubrica- caecilia monbaroni from the Early Creta- ceous in the presence of an interglenoid tubercle, the transverse orientation of the condylar facets, and the placement of a conspicuous neural foramen at the base of the pedicle (cf. Fig. 32; Evans and Sigog- neau-Russell, 2001, figs. 4, 5A—C). Evans and Sigogneau-Russell (2001: 268, fig. 5A) interpreted the articular surfaces of the condyles in R. monbaroni as extending across the midline, implying an unusual condition in which the occipital condyles would be spaced very closely together. In E. micropodia, the facets are narrowly sep- arated dorsally, but more widely separated ventrally by a distinct notch (Fig. 32B). Vertebral Regions. Vertebrae and ribs are associated with many specimens of Eocaecilia micropodia, but relatively few are articulated in series (MCZ 9169; MNA V8055, V8062). Although no specimen preserves a complete presacral column, there is sufficient evidence from associated elements (the atlas, pectoral girdle and forelimb, and hindlimb) that five vertebral regions are identifiable: postatlantal, dorsal (including possibly middorsal), posterior dorsal, sacral, and caudal. With the excep- tion of the caudal series, regional varia- tions in structure are subtle. Postatlantal Region. The holotype of Eocaecilia micropodia (MNA V8066, Fig. 34) preserves as an associated series the atlas, axis, and three additional postatlantal vertebrae (PA3—PA5). The vertebrae are damaged and slightly disarticulated post- mortem; the left half of the neural arches are missing, probably as the result of the quarrying process. The length of the atlas centrum is 1.5 mm. Other centra are slightly longer; the axis and PA4 are 1.6 mm, and PA5 is 1.7 mm (the length of PA3, as preserved, is 1.5 mm, but this is probably an artifact). Overall vertebral lengths (measured from the rostral ex- tremity of the prezygapophysis to the cau- dal extremity of the postzygapophysis) could only be estimated on PA2 and PA4 and are in the range of 2.2-2.3 mm. 334 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Figure 34. (A) The holotype skull, atlas and associated postatlantal vertebrae of Eocaecilia micropodia (MNA V8066) at the completion of the initial stage of preparation and before removing the skull from the matrix. (B) Enlarged view of the vertebrae after detailed preparation; rostral is to the left. The loss of the left half of the neural arches exposes the right pedicles in medial view, which reveals an internal process on the medial surface of each pedicle (p i p, white arrows). Anteroventral to each process is the internal ostium of the intravertebral foramen (stereophotographs). The loss of the left half of the neural arches of the atlas, axis (PA2), PA4, and PA5 (on PAS the arch is completely broken away) exposes the internal surface of the neural canal. The medial openings of in- travertebral foramina are evident on all pedicles. Bilaterally, along the pedicle— centrum junctions, the floor of the neural canal of the atlas is excavated by a deep, longitudinal sulcus. The rostral end of the sulcus begins at the transverse level of the medial opening of the intravertebral fora- men; the sulcus extends caudally to the posterior end of the atlantal centrum. The two sulci together compose about two- thirds of the breadth of the neural canal: ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll in the midline, the floor of the neural canal is slightly convex, except in the region of the interglenoid tubercle where the sur- face is slightly concave. Comparable sulci are exhibited on the axis—PA5, although 335 ni (MCZ 3424), Gymnopis multiplicata (MCZ 29265), Hypogeophis rostratus (MCZ 48935), and Oscaecilia ochrocepha- la (MCZ 14817). In species of the caeciliid genus Schistometopum, however, process- they differ by being transversely narrower and by extending the full length of the pedicle—centrum junction. The most extraordinary feature of the neural canal is the presence of a bony pro- cess that projects medially from the inter- nal surface of each pedicle (Fig. 34B). On the atlas, the process is situated directly posterior to the intravertebral foramen and near the pedicle—centrum junction; the at- las from another specimen shows the same feature (Fig. 15D). On the axis (PA2), the process is likewise posterior to the intra- vertebral foramen, but both the process and foramen are more dorsally positioned, approximately halfway up the internal face of the pedicle; the process, which is 0.2 mm in length, tapers to an apex and is di- rected medially and somewhat ventrally. The width of the neural canal of the axis is estimated to be 0.9 mm, but the two processes would reduce the space avail- able for passage of the spinal cord to about 0.5 mm. On PA4 and PAS, the intraver- tebral foramina lie close to the pedicle— centrum junction, and the processes are situated posterodorsal to the foramina. Internal processes within the neural ca- nal of anterior vertebrae have not previ- ously been reported in any Recent gym- nophionan. A review of atlantal specimens in the MCZ Herpetology collection reveals that comparable processes do occur in some taxa but are apparently not common. Representatives of Ichthyophiidae (Ichth- yophis glandulosus, MCZ 14003), Typhlo- nectidae (Typhlonectes compressicauda, MCZ 24524), and Scolecomorphidae (Sco- lecomorphus kirkii, MCZ 12234, 27120) exhibit pedicles with smooth internal sur- faces. Similarly, a number of caeciliid spe- cies are without processes: Boulengerula boulengeri (MCZ 12309), Dermophis mex- icanus (MCZ 12122), Gegeneophis ramas- wamii (MCZ 29460), Geotrypetes seraphi- es are present. In S. gregorii (MCZ 20057, 20070) distinct digital processes are di- rected anteromedially, whereas in other specimens (MCZ 20055, 20146) the pro- cesses are less prominently developed and in some cases are low rugosities. Processes are also present in S. thomensis (MCZ 29450). In one specimen of S. gregorii (MCZ 20056), dry preservation of soft tis- sues reveals that the processes are anchor points for a connective tissue suspensory ligament (very likely the pia mater, as in the denticulate ligament) that passes to the ventral surface of the spinal cord. A spec- imen of the uraeotyphlid Uraeotyphlus ox- yurus (MCZ 9484) bears low tubercles that are somewhat rugose. Comparable structures occur in some snakes (e.g., boids, Python, Romer, 1956, fig. 1291; Boa constrictor, MCZ 13019; col- ubrids, Coluber constrictor, MCZ 160014; and elapids, Ophiophagus hannah, MCZ 67054) as a delicate, longitudinal ridge that runs anteroposteriorly along the me- dial aspect of the pedicle at a level com- parable to that of the caecilian process. I]- lustrations of the atlas of a Cretaceous cae- cilian, Rubricacaecilia monbaroni, and the Recent form Ichthyophis mindanaoensis (Evans and Sigogneau-Russell, 2001, fig. 7B, E) might be interpreted to depict sim- ilar structures, but in neither case are the features comparable (S. E. Evans, personal communication). MCZ 9169 preserves an atlas, axis (Fig. 33A), and at least four additional vertebrae in a more or less articulated series. The atlantal centrum is 1.3-1.4 mm in length, very slightly shorter than in MCZ 9231, but otherwise similar to the latter in al- most all details, including the presence of both spinal and vascular foramina. The ex- ception is the spinal region. In place of the single, median tuberosity observed in MCZ 9231, the spinous process is a very 336 narrow, low, elongate ridge (the nuchal keel of Recent gymnophionans; Wake, 1980a), flanked on either side by a small tuberosity. The axial centrum of MCZ 9169, 1.6 mm in length, is separated from the atlas by an intercentrum that bears a postero- laterally directed parapophysis. Although the first rib is missing, the facet for the rib head is round and relatively large (0.4 mm diameter). The posteroventral margin of the axial centrum bears a raised parapoph- ysial demifacet for the second rib (also missing). The axial diapophyses are more massive and project farther from the ped- icle than any of the more caudal diapoph- yses; oval in cross section (diameters 0.7, 0.3 mm), the long axis is oriented oblique- ly (the caudal end is superior, the rostral end inferior). Above the diapophysis is a foramen, probably serving the same func- tion as that in the atlas, but situated more dorsally on the pedicle. The spinous pro- cess, or nuchal keel, is a higher ridge than that on the atlas. A pair of tubercles, com- parable to those that flank the spinous pro- cess anteriorly on the atlas, are posteriorly positioned on the dorsum of the postzyg- apophyses. The articulation between the atlas and axis confirms that the midline lamina of bone between the atlantal post- zygapophyses intimately overlies the ante- rior part of the axial neural arch. The third and fourth postatlantal verte- brae, disarticulated and somewhat dis- placed, are obscured by fractures and overlying bone fragments. Features that are evident (very low nuchal keel, bony lamina linking the postzygapophyses, ori- entation of the diapophyses) appear to be comparable to those of the axis. The cen- tra, however, are longer (1.7—1.8 mm). An intercentrum is preserved along the anter- oventral margin of PA3. A limb bone with a shaft diameter 0.4 mm and length of over 2 mm is associated with PA4; al- though the ends are somewhat crushed, the appearance most closely resembles a radius. Proximal rib fragments associated with Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 a skull (MCZ 9242, Fig. 12) are likely to be derived from the postatlantal region. The capituli and tuberculi, which are well separated, expand toward each terminus and bear a circular facet. The postatlantal region, reconstructed from MNA V8066 and MCZ 9169, can only be distinguished on the basis of a sin- gle feature: spinal nerves traverse foramina through the pedicles on the atlas, axis, and at least the next three vertebrae, whereas on anterior dorsal vertebrae, the egress of spinal nerves appears to have been around the posterior margin of the pedicle. Oth- erwise, postatlantal vertebrae of MCZ 9169 are structurally gradational with those of the anterior dorsal region. Rostro- caudally, the length of centra increases from 1.4 mm on the atlas to 2.1 mm on the 11th vertebra, the angle of zygapophy- sial facets shifts from nearly horizontal to about 45°, diapophyses are reduced, and ribs become shorter and more slender, with more closely spaced capituli and tu- berculi. Anterior Dorsal Region. The next two vertebrae in the MCZ 9169 series are ar- ticulated and well preserved (Fig. 33B) and are separated from PA4 by a gap of approximately one vertebral length. Two lines of evidence support our interpreta- tion that the gap represents a missing ver- tebra (or possibly vertebrae), and that the two next vertebrae are probably the sixth and seventh. First, the two vertebrae are lying on their left side, whereas PA4 is ly- ing on its right side, indicating that the continuity of the series has been substan- tially disrupted. Second, the anterior ver- tebra of the pair is unlikely to represent PA5 because it lacks a spinal foramen in the pedicle, which is known to be present in PA5 of the holotype. For purposes of the present description, the pair is designated as the sixth and sev- enth vertebra (Fig. 33B). The notochordal centra are 1.7—1.8 mm in length, with con- cave lateral and ventral surfaces. The ped- icles are mounted anteriorly on the cen- trum. Both lack spinal foramina traversing ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll the pedicles. The posterior margins of the pedicles are deeply incised for passage of the spinal nerves: the intervertebral “fo- ramen” is therefore situated over a single centrum, rather than being truly interver- tebral. The transition from the condition in which spinal nerves pass through foram- ina in the pedicles to the condition in which they pass behind the pedicles ap- pears to take place at vertebra 6. Variabil- ity of this feature is known in Recent taxa, with the transition occurring between the 15th and 21st vertebra in Dermophis mex- and between the fifth and 11th in icanus, Typhlonectes compressicauda (Wake. 1980a). A small tubercle (anapophysis) along the posterior margin of the pedicle lies in close relation to the prezygapophy- sis of the following vertebra, and would seem to support the zygapophysis from be- low. The diapophysial facets are compara- ble in orientation and dimensions (0.8 and 0.4 mm along the long and short axes, re- spectively) to those on the axis, but they are not as protuberant. Rib heads in this region appear to have articulated with par- apophysial hemifacets developed on the margins of adjoining centra anteroventral to the diapophysis. The postzygapophyses are interconnected by a bony lamina; zyg- apophysial facets appear to be less hori- zontally inclined than in more rostral ver- tebrae. The length of the vertebrae, mea- sured between the tips of the pre- and postzygapophyses, is 2.7 mm. Nuchal keels appear to be low: the spinous process is represented primarily by an elevated area between the dorsal surfaces of the post- zygapophyses. Three ribs preserved on the right side of the sixth and seventh vertebrae appear to be associated with these vertebrae, but they are displaced from their articulations (Fig. 33B). The most anterior rib, which is fractured but complete, measures 3 mm from head to distal tip; the proximal shaft has a diameter of 0.38 mm and the centers of the capitular and tubercular facets are 0.8 mm apart. The successive ribs appear to be slightly more slender, but the prox- JSo7 imal ends are unexposed, and the distal ends broken. An isolated vertebra, lying less than 2 mm away from the atlas axis of the MCZ 9169 series, appears to have characteristics intermediate between the sixth and sev- enth vertebrae and another articulated se- ries of four vertebrae. The isolated verte- bra, tentatively identified as the eighth, has a centrum length of 1.5 mm, a pre- to postzygapophysial length of 2.8 mm, and a smaller diapophysial facet (0.5 and 0.2 mm along the long and short axes, respectively: the long axis is less vertically inclined). The pre- and postzygapophysial facets are in- clined with respect to the horizontal by an estimated 20°. A series of four articulated vertebrae, tentatively identified as the ninth through 12th, is characterized by longer centra (2.1 mm) and very faint nuchal keels, with a slight excrescence between the dorsal sur- faces of the postzygapophyses representing the spinous process. Crescentic intercen- tra are present (anteroposterior thickness, 0.3 mm). The rib heads appear to have ar- ticulated with demifacets on adjacent su- perolateral margins of the ends of the cen- tra. There are no diapophysial processes as such. Rather, tuberculi appear to have ar- ticulated in small. circular depressions (0.2 mm diameter) on the side of the pedicle, the centers of which are located 0.5 (on vertebra 11) to 0.6 mm (on vertebra 8) dorsocaudally from the capitular demifa- cets. Immediately posterior to each de- pression is a low tuberosity. Other features are comparable to those seen in PAS and vertebrae 6 and 7, with the exception of the inclination of the zygapophysial facets. Crushing and distortion obviate precise measurement of facet angle, which is es- timated to be at least 30° but not more than 45°. The ribs are shorter and more slender than those of more anterior ver- tebrae, and the capitulum and tuberculum are closer together. Ribs 10 and 11, the most completely preserved im the series, are 2.3 and 2.45 mm long and have prox- imal shaft diameters of about 0.3 mm: the 338 centers of the capitular and tubercular fac- ets of rib 11 are separated by a distance estimated to be about 0.5 mm. Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 symmetrical vertebrae, the facets lie at about 30° to horizontal. Spinous processes are absent; a narrow, low median crest ex- Two additional isolated vertebrae lie within several centimeters of the anterior dorsal series, but neither is sufficiently well preserved to be informative. Two centimeters from the anterior dor- sal vertebrae described above, a coiled but articulated series of 13 vertebrae is asso- ciated with shoulder girdle and forelimb elements (Fig. 35A). The principal fea- tures of more anterior dorsal vertebrae are maintained in this series, with only slight changes in proportions. The series can thus reasonably be interpreted as a contin- uation of the dorsal column, thus repre- senting vertebrae 13 through 25. Mea- sured lengths (in some cases estimated) of 10 centra range from 1.9 to 2.3 mm, with a mean of 2.12 mm, which is comparable to that observed in the anterior dorsal se- ries; variations in length are random along the series and therefore appear to be due to preservational artifact. Total vertebral length, measured from the anterior end of the prezygapophyses to the posterior end of the postzygapophyses, appears to in- crease slightly. Measured or estimated lengths of seven specimens range from 3 to 3.4 mm, with a mean of 3.2 mm (the greatest overall length that can be mea- sured in the anterior dorsal series is 2.9 mm at vertebra 8). An increase in size of the crescentic intercentra intercalated be- tween the ventral margins of the noto- chordal centra appears to compensate for the increase in overall vertebral length when the length of the centra remains the same. Intercentra are 0.4—0.5 mm in an- teroposterior thickness (versus 0.3 mm in the anterior dorsal region). As in more an- terior dorsal vertebrae, the lateral and ven- tral aspects of the vertebral centra are con- cave; a faint median crest extends longi- tudinally along the ventral surface but does not reach the anterior or posterior ends. The orientation of zygapophysial fac- ets in many cases is altered by postmortem plastic deformation; on well-preserved, tends longitudinally from a slight tuber- osity between the bases of the prezyga- pophyses to a more prominent tuberosity between the postzygapophyses. The ribs associated with the fifth and twelfth ver- tebra of this series are slightly longer (length 2.65-2.7 mm) than those of more anterior dorsal ribs but are comparable in proximal shaft diameter (ca. 0.3 mm) and in the manner with which they articulate with the centra. Middorsal Region. In addition to the postatlantal and anterior dorsal vertebrae described above, MCZ 9169 also includes two isolated series of more or less articu- lated vertebrae (on a separate block of ma- trix) that lack any direct evidence of axial position. The shorter series of four dam- aged vertebrae provides little useful infor- mation. The longer series represents 18 vertebrae, although the 12th was lost dur- ing the quarrying process. The vertebrae are morphologically similar, particularly in rib size and articulation, to those of the anterior dorsal series, with the exception that there is no indication of a median nu- chal keel. The lengths of the centra, how- ever, range from 1.9 to 2.0 mm (mean 1.95 mm), which is about 0.25 mm shorter than the lengths of the centra of the last several vertebrae in the anterior dorsal series. If MCZ 9169 comprises a single specimen, the middorsal region is morphologically similar to the anterior dorsal region save for shorter vertebrae and would be com- parable to the gradual decrease in centrum and neural arch length in the middorsal and posterior dorsal regions of Dermophis mexicanus (Wake, 1980a). Alternatively, the series of 18 could represent the dorsal region of a slightly smaller individual. Posterior Dorsal, Sacral, and Caudal Vertebrae. Right and left hindlimb ele- ments are closely associated with a series of vertebrae in MNA V8062 (Fig. 36) and thus provide the evidence for identifying posterior dorsal and caudal vertebrae. Var- ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 339 Figure 35. (A) Anterior dorsal vertebrae of Eocaecilia micropodia (MCZ 9169) associated with forelimb elements: scapulocor- acoid, humerus, and ulna. (B) An enlargement to show the intercentra and details of the vertebrae (stereophotographs). 340 # ae Be Rig eee Sab owe Figure 36. Posterior dorsal and caudal vertebrae, partial femur and ?fibula, and tibia and phalanges of Eocaecilia micropodia (MNA V8062; stereophotographs). iable crushing and distortion of the entire series, however, substantially obscures many structural details. Only one (incom- plete) rib is preserved with the posterior dorsals; this slender rib appears compara- ble to those of other dorsals, with the cen- ters of the tuberculum and _ capitulum spaced 0.4 mm apart and a shaft diameter of 0.2 mm. The two vertebrae that are im- Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 all length and narrower in interzygapophy- sial width. These features are preserved on the second sacral; the first sacral is largely crushed, and only posterior interzygapoph- ysial width is preserved. Sacral centra are approximately 1.4 mm in length, whereas the centrum of the penultimate posterior dorsal is 1.7 mm; the third vertebra ante- rior to the penultimate has a centrum mediately adjacent to hindlimb bones are tentatively identified as sacrals. A charac- teristic sacral feature is the large size of the diapophyses, in contrast to those on posterior dorsal vertebrae that precede them (only four of which are sufficiently preserved to be useful for comparative analysis). In addition to the large size of the tubercular (diapophysial) facets, sa- crals appear to differ from posterior dorsal vertebrae in being slightly shorter in over- length of 1.8 mm. In typical dorsal verte- brae, tubercular facets are flush with the sides of the pedicles; on the putative sa- crals, tubercular facets are elevated from the lateral surface of the pedicles and are thus borne on low diapophyses. On two posterior dorsals, the transverse distance between the lateral margins of the postz- ygapophyses is 1.4 mm, compared with about 1 mm on the two sacrals. Proximal caudals in MNA V8062 appear ANATOMY OF EOCAECILIA MICROPODIA ¢ Jenkins, Walsh, and Carroll 341 Figure 37. to have centrum lengths comparable to those of the sacrals (1.4 mm) but have zyg- apophyses that project dorsally, in contrast to more horizontally inclined processes of dorsal and sacral vertebrae. Distal caudal vertebrae are represented by a single, dis- articulated vertebra and an attenuating se- ries of approximately eight vertebrae. The single vertebra has a centrum length of 1.1 mm and bears a prominent, posteriorly re- curved, tapering spinous process. The ter- minal series of vertebrae is very damaged but exhibits a diminishing length gradient of the centra from about 0.7 to 0.4 mm. MNA V8055 preserves some 30 verte- brae, a number of which occur in articu- lated series. One series of 12 vertebrae ap- pears to be continuous, although the con- tinuity is disrupted between the sixth and seventh by a disarticulation to a right-angle bend (Fig. 37A). A few slender rib frag- ments are associated, and the lengths of the centra diminish from about 2 mm an- teriorly to 1.55 mm posteriorly; this series (A) Dorsal vertebrae, (B) caudal vertebrae of Eocaecilia micropodia (MNA V8055; stereophotographs). appears to represent the dorsal or very possibly posterior dorsal region. Five iso- lated vertebrae, including two that are ar- ticulated, also appear to be dorsals; other isolated vertebrae are too imperfectly pre- served to be useful. Caudal vertebrae (Fig. 37B) are represented in a more or less ar- ticulated series of 10, with the lengths of the centra diminishing from 1.3 mm on the most anterior to approximately Jt ramen on the penultimate. The zygapophyses on the anterior three are small and set close to the midline; distal to the fifth vertebra, zygapophysial articulations are reduced to a simple overlap of a lamina with that of the vertebra behind. Several well-devel- oped haemal arches are associated with this series. The haemals are conventional in structure, tapering from a broad, per- forate base (0.6 mm in width) to the distal apex (1.7-1.8 mm in overall length). Among living caecilians, haemal arches are known only in the rhinatrematid Epicrion- ops (Wake, 1987b, 2003, fig. 22A; Carroll 342 et al. 1999, fig. 6A). The last five vertebrae in this series bear a posteriorly recurved, tapering spinous process comparable to those present in the terminal series of cau- dals in Epicrionops bicolor (Wake, 2003, fig. 22A). The caudal series of MNA V8055 is incomplete, however. The terminal ver- tebrae of Eocaecilia micropodia, known only in MNA V8062 (Fig. 36), exhibit re- curved processes that decrease in size and are lost altogether as the vertebrae are re- duced to tiny cylindrical structures. Comparative Anatomy of the Vertebral Column. The vertebral column of Eocae- cilia micropodia differs from those of Re- cent gymnophionans in the presence of in- tercentra, an interglenoid tubercle on the atlas, and most notably in the absence of the various elaborate processes that occur in modern forms. Gymnophionans typical- ly possess a longitudinal keel or process along the ventral aspect of the centra, an- teriorly projecting parapophysial process- es, and postzygapophyses that are inter- connected by an extension of the laminae which overlay the following vertebra; neu- ral spines are absent except on anterior vertebrae, where there is a longitudinal nuchal keel. Of these features, E. micro- podia possesses the postzygapophysial in- terconnection and overlap, but only the faintest representations of ventral and nu- chal keels. With the exception of the post- atlantal region, in which there are distinct diapophyses, costal tubercles articulate with facets that are only slightly raised from the surface of the pedicles. The first rib has a capitular articulation on the at- lantoaxial intercentrum, but other capituli appear to articulate with pairs of hemifa- cets developed on adjacent centra, with no elaboration of parapophyses or other pro- cesses. Intravertebral neural foramina are present on the atlas and at least the fol- lowing four vertebrae. Last, the promi- nent, posterodorsally reflected spinous processes on distal caudal vertebrae of Eocaecilia are a feature shared with the rhinatrematid Epicrionops (Wake, 2003, fig, 2ZA)y Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 The vertebral structure of Eocaecilia micropodia appears to be generally more primitive than that represented by eight isolated vertebrae of an early Cretaceous caecilian, Rubricacaecilia monbaroni, de- scribed by Evans and Sigogneau-Russell (2001). Eocaecilia micropodia does share certain features with R. monbaroni: am- phicoelous centra, an increase in centrum length from the postatlantal into the dorsal series, a low nuchal keel on anterior ver- tebrae, and circular, low diapophyses. However, E. micropodia does not exhibit any parapophysial processes or basapoph- yses, which in R. monbaroni would seem to represent an incipient stage in the dis- tinctively gymnophionan elaboration of these structures. Furthermore, the mid- ventral region of the centra in R. monba- roni is narrowed to a keellike ridge, of which there is only a faint indication in E. micropodia. Had Eocaecilia micropodia attained the distinctively high vertebral counts of mod- ern caecilians? Published estimates of the range of vertebral counts across living taxa vary slightly. Without citing specific taxa, Nussbaum and Naylor (1982) state that vertebral numbers in gymnophionans range from 70 to 283, whereas Wake (1980a) and Duellman and Trueb (1986) cite postatlantal vertebral counts of 95— 285. Most recently, Wake (2003) cited S6— 285. Taylor’s (1968) monograph, however, reports specific counts for individual spe- cies; the lowest occur among various spec- imens of Epicrionops (e.g., E. bicolor sub- caudalis, 75-78: E. lattivittatus, 78) and the highest in Oscaecilia bassleri (273). Given that vertebral numbers vary intra- specifically (Taylor, 1968), a more precise determination of the range would seem to offer little utility, especially in the context of a comparison with E. micropodia, for which a vertebral count can only be esti- mated. The most complete specimen (MCZ 9169) comprises five series of more or less articulated vertebrae, representing the postatlantal, anterior dorsal, and most probably, dorsal regions. The total number ANATOMY OF EOCAECILIA MICROPODIA ¢ Jenkins, Walsh, and Carroll of vertebrae is 49 if several associated but isolated vertebrae are included. MNA V8062 provides an imperfect representa- tion of the sacral and caudal regions. On the assumption that the identification of two sacral vertebrae is correct (these ver- tebrae are most closely associated with the hindlimb elements), there are at least 13 caudals. A minimum estimate of the total vertebral count in E. micropodia is there- fore 64. Although the anterior and poste- rior ends of the column can be enumer- ated, there is no specimen in which a ver- tebral series links the fore- and hindlimbs, and thus the count in the dorsal region is uncertain. Our conclusion is that vertebral numbers in E. micropodia certainly ap- proached, and very possibly nested within, the lower end of the range of variation known among modern caecilians. Appendicular Skeleton Anatomical Location of the Forelimb. As noted above, shoulder girdle and forelimb elements in MCZ 9169 are associated with an articulated series of 13 dorsal vertebrae (Fig. 35A) that likely represent the 13th through 25th vertebrae. The limb bones, although clearly associated, are rather completely disordered (Fig. 38); thus, they provide no definitive evidence that the forelimb was anatomically situated at this point in the dorsal series. The occurrence of an isolated radius lying on PA4 of MCZ 9169 confirms that limb bones were dis- placed in this specimen. Although the ac- tual position of the girdle and forelimb re- mains uncertain, the scapulocoracoid is here reconstructed in relation to more an- terior dorsal vertebrae (Fig. 31). The oc- currence of various shoulder and forelimb elements with numerous cranial speci- mens (MCZ 9163, 9167, 9171, 9237, 9242: MNA V9056, V8065) is suggestive evi- dence that the forelimb was positioned closer to the head than the association of MCZ 9169 might imply. Scapulocoracoid (MCZ 9237, 9238, 9169, 9171: MNA V8056, VS059, V8064, V8065). The two most completely known 343 scapulocoracoids of Eocaecilia micropodia (MNA V8056, MCZ 9237, Fig. 39B, C) are 3 mm in height, measured from the dorsal margin of the scapula to the posteroventral margin of the coracoid. The scapular blade is narrowly constricted at its midpoint, which is oval in cross section (Fig. 39). Dorsally, the blade is expanded and ends as a convex margin of trabecular bone in- dicative of its continuity with a suprascap- ular cartilage (Fig. 39A). The base of the scapula bears an oval glenoidal facet that faces posteroventrally and slightly laterally (Fig. 38) and an anteriorly directed pro- cess that is incised ventrally. The margin of the anterior process, like the dorsal margin of the scapular blade, appears to have been continued in cartilage; thus, the incisure was probably enclosed to form a coracoid foramen. The suture between the scapula and coracoid is preserved as a faint lineation that passes from the incisure across the glenoid. The coracoid bears an approximately circular, dorsolaterally fac- ing glenoidal facet (Fig. 38); the remainder of the coracoid is an approximately rect- angular plate of very thin bone and is pre- served, in crushed condition, in only two specimens (Fig. 40). The scapular and cor- acoidal ends of the glenoid facet, which have very different orientations, are con- nected by a narrow, intermediate part of the glenoid that turns a spiral from one end to the other. The spiral configuration of the glenoid in E. micropodia is thus similar to the pattern common among Pa- leozoic tetrapods. Ventral to the anterior half of the glenoid is a circular fossa of unknown function. A fossa in a similar po- sition occurs in a few microsaurs (e.¢., Asaphestra, Carroll and Gaskill, 1978: 173-174, fig. 8C; Batropetes, Carroll, 1991: 238, fig. 6A), but there is no com- pelling evidence that these features are strictly comparable. Humerus (MCZ 9163, 9166, 9167, 9171, 9237, 9238; MNA V8056, VS068). Com- plete humeri vary in length from 4.25 mm (MCZ 9169) to 4.4 mm (MCZ 9163, Fig. 41). The bulbous head extends from the 344 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Figure 38. Forelimb elements of Eocaecilia micropodia (MCZ 9169), an enlargement of Figure 35A to show details of the scapulocoracoid, glenoid, humerus, and partial ulna. The upper and lower arrows indicating the glenoid (gl) point, respectively, to the scapular and coracoidal components of the glenoid facet (stereophotographs). dorsolateral to the ventromedial surfaces of the proximal end in a spiral pattern common among Paleozoic amphibians (Fig. 42A—D; also present in MCZ 9166). The deltopectoral crest in an apparently undistorted humerus is reflected ventrally (Fig. 38) and is continuous with a low ridge that extends proximally. A small tu- berosity on the dorsomedial side of the proximal diaphysis (Fig. 42C, D) is com- parable in position to that in some micro- saurs (e.g., Cardiocephalus, Pantylus, Ric- nodon, Carroll and Gaskill, 1978, figs. 122A, B, 123D). In Recent Salamandra, a similar tuberosity serves as the site of in- sertion of subscapular musculature (Fran- cis, 1934). Complete humeri (Figs. 38, 41) exhibit a simple shaft that is approximately circular in cross section. The distal end of the humerus bears a bulbous, hemispher- oidal capitulum for the proximal radius and a broad but distinct trochlea for the proximal ulna (Fig. 42E; see also MCZ 9169, 9172). The ectepicondyle is repre- sented only as a low ridge. The entepicon- dyle, in contrast, is a relatively robust pro- tuberance that is prolonged distally; there is no entepicondylar foramen. Along the margin of the proximal border of the ca- pitulum, the diaphysis is incised with a deep, hemicircular sulcus that presumably accommodated the radius in its most flexed position. MNA V8056 includes an associated distal humerus and a proximal radius and ulna that lie in nearly articulat- ed position. Radius (MCZ 9242, 9169; MNA V8056). A radius presumptively associated with an ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 345 Figure 39. Left scapulocoracoids of Eocaecilia micropodia in lateral view: (A) MCZ 9238, (B) MNA V8056, (C) MCZ 9237. The coracoids are incomplete in all three specimens (stereophotographs). 346 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Figure 40. Scapulocoracoids of Eocaecilia micropodia with complete, although crushed, coracoids. (A) A left scapulocoracoid in lateral view (MNA V8064); the glenoid region has been damaged postmortem. (B) A left scapulocoracoid (in a medial view) and associated skeletal elements in MCZ 9171 (stereophotographs). ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll Figure 41. 347 1mm A complete left humerus of Eocaecilia micropodia (MCZ 9163) in ventral view. The specimen has been slightly crushed, with the deltopectoral crest deflected from the normal ventral orientation (stereophotographs). ulna (MCZ 9242, Fig. 43A) is 2.16 mm in overall length. The surface of the proximal facet is set at an angle of about 20° to a plane normal to the shaft. The length of the radius measured from the most distal margin of the articular facet is 1.9 mm. Diaphyseal width narrows at midshaft to 0.39 mm, with the proximal end of the ra- dius being more expanded than the distal end. The smaller distal articular facet is oval in outline and is set almost perpen- dicular to the shaft. Ulna (MCZ 9163, 9167, 9169, 9238, 9242; MNA V8054, VS065). Complete ule nae vary in length from 2.1 mm (MCZ 9163) to 2.2 mm (MNA VS8065, MCZ 9242). The proximal articular facet is asymmetrically biplanar, with the larger of the two demifacets being on the radial side (Fig. 43B). The olecranon process is short but distinct. The ulna is relatively broad, both mediolaterally and anteroposteriorly at its proximal end, and tapers to its nar- rowest point at midshaft. The distal end is only slightly expanded and bears a circular articular facet set perpendicularly to the shaft. Manus. Although various mesopodial, metapodial, and phalangeal elements are associated with several disarticulated spec- imens of Eocaecilia micropodia, none can be confidently attributed to the manus. Pelvis. No pelvic elements can be posi- tively identified. MCZ 9171 is a disaggre- gated assemblage that includes jaws, an at- las, a scapulocoracoid, and several caudal vertebrae (Fig. 40B). Adjacent to the cau- dal vertebrae is a bladelike bone that is not comparable to any other known skeletal el- ement in Eocaecilia. The bone (?pel, Fig. 40B) widens at one end, is narrower in the middle, and bears a concave articular fac- et, as well as a margin that is broken, at the other end, inviting speculation that the facet could be acetabular. The bladelike, rather than platelike, nature of the ele- ment is suggestive of an ilium, but in view of the lack of any comparative data on the pelvis of primitive caecilians, such an in- ference is entirely conjectural. Some crushed bone associated with the hind- limbs of MNA V8062 (Fig. 36) might rep- resent a pelvis, but the state of preserva- tion is uninformative. s ie) oy) Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 1mm Figure 42. The humeral articular surfaces of Eocaecilia micropodia. (A) A left humeral head in proximal view (MCZ 91771). (B) A left humeral head in proximal view and in slightly lateral aspect (MCZ 9237). (C) Dorsal and (D) ventral views of a left proximal humerus (MCZ 9171). (E) A left distal humerus (MCZ 9237) in end view (stereophotographs). ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll 349 1mm Figure 43. The antebrachial bones of Eocaecilia micropodia. (A) A presumptively associated left radius and ulna (MCZ 9242). (B) Proximal left ulna (MCZ 9238) in anterior view (stereophoiographs). Femur (MNA V8055, VS062). A com- plete left femur (Fig. 44) is 4.4 mm in length and has a midshaft width of 0.4 mm. The bulbous, approximately oval femoral head is oriented primarily dorso- ventrally but is slightly skewed, such that the dorsal half is more anteriorly situated than the ventral half. On the medial and lateral sides of the head are shallow de- pressions, comparable to the foveae of some salamanders that represent the at- tachments of acetabular ligaments from the pubis and ilium (Francis, 1934, pl. V, fig. 32). A prominent, triangular trochanter is situated on the medioventral side of the proximal shaft. The expanded distal end, which is somewhat damaged (estimated width 1 mm), bears two condy ‘les separat- ed by an intercondylar groove. The lateral condy Je is wider and more protuberant and possesses a larger radius of curvature than the medial condyle (Fig. 44C). The lateral epicondylar region is Hastanctly con- vex and facetlike, features that probably represent the articular surface for the fib- ula (Fig. 44B). In overall proportions and, specifically. in the configuration of the femoral head, the foveae, and the trochan- ter, the femur of Eocaecilia micropodia is similar to that in certain modern salaman- ders (for a comparative illustration, see Jenkins and Walsh, 1993, fig. lh, i). The femoral head distinctly differs from those of microsaurs and other Paleozoic amphib- ians, which are anteroposteriorly elongate. Tibia (MCZ 9237: MNA V8055, V. S062). A complete tibia associated with MNA V8062 (Fig. 36) is 2.1 mm in length with a midshaft diameter of 0.4 mm: the tibia associated with MNA V8055 (Fig. 45B) is 1.85 mm in length. The transv ersely ex- panded proximal end bears a slightly con- 350 Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 Figure 44. A complete left femur of Eocaecilia micropodia (MNA V8062) in (A) dorsal, (B) ventrolateral, and (C) ventral views (stereophotographs). vex articular facet that accommodates both femoral condyles; the anterior edge of the facet slopes anteriorly and is bordered by a raised lip. A slightly raised area on the proximal, anterior surface of the shaft ap- pears to represent a cnemial tuberosity (Fig. 45A). The distal articular facet is cir- cular in outline and convex. Fibula (MCZ 9237; MNA_ V8055, V8062). Fibular lengths range from 1.9 mm (MCZ 9237) to 2.1 mm (MNA V8055); midshaft diameter of the former is 0.3 mm. In both specimens, the fibula appears to have been preserved in an ar- ticular relationship with the tibia (Fig. 45). In both cases, the proximal articular facet, the plane of which is set at an angle of about 45° to the shaft, lies proximal to the ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll Boll Figure 45. A tibia, fibula, and incomplete tarsus of Eocaecilia micropodia. (A) Anterior view of a right tibia and fibula (MCZ 9237) preserved in normal anatomical position, associated with the distal end of the femur (above) and three tarsals (below). (B) A presumptively right tibia in posterior view (MNA V8055). The specimen includes a distal fragment of the associated femur, a proximal fibula preserved in normal anatomical position (the remainder of the fibula, lost during excavation, is represented by a matrix mold), four tarsals, and several metatarsals (stereophotographs). 302 end of the tibia, evidence that the fibula articulated with the lateral epicondylar re- gion of the femur. The fibular diaphysis is bowed laterally. Distally, the expanded end of the fibula terminates well short of the distal end of the tibia and bears two facets for contact with a fibulare and interme- dium. Pes (MCZ 9237; MNA V8055, V8056, V8062, V8073). MCZ 9237 and MNA V8055, taken together, demonstrate that the proximal tarsal row comprised a tibia- le, intermedium, and fibulare (Fig. 45A, B). A partial pes associated with MNA V8062 (Fig. 36) shows that at least some digits possessed three phalangeal bones; the most proximal phalanx of the pre- sumptively middle digit has a length of 0.89 mm and an estimated midshaft di- ameter of 0.25 mm. It is likely that Eocae- cilia micropodia possessed only three dig- its because of the occurrence in several specimens (MNA V8056, V8062, V8073) of triads of phalanges. DISCUSSION Jaw Mechanics. Eocaecilia micropodia already possessed many of the major skel- etal features of the jaw apparatus that are retained in Recent gymnophionans (e.g., prominent retroarticular process, pseu- doangular and pseudodentary, internal process of the pseudoangular, mandibular fossa in close proximity to the jaw joint). These structural similarities invite the in- ference that the basic pattern of jaw mus- cles known from living gymnophionans had been developed as well. Bemis et al. (1983) described various as- pects of the feeding apparatus in Dermo- phis mexicanus (skull, lower jaw and artic- ulation, teeth, hyobranchial apparatus, oral cavity and tongue, and jaw muscles) and confirmed the mechanics of the jaw through electromyography. Jaw closure is achieved not only by conventional action of the adductors, but through a gular mus- cle, the interhyoideus posterior, that acts to lower the retroarticular process (Fig. 46A; cf. Nussbaum, 1977). Jaw opening is Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 accomplished by the depressor mandibu- lae, which passes from the back of the skull to an insertion on the retroarticular process. Nussbaum (1983) described the relative size of individual jaw muscles in representatives of four families (Rhinatre- matidae, Ichthyophiidae, Caeciliidae, and Scolecomorphidae) and presented an anal- ysis of the uniquely caecilian feeding ap- paratus. To provide a more specific basis for in- terpreting the jaw mechanics of Eocaecilia, we review here additional details of the cranial muscles in an extant caecilian, Ichthyophis glutinosus, which was selected for this purpose as a relatively basal taxon. Ichthyophis glutinosus resembles other gymnophionans, with the exception of rhinatrematids, in the relatively large size of the interhyoideus posterior and the rel- ative reduction of the adductors (Nuss- baum, 1983, fig. 2). The retention of pre- and postfrontals in ichthyophiids is a prim- itive feature; they are lost or fused in most other gymnophionans. The derivation of the family Ichthyophiidae as being close to the base of the crown group gymnophion- ans is supported by morphological (Nuss- baum, 1979) as well as molecular data (Hedges et al., 1993; San Mauro et al., 2004). Four divisions of the adductor mandib- ulae can be recognized: adductor mandib- ulae externus, adductor mandibulae inter- nus, adductor mandibulae posterior (which includes a levator quadrati), and pterygoideus (Fig. 46B—E). The first three are oriented essentially vertically and can be recognized primarily on the basis of their separation by branches of the trigem- inal nerve. The adductor mandibulae externus (AME) is the largest and most superficial in position, lying lateral to the maxillary branch of the trigeminal nerve (Fig. 46B). Originating on the ventral surface of the skull roof, the muscle converges ventrally to insert on a small area of the pseudoan- gular that includes the mandibular fossa. The adductor mandibulae posterior ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll A 0m 353 Wittintittiiisiiiseier AME Figure 46. The jaw muscles in /chthyophis glutinosus. (A) Lateral view of superficial muscles. (B, C) Lateral view of deep muscles, with the cranial vault resected (oblique hatching) to show origins within the adductor fossa. The muscles in plate B lie lateral to those in plate C. (D) Dorsal view of the adductor muscles with the cranial roof resected. (E) Isolated adductor muscles in posterior view. (AMP) occupies the most posterior por- tion of the adductor chamber and is sep- arated from the AME by V, (Fig. 46B, D). The AMP originates from two areas of the quadrate: the lateral surface of the ptery- goid ramus and the medial surface of the palatine ramus. The insertion is within the mandibular fossa of the pseudoangular, which also transmits the mandibular ramus of V; and the mandibular artery. In the upper portion of the adductor chamber, the adductor mandibulae inter- nus (AMI) is separated from the AME by branches of V,, and more ventrally, by con- nective tissue (Fig. 46C). The AMI origi- nates somewhat ventral to the AME, from a more medial position on the underside of the skull roof. Most of the AMI joins the AME to share the same insertion in the mandibular fossa. A slip of the AMI (the levator quadrati, Fig. 46E), which arises from a more inferior position within the adductor chamber, is separated from the AMI by a large blood vessel accom- panying the deep branch of V, and con- nective tissue. The slip converges to its at- tachment in a pit at the top of the ptery- goid ramus of the quadrate. 304 In contrast to the other adductors, the pterygoideus is oriented more horizontally and is directed posteroventrally from its origin along the anterior margin of the ad- ductor fossa. The muscle divides, with one part inserting on the anterior margin of the internal process of the mandible and the other extending across the ventral as- pect of the retroarticular processes to in- sert on its lateral surface (Fig. 46C). The two divisions are separated by a robust lig- ament that connects the internal process and the pterygoid. The interhyoideus posterior (IHP, Fig. 46A) is a large, fan-shaped muscle that arises from the dorsal and lateral aspects of the neck, and the ventral aspect of the gular and “cervical” regions. The fibers converge to insert on the retroarticular process. A general assessment of the function of some of these muscles can be made from their fiber orientation, relative size, and mechanical advantage (lever arm length) with respect to the jaw joint. The three adductors (AME, AMP, and AMI) insert on the mandible in close proximity to the jaw joint and thus have less mechanical ad- vantage than does the interhyoideus pos- terior that attaches to the long lever arm of the retroarticular process. Furthermore, the origin of the adductors is confined to a relatively small, enclosed adductor fossa, a common condition among extant gym- nophionans (Fig. 6B, C); the adductor fos- sa in rhinatrematids is larger (Fig. 6D). The diverging fiber directions of the in- terhyoideus posterior are evidence of mul- tiple functions. Recruitment of the supe- rior, dorsally directed fibers would raise the retroarticular process and thus initiate jaw opening. Conversely, the bulk of the muscle, with fibers directed ventrally or posteroventrally, would act to depress the retroarticular process and thus contribute to the uniquely caecilian jaw closure mechanism. The contribution of the pterygoideus is less clear and could involve differential re- cruitment of its two divisions during the Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 masticatory cycle. The oblique orientation of the division of the pterygoideus that in- serts on the internal process likely contrib- utes to jaw closure (a conventional func- tion for the pterygoideus among tetra- pods). The other division, which passes be- low the internal process of the pseudoangular and wraps around the ven- tral aspect of the retroarticular process, is of uncertain function. In some mandibles of Eocaecilia micropodia, a groove on the ventral aspect of the jaw below the inter- nal process records the passage of this part of the muscle to the retroarticular process (Fig. 27D). The line of action of this part of the pterygoideus passes across the jaw joint (Fig. 46C); therefore, the fibers pos- sess no lever arm for either jaw opening or closing. Two possibilities remain. This division could produce an anterodorsally directed compressive effect on the jaw joint. Such an action could contribute to joint stability and, in particular, serve as an antagonist to the posteroventrally directed force of the interhyoideus posterior during jaw closure. A second possibility is that this part of the pterygoideus everts (by longi- tudinal lateral rotation) the jaw upon opening; the depressor mandibulae would effect inversion upon jaw closing. The similarities between Eocaecilia mi- cropodia and Recent caecilians notwith- standing, the configuration of the stapes- quadrate, jaw joint, and internal process of the pseudoangular in E. micropodia is ev- idence of a divergence in masticatory me- chanics. The geometric complexity of the articular surfaces borne by the quadrate and pseudoangular in gymnophionans (Fig. 29B, C), which provides structural stability, contrasts to the nearly vertically oriented, planar facets of E. micropodia (Fig. 29A). Unlike the modern caecilian jaw joint, in which the quadrate facets fit securely between condylar processes of the pseudoangular, no interlocking mech- anism is present in E. micropodia, where the joint possesses no mechanical imped- iment to anteroposterior or dorsoventral translation. The dimensional differences in ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll the quadrate and pseudoangular facets are small, however. The evidence that pseu- doangular facets are about 45% longer along an anteroposterior axis than facets on the stapes-quadrate supports an inter- pretation of slight protrusion upon jaw opening and retraction upon closing. The dentition of E. micropodia—with teeth that are numerous but very small—would seem suitable for apprehending only the smallest of prey, and propalinal jaw move- ments could well have been employed in ingestion. With a jaw joint of little intrinsic stability, the large size and dorsomedial orientation of the internal process might reflect the employment of the pterygoi- deus as a jaw protractor. The fusion of the stapes-quadrate and the unusual orienta- tion of the jaw joint might represent a spe- cialization to withstand the medially com- pressive forces generated by the inter- hyoideus posterior. In extant caecilians, the same medially compressive forces would be resisted by the obliquity of the complex interlocking of the quadrate and pseudoangular facets. The Habitus of Eocaecilia micropodia. With the exception of aquatic species, gymnophionans are capable burrowers. Several anatomical features of Eocaecilia micropodia provide a basis for evaluating the degree to which this Early Jurassic caecilian can represent a stage in the evo- lution of gymnophionan fossoriality. In a review of tetrapod limblessness, Gans (1975) observed that the majority of limb- less forms are shelterers or burrowers, the exception being snakes (although uropel- tids among Serpentes are true burrowers). Lizards in which the limbs are reduced to various degrees (e.g., scincids, anguids, cordylids) are not effective burrowers but are principally shelterers in crevice-rich environments. Gans suggested that a uni- versal correlate of limblessness is body elongation and that “limb reduction fol- lowed, and was probably produced by, se- lective pressure established after bodily elongation had occurred” (Gans, 1975: 465). However, in a review of limb reduc- 355 tion in squamates, Greer (1991: 167-171) noted that 22 of the 53 extant lineages that exhibit various degrees of limb reduction are not patently elongate, although he agreed that “... a gross reduction in the number of limb bones is always associated with an elongate body.” Our estimate of the vertebral count in E. micropodia is suggestive of body elongation but does not demonstrate the feature unequivocally. The evidence that E. micropodia had un- dergone some reduction of the appendic- ular skeleton, however, is unambiguous (Jenkins and Walsh, 1993, table 1; Fig. 31). In addition to the retention of limbs, E. micropodia had not yet attained other characteristics that are likely to be related to the fossorial habits of gymnophionans. The mouth is not obviously subterminal as in gymnophionans. The orbits, although proportionately smaller than is typical for Paleozoic amphibians, are larger than those in gymnophionans in which the eyes are much reduced and in some taxa cov- ered by bone. The atlanto-occipital joint of E. micropodia is relatively small compared with those of gymnophionans in which the robustness of the joint is integral to a bur- rowing style that employs head and “neck” movements. We conclude that limited evidence fa- vors the interpretation that Eocaecilia mi- cropodia was probably not a burrower with the capabilities of Recent gymnophionans. The reduction in overall limb size and the apparent loss of two digits are features shared with various squamates that are ei- ther poor burrowers or sand swimmers or that shelter themselves in crevices and other cramped niches, such as dense veg- etation (Gans, 1975). If this analogy is ap- propriate, then we might envision the hab- itus of these Early Jurassic caecilians as presenting substrate parameters sufficient- ly dense or spatially constraining that limb reduction was selectively advantageous. Thus, E. micropodia plausibly represents a stage in the evolution of gymnophionan burrowing but was probably not fossorial per se. The correlate of such a supposition 356 is that the relatively small head size, spe- cializations of the masticatory apparatus, and other cranial features that characterize both E. micropodia and Recent gymno- phionans and which are commonly as- sumed to be associated with the fossorial habits of gymnophionans, in fact arose during a stage in caecilian evolution before their entry into the subterranean world. Zygokrotaphy and Stegokrotaphy. A persistently problematic issue in caecilian evolution is whether an open (zygokro- taphic) or closed (stegokrotaphic) skull roof in the temporal region is the primitive condition. Numerous studies, including those of DeBeer (1937), Goodrich (1930), Parsons and Williams (1963), and Wake Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 sible lines of evidence in support of his interpretation that stegokrotaphy is sec- ondary in caecilians. First, there are no temporal bones between the squamosal and parietal in living caecilians, whether they be zygo- or stegokrotaphic; he re- garded their loss as more likely the result of temporal fenestration (displaced by ex- panded adductor muscles) than by in situ fusion. Alternatively, we note that fusion of adjacent cranial bones is known to have occurred extensively in caecilians (Wake, 2003; Wake and Hanken, 1982). Second, Nussbaum (1983) noted that the arrangement of postorbital bones in stegokrotaphic caecilians differs from that in labyrinthodonts and considered this dif- and Hanken (1982), have supported the now prevalent view (reviewed by Wake, 2003) that one of the cranial specializa- tions in living gymnophionans is secondary closure of the temporal fenestra, a derived condition most likely related to fossoriality. Bemis et al. (1983), in a functional study of feeding in Dermophis mexicanus, took no position on whether stegokrotaphy is primitive or derived. They suggested that the reduction in skull size that accompa- nied the development of burrowing habits places a constraint on the size of the ad- ductor chamber and that the interhyoideus posterior/retroarticular process functions as a compensatory mechanism. Nussbaum (1983) observed that this novel mechanism is least well developed in rhinatrematids, which are relatively basal gymnophionans in which heads of the adductor mandibu- lae are exposed by a temporal fenestra. The temporal fenestrae of rhinatrematids and the inferred, secondarily closed fenes- trae of more derived groups would thus be a feature shared with frogs and salaman- ders. The discovery of stegokrotaphy in an Early Jurassic caecilian, which is patently primitive in many cranial and postcranial characters, revived the alternative possi- bility that zygokrotaphy could be derived (Jenkins and Walsh, 1993). We reconsider this problem in light of this study. Nussbaum (1983) presented four plau- ference as evidence that a solid temporal region has been secondarily reconstructed. However, the architecture of the cranial vault in Eocaecilia is now known to include such elements as a postparietal, jugal, quadratojugal, and tabular (or supratem- poral), all in comparable positions to those in the primitive tetrapod pattern. Third, Nussbaum (1977, 1983) pointed out that the skulls of rhinatrematids (which are zygokrotaphic) and ichthy- ophiids (in which zygokrotaphy is incipi- ently developed) are primitive in the great- est number of features. Furthermore, the highly derived skulls of most caeciliids are stegokrotaphic, an exception being Geo- trypetes (Wake, 2003). Alternatively, we note that the putatively primitive rhinatre- matid, Epicrionops, exhibits a number of derived cranial features; these include loss of the pre- and postfrontals, encirclement of the orbital margin by the maxillopala- tine, and specialization of the articulation between the squamosal and the lateral margin of the os basale. More primitive character states of these features are found in Ichthyophiidae and Uraeotyphlidae. Stegokrotaphy in Eocaecilia is associated with numerous primitive features of the skull and postcranial skeleton. Finally, Nussbaum (1983: 551) suggest- ed that the incomplete development of the interhyoideus jaw closure mechanism in ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll DM AMI(su) Figure 47. 357 Dorsal view of adductor muscles in (A) a salamander, Ambystoma maculatum; (B) a frog, Ascaphus truer; and (C) a caecilian, Epicrionops petersi. A and B from Carroll and Holmes (1980, fig. 18); C reconstructed from sections (LSUMZ 27324). rhinatrematids engendered a compensa- tory enlargement of “... the ancestral component [that] dominates by expansion of the mam |musculus adductor mandib- ulae] through a temporal fossa.” Alterna- tively, we raise the possibility that rhina- trematids (and, to a lesser extent, other caecilians with incipient zygokrotaphy) might exhibit a secondary reduction of the contribution of the interhyoideus posterior to jaw closure and a commensurate in- crease in the size of the adductors. Carroll and Holmes (1980) cited differ- ences between frogs and salamanders in the arrangement of the adductor muscles in the temporal fenestra as evidence that the fenestrae themselves were indepen- dently derived. This comparison may now be extended to include a caecilian. The dominant muscle at the level of the tem- poral fenestra in salamanders is the exter- nal adductor (Fig. 47A), which is associ- ated with loss of ossification between the maxilla and jaw suspension (Carroll and Holmes, 1980), whereas in frogs, the larg- est muscle is the posterior adductor (Fig. 398 47B). In Epicrionops, the dominant mus- cle is the external adductor (Fig. 47C). In each of these groups, different divisions of adductor mandibulae extend posteriorly over the back of the skull table and otic . capsule: the long head of the posterior ad- ductor in frogs, the superficial head of the internal adductor in salamanders, and the external adductor in Epicrionops. The di- vergent patterns of the occupation of the temporal opening in these three taxa might support the interpretation that the evolution of the openings occurred sepa- rately in the three groups. Alternatively, the relative size and positioning of the ad- ductor divisions could reflect divergences in muscle function that developed subse- quent to fenestration. At present there is no certain basis for making a definitive decision on stegokro- taphy versus zygokrotaphy in caecilian phylogeny. That the stegokrotaphic con- dition is primitive for amphibians is un- deniable—as is the fact that temporal fen- estration and other modifications of the skull roof are known to have developed in other groups (e.g., Paleozoic lysorophid amphibians). The lingering questions are whether fenestration in caecilians had al- ready been initiated in the common an- cestor of lissamphibians, or whether the condition was acquired subsequently in the ancestors of some living caecilians. Eocaecilia micropodia, which is primitive in many aspects of the cranial and post- cranial skeleton, supports the latter possi- bility. Eocaecilia as a Stage in Caecilian Phy- logeny. We concur with the phylogenetic hypotheses of Evans and Sigogneau-Rus- sell (2001, fig. 8) and Trueb and Cloutier (1991, fig. 9) that Eocaecilia micropodia is a basal caecilian that lies outside the crown group Gymnophiona. This Jurassic form nonetheless has significant implications for our understanding of a previously un- known stage in caecilian evolution and di- versity. Eocaecilia micropodia shares with extant gymnophionans such a substantial array of shared derived characters that Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 there can be little doubt that the evolu- tionary transition to gymnophionans had largely taken place prior to the Early Ju- rassic. The most distinctive gymnophionan features of Eocaecilia are: a sulcus along the anteroinferior margin of the orbital rim that could be interpreted as a tentac- ular sulcus; an os basale representing con- solidation of the supraoccipital, exoccipi- tals, basioccipital, basisphenoid, pleuro- sphenoid, and parasphenoid elements; an internal naris posterior to the premaxil- lary—maxillary suture and medial to the tooth rows on the vomer and palatine; en- larged nasal capsules that lie between the sphenethmoid and rostral margin of the skull; an olfactory eminence on the vomer; a lower jaw comprising a pseudodentary and pseudoangular that meet along an elongate, oblique suture; and an internal process of the pseudoangular. Internal processes of the pedicles, which serve to anchor tissue suspending the spinal cord, were first discovered in E. micropodia and subsequently found in some gymnophione taxa. On the basis of limited evidence, Eocaecilia appears to exhibit an elongation of the body comparable to that in primitive extant gymnophionans, but a detailed comparison cannot be made because of uncertainty over the precise number of vertebrae. In contrast to the foregoing gymno- phionan synapomorphies, Eocaecilia also presents primitive or transitional features, or both, that might be expected in forms representing a transition to a highly spe- cialized life style. The retention of jugal, quadratojugal, Ptabular, and postparietal bones is primitive. The presence of inter- centra and limbs is also primitive, but the reduction in the relative size of the limbs would appear to be transitional toward the gymnophionan condition. The discovery of an operculum in Eocaecilia micropodia is novel confirma- tory evidence that the opercular apparatus is a synapomorphy of Lissamphibia. The apparent absence of an operculum in ex- tant gymnophionans (at least as a separate ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll element) is best explained by the related loss of the shoulder girdle and muscular link between the girdle and operculum. The accessory ear ossicle described by Carroll and Gaskill (1978: 163, fig. 113) in Rhynchonkos and other microsaurs is con- sistently suspended above the stem of the stapes; in shape and position this ossicle appears to differ from the operculum of Eocaecilia, which is oval and, in one spec- imen at least, is preserved in close associ- ation with the fenestra ovalis. In contrast to the operculum, the presence of pedi- cellate teeth in a Jurassic caecilian is en- tirely expected. This dental specialization, once widely recognized as a lissamphibian synapomorphy, is now known to occur in two Paleozoic amphibians—Doleserpeton (Bolt, 1969, 1991) and Apateon (Schoch and Carroll, 2003)—which have been in- terpreted to be sister taxa of frogs and sal- amanders, respectively. With an admixture of primitive, transi- tional, and demonstrably gymnophionan features, Eocaecilia appears even more chimeric in possessing an array of special- izations that, on present knowledge, are autapomorphic. Many of these features present a perplexing challenge to function- al interpretation, for no structural compa- rability is to be found among living or po- tentially antecedent forms. The internal process of the lower jaw is very robust, and projects into the adductor chamber. The obliquely oriented, more or less planar jaw joint would appear to provide little stabil- ity, and is thus structurally and functionally unlike that known in any other caecilian. The stapes-quadrate is also unique. Although our knowledge of caecilian evolution and diversity now extends into the Early Jurassic, Eocaecilia micropodia does not provide sufficient evidence to se- curely recognize the origin of gymno- phionans among known Paleozoic amphib- ians, or even contribute substantially to the unresolved issue of liassamphibian mono- phyly. Carroll and Currie (1975) proposed microsaurs as possible caecilian ancestors, but their hypothesis has been criticized by 359 Bolt (1991), among others, and is not widely accepted. Nonetheless, the third author of this monograph persists in this viewpoint (Carroll, 2000: 1410), from which the remaining authors must demur. We therefore address the list of putative synapomorphies that have been proposed as uniting Eocaecilia and the tuditano- morph microsaur Rhynchonkos |Gonio- rhynchus|, many of which have already been critiqued by Bolt (1991). 1. “far anterior jaw articulation.” The relative position of the jaw articulation in the reconstructed skulls of Rhynchonkos (Carroll, 2000, fig. 2A, B; Carroll and Gas- kill, 1978, figs. 63, 64) is not certainly com- parably displaced as in Eocaecilia. The central issue is the position of the jaw joint relative to the occiput. To the degree that the central third of the occiput is protract- ed posteriorly relative to the two lateral thirds, the jaw joint will seem anteriorly displaced. The senior author examined the original material of Rhynchonkos stovalli (FMNH-UR 1039 and 1040), and both specimens are deformed and crushed (as shown in Carroll and Gaskill, 1978, figs. 63A, 64, top row), particularly in the oc- cipital region. Not surprisingly, Carroll and Gaskill’s two reconstructions (1978, figs. 63B, 64, bottom row) are slightly different. Thus, with the reliability of the reconstruc- tions uncertain, the relative position of the jaw joint in Rhynchonkos is moot. 2. “conspicuous retroarticular process.” Without attempting to define conspicuous as a quantifiable morphological term, we note that the retroarticular process of Rhynchonkos is short (estimated at 5-6% of jaw length) relative to that in Eocaecilia (17-18% of jaw length). Retroarticular processes occur widely throughout tem- nospondyls and are not uniquely shared by Rhynchonkos and Eocaecilia. In addition to Rhynchonkos, retroarticular processes in microsaurs are also found in ostodole- pids and some gymnarthrids (Carroll and Gaskill, 1978). 3. “loss of intertemporal and supratem- poral.” This character has previously been 360 criticized by Bolt (1991) who pointed out that absence of the intertemporal and su- pratemporal does not constitute a unique synapomorphy with any of the lissamphi- bian orders. “The intertemporal is absent in numerous fossil amphibian groups, and the absence of both intertemporal and su- pratemporal is shared at least with ad- vanced aistopods, lysorophoids and adelo- gyrinids ... (Bolt, 1991: 208—209). 4. “medial rows of teeth on the palate and lower jaw.” Bolt (1991: 209) observed that “Many fossil amphibians have a row of palatal teeth disposed like that of Rhyn- chonkos—this character has evolved inde- pendently a number of times, on almost any hypothesis of amphibian relation- ships.” As Bolt (1991: 210) further pointed out, Rhynchonkos is hardly unique in the possession of an internal row of mandib- ular teeth, which occurs in various tem- nospondyls (Jupp and Warren, 1986), in- cluding Colosteus (Hook, 1983). 5. “closely integrated posterior portion of the braincase.” Without attempting to define close integration, we note that Bolt (1991: 209) previously rejected Carroll and Currie’s (1975: 237) use of the character of “an extensive pleurosphenoid [that] joins the otico-occipital portion of the braincase with the sphenethmoid.” Lateral ossification of the braincase between the otic capsules and sphenethmoid is not unique to microsaurs. 6. “closure of the region anterior to the opening of the Vth nerve.” Enclosure of the exit of the trigeminal nerve by a bony pleurosphenoid is simply an expression of the degree of ossification of the braincase and is not uniquely shared by Rhynchon- kos and caecilians. 7. “holospondylous centra.” Holospon- dylous vertebrae are characteristic of lis- samphibians and lepospondyls generally and are not shared just with microsaurs. The general trend of vertebral evolution among Paleozoic amphibians was a reduc- tion of one of two circumnotochordal el- ements (intercentrum, pleurocentrum) in Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 favor of establishing the other as the dom- inant, or sole, element. 8. “double occipital condyle.” Paired occipital condyles occur in many derived temnospondyls and other lissamphibian groups and are not uniquely shared by Rhynchonkos (and microsaurs) and Eocae- cilia (and gymnophionans). 9. “elongation of the [vertebral] col- umn” and 10. “reduction of limbs.” Lengthening of the body through an increase in verte- bral number has occurred repeatedly and independently in amphibian evolution— salamanders, lepospondyls, and aistopods being several examples. Likewise, limb re- duction or loss has developed convergently in various aquatic or burrowing amphibi- ans. Both characters denote a general re- semblance, but neither is uniquely shared by Rhynchonkos (and microsaurs) and Eocaecilia (and gymnophionans). 11. “a more general similarity is small size.” Small size is indeed a general simi- larity, but diminutive taxa are found across many amphibian groups. Size is more like- ly a consequence of convergence and pro- vides no persuasive indicator of relation- ship between microsaurs and Eocaecilia. We thus conclude, with our third author dissenting, that on available evidence, Eocaecilia and gymnophionans have no demonstrable relationship to Microsauria. The authors do concur that a substantial morphological and temporal gap still in- tervenes in the identification of the origins of caecilians, and in fact we might not even be in possession, among all the Paleozoic taxa now known, of the fossil record of caecilian origins. Although Bolt and Lom- bard (1985: see also Lombard and Bolt, 1988) have produced the most credible ev- idence yet of an anuran relationship to temnospondyls, and Schoch and Carroll (2003) have identified developmental sim- ilarities between branchiosaurids and sal- amanders, the concept of Lissamphibia still remains at best a crown group taxon, without a securely rooted stem in the Pa- leozoic. ANATOMY OF EOCAECILIA MICROPODIA * Jenkins, Walsh, and Carroll ACKNOWLEDGMENTS Paleontological exploration of the Kay- enta Formation on lands of the Navajo Na- tion was made possible through permis- sion of the Navajo Tribal Council and the Coal Mine Mesa Chapter. The discovery of the microvertebrate quarry at Gold Springs is directly attributable to the pros- pecting and collecting talents of the late William R. Downs, with the assistance of James Kirkland. The senior author ac- knowledges with warm thanks W. W. Amaral, J. Attridge, the late W. R. Downs, H. E. Jenkins III, J. Kirkland, S. Madsen, D. K. McClearn, R. J. O'Hara, T. B. Rowe, C. R. Schaff, N. H. Shubin, K. K. Smith, and H.-D. Sues for their important partic- ipation in fieldwork. The Museum of Northern Arizona, Flagstaff, offered sub- stantial logistical support to the Kayenta project. The extensive collection of specimens of Eocaecilia micropodia was deftly prepared by William W. Amaral, Manager of the Preparation Laboratory in the Museum of Comparative Zoology. We thank E. Selig and R. Pinto for electron microscopy and A. H. Coleman, C. Kenaley, and A. Holmes for stereophotography. Drawings of cranial material were originally drafted by D. Dilkes. Most of the line drawings are the admirable work of the late P. Gas- kill (Redpath Museum, McGill) who also undertook preliminary reconstructions of the yskull? Figures I, 17, 29, 31, and 33 were skillfully executed by L. Laszlo Mesz- oly (MCZ, Harvard). J. P. O. Rosado of the Herpetology Department, Museum of Comparative Zoology, kindly made avail- able skeletal specimens of gymnophionans. R.L.C. warmly thanks Marvalee H. Wake for providing access to her collection of caecilian materials (skeletal, serial sec- tioned, and cleared and stained), and for sharing her knowledge and advice during his sabbatical visit to the University of Cal- ifornia, Berkeley. The curators of the Field Museum of Natural History and at Mich- igan State University were generous in 361 providing additional caecilians for exami- nation and dissection. R.L.C. also thanks E. Roman and M.-A. Lacey for processing various figures and other editorial assis- tance. We are the grateful beneficiaries of de- tailed reviews and thoughtful advice from major authorities in the field of herpetol- ogy. We warmly thank Professor Susan E. Evans (University College London) for providing an array of helpful guidance on many comparative anatomical points; Pro- fessor Ronald A. Nussbaum (University of Michigan) for insightful suggestions on in- terpreting Eocaecilia, particularly on the issue of fossorialism, and for permission to reproduce a number of his published il- lustrations; and Professor Marvalee H. Wake (University of California, Berkley) for generously sharing her knowledge of caecilian biology and the literature in this Hel daa eaGlelien isechatenl sto Davids © Blackburn for numerous spirited discus- sions of matters herpetological, and espe- cially for pointing out the existence of ver- tebral structures in certain snakes that are comparable to the internal processes of the pedicles in Eocaecilia micropodia. Our work was supported by grants from the National Science Foundation and National Geographic Society (F.A.J.Jr.); the Bedford Fund for Zoological Research, King’s Col- lege, Cambridge (D.M.W.); and the Nat- ural Sciences and Engineering Research Council of Canada (R.L.C.). APPENDIX: A List of Eocaecilia micropodia Specimens SPECIMENS ACCESSIONED IN THE MUSEUM OF COMPARATIVE ZOOLOGY MCZ 9011 MCZ 9015 anterior two-thirds of skull and jaws crushed skull and jaws; a second skull crushed and disarticulated, including tooth crowns disarticulated jaws, fragmentary verte- brae left lower jaw, vertebra crushed and disarticulated skull right lower jaw mounted on SEM disc, partial left lower jaw, and Pskull frag- ments crushed skull and jaw, vertebrae and MCZ 9095 MCZ 9152 MCZ 9156 MCZ 9158 MCZ 9163 362 various limb bones including complete humerus (mounted on SEM disc) and ulna incomplete mandibles and _postcranial bones; proximal humerus mounted on SEM disc partial skull roof with otic region, atlas, partial lower jaw, distal humerus, proxi- mal ulna block A: articulated vertebral series with shoulder girdle and forelimb elements, prepared and mounted on an SEM disc; block B: disarticulated vertebral series including caudals; blocks C, D: articu- lated vertebral series with crushed skull and jaws; isolated teeth mounted on SEM discs proximal humerus, partial jaws, atlas, vertebrae, scapulocoracoid, and crushed, disarticulated skull fragmentary vertebra atlas and other fragmentary vertebrae, partial jaw disarticulated jaws crushed, disarticulated skull with stapes- quadrate, jaw, and vertebrae crushed skull and mandible, humeri, scapulocoracoid, Pulna, and associated tibia and fibula with two tarsals scapulocoracoid, ulna, and proximal hu- merus mounted on SEM disc; vertebrae fragmentary jaw and vertebrae disarticulated skull and jaws, isolated pterygoid, vertebrae, and associated ra- dius and ulna mounted on SEM disc MCZ 9166 MCZ 9167 MCZ 9169 MCZ 9171 MCZ 9173 MCZ 9231 MCZ 9233 MCZ 9235 MCZ 9237 MCZ 9238 MCZ 9241 MCZ 9242 SPECIMENS ACCESSIONED IN THE MUSEUM OF NORTHERN ARIZONA MNA V8053 MNA V8054 left lower jaw and vertebra vertebrae, stapes-quadrate, fragmen- tary right jaw, and ulna; posterior part of jaw and stapes-quadrate prepared several series of vertebrae on different blocks: an articulated posterior dorsal series and an incomplete caudal series; tibia, tarsals, metatarsals incomplete skull and jaws, scapulocor- acoid on SEM disc, distal humerus, proximal radius and ulna, and frag- mentary pedal elements vertebrae and ribs partial lower jaw incomplete skull and jaws with asso- ciated postcranial bones disarticulated skull and jaws and ver- MNA V8055 MNA V8056 MNA V8057 MNA V8058 MNA V8059 MNA V8060 tebrae MNA V8062 articulated posterior dorsal vertebrae, Psacrals, and caudal vertebrae, crushed skull and jaws, hindlimb ele- ments including phalanges; right fe- mur mounted on SEM disc Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 MNA V8063 MNA V8064 otic capsule with braincase vertebrae, right scapulocoracoid (in medial view) partial skull roof with otic region, atlas, partial lower jaw, distal humerus, prox- imal ulna, and partial scapulocoracoid [holotype] skull and jaws with associ- ated atlas and postatlantal vertebrae articulated vertebral series mandible and distal humerus vertebrae disarticulated skull, mandible, and sta- pes fragmentary skull showing palate, side of os basale, and ethmoid portion of braincase; vertebrae fragmentary lower jaw and stapes vertebrae and limb elements, includ- ing fragmentary pedal bones incomplete skull and jaws originally associated with MNA V8059 MNA V8065 MNA V8066 MNA V8067 MNA V8068 MNA V8069 MNA V8070 MNA V0871 MNA V8072 MNA V8073 MNA V9346 LITERATURE CITED BADENHORST, A. 1978. The development and the phylogeny of the organ of Jacobson and the ten- tacular apparatus of Ichthyophis glutinosus (Lin- né). Annale Universiteit van Stellenbosch, Serie AD. Mle N—BX6), BEMIS, W. E., K. SCHWENK, AND M. H. WAKE. 1983. Morphology and function of the feeding appa- ratus in Dermophis mexicanus (Amphibia: Gym- nophiona). Zoological Journal of the Linnean So- ciety, 77: 75-96. BILLO, R., AND M. H. WAKE. 1987. Tentacle devel- opment in Dermophis mexicanus (Amphibia, Gymnophiona) with an hypothesis of tentacle or- igin. Journal of Morphology, 192: 101-111. BOLT, J. R. 1969. Lissamphibian origins: possible pro- tolissamphibian from the Lower Permian of Oklahoma. Science, 166: 888—891. . 1991. Lissamphibian origins, pp. 194-222. In H.-P. Schultze and L. Trueb (eds.), Origins of the Higher Groups of Tetrapods: Controversy and Consensus. Ithaca, New York: Cornell University Press. BOLT, J. R., AND R. E. LOMBARD. 1985. Evolution of the amphibian tympanic ear and the origin of frogs. Biological Journal of the Linnean Society, 24: 83-99. BOULENGER, G. A. 1909. A list of the freshwater fish- es, batrachians, and reptiles obtained by Mr. J. Stanley Gardiner’s expedition to the Indian Ocean. Transactions of the Linnean Society of London, Second Series, 12: 291—300. BRAMBLE, D. M. 1978. Origin of the mammalian feeding complex: models and mechanisms. Pa- leobiology, 4: 271-301. BRAND, D. J. 1956. The cranial morphology of Sco- lecomorphus uluguruensis (Barbour and Lover- ANATOMY OF EOCAECILIA MICROPODIA ° Jenkins, Walsh, and Carroll idge). Annals of the University of Stellenbosch, Section A, 32: 1—25. CARROLL, R. L. 1991. Batropetes from the Lower Permian of Europe—a microsaur, not a reptile. Journal of Vertebrate Paleontology, 11: 229-242. . 2000. Eocaecilia and the origin of caecilians, pp. 1402-1411. In H. Heatwole and R. L. Carroll (eds.), Amphibian Biology. Vol. 4. Palaeontology, The Evolutionary History of Amphibians. Chip- ping Norton, United Kingdom: Surrey Beatty & Sons. CARROLL, R. L., AND P. J. CURRIE. 1975. Microsaurs as possible apodan ancestors. Zoological Journal of the Linnean Society, 57: 229-247. CARROLL, R. L., AND P. GASKILL. 1978. The Order Microsauria. Memoirs of the American Philo- sophical Society, 126: 1-211. CARROLL, R. L., AND R. HOLMES. 1980. The skull and jaw musculature as guides to the ancestry of salamanders. Zoological Journal of the Linnean Society, 68: 1—40. CARROLL, R. L., A. KUNTZ, AND K. ALBRIGHT. 1999. Vertebral development and amphibian evolution. Evolution & Development, 1: 36—48. DE BEER, G. R. 1937. The Development of the Ver- tebrate Skull. Oxford, United Kingdom: Claren- don Press. xxiii + 552 pp. DE JAGER, E. F-. J. 1939a. Contributions to the cranial anatomy of the Gymnophiona. Further points re- garding the cranial anatomy of the genus Der- mophis. Anatomischer Anzeiger, 88: 193-222. . 1939b. The Gymnophione quadrate and its processes, with special reference to the processus ascendens in a juvenile Ichthyophis glutinosus. Anatomischer Anzeiger, 88: 223-232. DE VILLIERS, C. G. S. 1936. Some aspects of the amphibian suspensorium, with special reference to the paraquadrate and quadratomaxillary. An- atomischer Anzeiger, 81: 225-247. . 1938. A comparison of some cranial features of the East African Gymnophiones Boulengerula boulengeri, Tornier and Scolecomorphus ulugur- ensis Boulenger. Anatomischer Anzeiger, 86: 1— 26. DuBols, A. 2004. The higher nomenclature of recent amphibians. Alytes, International Journal of Ba- trachology, 22: 1-14. DUELLMAN, W. E., AND L. TRUEB. 1986. Biology of Amphibians. New York: McGraw-Hill Book Company. xvii + 670 pp. EDGEWORTH, F. H. 1935. The Cranial Muscles of Vertebrates. Cambridge: Cambridge University Press, United Kingdom. viii + 493 pp. ESTES, R., AND M. H. WAKE. 1972. The first fossil record of caecilian amphibians. Nature, 239: 220-231, EVANS, S. E., A. R. MILNER, AND C. WERNER. 1996. Sirenid salamanders and a gymnophionan am- phibian from the Cretaceous of the Sudan. Pa- laeontology, 39: 77-95. EVANS, S. E., AND D. SIGOGNEAU-RUSSELL. 2001. A 363 stem-group caecilian (Lissamphibia: Gymno- phiona) from the Lower Cretaceous of North Af- rica. Palaeontology, 44: 259-273. FRANCIS, E. T. B. 1934. The Anatomy of the Sala- mander. Oxford, United Kingdom: Clarendon Press. xxxi + 381 pp. FROST, D. R., T. GRANT, ie FAIVOVICH, R. H. BAIN, A. HAAS, C. F. B. HADDAD, R. O. DE SA, A. CHANNING, M. WILKINSON, S.C. DONNELLAN, C. J. RAXWORTHY, J. A. CAMPBELL, B. L. BLOT- TO, P. MOLER, R. C. DREWES, R. A NUSSBAUM, J. D. LYNCH, D. M. GREEN, AND W. C. WHEEL- ER. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History, 297: 1-370. GANS, C. 1975. Tetrapod limblessness: evolution and functional corollaries. American Zoologist, 15: 455-467. GOODRICH, E. S. 1930. Studies on the Structure and Development of Vertebrates. London: MacMil- lan and Company. xxx + 837 pp. GREER, A. E. 1991. Limb reduction in squamates: identification of the lineages and discussion of the trends. Journal of Herpetology, 25: 166-173. GREVEN, H. 1984. The dentition of Gegeneophis ra- maswamii Taylor, 1964 (Amphibia, Gymno- phiona), with comments on monocuspid teeth in the Amphibia. Zeitschrift ftir zoologische Syste- matik und Evolutionsforschung, 22: 342-348. HECHT, M. K., AND T. C. LADUKE. 1997. Limbless tetrapods, pp. 95-99. In R. F. Kay, R. H. Mad- den, R. L. Cifelli, and J. J. Flynn (eds.), Verte- brate Paleontology in the Neotropics, The Mio- cene Fauna of La Venta, Colombia. Washington, DC: Smithsonian Institution Press. HEDGES, S. B., R. A. NUSSBAUM, AND L. R. MAXSON. 1993. Caecilian phylogeny and biogeography in- ferred from mitochondrial DNA sequences of the 12S rRNA and 16S rRNA genes (Amphibia: Gymnophiona). Herpetological Monographs, 7: 64—76. HOoK, R. W. 1983. Colosteus scutellatus (Newberry): a primitive temnospondyl amphibian from the Middle Pennsylvanian of Linton, Ohio. American Museum of Natural History Novitates, 2770: 1— 4]. HRAOUI-BLOQUET, S., AND J.-M. EXBRAYAT. 1996. Les dents de Typhlonectes compressicaudus (Amphibia, Gymnophiona) au cours du dével- oppement. Annales des Sciences naturelles, Zoologie, Paris, 13° Série, 17: 11-23. JENKINS, F. A., JR., AND N. H. SHUBIN. 1998. Pros- dlirus bitis and the anuran caudopelvic mecha- nism. Journal of Vertebrate Paleontology, 18: 495-510. JENKINS, F. A., JR., AND D. M. WALSH. 1993. An Early Jurassic caecilian with limbs. Nature, 365: YAB—250. Jupp, R., AND A. A. WARREN. 1986. The mandibles of the Triassic temnospondyl amphibians. Al- cheringa, 10: 99-124. 364 LAKJER, T. 1926. Studien iiber die Trigeminus—Ver- sorgte Kaumuskulatur der Sauropsiden. Kopen- hagen: C. A. Reitzel. 154 pp. LAWSON, R. 1963. The anatomy of Hypogeophis ros- tratus Cuvier (Amphibia: Apoda or Gymno- phiona). Part I, the skin and skeleton. Proceed- ings of the University of Durham Philosophical Society, Series A (Science), 13: 254-273. LEssA, E. P., AND M. H. WAKE. 1992. Morphometric analysis of the skull of Dermophis mexicanus (Amphibia: Gymnophiona). Zoological Journal of the Linnean Society, 106: 1—15. LOMBARD, R. E., AND J. R. BOLT. 1988. Evolution of the stapes in Paleozoic tetrapods: conservative and radical hypotheses, pp. 37-67. In B. Fritzsch, M. J. Ryan, W. Wilczynski, T. E. Heth- erington, and W. Walkowiak (eds.), The Evolu- tion of the Amphibian Auditory System. New York: John Wiley & Sons. Marcus, H. 1935. Zur Entstehung der Stapesplatte bei Hypogeophis. Anatomischer Anzeiger, 80: 142-146. MARCUS, H., E. STIMMELMAYR, AND G. PORSCH. 1935. Die Ossifikation des Hypogeophisschiadels. Beitrag zur Kenntnis der Gymnophionen XXV. Morphologisches Jahrbuch, 76: 375—420. McGowan, G. J. 1998. The development and func- tion of the atlanto-axial joint in albanerpetontid amphibians. Journal of Herpetology, 32: 116— OR MILNER, A. R. 1993. The Paleozoic relatives of lis- samphibians. Herpetological Monographs, 7: 8— DH, MONATH, T. 1965. The opercular apparatus of sala- manders. Journal of Morphology, 116: 149-170. MULLER, H., O. V. OOMMEN, AND P. BARTSCH. 2005. Skeletal development of the direct-developing caecilian Gegeneophis ramaswamii (Amphibia: Gymnophiona: Caeciliidae). Zoomorphology, 124: 171-188. NUSSBAUM, R. A. 1977. Rhinatrematidae: a new fam- ily of caecilians (Amphibia: Gymnophiona). Oc- casional Papers of the Museum of Zoology, Uni- versity of Michigan, 682: 1-30. . 1979. The taxonomic status of the caecilian genus Uraeotyphlus Peters. Occasional Papers of the Museum of Zoology, University of Michigan, 687: 1-20. 1983. The evolution of a unique dual jaw- closing mechanism in caecilians (Amphibia: Gymnophiona) and its bearing on caecilian an- cestry. Journal of Zoology, London, 199: 545— 554. 1984. Amphibians of the Seychelles, pp. 379-415. In D. R. Stoddart (ed.), Biogeography and Ecology of the Seychelles Islands. The Hague: Dr. W. Junk Publishers. . 1985. Systematics of caecilians (Amphibia: Gymnophiona) of the Family Scolecomorphidae. Occasional Papers of the Museum of Zoology, University of Michigan, 713: 1-49. Bulletin Museum of Comparative Zoology, Vol. 158, No. 6 NUSSBAUM, R. A., AND B. G. NAYLOR. 1982. Variation in the trunk musculature of caecilians (Amphibia: Gymnophiona). Journal of Zoology, London, 198: 383-398. NUSSBAUM, R. A., AND M. WILKINSON. 1989. On the classification and phylogeny of caecilians (Am- phibia: Gymnophiona), a critical review. Herpe- tological Monographs, 3: 1-42. PARKER, H. W. 1941. The caecilians of the Sey- chelles. The Annals and Magazine of Natural History, series 11, 12: 1-17. PARSONS, T. S., AND E. E. WILLIAMS. 1963. The re- lationships of the modern Amphibia: a re-exam- ination. Quarterly Review of Biology, 38: 26-53. PETER, K. 1898. Die Entwicklung und funktionelle Gestaltung des Schiidels von Ichthyophis gluti- nosus. Morphologisches Jahrbuch, 25: 555-628. RAGE, J.-C. 1986. Le plus ancien Amphibien apode (Gymnophiona) fossile. Remarques sur la répar- tition et Vhistoire paléobiogéographique des Gymnophiones. Centre Researche Académie des Sciences, Paris, 302(serie II, 16): 1033-1036. REIss, J. 1996. Palatal metamorphosis in basal cae- cilians (Amphibia: Gymnophiona) as evidence for lissamphibian monophyly. Journal of Herpetolo- gy, 30: 27-39. RIEPPEL, O. 1980. The phylogeny of anguinomorph lizards. Denkschriften der Schweizerischen Na- turforschenden Gesellschaft, 94: 1—86. ROMER, A. S. 1956. Osteology of the Reptiles. Chi- cago: University of Chicago Press. xxi + 772 pp. SAN MAURO, D., J. GOWER, O. V. OOMMEN, M. WIL- KINSON, AND R. ZARDOYA. 2004. Phylogeny of caecilian amphibians (Gymnophiona) based on complete mitochondrial genomes and nuclear RAG1. Molecular Phylogenetics and Evolution, 33: 413-427. SARASIN, P., AND F. SARASIN. 1887-1890. Ergebnisse naturwissenschaftlicher forschungen auf Ceylon in den Jahren 1884-1886. Vol. 2. Zur Entwick- lungsgeschichte und Anatomie der ceylonesisch- en Blindwiihle, Ichthyophis glutinosus, L. Wies- baden, Germany: C. W. Kreidel’s Verlag. 252 pp. SAVE-SODERBERGH, G. 1945. Notes on the trigeminal musculature in non-mammalian tetrapods. Nova Acta Regiae Societatis Scientiarum Upsaliensis, Series 4, 13: 1—59. SCHMIDT, A., AND M. H. WAKE. 1990. Olfactory and vomeronasal systems of caecilians (Amphibia: Gymnophiona). Journal of Morphology, 205: 955-268. SCHOCH, R. R., AND R. L. CARROLL. 2003. Ontoge- netic evidence for the Paleozoic ancestry of sal- amanders. Evolution and Development, 5: 314— 324. SHUBIN, N. H., AND F. A. JENKINS, JR. 1995. An Early Jurassic jumping frog. Nature, 377: 49-52. TAYLOR, E. H. 1968. The Caecilians of the World, A Taxonomic Review. Lawrence: The University of Kansas Press. viii + 845 pp. . 1969. Skulls of Gymnophiona and their sig- ANATOMY OF EOCAECILIA MICROPODIA ¢ Jenkins, Walsh, and Carroll nificance in the taxonomy of the group. The Uni- versity of Kansas Science Bulletin, 48: 585-687. TRUEB, L. 1993. Patterns of cranial diversity among the Lissamphibia, pp. 255-343. In J. Hanken and B. K. Hall (eds.), The Skull. Vol. 2. Patterns of Structural and Systematic Diversity. Chicago: Chicago University Press. TRUEB, L., AND R. CLOUTIER. 1991. A phylogenetic investigation of the inter- and intrarelationships of the Lissamphibia (Amphibia: Temnospondyli), pp. 223-313. In H.-P. Schultze and L. Trueb (eds.), Origins of the Higher Groups of Tetra- pods: Controversy and Consensus. Ithaca, New York: Cornell University Press. VISSER, M. H. C. 1963. The cranial morphology of Ichthyophis glutinosus (Linné) and Ichthyophis monochrous (Bleeker). Annale Universiteit van Stellenbosch, Serie A, 38: 67—102. WAKE, D. B. 1970. Aspects of vertebral evolution in the modern Amphiba. Forma et Functio, 3: 33— 60. WAKE, M. H. 1976. The development and replace- ment of teeth in viviparous caecilians. Journal of Morphology, 148: 33-64. . 1980a. Morphometrics of the skeleton of Dermophis mexicanus (Amphibia: Gymno- phiona). Part I. The vertebrae, with comparisons to other species. Journal of Morphology, 165: 117-130. . 1980b. Fetal tooth development and adult replacement in Dermophis mexicanus (Amphib- ia: Gymnophiona): fields versus clones. Journal of Morphology, 166: 203-216. . 1987a. A new genus of African caecilian (Amphibia: Gymnophiona). Journal of Herpetol- ogy, 21: 6-15. 1987b. Haemal arches in amphibians: a problem in homology and phylogeny. American Zoologist, 27: 33A. . 1992. “Regressive” evolution of special sen- sory organs in caecilians (Amphbia: Gymno- phiona): opportunity for morphological innova- tion. Zoologische Jahrbiicher Abteilung fiir An- atomie und Ontogenie der Tiere, 122: 325-329. . 2003. The osteology of caecilians, pp. 1809— 1876. In H. Heatwole and M. Davies (eds.), Am- 365 phibian Biology. Vol. 5. Osteology. Chipping Norton, United Kingdom: Surrey Beatty & Sons. WAKE, M. H., AND J. HANKEN. 1982. Development of the skull of Dermophis mexicanus (Amphibia: Gymnophiona), with comments on skull kinesis and amphibian relationships. Journal of Mor- phology, 173: 203-223. WAKE, M. H., AND G. Z. WuRST. 1979. Tooth crown morphology in caecilians (Amphibia: Gymno- phiona). Journal of Morphology, 159: 331-341. WERNER, C. 1994. Der erste Nachweis von Gymno- phionen (Amphibia) in der Kreide (Wadi-Milk- Formation, Sudan). Neues Jahrbuch fiir Geolo- gie und Paliontologie, Monatshefte, 1994: 633— 640. WEVER, E. G. 1973. The ear and hearing in the frog, Rana pipiens. Journal of Morphology, 141: 461— A477. . 1975. The caecilian ear. Journal of Experi- mental Zoology, 191: 63-71. WEVER, E. G., AND C. GANS. 1976. The caecilian ear: further observations. Proceedings of the National Academy of Sciences U.S.A., 73: 3744-3746. WIEDERSHEIM, R. 1879. Die Anatomie der Gymno- phionen. Jena: Gustav Fischer Verlag. vii + 100 Witkin SON, M. 1991. Adult tooth crown morphology in the Typhlonectidae (Amphibia: Gymno- phiona): a reinterpretation of variation and its significance. Zeitschrift fiir zoologische Syste- matik und Evolutionsforschung, 29: 304-311. . 1992. The phylogenetic position of the Rhin- atrematidae (Amphibia: Gymnophiona): evi- dence from the larval lateral line system. Am- phibia-Reptilia, 13: 74-79. . 1996. The heart and aortic arches of rhina- trematid caecilians (Amphibia: Gymnophiona). Zoological Journal of the Linnean Society, 118: 135-150. WILKINSON, M., AND R. A. NUSSBAUM. 1996. On the phylogenetic position of the Uraeotyphlidae (Amphibia: Gymnophiona). Copeia, 1996: 550— DOr . 1997. Comparative morphology and evolu- tion of the lungless caecilian Atretochoana eiselti (Taylor) (Amphibia: Gymnophiona: Typhlonecti- dae). Biological Journal of the Linnean Society, 62: 39-109. wie eons Ai eg? ( cnt vthteeataalinet ornare | inotK at fick emilee ee geet a : oT eer, a ai) , 4 Poh ti , is : wren : tila Lar cals 5 ht, Yycon val pee. HM ‘mae as ahaaNe tt ie as ey iaiwe Wi Naan brat tas ote: “alll YI iy) bee: DUCA ae ih a any, bert 7 Ti) hii 4 ok wey Aanettp ps Ady es yee me tT hited . oe 2h A vai. ealtbepard, : = yal Pol ee ee datales ei Lb = lhe et Tl Law, sy i eae Ae calli is eee ; hy 7 J ma. yt a —— eas visa ee TO OPE ay ai pe ays ish 5 ior h Ate ay ba) ‘i ie Ww dnp Lubitsaad ha ake ; Oa D paar ee wy is 1 kale 9 of bs , Me nail isa s| tah, Len co aes th ‘1 Bais tecky t sip ea ‘ = i re f ae a * p D ~ a S a ; med a ( i mie . i - - = ee Y \ ae # ~ a 1 . " To “ ta é “ : - : ; ; “, 2s ae | - a ~~ act ~ + ae ior A ; Lie ae “e ‘f ae 6. 4’ t RS a Ru lletin fe) ; THE - | Lool ogy =. The Cerion (Mollusca: Gastropoda: Pulmonata: Cerionidae) Taxa of Charles Johnson Maynard and Their Type Specimens _ M. G. HARASEWYCH, ADAM J. BALDINGER, YOLANDA VILLACAMPA, AND PAUL GREENHALL HARVARD UNIVERSITY _ VOLUME 158, NUMBER 7 __ CAMBRIDGE, MASSACHUSETTS, U.S.A. _ 7 September 2007 (US ISSN 0027-4100) PUBLICATIONS ISSUED OR DISTRIBUTED BY THE MUSEUM OF COMPARATIVE ZOOLOGY HARVARD UNIVERSITY Breviora 1952— BULLETIN 1863— Memoirs 1865-1938 Jounsonia, Department of Mollusks, 1941-1974 OCCASIONAL PAPERS ON MOLLUSKS, 1945— SPECIAL PUBLICATIONS. 1. Whittington, H. B., and W. D. I. Rolfe (eds.), 1963 Phylogeny and Evolution of Crustacea. 192 pp. 2. Turner, R. D., 1966. A Survey and illustrated Catalogue of the Tere- dinidea (Mollusca: Bivalvia). 265 pp. 3. Sprinkle, J., 1973. Morphology and Evolution of Blastozoan Echino- derms. 284 pp. 4, Eaton, R. J., 1974. A Flora of Concord from Thoreau’s Time to the Present Day. 236 pp. 5. Rhodin, A. G. J., and K. Miyata (eds.), 1983. Advances in Herpetology and Evolutionary Biology: Essays in Honor of Emest E. Williams. 729 pp. 6. Angelo, R., 1990. Concord Area Trees and Shrubs. 118 pp. Other Publications. Bigelow, H. B., and W. C. Schroeder, 1953. Fishes of the Gulf of Maine. Reprinted 1964. Brues, C.T., A. L. Melander, and F. M. Carpenter, 1954. Classification of Insects. (Bulletin of the M. C. Z., Vol. 108.) Reprinted 1971. Creighton, W. S., 1950. The Ants of North America. Reprinted 1966. Lyman, C. P., and A. R. Dawe (eds.), 1960. Proceedings of the First In- ternational Symposium on Natural Mammalian Hibernation. (Bulletin of the M. C. Z., Vol. 124.) Orinthological Gazetteers of the Neotropics (1975-). Peter’s Check-list of Birds of the World, vols. 1-16. ei ror 2 of the New England Zoological Club 1899-1947. (Complete sets only.) Price list and catalog of MCZ publications may be obtained from Publica- tions Office, Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 02138, U.S.A. This publication has been printed on acid-free permanent paper stock. © The President and Fellows of Harvard College 2007. = eae ee ee = ——~ a THE CERION (MOLLUSCA: GASTROPODA: PULMONATA: CERIONIDAE) TAXA OF CHARLES JOHNSON MAYNARD AND THEIR TYPE SPECIMENS M. G. HARASEWYCH,' ADAM J. BALDINGER,? YOLANDA VILLACAMPA,' AND PAUL GREENHALL' ABSTRACT. Charles Johnson Maynard (1845-1929) was a self-educated naturalist, teacher, and dealer in natural history specimens and materials who con- ducted extensive field work throughout Florida, the Bahamas, and the Cayman Islands. He published prolifically on the fauna, flora, and anthropology of these areas. His publications included descriptions of 248 of the 587 validly proposed species-level taxa within Cerionidae, a family of terrestrial gastropods endemic to the islands of the tropical western Atlan- tic. After his death, his collection of Cerionidae was purchased jointly by the Museum of Comparative Zo- ology (MCZ) and the United States National Muse- um, with the presumed primary types remaining at the MCZ and the remainder of the collection divided between these two museums and a few other insti- tutions. In this work, we provide 1) a revised collation of Maynard's publications dealing with Cerionidae, 2) a chronological listing of species-level taxa proposed in these works, 3) a determination of the number and status of name-bearing type specimens for each taxon within the context of the most recent edition of the International Code of Zoological Nomenclature, and 4) when necessary, the selection of a lectotype (for 185 taxa) or the designation of a neotype (for eight taxa) from among specimens in these museums to restrict the name-bearing type for each taxon to a single specimen, which is illustrated. Type material could not be located for three of the 248 species-level taxa proposed by Maynard. In these instances, neo- types are proposed to make these three taxa objective junior synonyms of other taxa. Strict application of the International Code of Zoological Nomenclature contradicts the authorship, publication date, and type status of name-bearing types for a significant number of Maynard’s taxa that had been previously reported in the literature. ' Department of Invertebrate Zoology, MRC 163, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560. * Curatorial Associate, Malacology, Museum of Comparative Zoology, Harvard University, 26 Oxford Street, Cambridge, Massachusetts 02138-2902. Bull. Mus. Comp. Zool., 158(7): 367-528, September, 2007 INTRODUCTION The family Cerionidae comprises a group of terrestrial pulmonate gastropods that are endemic to the tropical western Atlantic, ranging from southern Florida throughout the Bahamas, Greater Antilles, Cayman Islands, western Virgin Islands, and the Dutch Antilles, but are absent in Jamaica, the Lesser Antilles, and coastal Central and South America. These snails are halophilic, occurring on terrestrial veg- etation, generally within 100 m of the shore, but occasionally 1 km or more from the sea, presumably in areas where salt spray can reach them from one or more directions (Clench, 1957: 121). Cerionid snails generally occur in dense but patchy populations, often with 10,000 or more in- dividuals. They can be conspicuously abundant when living on open vegetation and less so when living in leaf litter. Al- though individual populations tend to be fairly uniform in the size and morphology of their shells, size, shape, color, and in- ternal as well as external sculpture of the shell vary enormously among populations. Such variation is evident not only through- out the range of Cerionidae, but even among neighboring populations separated by less than 100 m. This lavish yet geo- graphically circumscribed diversity has led to an extensive body of literature dealing with cerionids. Much of the early work was primarily taxonomic and focused on pars- ing the various phenotypes among roughly 600 nominal species and nearly two dozen genus-level taxa. More recent research, 367 368 particularly the work of Mayr, Gould, Woodruff, and Goodfriend, as well as their students and collaborators (see Woodruff, 1978, for an overview), addressed more basic biological questions: among them, the origins as well as the geographic and temporal stability of morphological and ge- netic diversity and the dynamics of hybrid zones and biogeographic patterns, both Holocene and Recent. Yet, as noted by Woodruff (1978: 224), “the evolutionary importance of these remarkable animals has been buried under an all but impen- etrable taxonomic thicket.” Much of the breadth and a substantial amount of the impenetrability of this “tax- onomic thicket” can be attributed to the work of Charles Johnson Maynard (1845— 1929), who published 248 of the 587 val- idly proposed species-level taxa (excluding obvious typographical errors), and seven of the 23 genus-level taxa within Cerionidae. Maynard was a self-educated naturalist and teacher who conducted extensive field work throughout Florida, the Bahamas, and the Cayman Islands (Abele, 2002; French, 1930; Johnson, 1930; Townsend, 1930; Turner, 1957). His bibliography con- sists of 277 publications (Batchelder, 1951), the overwhelming majority on birds, but includes works on butterflies, sponges, land snails, and many other areas of natural history and anthropology. Batch- elder (1951: 227) pointed out that many of Maynard’s publications, including all of his work on Cerionidae, were “not only the work of his active mind but of his own hands. He took pride in setting the type, making wood-cuts, drawing lithographs, and doing the press work.” Maynard’s publications were printed in limited numbers and most are very rare. For example, the prospectus on the inside front covers of each of the 12 parts of his Contributions to the History of the Cer- ionidae with Descriptions of Many New Species and Notes on Evolution in Birds and Plants (1919-26; hereafter Contribu- tions to the History of the Cerionidae) states, “The edition will consist of 150 cop- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 ies.” Maynard’s writings on the Cerionidae were confined to eight publications, sey- eral of which appeared in multiple parts (Table 1). Six of these publications contain descriptions of new taxa, and a seventh contains a nomen nudum. One is an ad- vertisement. Although Maynard’s publications are rare, specimens of the majority of his taxa are not. In his undated sales catalog, May- nard (1924b?) mentioned having 200,000 specimens of Cerion, whereas the supple- ment to this catalog noted that an addi- tional 44,000 specimens, representing 30 species, were collected in 1924. Nearly all of Maynard’s species descriptions mention the number of specimens examined. In his description of Strophia extensa, as an ex- treme example, Maynard (1924c: 2-3) commented that 23,000 specimens were collected, of which only about 3,000 were living. Shortly after Maynard’s death (October 15, 1929), his daughter offered his large collection of Cerion for sale to the Muse- um of Collective Zoology (MCZ). Turner (1957: 151) reported that, “As the collec- tion was so large, and the specimens so numerous, it was decided to buy it jointly with the United States National Museum [USNM, now the National Museum of Natural History, Smithsonian Institution] for $500. The collection was divided equally between the two institutions, with the holotypes being retained at the MCZ. A few duplicate sets of the larger series of Maynard’s types were sold to various insti- tutions, so that much of Maynard’s type material is now well distributed.” Records at the MCZ (Mollusk Department Acces- sion file 766) indicate that 475 lots of “mostly cerions from the Maynard collec- tion, including all of his species” were pur- chased by Dr. T. Barbour for the Museum and were received in the Mollusk Depart- ment in October 1931. These were as- signed catalog numbers ranging from MCZ 76001 to 76475. Specimens with these catalog numbers were in Maynard's collection at the time of his death. Com- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. menting on the condition of the collection, Turner (1957: 151) wrote “the collection, at the time it was sold, was in rather sad shape, for the cabinet drawers had warped and once pulled open could not be closed. The top drawers of the cabinets became a parade ground for pigeons, and their drop- pings cemented the shells together in a solid mass! After transferring the collec- tion in its original drawers to the MCZ, each lot was sorted, cleaned and properly labeled—a truly gigantic task.” Because Maynard was a dealer and col- lector who for decades sold or exchanged specimens to support his research and ex- peditions, an unquantified amount of type material was also distributed during his lifetime. A series of cerions was donated by Maynard to the MCZ in 1895 and was catalogued with the numbers MCZ 10242 to 10360. Correspondence and invoices in the Smithsonian Archives indicate that Dr. Paul Bartsch purchased for the National Museum of Natural History specimens representing 121 of Maynard’s species Prom chem neype localities: imei 923 (USNM 359387—359518), and an addition- al 57 specimens in 1925. Another series of Maynard's cerion specimens, originally in the collections of the Boston Society of Natural History, was transferred to the MCZ in February 1930 and bear the cat- alog numbers MCZ 39713-39719. Al- though many of Maynard’s taxa are rep- resented in museum collections, particu- larly in the eastern United States, the type status of many, if not most, of these spec- imens was undetermined or incorrectly la- beled at the time of this writing. Although Turner (1957: 151) reported that the holotypes of Maynard’s taxa were retained at the MCZ, it is far from clear how many and which of Maynard’s taxa are based on holotypes and which on syntypes. If application of the International Code of Zoological Nomenclature (ICZN) is re- stricted to the descriptions of individual species, one conclusion can be reached. If other portions of the original publication are considered, this conclusion could be 369 contradicted. By way of example, the orig- inal description of Strophia scripta May- nard, 1896 (p. 4), provided a detailed de- scription that lists “size of type, 1.30 by A7” and identifies the illustration of a sin- gle specimen as “fig. 3, front, fig. 4, side view of type.” According to Article 73.1.1 (ICZN, 1999: 79), this specimen would be the holotype. However, earlier in the same publication, Maynard (1896: 1) states, “The first portion of the present install- ment of this monograph is based upon specimens belonging to the MCZ which I have been permitted to examine.... I have returned the types figured to the Mu- seum, but have retained a set of co-types in my own collection.” This indicates that the figured specimen returned to the MCZ and the specimen retained by Maynard for his own collection are syntypes because they can be considered expressly designat- ed as such in the original publication (Ar- ticle 73.2.1, ICZN, 1999: 81). Most of the taxon descriptions pub- lished in Maynard’s first work on the Cer- ionidae, the Monograph of the Genus Stro- phia, a Group of Tropical and Sub-Tropi- cal Land Shells that appeared in seven is- sues of the Contributions to Science (1889-96), were exemplary for the period. Each included a detailed description and illustrations (usually with two views), pro- vided dimensions of the “type” or “types,” as well as of the largest and smallest spec- imens, and had sections with observations on variation within the taxon and on its distribution and habitats. In contrast, taxonomic descriptions in most subsequent publications (e.g., May- nard, 1913a,b,c, 1919-26 [Contributions to the History of the Cerionidae], 1924b?,c; Maynard and Clapp, 1914, 1915) were very brief and written in tele- graphic style. Some were later supple- mented with more detailed descriptions of the shell and anatomy (Maynard, 1924a, 1925, 1926). Each intended taxonomic de- scription typically included a measurement and gave the number of specimens exam- ined but did not contain any version of the 370 Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 TABLE 1. PUBLICATIONS BY CHARLES JOHNSON MAYNARD CONTAINING ARTICLES ON CERIONIDAE. Maynard, C. J. 1889-96. Monograph of the Genus Strophia, a group of tropical and sub-tropical land shells. Contributions to Science, 1(1, April 1889): 1-29, pls. 1-2; 1(2, July 1889): 68-79, pl. 7; 1(3, October 1889): 125-135, pl. 16; 1(4, January 1890 [dated “January, 1889” on page 1]): 188-197; 2(1 [July 1893 on cover, November 1893 on page 1]): 1-48; 2(2 [November 1894 on cover]): 49-104; 2(3 [December 1894 on cover]): 105-152; 2(4 [December 1894 on cover]): 153-200; 3(1, March 1896): 1-40, pls. 1-7. (Privately published, Newtonville, Massachusetts) Maynard, C. J. 1913. Descriptions of some species of the family Cerionidae. Records of Walks and Talks with Nature. Appendix to Volume 5: 177—200 (illustrations on paper covers; pp. 177—182 are dated January 1913; pp. 183-200, February 1913. Footnote on p. 186: “Pages 183-186 were issued Feb. 1, 1913.” Footnote on p. 199: “Pages 187-199 were issued Feb. 9, 1913.” Text is complete on p. 199. Page 200 contains figures of three species reproduced from Contributions to Science). (Privately published, West Newton, Massa- chusetts) Maynard, C. J. 1914. A species of land shell of the family Cerionidae redescribed. Records of Walks and Talks with Nature. Appendix to Volume 6: 177-179 [illustrations on paper covers]. [Dated May 1914 at top of each page, but footnote on p. 179 states “Actual date of publication of this sheet, Aug 18, 1914.”] (Privately published, Newtonville, Massachusetts) Maynard, C. J., and N. A. Clapp. 1914-15. Descriptions of new Strophias. Records of Walks and Talks with Nature. Appendix to Volume 6: 180 [August 18, 1914], 179 bis, 180 bis, 181, 182 [February 8, 1915]. [Page 180 has same date as Maynard, 1914; footnote on p. 182 states, “Actual date of publication, Feb. 8, 1915.”] (Privately published, Newtonville, Massachusetts) Maynard, C. J. 1919. A New Book on Evolution. Contributions to the History of Cerionidae. 4 unnumbered pp. (Privately published, Newtonville, Massachusetts) Note: This is a four-page advertisement for the Contributions to the History of the Cerionidae [see below] that provided a brief description of the intended work, which was to contain “upward of 400 pages, and over 20 plates of shells, birds, etc” and be limited to 150 copies. Versions with either plain or hand-colored plates were advertised. This advertisement was bound within the covers of Number 1 of Contributions to the History of the Cerionidae in the ANSP copy, with a Library Stamp dated October 22, 1919. Batchelder (1951: 256) noted, “Issued first perhaps in 1921, reprinted, most of it verbatim, but with some changes, in 1924.” Maynard, C. J. 1919-26. Contributions to the history of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature. Appendix to Volume 10. 1[October 16, 1919]: 1-24, pls. 5, 6 [illustrations on paper covers]; 2[ November 6, 1919]: 25-52, pls. 7, 8 [illustrations on paper covers]; 3[ December 31, 1919]: 53-68, pls. 9, 10 [illustrations of paper covers]; 4|February 29, 1920]: 69-80, pls. 10*, 11 [illustrations on paper covers]; 5[April 8, 1920]: 81-92, pls. 12, 13 [illustrations on paper covers]; 6[July 10, 1920]: 93-110, pls. 14-19 [illustrations on paper covers]; 7|September 30, 1920]: 111-126, pls. 1-4, 20-25 [illustrations on paper covers]; 8|May 2, 1921]: 127-138, pls. 26-33 [illustrations on paper covers]; 9[July 15, 1921]: 139-154, pls. 34-37 [illustrations on paper covers]; 10[October 25, 1924]: 155-170, pls. 38-39 [illustrations on paper covers]; 11|July 18, 1925]: 171— 194, pls. 40-41 [illustrations on paper covers]; 12[March (10 printed, overwritten in ink to 24) 1926]: 195— 218, pls. 42-43 [illustrations on paper covers]. (Privately published, Newtonville, Massachusetts) Note: Collations and dates of individual parts of this work are based on copies in the libraries of the Smithsonian Institution and the Academy of Natural Sciences of Philadelphia, which were bound as issued. The dates and the apportionment of pages, especially of plates to the individual numbers, differ considerably from the collation given in Batchelder (1951: 254-256), which, like the copies in the libraries of the MCZ and Mr. Richard Goldberg, have the plates in numerical order. The collation of this work in the library of the Field Museum of Natural history differs in that 1) pages 49-52 are bound with number 3 instead of number 2; 2) number 3 is dated (in pencil) December 4 instead of December 31; 3) plates 40 and 41 are bound between the back cover of number 10 and the front cover of number 11, rather than within the covers of number 11. Publication dates are based on: “Number 1 Published Oct. 6 1919” printed on front cover. * “Number 2 Published Nov. 6 1919” printed on front cover. * “Number 3 Published Dec. 31 1919” printed on front cover but numbers in bold handwritten in ink. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 371 TABLE 1. CONTINUED. “Number 4 Date of issue on last page.” inclusive. issued Feb. 29, 1920.” “Number 5 Date of issue on last page.” inclusive. issued April 8, 1920.” “Number 6 Date of issue on last page.” 110, inclusive. issued July 10, 1920.” “Number 7 Date of issue on last page.” 126, inclusive. issued Sept. 30, 1920.” “Number 8 Date of issue on last page.” 138, inclusive. Issued May 2, 1921.” “Number 9 Date of issue on last page.” 154, inclusive. issued July 15, 1921.” “Number 10 Date of issue on last page.” 170, inclusive. issued Oct. 25, 1924.” “Number 11 Date of issue on last page.” 194, inclusive. Issued July 18, 1925.” “Number 12 Date of issue on last page.” printed on front cover. Footnote on p. 80, “No. 4. Pages 69-80, printed on front cover. Footnote on p. 92, “No. 5. Pages 81-92, printed on front cover. Footnote on p. 110, “No. 6. Pages 93— printed on front cover. Footnote on p. 126, “No. 7. Pages 111— printed on front cover. Footnote on p. 138, “No. 8. Pages 127— printed on front cover. Footnote on p. 154, “No. 9. Pages 139— printed on front cover. Footnote on p. 170, “No. 10. Pages 155— printed on front cover. Footnote on p- 194, “No. 11. Pages 171— printed on front cover. Footnote on p. 218, “No. 12. Pages 195— 218, inclusive. Issued March 10 [overwritten in ink to 24], 1926.” Maynard, C. J. [Undated, referred to herein as 1924b?]. Catalogue of Specimens of the Family Cerionidae for Sale by Charles J. Maynard Containing Many New Species. Paper cover and 8 unnumbered pages. (Privately published, Newtonville, Massachusetts) Note: Batchelder (1951: 256) listed 1920 as the date for this work, whereas Clench (1957: 169) cited a six- page catalog with identical title as published in 1924. Maynard began advertising specimens and publications in The Nautilus, beginning in July 1920 and continued until October 1929 (volume 34, number 1, through volume 43, number 2), the advertisements appearing at the top of the inside front cover. Beginning July 1921 and continuing until Maynard’s death (October 15, 1929), these advertisements offered a “Catalog of Cerion for sale now ready, over 100 species.” The inside front paper cover of the Catalog in the Smithsonian Institution’s library lists “Appendix to Records Vol. X, 1921,” and several taxa described in 1921 are offered for sale in this catalogue. Also listed [page 3] is Strophiops albicostata, the description of which appears in the supplement to this catalog (dated December 10, 1924). These data suggest that the catalog first ap- peared on or before July 1921 but that it might have undergone periodic revision and updating, the last appearing on or before December 10, 1924. The copy in the Academy of Natural Sciences Library is hand dated “1921,” but the Library Stamp reads January 31, 1929. Maynard, C. J. 1924 [10 December]. Supplement to Sale Catalogue of Cerionidae with Descriptions of New Species Collected in the Bahama Islands in Summer of 1924. pp. 1-6. [Footnote on p. 6 “Issued December 10, 1924] (Privately published, Newtonville, Massachusetts) * Asterisk denotes a date different from that cited by Batchelder (1951: 254). word “type” in the description or in figure captions for those taxa that were illustrat- ed. International Code of Zoological No- menclature Article 73.2 (1999: 81) states that, “for a nominal species-group taxon established before 2000 all specimens of the type series are automatically syntypes if neither a holotype nor a lectotype has been fixed.” Thus, most of Maynard’s taxa published after 1896, including the taxa of Maynard and Clapp, can be considered as being typified by multiple syntypes (23,000 syntypes in the case of Strophia extensa Maynard, 1924; see above). In the preamble to the species descrip- tions published in Contributions to the History of the Cerionidae under the head- ing “Typical Specimens,” Maynard wrote, It has long been the custom with naturalists when describing new species to select some individual specimen that as nearly as possible combines all of the characters which are possessed by the species of which it is a member. This specimen is called a type. As will be seen, however, by the previous re- marks and the matter which follows, that groups occur in all species composed of members which have similar characters. From some one of these groups the type must be selected. This is usually selected in the species described in this volume, from the group that has the largest number of spec- imens. This group I have called typical. (Maynard, 1919b: 43-44) 372 This indicates that Maynard understood the concept of type specimen and defined it explicitly. Maynard based his taxonomic descriptions on characteristics of popula- tions, then selected as “type” an individual that “as nearly as possible combines all of the characters possessed by the species of which it is a member.” This differs con- ceptually from the definition of Holotype, which is “the single specimen upon which a new nominal species group taxon is based” (Article 73.1, ICZN, 1999: 79). Maynard’s statement that “this [the type] is usually selected in the species de- scribed in this volume” not only excludes species not described in that volume (i.e., Maynard, 1913a, b, c, 1924b, c; Maynard and Clapp, 1914, 1915) but does not spec- ify for which of the species in Maynard (1919-26) such specimens were selected. Even with no mention of the term “type” in the original descriptions, many of the specimens now in the collections of the MCZ that are derived from the May- nard collection are labeled “Holotype” and are accompanied by a slip of paper, appar- ently in Maynard’s handwriting, that in- cludes the term “type.” These labels also contain the plate and figure numbers for the illustrations of the taxon in Maynard (1919-26) in the same handwriting and the same ink, suggesting that the slips were written at the time of preparation of the plates for this work. Such labels can be construed as evidence that Maynard con- sidered these specimens to be the types of their respective taxa, some of which had been published years or decades earlier than their illustrations. However, it is not possible to interpret them as holotype des- ignations within the Code, which states that “the holotype of a new nominal spe- cies-group taxon can only be fixed in the original publication by the original author” (Article 73.1.3, ICZN, 1999: 79). In the vast majority of cases, these “type” labels accompany a single speci- men, but in over a dozen cases, they ac- company a lot containing multiple speci- mens. Given the condition of the collec- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 tion at the time of Maynard’s death (see above; Turner, 1957: 151), it is entirely possible that specimens or labels moved within the drawers. When such specimens, or one of such specimens, can be reason- ably construed as approximating the pub- lished measurements or, for taxa that are illustrated, the original illustration, even though neither are associated with the term “type” within the publication, these specimens are designated as the lectotypes for their respective species-level taxa here- in. Many researchers have conjectured that perhaps only 10-20% of the species-level taxa within Cerionidae will eventually be found to represent valid species (e.g., Clenech, 1957: 121.) Turmer 1954-152: Woodruff, 1978: 235). The determination of whether the nomina proposed by May- nard represent valid species, subspecies, demes, or distinctive combinations of al- leles is beyond the scope of this work. Rather, our objective here is to enumerate chronologically and in order of publication all of the species-level taxa within the fam- ily Cerionidae that were validly proposed by Maynard, either individually or in col- laboration with Clapp; to determine the dates on which each of these taxa became available; and to locate and to restrict the name-bearing type for each taxon to a sin- gle specimen, when necessary by the se- lection of a lectotype or designation of a neotype. As noted in ICZN Article 61.1 (1999: 63), “The fixation of a name-bear- ing type of a nominal taxon provides the objective standard of reference for the ap- plication of the name it bears.” Providing objective standards for all of Maynard's taxa is a necessary first step toward prun- ing the “all but impenetrable taxonomic thicket” that has been a hallmark of, and an impediment to, studies of the Cerioni- dae. MATERIALS AND METHODS The publications listed in Table 1 were surveyed for descriptions of species-level taxa. The species-level taxa are listed chro- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. nologically according to the date the taxa became available and in order of publica- tion to the extent possible. For cases in which a taxon name appeared only on a plate caption but not in the text, it is listed at the end of the taxa published in the same issue or on the same date. This list was compared with similar compilations in Batchelder (1951) and in Clench’s (1957) catalog of the Cerionidae. We do not in- clude misspellings of Maynard’s taxa by subsequent authors (e.g., Batchelder, 1951; Clench, 1957; Pilsbry, 1901-02) or Maynard's uses or misspellings of other au- thors’ taxa. Maynard’s not infrequent mis- spellings of his own taxa, likely introduced when he set the type, are reconciled, usu- ally with the accepted spelling of the taxon (see however, S. nivia, taxa 26 and 72), and are also included in the alphabetical index of Maynard's taxa. Each validly introduced taxon is num- bered sequentially and is followed by one or more citations, including the valid taxon description (in bold), which for several taxa differs from the intended taxon de- scription. Also included are details of spec- imens associated with the term “type” within the intended taxon description, published measurements, and the number of specimens examined. The number and the status of name- bearing type specimens for each taxon are determined within the context of the most recent edition of the ICZN (1999). This determination is based on information in the original description, other portions of the same publication, subsequent publi- cations, and works of other authors, as well as on specimens traceable to Maynard's collection now in the collections of the MCZ. Several of Maynard’s taxa were not represented in his collection when it reached the Museum, as confirmed by an examination of the MCZ catalog. For these taxa, efforts were made to locate speci- mens traceable to Maynard in the collec- tions of the MCZ and the National Mu- seum of Natural History, Smithsonian In- stitution (USNM), Academy of Natural 373 Sciences of Philadelphia (ANSP), the Field Museum of Natural History (FMNH), Florida Museum of Natural History (FLMNH), and Museum of Bio- logical Diversity, The Ohio State Univer- sity (OSU). When necessary, a lectotype is selected or a neotype is designated from among specimens in these museums to re- strict the name-bearing type for each taxon to a single specimen. This is followed by measurements of the single name-bearing type, the type locality for the taxon, and remarks pertaining to the taxon. Images of the apertural and right lateral views are also included. Comparisons with published measure- ments and figures have been made when selecting or verifying primary type speci- mens (Recommendation 74B, ICZN, 1999: 83). It should be noted from the on- set that some variation is to be expected in measurements of length and far more in measurements of width, as these de- pend not only on the accuracy of the in- strument used (ruler vs. digital caliper), but also on how the measurements were made (length measured parallel to the axis of coiling vs. maximum length; width of the cylindrical portion of the shell vs. max- imum width including flaring aperture). Even the best of Maynard’s illustrations, while generally adequate for distinguishing taxa, are rarely precise enough to differ- entiate among the often numerous speci- mens in the type series. Turner (1957: 140) quoted from Maynard’s unpublished autobiography, “When I was a child I took a great liking to the art of wood engraving. Not because I had ever seen any of the work done, for I had not, but with the idea in mind that I should like the work, I con- sulted a neighbor who was a wood engrav- er. After questioning me as to my ability to make drawings and after not finding me at all proficient in that art, he said em- phatically that I would not succeed as an engraver, so I abandoned that project.” Turner went on to write “It is interesting to see that in later years he made all of his own illustrations and wood engravings, as 374 well as hand coloring many of his plates. These illustrations are remarkably well done considering his complete lack of training.” Contributions to the History of the Cer- ionidae, published in 12 numbers between 1919 and 1926, presents a variety of no- menclatural complexities for several of the taxa proposed within its pages. Most in- volve the consequences of unintended no- menclatural acts upon the authorship of the taxa, the dates on which names be- came available, and the determination of their primary types. Strict interpretation within the context of the International Code of Zoological Nomenclature (ICZN, 1999) clearly contradicts the authors’ in- tentions in most of these cases. In the preface to this work, Maynard (1919a: 1) stated that he had “written Parts One and Three and the descriptions of the animals and their habitats in Part Two, but that the descriptions of the shells in this part have been written with the collabo- ration of Mrs. William F. [Nellie A.] Clapp, and the names of new species should be credited to us jointly.” Part One is explicitly delimited as spanning pages 3— 112, but also included figures 1—89, and was published with plates 5-10, 10*, and 11-19. Part Two is labeled as beginning on page 113 and would appear, on the basis of content and style, to end on page 153. It contains one map and no figures and includes plates 1-4 and 20-37 but also re- fers to illustrations on plates 38—48. Plates 3843 were published later, together with the text portions of Part Three, whereas plates 44-48 were never published. What appear, on the basis of content and style, to be continuations of Part One span pages 154-163 and 178-180. A more detailed “Description of Species” that begins on page 164 and, except for pp. 178-180, continues until the last page (p. 218), is presumed to be Part Three, although it is nowhere identified as such. It would ap- pear that Maynard intended to publish ad- ditional text and plates. In the introduction to Part Two, May- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 nard (1920d: 113) again confirmed that new species “should be credited to May- nard and Clapp.” However, it is clear that Maynard retained authorship of the work in the preface (Maynard, 1919a: 1). He subsequently (Maynard, 1924c: 1) referred to this work as “my Contributions to the History of the Cerionidae began in 1919.” Thus, the authorship of taxa proposed in Part Two should be cited as Maynard and Clapp in Maynard, 1920-25. Taxa inadver- tently introduced in other portions of this work are attributed to Maynard, unless there was an indication to the contrary (e.g., taxon name followed by “M. & C.” in the figure caption). The first six issues of Contributions to the History of the Cerionidae and a por- tion of the seventh (Maynard, 1919a— 1920d) comprise Part One and contain ex- tensive notes on Cerionidae, including comments on the age and ecology of living and fossil Cerions, postulated laws govern- ing the evolution of the family, and a list of characteristics deemed useful for differ- entiating groups or species. Unfortunately, Maynard used the names of several of the taxa intended to be described in Part Two of this work to illustrate particular mor- phological features. These names often ap- peared in figure captions or in conjunction with references to illustrations that would be published later, but still before the in- tended taxon descriptions. In some cases, this resulted in the creation of nomina nuda, in others, to inadvertent but nomen- claturally valid species descriptions. Al- though these inadvertent species descrip- tions rarely alter the intended concept of the taxon, they do affect the date, author- ship, and type designations for these taxa. Supplemental information for each of Maynard's Cerion taxa, including the text of the original description and images of the primary types in color, can be found on the Cerion website (http://nvertebrates.si.edu/ cerion). ACKNOWLEDGMENTS We are grateful to Paul Callomon at the Academy of Natural Sciences of Philadel- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. phia and to Jochen Gerber of the Field Museum of Natural History for loan of specimens from the collections in their care and for research into the collations of Maynard's publications housed in the li- braries of their institutions. We thank Richard Goldberg for information on the collation of his copy of Maynard’s Contri- butions to the History of the Cerionidae. Richard E. Petit, Philippe Bouchet, and Stephen Cairns kindly provided helpful discussions and opinions on the interpre- tation of the International Code of Zoo- logical Nomenclature in numerous in- stances. We are grateful to the editor and especially to the referees, Richard E. Petit, Alan Kabat, and Riidiger Bieler, for their considerable efforts and for the many im- provements that they have suggested. This research was supported in part by NSF Grant EAR 0106936 to Goodfriend, Gould, and Harasewych. THE CERION TAXA OF CHARLES JOHNSON MAYNARD All species-level taxa within the family Cerionidae that were validly proposed by Maynard, either individually or in collab- oration with Clapp, are listed chronologi- cally according to the date the taxa became available and in order of publication to the extent possible. In cases in which a taxon name appeared only on a plate caption but not in the text, it is listed at the end of the taxa published in the same issue or on the same date. An alphabetical listing of the taxa (as well as their misspellings) is pro- vided in the index at the end of this work. Each validly introduced taxon is num- bered sequentially and is followed by one or more citations, including the valid taxon description (in bold), which for several taxa differs from the intended taxon de- scription. Also included are details of spec- imens associated with the term “type” within the intended taxon description, published measurements, and the number of specimens examined. These details are provided as they appeared in the original publications and typographical errors have 3795 not been corrected. Type localities are also provided as originally published, with sup- plemental information included in square brackets. Taxon 1. Strophia pannosa Maynard, 1889a [April]: 10-11, pl. 1, figs. 2, 13, pl. 2, figs. 1, 1B, 1C, 1D. Exam- ined 400 specimens. Original descrip- tion lists sizes of two “types” given as 1.27 by .57 and 1.23 by .50 [inches; 32.3 by 14.5 and 31.2 by 12.7 mm]. Caption to plate 2, figure 1 states “Strophia pan- nosa, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining 398 specimens are, according to Article) 7274-6 (IGZN, 11999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- mens labeled as such are mislabeled. Clench (1964: 368, pl. 63, fig. 1) designat- ed and figured as lectotype MCZ 246343, noting, “In several cases Maynard failed to select a holotype, so lectotypes have been selected here which agree as closely as can be determined with his descriptions and measurements” (Clench, 1964: 370). This lectotype approximates the measurements of the smaller of the two syntypes. The re- maining syntype became the paralecto- type. Lectotype Measurements. Length See) mm, diameter (excluding lip) 13.1 mm; ap- erture height (including lip) 11.6 mm, ap- erture width (including lip and peristome) 10.1 mm. Type Locality. West end of Little Cay- man, Cayman Islands. Remarks. Clench (1964: 368) regarded this taxon as a valid species and provided an extensive synonymy. Taxon 2. Strophia levigata Maynard, 1889a [April]: 12-13, pl. 2, figs. 2, 2B. Examined 150 specimens. Original description lists sizes of two “types” giv- en as 1.25 by .52 and 1.15 by .48 [inch- 376 es; 31.8 by 13.2 and 29.2 by 12.2 mm]. Caption to plate 2, figure 2, “Strophia levigata, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining 148 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- mens labeled as such are mislabeled. Clench (1964: 368, pl. 63, fig. 2) designat- ed and figured as lectotype MCZ 247021, noting, “In several cases Maynard failed to select a holotype, so lectotypes have been selected here which agree as closely as can be determined with his descriptions and measurements” (Clench, 1964: 370). This specimen is within 5% of the length of ei- ther of the syntypes but closer to the larger syntype in width. The remaining syntype became the paralectotype. Lectotype Measurements. Length 30.9 mm, diameter (excluding lip) 14.3 mm; ap- erture height (including lip) 13.1 mm, ap- erture width (including lip and peristome) AS rraniae Type Locality. West end of Little Cay- man, Cayman Islands. Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Taxon 3. Strophia intermedia Maynard, 1889a [April]: 13-15, pl. 2, figs. 3, 3B. Examined 3,000 specimens. Origi- nal description lists sizes of two “types” given as .90 by .40 and .88 by .48 [inch- es; 22.9 by 10.2 and 22.4 by 12.2 mm]. Caption to plate 2, figure 3, “Strophia intermedia, front view of type shell.” Type Material. The original description Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining 2,998 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- mens labeled as such are mislabeled. Clench (1964: 368, pl. 63, fig. 3) listed and figured as Holotype MCZ 76067. This specimen, which is closer to the larger syn- type in length and to the smaller syntype in width, cannot be the holotype, as two syntypes had been listed. Clench’s listing cannot be accepted as a lectotype desig- nation. Article 74.5 (ICZN, 1999: 83) states, “When the original work reveals that the taxon had been based on more than one specimen, a subsequent use of the term ‘holotype’ does not constitute a valid lectotype designation unless the au- thor, when wrongly using that term, ex- plicitly indicated that he or she was se- lecting from the type series that particular specimen to serve as the name-bearing type.” This specimen (MCZ 76067) is here selected as the lectotype. The remaining syntype becomes the paralectotype. Lectotype Measurements. Length 24.8 mm, diameter (excluding lip) 12.0 mm; ap- erture height (including lip) 10.1 mm, ap- erture width (including lip and peristome) 8.8 mm. Type Locality. Coast along the south side of Little Cayman, Cayman Islands. Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Taxon 4. Strophia acuta Maynard, 1889a [April]: 15-17, pl. 2, figs. 4, 4B. Ex- amined 500 specimens. Sizes of two “types” given as .94 by .40 and .94 by .38 [inches; 23.9 by 10.2 and 23.9 by 9.7 — Figures 1-12. Taxon 1. Strophia pannosa Maynard, 1889. Lectotype MCZ 246343. West end of Little Cayman, Cayman Islands. Taxon 2. Strophia levigata Maynard, 1889. Lectotype MCZ 247021. West end of Little Cayman, Cayman Islands. Taxon 3. Strophia intermedia Maynard, 1889. Lectotype MCZ 76067. South side of Little Cayman, Cayman Islands. Taxon 4. Strophia acuta Maynard, 1889. Lectotype MCZ 247022. Field inland from the south coast of Little Cayman, Cayman Islands. Taxon 5. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 377 Strophia festiva Maynard, 1889. Lectotype MCZ 247023. Fields along path, near center of Little Cayman, Cayman Islands. Taxon 6. Strophia picta Maynard, 1889. Lectotype MCZ 247019. West end of Little Cayman, Cayman Islands. Taxon 7. Strophia lineota Maynard, 1889. Lectotype MCZ 246346. South side of Little Cayman, Cayman Islands. Taxon 8. Strophia copia Maynard, 1889. Lectotype MCZ 76065. West end of Cayman Brac, Cayman Islands. Taxon 9. Strophia parva Maynard, 1889. Lectotype MCZ 246423. West end of Cayman Brac, Cayman Islands. Taxon 10. Strophia glaber Maynard, 1889. Lectotype MCZ 246344. West end of Cayman Brac, Cayman Islands. Taxon 11. Strophia nana Maynard, 1889. Lectotype MCZ 246737. West end of Little Cayman, Cayman Islands, near intersection of two paths. Taxon 12. Strophia nuda Maynard, 1889. Lectotype MCZ 356977. Near Clarence Harbor, Long Island, Bahamas. 378 mm]. Caption to plate 2, figure 4, “Stro- phia acuta, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining 498 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- mens labeled as such are mislabeled. Clench (1964: 368, pl. 63, fig. 4) designat- ed and figured as lectotype MCZ 247022, noting, “In several cases Maynard failed to select a holotype, so lectotypes have been selected here which agree as closely as can be determined with his descriptions and measurements’ (Clench, 1964: 370). This specimen approximates the narrower of the two syntypes in width. The remaining syntype became the paralectotype. Lectotype Measurements. Length WD, mm, diameter (excluding lip) 9.2 mm; ap- erture height (including lip) 8.6 mm, ap- erture width (including lip and peristome) To) ianien, Type Locality. Field inland from the south coast of Little Cayman, Cayman Is- lands. Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Taxon 5. Strophia festiva Maynard, 1889a [April]: 17-18, pl. 2, figs. 5, 5B, C. Examined 22 specimens. Sizes of two “types” given as 1.20 by .50 and 1.18 by .48 [inches; 30.5 by 12.7 and 30.0 by 12.2 mm]. Caption to plate 2, figure 5, “Strophia festiva, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining 20 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 mens labeled as such are mislabeled. Clench (1964: 368, pl. 63, fig. 5) designat- ed and figured as lectotype MCZ 247023, noting, “In several cases Maynard failed to select a holotype, so lectotypes have been selected here which agree as closely as can be determined with his descriptions and measurements’ (Clench, 1964: 370). This specimen approximates the smaller of the two syntypes in length. The remaining syn- type became the paralectotype. Lectotype Measurements. Length 29.7 mim, diameter (excluding lip) 12.3 mm; ap- erture height (including lip) 12.1 mm, ap- erture width (including lip and peristome) 10.2 mm. Type Locality. Fields along path, near center of Little Cayman, Cayman Islands. Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Taxon 6. Strophia picta Maynard, 1889a [April]: 18-20, pl. 2, figs. 6, 6B. Ex- amined 75 specimens. Sizes of two “types” given as .96 by .40 and .86 by .39 [inches; 24.4 by 10.2 and 21.8 by 9.9 mm]. Caption to plate 2, figure 6, “Stro- phia picta, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining TS specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- mens labeled as such are mislabeled. Clench (1964: 368, pl. 63, fig. 6) designat- ed and figured as lectotype MCZ 247019, noting, “In several cases Maynard failed to select a holotype, so lectotypes have been selected here which agree as closely as can be determined with his descriptions and measurements’ (Clench, 1964: 370). This specimen approximates the larger of the two syntypes in length. The remaining syn- type became the paralectotype. Lectotype Measurements. Length 24.4 _ THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. mm, diameter (excluding lip) 9.9 mm; ap- erture height (including lip) 9.3 mm, ap- erture width (including lip and peristome) 7.8 mm. Type Locality. West end of Little Cay- man, Cayman Islands. Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Taxon 7. Strophia lineota Maynard, 1889a [April]: 20-22, pl. 2, figs. 7, 7B. Examined 1,000 specimens. Sizes of two “types” given as 1.04 by .40 and .98 by .42 [inches; 26.4 by 10.2 and 24.9 by 10.7 mm]. Caption to plate 2, figure 7, “Strophia lineota, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining 998 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- mens labeled as such are mislabeled. Clench (1964: 368, pl. 63, fig. 7) designat- ed and figured as lectotype MCZ 246346, noting, “In several cases Maynard failed to select a holotype, so lectotypes have been selected here which agree as closely as can be determined with his descriptions and measurements’ (Clench, 1964: 370). This specimen approximates the larger of the two syntypes in length. The remaining syn- type became the paralectotype. Lectotype Measurements. Length 26.1 mim, diameter (excluding lip) 11.3 mm; ap- erture height (including lip) 11.0 mm, ap- erture width (including lip and peristome) 9.0 mm. Type Locality. Coconut grove, south side of Little Cayman, Cayman Islands. Maynard (1889a: 21) noted that this taxon was also found near the boat landing on Cayman Brac, but was probably intro- duced there from Little Cayman. Clench (1957: 151, 1964: 368) listed both locali- 379 ties. The label accompanying the lectotype gives the locality as “S.E. side of Little Cayman Id., Cayman Ids.” Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Taxon 8. Strophia copia Maynard, 1889a [April]: 22-24, pl. 1, figs. 1, 3, 7-12, pl. 2, figs. 8, 8B. Examined 10,000 specimens. Sizes of two “types” given as .90 by .40 and .95 by .40 [inches; 22.9 by 10.2 and 24.1 by 10.2 mm]. Caption to plate 2, figure 8, “Strophia copia, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining 9,998 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- mens labeled as such are mislabeled. Clench (1964: 368, pl. 63, fig. 8) designat- ed and figured as lectotype MCZ 76065, noting, “In several cases Maynard failed to select a holotype, so lectotypes have been selected here which agree as closely as can be determined with his descriptions and measurements” (Clench, 1964: 370). This specimen approximates the smaller of the two syntypes in length. The remaining syn- type became the paralectotype. Lectotype Measurements. Length DDD mm, diameter (excluding lip) 9.6 mm; ap- erture height (including lip) 9.0 mm, ap- erture width (including lip and peristome) WeOnmann, Type Locality. West end of Cayman Brac, Cayman Islands. Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Hummelinck (1980: 59) recognized it as a distinct species. Taxon 9. Strophia parva Maynard, 1889a [April]: 24-25, pl. 2, figs. 9, 9B. Ex- amined about 1,000 specimens. Sizes of two “types” given as .60 by .30 and .63 380 by .32 [inches; 15.2 by 7.6 and 16.0 by 8.1 mm]. Caption to plate 2, figure 9, “Strophia parva, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining 998 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- mens labeled as such are mislabeled. Clench (1964: 368, pl. 63, fig. 9) designat- ed and figured as lectotype MCZ 246423, noting, “In several cases Maynard failed to select a holotype, so lectotypes have been selected here which agree as closely as can be determined with his descriptions and measurements’ (Clench, 1964: 370). This specimen approximates the larger of the two syntypes in length. The remaining syn- type became the paralectotype. Lectotype Measurements. Length 19.1 mm, diameter (excluding lip) 9.0 mm; ap- erture height (including lip) 7.6 mm, ap- erture width (including lip and peristome) 6.6 mm. Type Locality. West end of Cayman Brac, Cayman Islands. Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Taxon 10. Strophia glaber Maynard, 1889a [April]: 25-26, pl. 2, figs. 10, 10B. Examined 16 specimens. Sizes of two “types” given as .62 by .33 and .65 by .30 [inches; 15.7 by 8.4 and 16.5 by 7.6 mm]. Caption to plate 2, figure 10, “Strophia glaber, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining 14 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 there can be no paratypes, and any speci- mens labeled as such are mislabeled. Clench (1964: 368, pl. 63, fig. 10) desig- nated and figured as lectotype MCZ 246344, noting, “In several cases Maynard failed to select a holotype, so lectotypes have been selected here which agree as closely as can be determined with his de- scriptions and measurements” (Clench, 1964: 370). This specimen approximates the larger of the two syntypes in length. The remaining syntype, MCZ 39715, be- came the paralectotype. Lectotype Measurements. Length 17.6 mm, diameter (excluding lip) 7.7 mm; ap- erture height (including lip) 6.3 mm, ap- erture width (including lip and peristome) 5.9 mm. Type Locality. West end of Cayman Brac, Cayman Islands. Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Taxon 11. Strophia nana Maynard, 1889a [April]: 27-29, pl. 2, figs. 11A, B, C, D. Examined 2,000 specimens. Sizes of two “types” given as .62 by .25 and .60 by .18 [inches; 15.7 by 6.4 and 15.2 by 4.6 mm]. Caption to plate 2, fig- ure 11, “Strophia nana, A, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. The re- maining 1,998 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- mens labeled as such are mislabeled (e.g., Hummelinck, 1980, pl. 11, figs. g—k). Clench (1964: 373, pl. 63, fig. 14) desig- nated and figured as lectotype MCZ 246737, noting, “In several cases Maynard failed to select a holotype, so lectotypes have been selected here which agree as closely as can be determined with his de- scriptions and measurements” (Clench, _ THE CERION TAXA OF CHARLES JOHNSON MAYNARD °* Harasewych et al. 1964: 370). This specimen approximates the smaller of the two syntypes in length. The remaining syntype became the para- lectotype. Lectotype Measurements. Length 14.4 mm, diameter (excluding lip) 5.4 mm; ap- erture height (including lip) 5.2 mm, ap- erture width (including lip and peristome) Al mm. Type Locality. West end of Little Cay- man, Cayman Islands, near intersection of two paths. Maynard (1889a: 28) noted “the Dwarf Strophias [Strophia nana] occur in a space which is only five or six yards wide by twenty long... I consider that this spe- cies has the most restricted range of any animal with which I am acquainted.” Remarks. Clench (1964: 373) regarded this taxon to be a valid species. Taxon 12. Strophia nuda Maynard, 1889a [April]: 29, pl. 2, figs. 12, 12B. Examined three specimens. Sizes of two “types” given as 1.00 by .39 and .82 by .32 [inches; 25.4 by 9.9 and 20.8 by 8.1 mm]; other specimen .89 by .38 [inches; 22.6 by 9.6 mm]. Caption to plate 2, fig- ure 12, “Strophia nuda, front view of type shell.” Type Material. The original description provided measurements for two “type” specimens, which are thus syntypes. One of these syntypes was illustrated. Of the three lots at the MCZ that were possible types [MCZ 39713, MCZ 76267, MCZ, 87985], one, MCZ 76267, contained six specimens, the other two lots have one specimen each. Maynard (1889a: 29) wrote, “... I have only three of this sin- gular Strophia ...” Nevertheless, MCZ 76267 contains Maynard's label with the term “type,” and a later MCZ label that is annotated on the back, “Only three spec- imens in original series given by May- nard—He has apparently added to them!!!” The length and width of each of the specimens in each of these three lots were measured. None of the specimens in any 381 of the lots matched Maynard’s published measurements exactly, although the largest specimen [24.9 by 9.7 mm] came closest to the measurements of the larger of the two syntypes listed by Maynard. When the ratios of length to width were computed and compared with ratios of published type measurements, this same specimen matched to within 1%. None of the remaining specimens came as close to matching either the linear mea- surements or the length to width ratio of the other type, although the single speci- men in MCZ 39713 came within 5%. A specimen in MCZ 76267 sectioned to ex- pose the columellar axis comes closest to the measurements given for the “other specimen.” The largest specimen (24.9 mm) from MCZ 76267 was therefore se- lected as the lectotype and recatalogued as MCZ 356977. The specimen in MCZ 39713 becomes the paralectotype. The re- maining specimen is, according to Article 72.4.6 (ICZN, 1999: 77), expressly exclud- ed from the type series. Therefore, there can be no paratypes and any specimens la- beled as such are mislabeled. Lectotype Measurements. Length 24.9 mm, diameter (excluding lip) 9.7 mm; ap- erture height (including lip) 8.7 mm, ap- erture width (including lip and peristome) (oi nansie Type Locality. Near Clarence Harbor, Long Island, Bahamas. Taxon 13. Strophia ianthina Maynard, 1889a [April], pl. 2, figs. 13, 13B. Cap- tion to plate 2, figure 13, “Strophia ian- thina, front view of type shell”; Maynard, 1889b | July]: 69-70. Text noted 300 spec- imens examined. Sizes of two “types” giv- en as 1.15 by .37 and 1.00 by .36 [inches; 29.2 by 9.4 and 25.4 by 9.1 mm]. Nomenclatural Remarks. The _ illustra- tions (Maynard, 1889a, pl. 2, figs. 13, 13B) and caption were published prior to the text. The taxon thus dates from the pub- lication of the name in the caption accom- panying the illustrations, not from the sub- 382 sequently published text (Article 12.2.7, ICZN, 1999: 17). Type Material. The specimen represent- ed in the figure was identified as the “type shell” and is therefore the holotype (Arti- cle 73.1.4, ICZN, 1999: 80). Although there was no mention of other specimens in the plate caption, the remaining 299 specimens enumerated in the text portion published 3 months later were likely avail- able to Maynard at the time the plate was prepared and are thus part of the type se- ries [Article 72.4.1.1, ICZN, 1999: 76], serving as paratypes of Strophia ianthina. There was no specimen labeled “Holo- type” at the MCZ. Lot number MCZ 39714 contains one specimen labeled “paratype,” and MCZ 10317 contained two adult specimens, also labeled “paratypes.” The larger of these two specimens closely approximates the figured specimen in size and proportions. This is here considered to be the figured specimen and therefore the holotype. Holotype Measurements. Length 27.6 mim, diameter (excluding lip) 10.1 mm; ap- erture height (including lip) 9.8 mm, ap- erture width (including lip and peristome) 8.1 mm. Type Locality. Between southern shore of Inagua | Bahamas] and the extensive salt lake about 25 miles from Mathewstown. Remarks. Application of Article 12.2.7 (ICZN, 1999: 17) does not alter the au- thorship of this taxon, but accelerates its publication by 3 months. Clench (1959: 45) listed this taxon as a synonym of Cer- ion (Diacerion) rubicundum (Menke, 1829). Taxon 14. Strophia palida Maynard, 1889a [April], pl. 2 figs. 14, 14B. Caption to plate 2, figure 14, “Strophia Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 palida, front view of e shell.” Stro- phia pallida Maynard, 1889b [July]: 70— 71. Text noted 25 specimens examined and mentioned the sizes of two “types” given as 1.03 by .36 and 1.03 by .33 [inches; 26.2 by 9.1 and 26.2 by 8.4 mm]. Nomenclatural Remarks. The illustra- tions [pl. 2, figs. 14, 14B] and caption were published prior to the text. The taxon thus dates from the publication of the name in the caption accompanying the illustrations, not from the subsequently published text (Article 12.2.7, ICZN, 1999: 17). Type Material. The specimen represent- ed in the figure is identified as the “type shell” and is therefore the holotype (Arti- cle 73.1.4, ICZN, 1999: S80). Although there was no mention of other specimens in the plate caption, the remaining 24 specimens enumerated in the text portion published 3 months later were likely avail- able to Maynard at the time the plate was prepared and are thus part of the type se- ries [Article 72.4.1.1, ICZN, 1999: 76], serving as paratypes of Strophia palida. Lot number MCZ 76250 contained two specimens, one adult and the other juve- nile, as well as multiple labels that state “type,” “Holotype,” and “paratype.” The adult specimen is a close approximation to the illustration and is here considered to be the figured specimen and therefore the holotype. The juvenile specimen is a para- e. Holotype Measurements. Length 26.1 mm, diameter (excluding lip) 8.8 mm; ap- erture height (including lip) 8.7 mm, ap- erture width (including lip and peristome) 6.9 mm. Type Locality. In cultivated fields on slopes of hills bordering the southern —_ Figures 13-24. Taxon 13. Strophia ianthina Maynard, 1889. Holotype MCZ 10317. Between southern shore of Inagua [Baha- mas] and the extensive salt lake about 25 miles from Mathewstown. Taxon 14. Strophia palida Maynard, 1889. Holotype MCZ 76250. Hills bordering the southern shore of Inagua, Bahamas, 15-20 miles from Mathewstown. Taxon 15. Strophia perplexa Maynard, 1889. Holotype MCZ 246345. About 2 miles from the western end of Cayman Brac, Cayman Islands. Taxon 16. Strophia nitela Maynard, 1889. Holotype MCZ 247020. Path, west end of Little Cayman, Cayman Islands. Taxon 17. Strophia THE CERION TAXA OF CHARLES JOHNSON MAYNARD ° Harasewych et all. 383 Ss \ eet. 5 —— alba Maynard, 1889. Holotype MCZ 76150. West coast of Rum Cay, Bahamas, near salt pond. Taxon 18. Strophia lentiginosa Maynard, 1889. Holotype MCZ 83132. Interior of Rum Cay, Bahamas. Taxon 19. Strophia fusca Maynard, 1889. Holotype MCZ 76069. West end of Little Cayman, Cayman Islands. Taxon 20. Strophia inflata Maynard, 1889. Holotype MCZ 76400. Salina Point, Auklin Islands, Bahamas. Taxon 21. Strophia dallii Maynard, 1889. Holotype MCZ 76294. Inagua. Taxon 22. Strophia brownei Maynard, 1889. Neotype USNM 58085. Rum Key, Bahamas, near the north side. Taxon 23. Strophia viola Maynard, 1889. Holotype MCZ 76407. Type locality not published, but Inagua, Bahamas, on the original label. Taxon 24. Strophia orbi- cularia Maynard, 1889. Holotype MCZ 76408. Inagua, Bahamas. 384 shore of Inagua [Bahamas], 15-20 miles from Mathewstown. Remarks. Application of Article 12.2.7 (ICZN, 1999: 17) does not alter the au- thorship of this taxon but accelerates its publication by 3 months. The publication of the plate and its caption, on which the taxon name is spelled palida (one “1”) pre- ceded the publication of the text, in which the taxon name is spelled pallida (two APO, Although it seems clear that Maynard intended the taxon to be known as pallida, it is not possible to consider palida as an “incorrect original spelling” within the confines of Article 32.5.1 (ICZN, 1999: 39) because there is no clear evidence to sug- gest an alternative spelling within the orig- inal publication itself. This taxon was listed as Saapalliida spya@lenem \GGoieaaltao)? Clench (1959: 45) included this taxon [as pallida| in the synonymy of Cerion (Di- acerion) rubicundum (Menke, 1829). Taxon 15. Strophia perplexa Maynard, 1889b [July]: 71-72, pl. 7, figs. 15, 1Aa, text figs. 7A, B [plate and text fig- ures identical]. Examined 300 speci- mens. Size of type given as .90 by .40 linches; 22.9 by 10.2 mm]. Caption to plate 7, figure 15, “Strophia perplexa, front view of type shell.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. The remaining 299 specimens are paratypes. Clench (1964: 368, pl. 63, fig. 11) designated and figured as lectotype MCZ 246345. This specimen is here con- sidered to be the originally designated ho- lotype. Holotype Measurements. Length 23.7 mm, diameter (excluding lip) 10.2 mm; ap- erture height (including lip) 9.2 mm, ap- erture width (including lip and peristome) 7.9 mm. Type Locality. Cayman Brac, [Cayman Islands] barren rocky section about 2 miles from the western end of the key % mile from the south shore. Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Taxon 16. Strophia nitela Maynard, 1889b [July]: 73-74, pl. 7, figs. 16, 16A, text figs. 8A, B [plate colored and text fig- ures uncolored, otherwise identical]. Examined 300 specimens. Size of type given as 1.10 by .50 [inches; 27.9 by 12.7 mm]. Caption to plate 7, figure 16, “Strophia nitela, front view of type shell; 16A, left side of same individual.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. The remaining 299 specimens are paratypes. Clench (1964: 368, pl. 63, fig. 12) designated and figured as lectotype MCZ 247020. This specimen is here con- sidered to be the originally designated ho- lotype. Holotype Measurements. Length 27.5 mm, diameter (excluding lip) 11.8 mm; ap- erture height (including lip) 11.1 mm, ap- erture width (including lip and peristome) 9.2 mm. Type Locality. Margins of path, west end of Little Cayman, Cayman Islands. Remarks. Clench (1964: 368) regarded this taxon as a synonym of Cerion pannos- um (Maynard, 1889). Taxon 17. Strophia alba Maynard, 1889b [July]: 74-75, pl. 7, figs. 17, 17A, B, text figs. 9A, B, text fig. 11 (sic) (11 in caption, 10 on figure) [plate and text figures identical]. Examined 250 specimens. Size of “type” given as 1.42 by .55 [inches; 36.1 by 14.0 mm]. Cap- tion to plate 7, figure 17, “Strophia alba, front view of type shell; 17a, left side of same individual.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. The remaining 249 specimens are paratypes. There was no specimen labeled “Holotype” at the MCZ. Lot MCZ 76150 THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. was labeled “Paratypes” and contained four specimens: two intact adults, a sec- tioned adult, and a juvenile. The largest intact shell is 33.0 by 14.0 mm. Although this is shorter than the published length of the type, it does match closely the size and proportions of the figured specimen, which the caption identifies as the type. This specimen (MCZ 76150) is here con- sidered to be the originally designated ho- lotype. Paratypes include MCZ 356978 (three remaining specimens from MCZ 76150), MCZ 39718 (one specimen), MCZ 10243 (two specimens), and USNM 420067 (nine specimens). Holotype Measurements. Length 33.0 mm, diameter (excluding lip) 14.0 mm; ap- erture height (including lip) 12.6 mm, ap- erture width (including lip and peristome) 10.2 mm. Type Locality. West coast of Rum Cay [Bahamas], near salt pond. Remarks. Clench (1934: 208) listed this taxon as a junior synonym of Cerion len- tiginosum (Maynard, 1889). Taxon 18. Strophia lentiginosa Maynard, 1889b [July]: 75-76, pl. 7, figs. 18, 18a, text figs. lla, b [plate colored, text figures uncolored, otherwise iden- tical]. Examined 150 specimens. Size of types [sic] given as 1.23 by .54 [inches; 31.2 by 17.7 mm]. Caption to pl. 7, fig. 18, “Strophia lentiginosa, front view of type shell; 18a, left side of the same in- dividual.” Type Material. The original description provided a single set of measurements, but states “types” rather than type. This is pre- sumed to be a typographical error, with the measurements applying to a single specimen, the holotype, which was illus- trated. The remaining 149 specimens are paratypes. Clench (1934: 209, pl. 2, fig. D) listed and illustrated MCZ 83132 as the holotype. This specimen is the originally designated holotype. Paratypes include: MCZ 10294, one paratype; MCZ 86701, one paratype; MCZ 76073, 13 paratypes. 385 Holotype Measurements. Length 30.9 mm, diameter (excluding lip) 14.2 mm; ap- erture height (including lip) 13.4 mm, ap- erture width (including lip and peristome) 10.4 mm. Type Locality. Interior of Rum Cay [Ba- hamas]. Remarks. Clench (1934: 208) regarded S. lentiginosa to be the senior synonym of S. alba Maynard, 1889 (Taxon 17), and of S. brownei Maynard, 1890 (Taxon 22). Taxon 19. Strophia fusca Maynard, 1889b [July]: 77-78, pl. 7, figs. 19, 19a, text figs. 12A, B [plate colored and text figures uncolored, otherwise identical]. Examined 46 specimens. Size of type given as 1.25 by .45 [inches; 31.8 by 11.4 mm]. Caption to plate 7, figure 19, “Strophia fusca, front view of type shell: 18a, left side of same individual.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. The remaining 45 specimens are paratypes. Clench (1964: 368, pl. 63, fig. 13) listed and illustrated MCZ 76069 as the holotype. This specimen is the origi- nally designated holotype. Holotype Measurements. Length 28.48 mm, diameter (excluding lip) 11.51 mm; aperture height (including lip) 10.41 mm, aperture width (including lip and _peri- stome) 8.97 mm. Type Locality. West end of Little Cay- man, Cayman Islands. Remarks. Clench (1964: 368) regarded this taxon to be a synonym of Cerion pan- nosum (Maynard, 1889). Taxon 20. Strophia inflata Maynard, 1889b [July], pl. 7, figs. 21, 21a. Cap- tion to plate 7, figure 21 “Strophia infla- ta, front view of type shell; 21a, left side of same individual”; Maynard, 1889c [October]: 126-127, text fig. 30A, B [plate colored and text figures unco- lored, otherwise identical]. Text noted 25 specimens examined. Size of “types” 386 [sic] given as .90 by .35 [inches; 22.9 by 8.9 mm]. Type Material. The specimen represent- ed in the figure is identified as the “type shell” and is therefore the holotype (Arti- cle 73.1.4, ICZN, 1999: 80). Although there was no mention of other specimens in the plate caption, the remaining 24 specimens enumerated in the text portion published 3 months later were likely avail- able to Maynard at the time the plate was prepared and are thus part of the type se- ries [Article 72.4.1.1, ICZN, 1999: 76], serving as paratypes of Strophia inflata. Lot MCZ 76400, labeled “Holotype,” con- taining a single specimen that closely ap- proximates the original figure, and accom- panied by a note in Maynard’s handwrit- ing, is here recognized as the holotype. Holotype Measurements. Length 22.9 mm, diameter (excluding lip) 9.2 mm; ap- erture height (including lip) 8.5 mm, ap- erture width (including lip and peristome) 7.9 mm. Type Locality. Salina Point, Auklin [Acklins] Islands, Bahamas. Remarks. Application of Article 12.2.7 (ICZN, 1999: 17) does not alter the au- thorship of this taxon but accelerates its publication by 3 months. Maynard (1889c: 127) noted that the specimens on which he based this taxon were from the collec- tion of the Boston Society of Natural His- tory. Maynard (1919b, pl. 8, figs. 5, 6) reil- lustrated the holotype. Clench (1963: 408) included this taxon in the synonymy of Cerion (Multistrophia) marmoratum (Pfeiffer, 1847). Taxon 21. Strophia dallii Maynard, 1889c [October]: 128-135, pl. 13, figs. 23, 23a [plate labeled as pl. 16, but there are no plates 13-15, two figs. each 1B* on plate], text figs. 32, A, 0, e, fig. 33 [plate colored, text fig- ures uncolored, otherwise identical ex- cept that text figure has b and d la- beled]. Examined 25 specimens. Size of type given as 1.20 by .42 |[inches; 30.5 Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 by 10.7 mm]. Caption to plate 16, figure 1b*, “Strophia dalli, right hand figure, front view of type; left, right view of same. Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. The remaining 24 specimens are paratypes. No specimen labeled as the ho- lotype was found at the MCZ. Lot MCZ 76294 was labeled “Paratype” and con- tained three adult specimens together with a note in Maynard's hand containing the term “type.” Of the three specimens only one closely approximates both the mea- surements (including proportions) and the figures. This specimen is here considered to be the originally designated holotype. The remaining two specimens (now MCZ 306979) and a single specimen (MCZ 39707) are paratypes. Holotype Measurements. Length 30.0 mm, diameter (excluding lip) 10.7 mm; ap- erture height (including lip) 10.4 mm, ap- erture width (including lip and peristome) 8.6 mm. Type Locality. Maynard (1889c: 135) wrote, “The first specimen of Dall’s Stro- phia that I ever saw, I found in my Baham- ian collection of shells, but unfortunately labeled so that it was uncertain whether it came from Inagua or not. Later I found a few of this species in the collection of Mr. James A. Southwick, but again I was un- fortunate in not getting the locality. It was only upon receiving a series of the Smith- sonian Strophias, kindly forwarded to me by Dr. Dall, that I found this species la- beled as coming from Inagua.” Remarks. Clench (1934: 217) reported on specimens from Sheep Cay, western Great Inagua, but commented that these differed from Maynard’s types. Clench (1959: 46) recognized this taxon as Cerion (Diacerion) dalli (Maynard, 1889). Taxon 22. Strophia brownei Maynard, 1889c [October], pl. 16, figs. 4A, B. Caption to plate 16, figure 4A, “Strophia THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. brownei, front view of type shell; 4B, right side of same”; Maynard, 1890 [Jan- uary]: 196-197, text fig. 53c, d [text fig- ures uncolored, otherwise identical to figures on pl. 16]. Text noted 25 speci- mens examined. Size of type given as 1.10 by .50 [inches; 27.9 by 12.7 mm]. Type Material. The illustrations and caption were published prior to the text. The taxon thus dates from the publication of the name in the caption accompanying the illustrations, not from the subsequent- ly published text (Article 12.2.7, ICZN, 1999: 17). The specimen represented in the figure is identified as the “type shell” and is therefore the holotype (Article 73.1.4, ICZN, 1999: 80). Although there was no mention of other specimens in the plate caption, the remaining 24 specimens enumerated in the text portion published 3 months later were likely available to Maynard at the time the plate was pre- pared and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76] serving as paratypes of Strophia brownei. Maynard noted (1890: 197) that the specimens were from the collection of Mr. Frank C. Browne and commented that he [Maynard] saw a number of specimens as he rode along the road and “fully intended to collect them as I returned, but when I passed the place in the evening, darkness prevented me from securing any, and I did not have the op- portunity of visiting the locality again.” This species was not listed for sale in May- nard’s catalog. The MCZ Mollusk catalog shows that two “paratypes” of Strophia brownei were present in Maynard's collection at the time of his death and that these specimens had been catalogued as MCZ 76288. These specimens could not be located in the MCZ collection, despite repeated search- es. It is possible that most of the speci- mens on which Maynard based his de- scription were returned to Browne. In an unsigned obituary of Francis C. Browne, Pilsbry (1901: 132) commented that Browne had frequently sent him specimens for 387 identification or verification. A survey of the collections at the Academy of Natural Sciences in Philadelphia revealed that they did not contain any specimens of Cerion brownei nor any specimens from Rum Cay that came from Browne. Two specimens in the USNM collec- tions (USNM 58085) are annotated “S. brownei Maynard fide Maynard.” These specimens were collected by Rawson on Rum Key and catalogued [as Pupa mumia Brug.] on February 4, 1886, well before S. brownei was published. However, a sub- sequent annotation on the label indicated that Maynard had identified one of these specimens as Strophia brownei. This spec- imen (USNM 58085) is here designated as the neotype of Strophia brownei to pro- vide an objective standard of reference for this species-group taxon that is consistent with Maynard's concept of it. The remain- ing specimen (now USNM 1093788) is not S. brownei and is not a type. Neotype Measurements. Length Ds mim, diameter (excluding lip) 12.8 mm; ap- erture height (including lip) 11.5 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. Rum Key [Bahamas] near the north side. Remarks. Application of Article 12.2.7 (ICZN, 1999: 17) does not alter the au- thorship of this taxon but accelerates its publication by 3 months (from 1890 to 1889). Clench (1934: 208) listed this taxon as a synonym of Cerion lentiginosum (Maynard, 1889). Taxon 23. Strophia viola Maynard, 1889c [October], pl. 16, figs. SA, B [no de- scription or mention in text]. Caption to plate 16, figure 5A, “Strophia viola, front view of type; 5B, right side of same.” Type Material. The caption to the fig- ures identified a single type specimen, the holotype. Because no other specimens were mentioned, there are no paratypes. Clench (1959, pl. 1, fig. 2) illustrated this holotype, MCZ 76407, as Cerion rubicun- dum viola (Maynard, 1889). 388 Holotype Measurements. Length 16.6 mm, diameter (excluding lip) 7.1 mm; ap- erture height (including lip) 6.4 mm, ap- erture width (including lip and peristome) Dy idan. Type Locality. Clench (1957: 168) noted “no locality given, but Inagua, Bahamas on original label.” Remarks. Pilsbry and Vanatta (1896: 328) listed this taxon as an undescribed species, whereas Pilsbry (1902: 281) re- garded this as an unrecognizable form. Clench (1933: 99) considered this to be a valid species related to Cerion bryanti Pfeiffer. Later, Clench (1959, pl. 1, fig. 2) reduced this taxon to a subspecies, C. rub- icundum viola (Maynard, 1889). Taxon 24. Strophia orbicularia Maynard, 1889c [October], pl. 16, figs. 6A, B [no description or mention in text]. Caption to plate 16, figure 6A, “Strophia orbicularia, front view of type; 6B, right side of same.” Type Material. The caption to the fig- ures identified a single type specimen, the holotype. Because no other specimens were mentioned, there are no paratypes. Clench (1959: 45) listed MCZ 76408 and MCZ 76409 as “type specimens.” Lot MCZ 76408 is labeled “holotype,” closely approximates the illustration, and contains a note in Maynard’s hand containing the Ler hyper This specimen is recognized as the originally designated holotype. The remaining specimen MCZ 76409 is not a paratype because it was not mentioned in the original description. Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Holotype Measurements. Length 22.7 mm, diameter (excluding lip) 8.1 mm; ap- erture height (including lip) 8.1 mm, ap- erture width (including lip and peristome) 6.2 mm. Type Locality. Clench (1957: 155) noted “Inagua, Bahamas, label on type speci- mens. Remarks. Pilsbry and Vanatta (1896: 328) listed this taxon as an undescribed species. Clench (1933: 99) listed this taxon as a synonym of Cerion bryanti Pfeiffer and later (Clench, 1959: 45) of C. (Di- acerion) rubicundum (Menke, 1829). Taxon 25. Strophia curtissii Maynard, 1894a [December]: 107-112, text figs. 33A, B, C, D, fig. 40a (map). Ex- amined 2,000 specimens. Size of type .98 by .40 linches; 24.9 by 10.2 mm]. Caption to text figure 33, “Strophia cur- tissii. ... B, side view of type; C, front view of same.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. No specimen was labeled “Holo- type” at the MCZ. Lot number MCZ 10274 was labeled “Paratype” and con- tained two specimens. The smaller of these specimens approximates the pub- lished measurements within 3% and more closely resembles figure 33C than 33B. Unlike the remaining specimen (now MCZ 356980), it has a minute lower tooth and lacks an upper tooth, as specified in the description (p. 108). This specimen (MCZ 10274) is here considered to be the —=> Figures 25-36. Taxon 25. Strophia curtissii Maynard, 1894. Holotype MCZ 10274, cemetery east of Nassau, New Providence Island, Bahamas. Taxon 26. Strophia curtissii nivia Maynard, 1894. Holotype MCZ 010279. Banyan tree near ruin in the cemetery where S. curtissii is found [large cemetery east of Nassau, New Providence Island, Bahamas]. Taxon 27. Strophia thorndikei Maynard, 1894. Holotype MCZ 76086. West of main path in cemetery east of Nassau, New Providence Island, Bahamas. Taxon 28. Strophia cinerea Maynard, 1894. Holotype MCZ 76254. Hog Island [Paradise Island], along the shore of Middle Bay, on the south side of the Key, New Providence Island, Bahamas. Taxon 29. Strophia cinerea robusta Maynard, 1894. Neotype MCZ 356982. North side of Hog Island [Paradise Island], directly back of the beach, New Providence Island, Bahamas. Taxon 30. Strophia cinerea tracta Maynard, 1894. Holotype MCZ 76081. Field near the extreme eastern point of Hog Island [Paradise Island], New Providence Island, Bahamas. Taxon 31. Strophia cinerea mutata Maynard, 1894. Holotype MCZ 76279. Northern shore of the western half of Long Key, about 1 mile east of Hog Island [Paradise Island], New Providence Island, Bahamas. Taxon 32. Strophia albea Maynard, 1894. Holotype MCZ 10242. South side of Spruce Key, due north of Long Key, about 1 mile THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 389 east of Hog Island [Paradise Island], New Providence Island, Bahamas. Taxon 33. Strophia coryi Maynard, 1894. Holotype MCZ 76079. Along bay on the extreme west end of New Providence Island, Bahamas, also on Spruce Key, N of Long Key, 1 mile east of Hog Island [Paradise Island], New Providence Island, Bahamas. Taxon 34. Strophia ritchiei Maynard, 1894. Holotype MCZ 76077. Highburn Key, Exuma Group, Bahamas. Taxon 35. Strophia grayi Maynard, 1894. Holotype MCZ 118169. Hill at northern end of Highburn Key, Exuma Group, Bahamas. Taxon 36. Strophia grayi gigantea Maynard, 1894. Holotype MCZ 10290, Hillside, middle part of Highburn Key, Exuma Group, Bahamas. 390 originally designated holotype. How many of the remaining 1,999 specimens are paratypes is uncertain (see Remarks). Holotype Measurements. Length 25.2 mm, diameter (excluding lip) 9.9 mm; ap- erture height (including lip) 9.4 mm, ap- erture width (including lip and peristome) 7.6 mm. Type Locality. “The type form occurs in the large cemetery to the eastward of Nas- sau, |New Providence Island, Bahamas] on the western side of a path which crosses the grounds |[map, fig. 40a]” (Maynard, 1894a: 109). Remarks. Maynard (1894a: 109) distin- guished five forms [Nos. 1—5] without naming them. The type series is defined (Article 72.4.1, ICZN, 1999: 76) as “all the specimens included by the author in the new nominal taxon, except any that the au- thor expressly excludes from the type se- ries, or refers to as distinct variants (e.g., by name, letter, or number).” Thus, spec- imens representing forms No. 1-5 are not paratypes of Strophia curtissii. Clench (1957: 142) followed Pilsbry (1902: 256) in listing this taxon as a synonym of Cerion varium (Bonnet, 1864). Gould and Wood- ruff (1986: 476) regarded this taxon to be “intermediate” between Cerion glans (Ktister, 1844) and C. gubernatorium (Crosse, 1869). Taxon 26. Strophia curtissii nivia May- nard, 1894a [December]: 112-116 [nivea in text], fig. 34A [nivia in cap- tion]. Examined 40 specimens. Size of type .90 by .37 [inches; 22.9 by 9.4 mm]. Caption to text figure 34A, “Strophia curtissii nivia, front view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. How many of the remaining 39 specimens are paratypes is uncertain (see Remarks). An examination of the collec- tions and catalog at the MCZ revealed that, at the time of his death, Maynard’s collection contained one lot of this taxon Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 [catalogued as MCZ 76097 “Paratypes” | that contained in excess of 200 specimens, as well as handwritten labels dated 1891 and 1897. These labels indicate that the majority of the specimens were added af- ter the taxon was described. There was no single, segregated specimen labeled “Ho- lotype” either in the collection or the cat- alog. Lot MCZ 10279 contained two spec- imens labeled “paratype” and had been purchased from Maynard in 1895. One of these specimens matches the figure of the holotype in having a columellar tooth, which is not evident in the remaining | specimen. This specimen is considered to be the holotype. Lot number USNM 420034 contains 100 specimens. Lot num- ber MCZ 10280 contained two specimens labeled “paratype” and are accompanied by a label indicating “Form No. 2,” which excludes these specimens from the type series (see Remarks). Holotype Measurements. Length YS, UL mm, diameter (excluding lip) 9.5 mm; ap- erture height (including lip) 8.6 mm, ap- erture width (including lip and peristome) 7.0 mm. Type Locality. Banyan tree near ruin in the cemetery where S. curtissii is found [large cemetery eastward of Nassau, New Providence Island, Bahamas]. Remarks. The taxon name originally ap- peared as Strophia curtissii nivea in the heading of the original description, but as S. curtissii nivia in the caption to the fig- ure within the body of the description. Maynard (1913b: 186) described S. varia- nivia as “Intermediate between varia and nivia.” With this use of nivia, Maynard (1913b: 186) acted as the First Revisor of spellings [Article 24.2.4, ICZN, 1999: 30- 31] and fixed the spelling of this species as nivia. Strophiops nivia again appeared in Maynard (1921b: 148) and in Maynard's [1924b?P] sales catalog. The spelling “ni- veda” appeared with a citation to the orig- inal description in Maynard (1920a: 79). Oddly, Maynard (1913b: 186) described S. nivia as a new taxon while incorrectly list- ing the preceding taxon, S. varia-nivia as THE CERION TAXA OF CHARLES JOHNSON MAYNARD °* Harasewych et al. previously described. This is almost cer- tainly an error in typesetting. Strophiops nivia Maynard (1913) is a junior secondary homonym of Strophia curtissii nivia May- nard (1894) as well as an objective junior synonym because they have the same type specimen [see Taxon 72]. Maynard (1894a: 113) distinguished two forms [Nos. 1 and 2] without naming them. The type series is defined in Article 72.4 (ICZN, 1999: 76) as “all the speci- mens included by the author in the new nominal taxon, ... except any that the au- thor expressly excludes from the type se- ries, or refers to as distinct variants (e.g., by name, letter, or number).” Thus, spec- imens representing forms No. 1 or 2 are not paratypes of Strophia curtissii nivia. In his discussion of S. nivia, Maynard (1921b: 148) provided larger measurements (1.00 by .40 [inches; 25.4 by 10.2 mm]), listed 829 specimens rather than the 40 in the original description, and included citations to illustrations (plate 44, figs. 3, 4) that were never issued. Clench (1957: 155) listed this taxon as nivea and considered nivia an error for ni- vea. Gould and Woodruff (1986: 476) re- garded this taxon to be “intermediate” be- tween Cerion glans (Kiister, 1844) and C. gubernatorium (Crosse, 1869). Taxon 27. Strophia thorndikei Maynard, 1894a [December]: 116-119, text figs. 34B, C, D, fig. 40t (map). Ex- amined 2,000 specimens. Size of type .70 by .30 [inches; 17.8 by 7.6 mm]. Caption to text fig. 34D, “Strophia thorndikei, front view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot number MCZ 76086 contains a single specimen labeled “Holotype,” which is 6% larger than Maynard’s pub- lished length but otherwise approximates the figure of the type and includes May- nard’s original label annotated “type.” This specimen is here recognized as the Holo- oul type. Lot number USNM 419958 contains 200 paratypes. How many of the remain- ing 1,799 specimens, are paratypes is un- certain (see Remarks). Holotype Measurements. Length 18.9 mm, diameter (excluding lip) 8.1 mm; ap- erture height (including lip) 7.9 mm, ap- erture width (including lip and peristome) 6.2 mm. Type Locality. West of main path in cemetery eastward of Nassau (map, fig. 40t) [New Providence Island, Bahamas]. Remarks. Maynard (1894a: 117-118) distinguished five forms [Nos. 1—5] with- out naming them. He illustrated form No. 3 (fig. 34B) and form No. 4 (fig. 34C). Specimens representing forms No. 1—5 are not paratypes of Strophia thorndikei |Ar- ticle 72.4, ICZN, 1999: 76]. This taxon was figured by Manard (1919b: 47, fig. 8 as S. thorndikeii Mayn.). Maynard (1921b: 146) provided descriptive notes, including ref- erence to a color illustration (plate 8, fig. 10) published in 1919, and commented that the taxon was represented by 2,500 specimens, at least 500 of which presum- ably were collected after the taxon was de- scribed. Maynard (1924c: 6) listed S. thorndikei as extinct in 1913 because of burning of vegetation prior to cultivation. Gould and Woodruff (1986: 476) regarded this taxon to be “intermediate” between Cerion glans (Kiister, 1844) and C. gub- ernatorium (Crosse, 1869). Taxon 28. Strophia cinerea Maynard, 1894a |December]: 119-121, text figs. 35A, B, fig. 40e (map). Examined 2,000 specimens. Size of type 1.15 by 45 [inches; 29.2 by 11.4 mm]. Caption to text figure 35, “B, front view of type. A, side view of same.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot number MCZ 76254 contains a single specimen labeled “Holotype,” which is within 5% of the published mea- surements and includes Maynard's original 392 label annotated “type.” This specimen is here recognized as the Holotype. Lot number USNM 420076 contains 200 para- types. How many of the remaining 1,799 specimens are paratypes is uncertain (see Remarks). Holotype Measurements. Length 28.0 mm, diameter (excluding lip) 11.3 mm; ap- erture height (including lip) 9.4 mm, ap- erture width (including lip and peristome) 7.6 mm. Type Locality. Hog Island [Paradise Is- land], along the shore of Middle Bay, on the south side of the key (map, fig. 40e) [New Providence Island, Bahamas]. Remarks. Maynard (1894a: 120-121) distinguished three forms | Nos. 1—3] with- out naming them. Specimens representing forms No. 1-3 are not paratypes of Stro- phia cinerea |Article 72.4, ICZN, 1999: 76]. Maynard (1921b: 144; 1925, pl. 40, figs. 3, 4) noted 2,500 specimens, which must have included subsequent collec- tions, and provided new, colored illustra- tions for this taxon. Taxon 29. Strophia cinerea robusta May- nard, 1894a [December]: 121-123, text figs. 36 A, B, fig. 40f (map). Ex- amined 75 specimens. Size of type 1.12 by .55 [inches; 28.4 by 14.0 mm]. Cap- tion to text figure 36, “Strophia cinerea robusta. A, front view, B, side view, of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. There is no specimen labeled “Ho- lotype” at the MCZ. An examination of the MCZ catalog showed that no representa- tives of this taxon were present in May- nard’s collection when purchased from his daughter. Lot number MCZ 10259 was la- beled “Paratype” and contained two spec- imens that were acquired from Maynard in 1895. The specimen in MCZ 10259 that more closely matches the published fig- ures and measurements was recatalogued as MCZ 356982 and is here designated as Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 the neotype. How many of the remaining 74 specimens are paratypes is uncertain (see Remarks). Neotype Measurements. Length 29.0 mm, diameter (excluding lip) 13.0 mm; ap- erture height (including lip) 11.3 mm, ap- erture width (including lip and peristome) 8.4 mm. Type Locality. North side of Hog Island [Paradise Island], directly back of the beach (map, fig. 40f) [New Providence Is- land, Bahamas]. Remarks. Maynard (1894a: 123) distin- guished a small form [No. 1] without nam- ing it. Lot MCZ 10260 contained two specimens labeled form No. 1. Specimens representing this form are not paratypes of Strophia cinerea robusta [Article 72.4, ICZN, 1999: 76]. Gould and Woodruff (1986: 475) regarded this taxon to be a synonym of Cerion glans (Kiister, 1844). Taxon 30. Strophia cinerea tracta May- nard, 1894a [December]: 123-125, text fig. 37A, fig. 40g (map). Examined 300 specimens. Size of type 1.05 by .45 [inches; 26.7 by 11.4 mm]. Caption to text figure 37A, “Strophia cinerea tracta, front view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot number MCZ 76081 contains a single specimen labeled “Holotype,” closely matches Maynard’s published length and figure of the type, and includes Maynard's original label annotated “type.” This specimen is here recognized as the Holotype. Lot number USNM 420042 contains paratypes. How many of the re- maining specimens are paratypes is uncer- tain (see Remarks). Holotype Measurements. Length 26.9 mm, diameter (excluding lip) 12.0 mm; ap- erture height (including lip) 9.6 mm, ap- erture width (including lip and peristome) 8.4 mm. Type Locality. Field near the extreme eastern point of Hog Island [Paradise Is- THE CERION TAXA OF CHARLES JOHNSON MAYNARD ° Harasewych et al. land] (map, fig. 40g) [New Providence Is- land, Bahamas]. Remarks. Maynard (1894a: 124—125) distinguished five forms [Nos. 1-5] with- out naming them. Specimens representing forms No. 1—5 are not paratypes of Stro- phia cinerea tracta [Article 72.4, ICZN, 1999: 76]. Maynard (1921b: 144—145) list- ed this taxon as a full species, and provided new, colored illustrations (Maynard, 1925, pls. 40, figs. 7, 8). Clench (1957: 165) fol- lowed Pilsbry (1902: 253) in synonymizing this taxon under Cerion varium (Bonnet, 1864), whereas Gould and Woodruff (1986: 475) considered it to be a synonym of C. glans (Kiister, 1844). Taxon 31. Strophia cinerea mutata May- nard, 1894a [December]: 125-128, text fig. 37B, fig. 40h (map). Exam- ined 100 specimens. Size of type 1.15 by .40 [inches; 29.2 by 10.2 mm]. Cap- tion to text figure 37B, “Strophia cinerea mutata, front view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot number MCZ 76279 contains a single specimen labeled “Holotype,” which is 11% shorter than Maynard’s pub- lished length and slightly wider but ap- proximates the figure of the type and con- tains Maynard’s original label annotated “type.” This specimen is here recognized as the Holotype. Lot number USNM 420132 contains paratypes. How many of the remaining specimens are paratypes is uncertain (see Remarks). Holotype Measurements. Length 26.0 mm, diameter (excluding lip) 11.0 mm; ap- erture height (including lip) 9.7 mm, ap- erture width (including lip and peristome) S.0 mm. Type Locality. Northern shore of the western half of Long Key, about 1 mile east of Hog Island [Paradise Island] (map, fig. 40h) [New Providence Island, Baha- mas]. Remarks. Maynard (1894a: 126) distin- 393 guished a single form |[No. 1] without naming it. Specimens representing this form are not paratypes of Strophia cinerea mutata [Article 72.4, ICZN, 1999: 76]. Maynard (1921b: 146) listed this taxon as a full species and referred to a previously published [1920a, pl. 10*, fig. 4] colored illustration but noted a different locality (Atoll Island) and a higher number of specimens (200). Taxon 32. Strophia albea Maynard, 1894a [December]: 128-129, text figs. 38A, B. Examined 40 specimens [only three living specimens]. Size of type 1.00 by .36 [inches; 25.4 by 9.1 mm]. Caption to text figure 38, “Stro- phia albea. A, front view, B, side view, of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. The remaining 39 specimens are paratypes. No specimen was labeled “Ho- lotype” at the MCZ. An examination of the MCZ catalog showed that no representa- tives of this taxon were present in May- nard’s collection when purchased from his daughter. Lot MCZ 10242 was labeled “Paratype” and contained two specimens that were acquired from Maynard in 1895. One of these specimens approximates the length in the original description to within 3% and agrees with the generalized illus- tration (more fig. 38B than 38A). This specimen is considered to be the Holo- type. The remaining specimen has been recatalogued as MCZ 356983. Maynard (1921b: 139) listed 590 specimens, indi- cating that 550 specimens had been add- ed. Some of the 100 specimens of USNM 420097 may be paratypes. Holotype Measurements. Length 26.0 mm, diameter (excluding lip) 10.1 mm; ap- erture height (including lip) 9.0 mm, ap- erture width (including lip and peristome) 6.8 mm. Type Locality. South side of Spruce Key, due north of Long Key, about 1 mile E of 394 Hog Island [Paradise Island] [New Provi- dence Island, Bahamas]. Remarks. The identical illustration ap- pears in Maynard (1913c: 200) as Stro- phiops albea Mayn. and again in a colored version (Maynard, 1919c, pl. 10, figs. 1, 2 [lower row]). The taxon was redescribed in Maynard (1921b: 139) on the basis of ex- amination of 590 specimens. It appeared twice in the sales catalog (Maynard, 1924b: [6] and [7]), the first time correctly (from Spruce Key), the second time from Rum Key, an error for alba. Clench (1957: 165) followed Pilsbry (1902: 253) in synonymiz- ing this taxon under Ceron varium Bon- net, 1864. Gould and Woodruff (1986: 474) had examined “paratypes” (presum- ably MCZ 10242) and synonymized this taxon with C. glans (Kiister, 1844). Taxon 33. Strophia coryi Maynard, 1894a [December]: 129-135, text figs. 39A, B. Examined 2,000 speci- mens. Size of type 1.00 by .40 |[inches; 25.4 by 10.2 mm]. Caption to text figure 39, “Strophia coryi, A, front view, B, side view, of type.” The same figures ap- pear (Maynard, 1913c: 200) as Stro- phiops coryi Mayn. Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Clench (1934: 214) mentioned ex- amining type specimens but did not fur- ther identify them. Lot number MCZ 76079 contains a single specimen labeled “Holotype,” which is 6% shorter and slightly wider than Maynard’s published measurements but approximates the figure of the type and contains Maynard's original label annotated “type.” This specimen is here recognized as the Holotype. Lot number USNM 419983 contains 100 para- types. How many of the remaining 1,899 specimens are paratypes is uncertain (see Remarks). Holotype Measurements. Length 23.9 mim, diameter (excluding lip) 10.4 mm; ap- erture height (including lip) 9.2 mm, ap- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 erture width (including lip and peristome) 7.9 mm. Type Locality. Along bay on the extreme west end of New Providence Island, Ba- hamas, also on Spruce Key, N of Long Key, 1 mile E of Hog Island [Paradise Is- land| [New Providence Island, Bahamas]. Remarks. Maynard (1894a: 131-132) distinguished five forms [Nos. 1-5] with- out naming them. Specimens representing forms No. 1—5 are not paratypes of Stro- phia coryi |Article 72.4, ICZN, 1999: 76]. This taxon was subsequently reillustrated (Maynard, 1919b, fig. 18, pl. 7, figs. 9, 10). Clench (1934: 214, 1952: 108) listed this taxon as a form of Cerion glans (Kiister, 1844). Taxon 34. Strophia ritchiei Maynard, 1894a [December]: 135-138, text figs. 41A, B. Examined 1,000 speci- mens. Size of type 1.37 by .57 [inches; 34.5 by 14.5 mm]. Caption to text figure 41, “Strophia ritchiei, A, front; B, side view of type.” [Illustration repeated on back cover of Maynard (1913) and the front covers of Maynard (1920a—1924a) as Strophiops ritchiei Mayn. | Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot number MCZ 76077 contains a single specimen labeled “Holotype,” which is within 0.1 mm of Maynard’s pub- lished measurements, approximates the figure of the type, and contains Maynard's original label annotated “type.” This spec- imen is here recognized as the Holotype. Lot number USNM 419967 contains 500 paratypes. How many of the remaining 499 specimens are paratypes is uncertain (see Remarks). Holotype Measurements. Length 34.6 mm, diameter (excluding lip) 14.6 mm; ap- erture height (including lip) 13.17 mm, ap- erture width (including lip and peristome) 10.12 mm. Type Locality. Highburn Key [Exuma Group, Bahamas]. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. Remarks. Maynard (1894a: 136-137) distinguished four forms [Nos. 1—4] with- out naming them. Specimens representing forms No. 1-4 are not paratypes of Stro- phia ritchiei [Article 72.4, ICZN, 1999: 76]. The name was misspelled as S. ritchei (Maynard, 1920a: 75). Taxon 35. Strophia grayi Maynard, 1894a [December]: 138-141, text figs. 422A, B. Examined 700 specimens. Size of type 1.25 by .47 [inches; 31.8 by 17.8 mm]. Caption to text figure 42, “Strophia grayi, A, front; B, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot number MCZ 118169 contains a single specimen labeled “co-type.” This specimen approximates the published length (within 5%) and is a reasonable ap- proximation of the original illustration. It is accompanied by a label identifying it as a co-type, initialed by Maynard and dated “12/14/15,” indicating that the label was added after the description of the species. This specimen is here recognized as the Holotype. Lot number USNM 420101 contains 100 paratypes. How many of the remaining 599 specimens are paratypes is uncertain (see Remarks). Holotype Measurements. Length 30.5 mim, diameter (excluding lip) 13.0 mm; ap- erture height (including lip) 11.0 mm, ap- erture width (including lip and peristome) 9.1 mm. Type Locality. Hill at northern end of Highburn Key [Exuma Group], Bahamas. Remarks. Maynard (1894a: 139-141) distinguished three forms [Nos. 1—3] with- out naming them and illustrated form No. 3 (fig. 43A). Specimens representing forms No. 1—3 are not paratypes of Strophia gra- yi [Article 72.4, ICZN, 1999: 76]. Taxon 36. Strophia grayi gigantea May- nard, 1894a [December]: 141-143, text fig. 44A. Examined 50 specimens. Size of type 1.56 by .55 [inches; 39.6 by 395 14.0 mm]. Caption to text figure 44A, “Strophia grayi gigantea.” Type Material. The original description provided measurements for a single “type” specimen, but the illustrated specimen was not referred to as type. The measurements are considered to designate a holotype. No specimen labeled holotype could be found at the MCZ. Two specimens of MCZ 10290 were labeled “paratype.” The larger of these specimens approximates the length and width in the original descrip- tion to within 4%. This specimen is con- sidered to be the Holotype. The remaining specimen (now MCZ 356984) is a para- type. How many of the remaining 48 spec- imens are paratypes is uncertain (see Re- marks). Holotype Measurements. Length 38.1 mm, diameter (excluding lip) 14.4 mm; ap- erture height (including lip) 12.6 mm, ap- erture width (including lip and peristome) 9.9 mm. Type Locality. Hillside, middle part of Highburn Key [Exuma Group], Bahamas. Remarks. Maynard (1894a: 139-141) distinguished a single form [No. 1] without naming it. Two specimens of MCZ 10291 were labeled “paratype.” However, May- nard’s printed label, which accompanies these two specimens, identifies them as form No. 1. Specimens representing this form are not paratypes of Strophia grayi gigantea |Article 72.4.1, ICZN, 1999: 76]. Taxon 37. Strophia grayi pumilia May- nard, 1894a [December]: 143-144, text fig. 44B. Examined 50 specimens. Size of type .90 by .42 [inches; 22.9 by 10.7 mm]. Caption to text figure 44B, “Strophia grayi pumilia.” Type Material. The original description provided measurements for a single “type” specimen, but the illustrated specimen was not referred to as a type. The measure- ments are sufficient to designate a holo- type. No specimen was labeled “Holotype” at the MCZ. The MCZ collections contain two lots of Strophia grayi pumilia, both 396 labeled “paratype.” One, MCZ 76325, contains 12 specimens, the other, MCZ 10292, contains two specimens. Although none match the measurements precisely, the larger of the two specimens in MCZ 10292 comes closest to approximating the size (within 5%) and proportions (within 1%) of the measurements as well as the illustration. This specimen is regarded as the originally designated Holotype, the re- maining specimen (now MCZ 356985) is a paratype. The remaining 48 specimens are paratypes. Holotype Measurements. Length 2370 mm, diameter (excluding lip) 11.1 mm; ap- erture height (including lip) 8.9 mm, ap- erture width (including lip and peristome) 7.9 mm. Type Locality. South of deep gorge, middle part of Highburn Key [Exuma Group], Bahamas. Remarks. The taxon appeared as “Stro- phia gray! pumilia” in the original descrip- tion and as Strophia grayi pumilia in the figure caption but as pumilla in the index. Taxon 38. Strophia eburnia Maynard, 1894a [December]: 144-148 [pages printed out of order], text figs. 45A, B. Examined 100 specimens [12 collected living, p. 147]. Size of type 1.15 by .45 linches; 29.2 by 11.4 mm]. Caption to text figure 45, “Strophia eburnia, A, front view, B, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. The remaining 99 specimens are paratypes. Lot MCZ 76224 was labeled “syntypes” and contained two specimens, as well as Maynard’s original label anno- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 tated “type.” One specimen is gray, has a red “Holotype” label affixed to it, and con- forms to the published figure as well as the measurements given for “the type.” The original catalog entry at the MCZ listed only one specimen and referred to it as the “holotype.” The gray, labeled specimen is here recognized as the holotype. The other dark brown specimen is a paratype and has been recatalogued as MCZ 356986. Lot USNM 420021 contains 30 paratypes. Holotype Measurements. Length 29.0 mm, diameter (excluding lip) 11.2 mm; ap- erture height (including lip) 9.9 mm, ap- erture width (including lip and peristome) ~ 8.3 mm. Type Locality. U Key [Exuma Group], Bahamas, single specimen tentatively identified as this species on Pimlico Key. Remarks. This taxon was misspelled as S. eburnea (Maynard, 1920a: 74, 78). Taxon 39. Strophia elongata Maynard, 1894a [December]: 148-150 [pages printed out of order], text figs. 46A, B. Examined 25 specimens. Size of type 1.33 by .50 [inches; 33.8 by 12.7 mm]. Caption to text figure 46, “Strophia elongata, A, front B, side view.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. The remaining 24 specimens are paratypes. There are no specimens of S. elongata in the collections of the MCZ, nor were there any entries corresponding to this taxon in the MCZ catalog either during Maynard's life or among the spec- imens purchased from his estate. Although it seems probable that Maynard retained the holotype in his collection, it might not SS Figures 37-48. Taxon 37. Strophia grayi pumilia Maynard, 1894. Holotype MCZ 10292. South of deep gorge, middle part of Highburn Key, Exuma Group, Bahamas. Taxon 38. Strophia eburnia Maynard, 1894. Holotype MCZ 76224. U Key, Exuma Group, Bahamas. Taxon 39. Strophia elongata Maynard, 1894. Neotype FMNH 42208. Key, 1 mile north of U Key at Allen’s Harbor, Exuma Group, Bahamas. Taxon 40. Strophia neglecta Maynard, 1894. Holotype MCZ 76376. One mile west of Fort Charlotte, New Providence Island, Bahamas. Taxon 41. Strophia neglecta agava Maynard, 1894. Holotype MCZ 76103. Sisal fields west of Nassau, New Providence Island, Bahamas. Taxon 42. Strophia carlotta Maynard, 1894. Holotype MCZ 10249. At the foot of the hill (north side) on which stands Fort Charlotte, New Providence Island, Bahamas. Taxon 43. Strophia glans THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 397 grisea Maynard, 1894. Holotype MCZ 356674. Fields north of Fresh Creek, about 1 mile from the settlement, Andros, Bahamas. Taxon 44. Strophia regula Maynard, 1894. Neotype MCZ 10299. Fields on both sides of Fresh Creek, near the settlement, Andros, Bahamas. Taxon 45. Strophia bimarginata Maynard, 1894. Neotype USNM 420095. Green Key, east coast of Andros, Bahamas. Taxon 46. Strophia bimarginata cera Maynard, 1894. Holotype MCZ 76379. Green Key, east coast of Andros, Ba- hamas. Taxon 47. Strophia pilsbryi Maynard, 1894. Holotype MCZ 10297. Eastern half of Goat Key, Middle Bight, Andros, Bahamas. Taxon 48. Strophia pilsbryi evolva Maynard, 1894. Holotype MCZ 76383. Western end of Goat Key, Middle Bight, Andros, Bahamas. 398 have been labeled prominently. At least some specimens of this taxon would have been retained at the MCZ had they been recognizably labeled in Maynard's collec- tion. Nor was this taxon present among the material sent to USNM after the acquisi- tion of Maynard’s collection. The Field Museum of Natural History has two spec- imens (FMNH 42208) that were collected by Maynard in 1893 and are paratypes. Because the holotype is clearly lost, the better preserved FMNH specimen, which also more closely approximates the mea- surements and illustration, is designated as the neotype to provide an objective stan- dard of reference for this species-group taxon that is consistent with Maynard's original concept of it. This neotype retains FMNH 42208, whereas the remaining specimen, a paratype, was recatalogued as FMNH 303191. Neotype Measurements. Length 34.3 mm, diameter (excluding lip) 12.2 mm; ap- erture height (including lip) 11.2 mm, ap- erture width (including lip and peristome) 8.7 mm. Type Locality. Key, 1 mile north of U Key at Allen’s Harbor [Exuma Group], Ba- hamas. Remarks. Maynard (1894a: 149) noted that no living specimens were found, and commented that the site had been inhab- ited by Lucayan Indians. Taxon 40. Strophia neglecta Maynard, 1894a [December]: 148-150 [pages printed out of order], text figs. 46A, B. Number of examined specimens not provided. Size of type .92 by .45 [inches; 23.4 by 11.4 mm]. Caption to text figure 45, “Strophia neglecta, A, front view, B, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Although the number of specimens examined was not mentioned in the de- scription, Maynard (1894a: 151) discussed variation in this taxon and noted that they Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 were abundant and that “all gathered in this locality were typical.” MCZ 76376 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and illustration of this taxon and is accompanied by a label in Maynard’s hand with the term “type” that includes refer- ences to figures published after this spe- cies was described. This specimen is here recognized as the holotype. Holotype Measurements. Length 22.60 mm, diameter (excluding lip) 11.2 mm; ap- erture height (including lip) 9.32 mm, ap- erture width (including lip and peristome) 8.44 mm. Type Locality. One mile west of Fort Charlotte, New Providence Island, Baha- mas. Remarks. Maynard (1894a: 151-152) commented that the field had been con- verted to agave cultivation and that after 9 years, S. neglecta “no longer exists,” having been replaced by S. neglecta agava. This taxon was illustrated as Strophiops neglec- ta (Maynard, 1913a: 177, 1919a: 4, fig. 3, pl. 5, figs. 4, 5). Maynard (1921b: 142) gave the number of specimens as 500 and commented that it was probably extinct. Maynard (1924c: 6) listed S. neglecta as extinct in 1913 from the burning of vege- tation prior to cultivation. Taxon 41. Strophia neglecta agava May- nard, 1894a [December]: 152, 1894b [December]: 153-154, text figs. 48A, B. Examined 500 specimens. Size of type as 1.21 by .47 [inches; 30.7 by 17.8 mm]. Caption to text figure 48, “Strophia neglecta agava, A, front, B, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot MCZ 76103, which is labeled “Holotype” and contains a single specimen accompanied by a note in Maynard’s hand- writing with the term “type,” is here rec- ognized as the Holotype (despite the 5% difference in length). The remaining 499 THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. specimens, including MCZ 76104 and MCZ 10296, are paratypes. Holotype Measurements. Length 29.2 mim, diameter (excluding lip) 12.6 mm; ap- erture height (including lip) 11.7 mm, ap- erture width (including lip and peristome) 9.6 mm. Type Locality. Sisal fields west of Nas- sau, New Providence Island, Bahamas. Remarks. Originally proposed as a sub- species, this taxon was elevated to a full species by Maynard (1913c: 191). The original illustration was subsequently re- published as Strophiops agava (Maynard, 1913c: 200, 1919¢, fig. 24, 1920c: 104, fig, 87, pl. 19, fig. 20 [apertural view only]). Clench (1957: 136) considered this taxon to be a synonym of Cerion coryi (Maynard, 1894). Gould and Woodruff (1986: 474) had examined the holotype and synony- mized it with C. glans (Kiister, 1844). Taxon 42. Strophia carlotta Maynard, 1894b [December]: 154-156, text figs. 49A, B. Examined 1,000 speci- mens. Size of type .85 by .40 [inches; 21.6 by 10.2 mm]. Caption to text figure 49, “Strophia carlotta, A, front, B, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. How many of the remaining 999 specimens are paratypes is uncertain (see Remarks). Lot MCZ 10249 was labeled “paratype” and contained two specimens that were acquired from Maynard in 1895. One of these specimens closely resembles the illustration, but both length and width are exactly 1 mm longer than the pub- lished measurements. This specimen is considered to be the figured specimen and the originally designated holotype. The re- maining specimen, a paratype, was reca- talogued as MCZ 356987. There were no MCZ catalog entries for this taxon in the material from Maynards estate. Holotype Measurements. Length 22.6 mm, diameter (excluding lip) 11.2 mm; ap- erture height (including lip) 8.7 mm, ap- erture width (including lip and peristome) 7.6 mm. Type Locality. At the foot of the hill (north side) on which stands Fort Char- lotte, New Providence Island, Bahamas. Remarks. Maynard (1894b: 156) distin- guished two forms [Nos. | and 2] without naming them. Two specimens of MCZ 10250 are labeled form No. 1, whereas MCZ 10251 contains four specimens la- beled form No. 2. Specimens representing forms No. | and 2 are not paratypes of Strophia carlotta [Article 72.4, ICZN, 1999: 76]. Maynard (1913c: 192) recog- nized that this is a synonym of S. glans (Kiister, 1844), as did Gould and Woodruff (1986: 474). Taxon 43. Strophia glans grisea Maynard, 1894b [December]: 159-161, text figs. SLA, B. Examined 200 specimens. Size of type 1.10 by .45 [inches; 27.9 by 11.4 mm]. Caption to text figure 48, “Strophia glans grisea, A, front view of type, B, margin of same.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. A search of the MCZ catalog indi- cated that only one lot containing 222 specimens was present in Maynard’s col- lection at the time it was acquired from his daughter. This lot, MCZ 76085, was catalogued as “paratypes” and did not con- tain a label in Maynard’s handwriting. These specimens were compared against the published measurements and illustra- tion. Although none matched exactly, a single specimen was identified that most closely approximated the figures and mea- surements. This specimen is considered to be the originally designated holotype and has been recatalogued as MCZ 356674. Holotype Measurements. Length DTS mm, diameter (excluding lip) 11.6 mm; ap- erture height (including lip) 11.2 mm, ap- erture width (including lip and peristome) 9.6 mm. 400 Type Locality. Fields north of Fresh Creek, about 1 mile from the settlement Andros, Bahamas. Remarks. Maynard (1894b: 161) noted that this taxon was also found south of the creek and that these specimens “were un- doubtedly transplanted from the north side through the agency of the inhabitants, who pass from one set of fields to the oth- er, carrying with them the plants of the casava on which the Strophias frequently OCCuline Taxon 44. Strophia regula Maynard, 1894b [December]: 161-164, text figs. 52A, B. Examined 50 specimens. Size of type 1.50 by .60 [inches; 38.1 by 15.2 mm]. Caption to text figure 52, “Regular Strophia. A, side, B, front view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. The remaining 49 specimens are paratypes. A review of the MCZ catalog revealed that Maynard’s collection con- tained one lot of 25+ specimens [MCZ 76381] that had been catalogued as “para- types.” Neither this lot nor any single spec- imen identified as the holotype could be found at the MCZ. Lot MCZ 10299 con- tained two specimens labeled “paratype” that had been purchased from Maynard in 1895. Both specimens have damage, indi- cating they had been occupied by hermit crabs, but neither closely matched the fig- ure or published measurements of the ho- lotype. The less damaged specimen retains MCZ 10299 and is here designated as the neotype of Strophia regula to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of it. The remaining specimen, a paratype, has been recata- logued as MCZ 356988. Neotype Measurements. Length 33.3 mm, diameter (excluding lip) 14.4 mm; ap- erture height (including lip) 12.5 mm, ap- erture width (including lip and peristome) 10.5 mm. Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Type Locality. Fields on both sides of Fresh Creek, near the settlement Andros, Bahamas. Remarks. Maynard (1894b: 162) report- ed that this taxon is known only from shells inhabited by hermit crabs and suggested that the species is extinct. Maynard (1919c: 68, caption to fig. SOEs IPAS Jee 1924?: [7]) referred to this taxon as S. reg- ular. Taxon 45. Strophia bimarginata Maynard, 1894b [December]: 164-168, text figs. 53A, B, C, D. Examined 2,000 specimens. Size of type 1.50 by .60- [inches; 38.1 by 15.2 mm]. Caption to text figure 53, “Strophia bimarginata. A, front view; B, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. How many of the remaining 1,999 specimens are paratypes is uncertain (see Remarks). No specimen labeled holotype could be found at the MCZ. Lot MCZ 10246 contains two specimens labeled “paratype.” Neither of these specimens matches the measurements nor has the “double margin” evident in figure 53B. The holotype is presumed to be lost. Lot USNM 420095 contains over 200 para- types. A specimen that closely approxi- mates the measurements and the pub- lished figures is here designated as neo- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of this taxon. This specimen retains USNM 420095. The remaining paratypes have been recatalogued under USNM 1093843. Other paratypes include MCZ 76105 (202 specimens), MCZ 10244 (two specimens), and MCZ 10245 (two specimens). Neotype Measurements. Length 27.5 mm, diameter (excluding lip) 11.4 mm; ap- erture height (including lip) 8.9 mm, ap- erture width (including lip and peristome) 7.4 mm. Type Locality. Green Key, east coast of Andros, Bahamas. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. Remarks. Maynard (1894b: 167) distin- guished two forms [Nos. 1 and 2] without naming them. He reported that form No. 1 also occurs on Little Galden Key. Spec- imens representing forms No. | and 2 are not paratypes of Strophia bimarginata | Ar- ticle 72.4, ICZN, 1999: 76]. Taxon 46. Strophia bimarginata cera May- nard, 1894b [December]: 168-170, text figs. 54A, B. Examined nine spec- imens. Size of type 1.00 by .40 [inches; 25.4 by 10.2 mm]. Caption to text figure 54, “Strophia bimarginata cera. A, front [sic, margin]; B, margin [sic, front] of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot MCZ 76379, which is labeled “Holotype” and contains a single specimen accompanied by a note in Maynard’s hand- writing with the term “type,” is here rec- ognized as the holotype. The remaining eight specimens, including MCZ 76380 and MCZ 10248, are paratypes. Holotype Measurements. Length 24.8 mm, diameter (excluding lip) 10.5 mm; ap- erture height (including lip) 9.1 mm, ap- erture width (including lip and peristome) 7.0 mm. Type Locality. Green Key, east coast of Andros, Bahamas. Taxon 47. Strophia pilsbryi Maynard, 1894b [December]: 170-172, text figs. 55A, B, text fig. 56 (map). Ex- amined 125 specimens. Size of type as 1.07 by .45 [inches; 27.2 by 11.4 mm]. Caption to text figure 55, “Strophia pils- bryi. A, front, B, side view, of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. The remaining 124 specimens are paratypes. No specimen labeled holotype could be found at the MCZ. Lot MCZ 10297 contained two specimens labeled “paratypes” that had been obtained from 401 Maynard in 1895. One of these specimens very closely matches the published mea- surements and figures of the holotype. This specimen is considered to be the originally designated holotype. The other specimen has been recatalogued as MCZ 356989. Holotype Measurements. Length 27.3 mm, diameter (excluding lip) 11.4 mm; ap- erture height (including lip) 10.1 mm, ap- erture width (including lip and peristome) 8.8 mm. Type Locality. Eastern half of Goat Key, Middle Bight, Andros, Bahamas (map, fig. HO, 4): Remarks. Maynard (1894b: 171) dis- cussed a single form but did not distin- guish it with either a name or number. Taxon 48. Strophia pilsbryi evolvua May- nard, 1894b [December]: 173-175, text figs. 57A, B, C. Examined 70 spec- imens. Size of type 1.12 by .43 [inches; 28.4 by 10.9 mm]. Caption to text figure 5, “Strophia pilsbryi evolva. A, front view, B, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot MCZ 76383 contains a single specimen labeled “Holotype” that closely approximates the illustration and the mea- surements of this taxon. This specimen is recognized as the holotype. The remaining 69 specimens are paratypes. Holotype Measurements. Length 28.4 mm, diameter (excluding lip) 11.5 mm; ap- erture height (including lip) 10.7 mm, ap- erture width (including lip and peristome) 9.0 mm. Type Locality. Western end of Goat Key, Middle Bight, Andros, Bahamas (map, fig. 56,*). Taxon 49. Strophia restricta Maynard, 1894b [December]: 175-177, text figs. 58A, B (map, p. 172, fig. 56). Ex- amined 75 specimens. Size of type .75 by .32 [inches; 19.1 by 8.1 mm]. Caption 402 to text figure 58, “Strophia restricta. A, front, B, side view.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. There is no specimen at the MCZ labeled “holotype.” MCZ 76384 contained 33 specimens and labels stating “para- type,” as well as six handwritten labels, seemingly by Maynard. All bear the taxon name, one contains the term “types,” the remainder are numbered 1, 3, 4, 5, 7. None of the specimens match the pub- lished measurements. However, neither do the published illustrations. The specimen most closely resembling the measure- ments, figures, and proportions is consid- ered to be the Holotype. The remaining 32 specimens have been recatalogued as MCZ 356990. Holotype Measurements. Length 20.2 mm, diameter (excluding lip) 9.3 mm; ap- erture height (including lip) 7.4 mm, ap- erture width (including lip and peristome) 6.6 mm. Type Locality. Western end of Goat Key, Middle Bight, Andros, Bahamas (Maynard, 1894b, fig. 56, }). Remarks. Maynard (1894b: 177) report- ed that this taxon was “confined to a single tree and the bushes that come in contact with it.” Maynard (1894b: 176) distin- guished a single form [No. 1] without naming it. Specimens representing this form [No. 1] are not paratypes of Strophia restricta [Article 72.4, ICZN, 1999: 76]. Two specimens of MCZ 10301 are labeled form No. 1 and therefore are not para- types. Taxon 50. Strophia eximea Maynard, 1894b [December]: 177-179, text figs. 59A, B, 61A, B. Examined 1,000 specimens. Size of type as 1.12 by .45 Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 [inches; 28.5 by 12.8 mm]. Caption to text figure 59, “Strophia eximia. A, front, B, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot MCZ 76243 contains a single specimen labeled “Holotype” and closely approximates the published measurements and figure. A scrap of cardboard labeled “type” by Maynard accompanies this spec- imen, which is here recognized as the ho- lotype. Lot USNM 420038 contains 15 paratypes. How many of the remaining ~ 984 specimens are paratypes is uncertain (see Remarks). Holotype Measurements. Length 28.8 mm, diameter (excluding lip) 11.3 mm; ap- erture height (including lip) 11.1 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. “Procured by Mr. Curtiss in Nassau, from a creole who brought them from Cat Island.” Remarks. Maynard (1894b: 179) distin- guished a single form [No. 1] without naming it. Specimens representing this form |No. 1] are not paratypes of Strophia eximea [Article 72.4, ICZN, 1999: 76]. This taxon serves as the type species of the genus Multistrophia Maynard, (1894b: 179) and is later (Maynard, 1920a: 69, 1920d: 126) included within Multicostata (likely errors for Multistrophia). Taxon 51. Strophia agrestina Maynard, 1894b [December]: 179-182, text figs. 60A, B, C. Examined 1,000 spec- imens. Size of type .87 by .35 [inches; 22.1 by 8.9 mm]. Caption to text figure 60, “Strophia agristina. A, front, B, side view of type.” Type Material. The original description — Figures 49-60. Taxon 49. Strophia restricta Maynard, 1894. Holotype MCZ 76384. Western end of Goat Key, Middle Bight, Andros, Bahamas. Taxon 50. Strophia eximea Maynard, 1894. Holotype MCZ 76243. Cat Island, Bahamas. Taxon 51. Strophia agrestina Maynard, 1894. Holotype MCZ 76083. South side of New Providence Island, Bahamas, opposite Nassau. Taxon 52. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 403 Strophia scripta Maynard, 1896. Lectotype MCZ 10332. Cardenas, Cuba. Taxon 53. Strophia scripta obliterata Maynard, 1896. Lectotype 10335. Matanzas, Cuba, rocks close to the sea. Taxon 54. Strophia fastigata Maynard, 1896. Lectotype MCZ 10314. Matanzas, Cuba. Taxon 55. Strophia eurystoma Maynard, 1896. Lectotype MCZ 10312. Havana, Cuba. Taxon 56. Strophia eurystoma ignota Maynard, 1896. Lectotype MCZ 10313. Havana, Cuba. Taxon 57. Strophia marmorata polita Maynard, 1896. Lectotype MCZ 189792. Cabo Cruz, Cuba. Taxon 58. Strophia media Maynard, 1896. Lectotype MCZ 10323. Cuba. Taxon 59. Strophia ferruginea Maynard, 1896. Lectotype MCZ 76230. Washed up by the tide near Jeremie, southwestern Haiti. Taxon 60. Strophia obscura Maynard, 1896. Lectotype MCZ 76411. Cayo Birde del Norte, Cuba. 404 provided measurements for a single “type” specimen, the holotype, which was illus- trated. Lot number MCZ 76083 contains a single specimen labeled “holotype,” as well as a label with the term “Type” in Maynard's hand. It approximates the orig- inal measurements and illustration and is here recognized as the Holotype. Lot number USNM 420134 contains 200 para- types. How many of the remaining 799 specimens are paratypes is uncertain (see Remarks). Holotype Measurements. Length 22.8 mm, diameter (excluding lip) 9.2 mm; ap- erture height (including lip) 8.9 mm, ap- erture width (including lip and peristome) (2) aaa Type Locality. South side of New Prov- idence Island, Bahamas, opposite Nassau. Remarks. The taxon name originally ap- peared as Strophia agrestina in the head- ing of the original description, but as S. agristina in the caption to the figure within the body of the description. The name ap- peared as S. agrestina in Maynard (1919b: 50, figs. 17, 17a, 1921b: 150, 1924c: 6) but as S. agsestina in Maynard (1919b, pl. 7, figs. 11-13). Maynard (1919b: 50) acted as the First Revisor of spellings [Article 24.2.4, ICZN, 1999: 30-31], fixing the spelling as agres- tind. Maynard (1894b: 179) distinguished three forms [Nos. 1-3] without naming them. Specimens representing forms No. 1-3 are not paratypes of Strophia agresti- na [Article 72.4, ICZN, 1999: 76]. May- nard (1921b: 150) noted slightly larger measurements .90 by .35 [inches; 22.9 by 8.9 mm], reported a larger number (2,000) of specimens, presumably from subse- quent collections, and cited previously published illustrations (Maynard, 1919b: 52, figs. 17, pl. 7, figs. 11-13). Maynard (1924c: 6) listed S. agrestina as extinct in 1924 from the burning of vegetation prior to cultivation. Taxon 52. Strophia scripta Maynard, 1896 [March]: 3-4, pl. 1, figs. 3, 4. Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Examined 29 specimens. Size of type 1.30 by .47 [inches; 33.0 by 11.9 mm]. Description noted, “Plate 1, fig. 3, front, fig. 4, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, which was illustrated. Maynard (1896: 1) stated that he returned this fig- ured type specimen to the MCZ but re- tained a co-type in his collection. Thus, these two specimens comprise the syntype series. Lot number MCZ 10332 contained two specimens labeled “holotype.” The | larger of the two specimens very closely approximates the originally published measurements and illustrations and is the figured specimen Maynard returned to the MCZ. This specimen is here designated as the lectotype to provide an objective stan- dard of reference for this species-group taxon that is consistent with Maynard's original concept of it. It is unclear whether the smaller specimen [25.8 by 9.2 mm] is the syntype retained by Maynard and re- acquired by the MCZ after his death or one of the 27 remaining, originally men- tioned specimens, which, according to Ar- ticle 72.4.6 (ICZN, 1999: 77) are expressly excluded from the type series. There are no paratypes for this taxon, and any spec- imens labeled as such are mislabeled. Lots MCZ 26498 (six specimens) and MCZ 76176 (five specimens) have no status as types. Lectotype Measurements. Length 32.5 mm, diameter (excluding lip) 11.9 mm; ap- erture height (including lip) 11.0 mm, ap- erture width (including lip and peristome) 9.1 mm. Type Locality. Cardenas, Cuba. . Remarks. Maynard (1896: 4) distin- guished a single form [No. 1], composing 20% of the sample, without naming it. Maynard (1919b, pl. 7, figs. 3, 4) reillus- trated this taxon. Taxon 53. Strophia scripta obliterata May- nard, 1896 [March]: 5-6, pl. 1, figs. 5, 6. Examined 102 specimens. Size of THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. type 1.08 by .40 [inches; 27.4 by 10.2 mm]. Description noted, “Plate I, fig. 5, front, fig. 6, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, which was illustrated. Maynard (1896: 1) stated that he returned this fig- ured type specimen to the MCZ but re- tained a co-type in his collection. Thus, these two specimens compose the syntype series. Lot MCZ 10335, which was labeled “holotype,” contains a single specimen ac- companied by a note in Maynard's hand- writing identifying the taxon and the term “type.” This specimen, which closely matches the published measurements and figures, is here designated as the lectotype to provide an objective standard of refer- ence for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 100 specimens, in- cluding MCZ 26502, MCZ 26503, MCZ 26504, and MCZ 26505, are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any speci- mens labeled as such are mislabeled. Lectotype Measurements. Length 27.5 mm, diameter (excluding lip) 10.1 mm; ap- erture height (including lip) 9.2 mm, ap- erture width (including lip and peristome) Tigoeemany Type Locality. Matanzas, Cuba, rocks close to the sea. Remarks. Maynard (1896: 4) distin- guished five forms [Nos. 1-5] without naming them. Taxon 54. Strophia fastigata Maynard, 1896 [March]: 6-7, pl. 2, figs. 1, 2. Examined 62 specimens. Size of type 1.20 by .40 [inches; 30.5 by 10.2 mm]. Description noted, “Plate II, fig. 1, front, fig. 2, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, which was illustrated. Maynard (1896: 1) stated that he returned this fig- ured type specimen to the MCZ but re- 405 tained a co-type in his collection. Thus, these two specimens compose the syntype series. Lot MCZ 10314 contains a single specimen labeled “Holotype,” with one la- bel annotated “id. C. J. Maynard” and an- other with the word “type,” initialed by Maynard. Another lot, MCZ 76293, con- tains a single specimen and was labeled “Paratype.” Although neither specimen closely matches the published measure- ments, the specimen in MCZ 10314 is closer and approximates the illustration of the type. This is the specimen returned to the MCZ by Maynard and is here desig- nated as the lectotype to provide an ob- jective standard of reference for this spe- cies-group taxon that is consistent with Maynard’s concept of this taxon. The sec- ond specimen (MCZ 76293) was part of Maynard’s collection at the time of his death and is the co-type that he retained. It becomes the sole paralectotype. The re- maining 60 specimens (including MCZ LOSTA LT MICZn16298. MEZ. 26513. and MCZ 26515) are, according to Article 72.4.6 (ICZN, 1999: 77), expressly exclud- ed from the type series. Therefore, there can be no paratypes, and any specimens labeled as such are mislabeled. Lectotype Measurements. Length 28.7 mm, diameter (excluding lip) 9.8 mm; ap- erture height (including lip) 10.4 mm, ap- erture width (including lip and peristome) 8.8 mm. Type Locality. Maynard (1896: 7) wrote, “This species I found in the collection in the same tray with the shells which I have named S. scripta obliterata, therefore it is to be presumed that they came from Ma- tanzas, Cuba, but as the shells of S. fasti- gata are deeply stained with a red earth of which those of S. s. obliterata bear no trace, it is evident that they did not come from “rocks close to the sea” as did the others, but probably from some neighbor- ing field. Another lot was labeled Punte Goade, Matanzas, and still another lot Chorrea.” Clench (1957: 145) listed the type locality as “Matanzas [Havana] Cuba.” The lectotype is from Matanzas, 406 Cuba, which is the type locality (Article 76.2, ICZN, 1999: 87). Remarks. Maynard (1896: 7) distin- guished a single form [No. 1] without naming it. Taxon 55. Strophia eurystoma Maynard, 1896 [March]: 7-9, pl. 2, figs. 3, 4. Examined 21 specimens. Size of typical specimen 1.35 by .40 [inches; 34.3 by 10.2 mm]. Description noted, “Plate II, fig. 3, front, fig. 4, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, which was illustrated. Maynard (1896: 1) stated that he returned this fig- ured type specimen to the MCZ but re- tained a co-type in his collection. Thus, these two specimens compose the syntype series, The remaining 19 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type se- ries. Therefore, there can be no paratypes, and any specimens labeled as such are mislabeled. Lot MCZ 10312 contains a single specimen labeled “Holotype,” with the label annotated “id. C. J. Maynard.” This specimen matches closely the mea- surements and figures of the specimen identified as the type. It is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s concept of this taxon. Lectotype Measurements. Length 34.4 mim, diameter (excluding lip) 11.5 mm; ap- erture height (including lip) 12.6 mm, ap- erture width (including lip and peristome) 10.5 mm. Type Locality. Havana, Cuba. Remarks. Maynard (1896: 8-9) distin- guished a single form [No. 1], which he provisionally named Strophia eurystoma ignota (see below). The taxon name also appeared as S. eryrostoma (Maynard, 1919c: 54). Taxon 56. Strophia eurystoma ignota May- nard, 1896 [March]: 8-9. Not illus- trated. Nine specimens [two from the Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 MCZ, seven from another source] listed in the description of form No. 1| of Stro- phia eurystoma, but none measured. Type Material. Maynard (1896: 1) stated that he returned figured type specimens to the MCZ and retained a co-type in his col- lection. Because this taxon was not figured, there is no indication that type material was designated. The original description lists a total of nine specimens, which are all syntypes. Lot MCZ 10313 contains a single specimen labeled “Holotype,” with the label annotated “id. by C. J. Maynard.” This specimen is here designated as the © lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of this taxon. The remaining eight speci- mens, including the three specimens in MCZ 76228, become paralectotypes. Lectotype Measurements. Length 35.9 mm, diameter (excluding lip) 10.6 mm; ap- erture height (including lip) 11.3 mm, ap- erture width (including lip and peristome) 9.3 mm. Type Locality. Havana, Cuba. Remarks. Maynard (1896: 8—9) distin- guished a single form [No. 1] and noted that the typical form is stained by red ochraceous earth, whereas form No. | is stained by brown earth or without staining. On this basis, Maynard supposed that the two did not occur together. He wrote (p. 9), “Should this prove true, form No. 1 must take subspecific rank, in which case it may be called Strophia eurystoma ig- nota.” Taxon 57. Strophia marmorata polita Maynard, 1896 [March]: 14-15, pl. 3, figs. 3, 4. Examined 25 specimens. Size of type 1.05 by .45 [inches; 26.7 by 11.4 mm]. Description noted, “Plate II, fig. 3, front, fig. 4, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, which was illustrated. Maynard (1896: 1) stated that he returned this fig- ured type specimen to the MCZ but re- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. tained a co-type in his collection. Thus, these two specimens compose the syntype series. The remaining 23 specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type se- ries. Therefore, there can be no paratypes, and any specimens labeled as such are mislabeled. Lot MCZ 189792 contains a single specimen labeled “Holotype” and matches both the published measurements of the type and its illustration. This spec- imen has a hole in the dorsal surface of the final whorl that reveals the details of the columellar and apertural teeth. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of this taxon. A second specimen, MCZ 76201, was labeled “paratype.” This spec- imen was part of Maynard's collection at the time of his death and is the remaining syntype, which becomes a paralectotype. Lectotype Measurements. Length 26.1 mm, diameter (excluding lip) 11.6 mm; ap- erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. Cabo Cruz, Cuba. Remarks. Maynard (1896: 15) distin- guished a single, larger form [No. 1] with- out naming it. Maynard (1919b, pl. 8, figs. 7, 8) reillustrated this taxon. Taxon 58. Strophia media Maynard, 1896 [March]: 18-19, pl. 4, figs. 3, 4. Ex- amined two specimens. Size of type 1.20 by .50 linches; 30.5 by 12.7 mm]. De- scription noted, “Plate IV, fig. 3, front, fig. 4, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, which was illustrated. Maynard (1896: 1) stated that he returned this fig- ured type specimen to the MCZ but re- tained a co-type in his collection. Thus, the two specimens comprise the syntype se- ries. Lot number MCZ 10323 contains a single specimen labeled “Holotype” with 407 the label annotated “ex. C. J. Maynard.” This specimen matches closely the mea- surements and figures of the specimen identified as the type. It is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s concept of this taxon. Lot number MCZ 76225 contains two specimens la- beled “paratype.” The MCZ catalog num- ber indicates that these specimens were among, those purchased from Maynard's daughter after his death. The dead speci- men is likely the remaining syntype and therefore a paralectotype. The live collect- ed specimen might have been added after the original description and would not have any status as a type. Lectotype Measurements. Length Oli mm, diameter (excluding lip) 12.2 mm; ap- erture height (including lip) 12.2 mm, ap- erture width (including lip and peristome) 10.6 mm. Type Locality. Cuba. Taxon 59. Strophia ferruginea Maynard, 1896 [March]: 19-21, pl. 4, figs. 5, 6. Examined nine specimens. Size of type .90 by .25 [inches; 22.9 by 6.4 mm]. De- scription noted, “Plate IV, fig. 5, front, fig. 6, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, which was illustrated. Maynard (1896: 1) stated that he returned this fig- ured type specimen to the MCZ but re- tained a co-type in his collection. Thus, the two specimens compose the Syntype se- ries. The remaining seven specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type se- ries. Therefore, there can be no paratypes, and any specimens labeled as such are mislabeled. Lot MCZ 76230 contains a single specimen labeled “Holotype” and approximates the published length (but not width, which is likely a typographical error) and illustrations of this taxon. This specimen is here designated as the lecto- 408 type to provide an objective standard of reference for this species-group taxon. Lectotype Measurements. Length 21.46 mm, diameter (excluding lip) 9.25 mm; ap- erture height (including lip) 8.61 mm, ap- erture width (including lip and peristome) Oevounmaiaie Type Locality. “Washed up by the tide near Jeremie, Cote de Fer.” Maynard (1896: 21) noted that “Jeremie is in Hayti, on the southwest portion,” and that “Cote de Fer” refers to iron shore. Taxon 60. Strophia obscura Maynard, 1896 [March]: 21-22, pl. 3, figs. 5, 6. Examined 50 specimens. Size of type 1.10 by .42 [inches; 27.9 by 10.7 mm]. Description noted, “Plate III, fig. 5, front, fig. 6, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, which was illustrated. Maynard (1896: 1) stated that he returned this fig- ured type specimen to the MCZ but re- tained a co-type in his collection. Thus, the two specimens compose the syntype se- ries. The remaining 4§ specimens are, ac- cording to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be no paratypes, and any specimens labeled as such are misla- beled. Lot number MCZ 76411 contains a single specimen labeled “holotype.” This specimen matches the length measure- ment more closely than the width mea- surement and approximates the figures of the specimen identified as the type. It is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Maynard’s concept of this taxon. Lots MCZ 10329 and MCZ 76410 each contain a specimen labeled “paratype.” Lot num- ber MCZ 10329 is the specimen that May- nard returned to MCZ during his lifetime. It was labeled “type” by Maynard but ap- proximates the published measurements and figure less closely than the lectotype. This specimen came from Cayo Piedras del Norte rather than Cayo Birde del Nor- te. It is here considered one of the two original syntypes and becomes a paralec- totype. Lot MCZ 76410 is from Cayo Bir- de del Norte and is likely one of the re- maining 48 specimens without type status. Lectotype Measurements. Length 27.9 mm, diameter (excluding lip) 11.1 mm; ap- erture height (including lip) 11.4 mm, ap- erture width (including lip and peristome) 9.6 mm. Type Locality. Cayo Birde del Norte, Cuba. Remarks. Maynard (1896: 22) distin- guished a form [No. 1] composing 20% of the sample without naming it. Specimens representing this form are not paratypes of Strophia obscura [Article 72.4, ICZN, 1999: 76]. Taxon 61. Strophia faxoni Maynard, 1896 [March]: 32-34, pl. 7, figs. 1, 2. S. faxsoni Maynard, 1920a: 73. Examined two specimens. Size of type 1.12 by .40 linches; 28.4 by 10.2 mm]. Description noted, “Plate Vite. le tires teas side view of type.” Type Material. The original description provided measurements for a single “type” specimen, which was illustrated. Maynard (1896: 1) stated that he returned this fig- — Figures 61-73. Taxon 61. Strophia faxoni Maynard, 1896. Lectotype MCZ 184649. Cuba. Taxon 62. Strophia cylindrica May- nard, 1896. Lectotype MCZ 10311, Mathewstown, Inagua, Bahamas. Taxon 63. Strophiops antiqua Maynard, 1913. Lectotype MCZ 76114. Eastern Nassau, New Providence Island, Bahamas. Taxon 64. Strophiops primigenia Maynard, 1913. Lectotype MCZ 76088. Fossil in limestone beneath a sand cliff on the east coast of Salt Key, New Providence Island, Bahamas. Taxon 65. Strophiops salinaria Maynard, 1913. Lectotype MCZ 76089. Salt Key, New Providence Island, Bahamas. Taxon 66. Stro- phiops larga Maynard, 1913. Lectotype MCZ 76139. Rose Island opposite Green Key, New Providence Island, Bahamas. Taxon 67. Strophiops affinis Maynard, 1913. Lectotype MCZ 76091. Sandy Key, New Providence Island, Bahamas. Taxon 68. Stro- phiops acceptoria Maynard, 1913. Lectotype MCZ 76095. Low Bay Key east of Rose Island, New Providence Island, Bahamas. Taxon 69. Strophiops cinereavaria Maynard, 1913. Lectotype MCZ 76395. East end of Hog Island [Paradise Island], New THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 409 Providence Island, Bahamas. Taxon 70. Strophiops variathorndikei Maynard, 1913. Lectotype USNM 359502. Cemetery east of Nassau, New Providence Island, Bahamas. Taxon 71. Strophiops varianivia Maynard 1913. Lectotype MCZ 76392. About Banyan tree in Eastern Cemetery west along Shirley St. to St. Paul Quarry, Nassau, New Providence Island, Bahamas. Taxon 72. Strophiops nivia Maynard, 1913. [Not illustrated.] The holotype (MCZ 010279) of Strophia curtissii nivia Maynard, 1894 [Taxon 26], is designated as the lectotype of Strophiops nivia Maynard, 1913, making Strophiops nivia Maynard, 1913, an objective junior synonym and junior homonym of Strophia curtissii nivia Maynard, 1894. Taxon 73. Strophiops rediviva Maynard, 1913. Lectotype MCZ 76093. In field just west of St. Paul Quarry, East Nassau, New Providence Island, Bahamas. 410 ured type specimen to the MCZ but re- tained a co-type in his collection. Thus, the two specimens compose the syn e se- ries. Clench and Aguayo (1952: 420, pl. 55, fig. 2) listed and illustrated MCZ 184649 as the “holotype.” This specimen matches closely Maynard’s published measure- ments and illustration and is accompanied by two MCZ labels. One label, with MCZ 10363 crossed out and replaced by 184649, is annotated “Type” and “id. C. J. Maynard.” The other label bears the num- ber MCZ 184649 and is annotated “ex. C. J. Maynard, Acc. 766” and labeled “Ho- lotype.” Article 74.5 (ICZN, 1999: 82-83) states, “When the original work reveals that the taxon had been based on more than one specimen, a subsequent use of the term “holotype” does not constitute a valid lectotype designation unless the au- thor, when wrongly using that term, ex- plicitly indicated that he or she was se- lecting from the type series that particular specimen to serve as the name-bearing type.” Thus, there is no holotype for this taxon, nor can Clench and Aguayo’ treat- ment of MCZ 184649 be considered a lec- totype designation. This specimen |MCZ 184649] is here designated as the lectotype to provide an objective standard of refer- ence for this species-group taxon that is consistent with Maynard’s original and Clench and Aguayo’s subsequent concepts of this taxon. Lectotype Measurements. Length 28.9 mm, diameter (excluding lip) 10.2 mm; ap- erture height (including lip) 9.6 mm, ap- erture width (including lip and peristome) 8.5 mm. Type Locality. Cuba. Remarks. Maynard (1896: 34) com- mented that “Although these specimens were simply labeled ‘Cuba, I have no doubt but what they came from the neigh- borhood of Gibara, Cuba.” Taxon 62. Strophia cylindrica Maynard, 1896 [March]: 34-36, pl. 7, figs. 3, 4. Number of specimens examined not specified. Size of type 1.10 by .40 [inch- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 es; 27.9 by 10.2 mm]. Description not- ed, “Plate VII, fig. 3, front, fig. 4, side view of type.” Type Material. The original description provided measurements for a single “type” specimen, which was illustrated. Maynard (1896: 1) stated that he returned this fig- ured type specimen to the MCZ but re- tained a co-type in his collection. Thus, these two specimens compose the syntype series. Of these, MCZ 10311, which was labeled “Holotype” and contains a single specimen accompanied by a note in May- | nard’s handwriting identifying the taxon but not using the term “type,” is here des- ignated as the lectotype to provide an ob- jective standard of reference for this spe- cies-group taxon that is consistent with Maynard’s original concept of it. The un- specified number of additional specimens are, according to Article 72.4.6 (ICZN, 1999: 77), expressly excluded from the type series. Therefore, there can be but one paralectotype. Lectotype Measurements. Length 27.2 mm, diameter (excluding lip) 9.8 mm; ap- erture height (including lip) 9.0 mm, ap- erture width (including lip and peristome) 7.7 mm. Type Locality. Mathewstown, Inagua [Bahamas], from town border to salina. Remarks. Maynard (1896: 35) distin- guished four forms [Nos. 1—4] without naming them. Maynard (1919b, fig. 13, pl. 7, figs. 1, 2) reillustrated this taxon. Clench (1959: 45) listed this taxon as a synonym of Cerion (Diacerion) rubicundum (Men- ke, 1829) but did not select a lectotype. Taxon 63. Strophiops antiqua Maynard, 1913b [February 1]: 183-184 [not il- lustrated in original publication]. Ex- amined 375 specimens. Size given as 1.25 by .50 [inches; 31.8 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 375 speci- mens examined, and there was no mention of type material in the other parts of this THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. publication. Thus, all 375 specimens are syntypes. Lot number MCZ 76114 con- tains a single specimen labeled “holotype.” Although the width of this specimen matches the published measurement, the length differs by nearly 10%. The speci- men is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consis- tent with Maynard’s original concept of it. The remaining 374 specimens, including MCZ 76115 and MCZ 117863, become paralectotypes. Lectotype Measurements. Length 29.0 mm, diameter (excluding lip) 12.7 mm; ap- erture height (including lip) 11.1 mm, ap- erture width (including lip and peristome) 9.9 mm. Type Locality. Eastern Nassau [New Providence Island, Bahamas]. Fossil. May- nard’s handwritten note accompanying the type specimen states “crab holes, Water- loo.” Remarks. Maynard (1913b: 183-184) surmised that, “Neither this nor the above [S. agassizii] appear to be ancestors of any living form thus discovered.” Maynard (1921b: 151) reported 400 specimens and provided citations to illustrations (pl. 47, figs. 3, 4) that were never issued. Gould and Woodruff (1986: 480) reported ex- amining the “holotype” and provisionally considered this taxon to be a synonym of Cerion agassizti Dall, 1894. Taxon 64. Strophiops primigenia May- nard, 1913b [February 1]: 184 [not illustrated in original publication]. Ex- amined 100 specimens. Size given as 1.50 by .50 [inches; 38.1 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 100 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 100 specimens are syntypes. Lot number MCZ 76088 con- tained 56 specimens and was labeled “paratype.” This lot also contained labels 41] in Maynard’s hand identifying the speci- mens as “typical,” but none use the word “type” nor refer to the subsequently pub- lished illustrations. A large specimen that approximates the published measurements and subsequently published illustration is selected as the lectotype to provide an ob- jective standard of reference for this spe- cies-group taxon that is consistent with Maynard's concept of it. The remaining 55 specimens were recatalogued as MCZ 356992 and are paralectotypes. Lectotype Measurements. Length 35.0 mm, diameter (excluding lip) 12.4 mm; ap- erture height (including lip) 11.2 mm, ap- erture width (including lip and peristome) 9.7 mm. Type Locality. Fossil in limestone be- neath a sand cliff on the east coast of Salt Key, New Providence Island, Bahamas. Remarks. Maynard (1913b: 184) consid- ered this to be the ancestor of all the spe- cies in the “Primigenia Group” [an “aggre- gate of species” sensu Article 10.4 (ICZN, 1999: 9) rather than a genus-group taxon]. Strophiops primigenia was subsequently illustrated (Maynard, 1924a, pl. 39, figs. 3, 4). Maynard (1921b: 142) provided slightly smaller measurements and a different lo- cality (east of Rose Island). Gould and Woodruff (1986: 481) doubted that this taxon was truly a fossil, and regarded it to be a synonym of Cerion glans (Kiister, 1844). Taxon 65. Strophiops salinaria Maynard, 1913b [February 1]: 184 [not illus- trated in original publication]. Exam- ined 500 specimens. Size given as 1.35 by .45 [inches; 34.3 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 500 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 500 specimens are syntypes. Lot number MCZ 76089 con- tains a single specimen that was cata- logued as the “Holotype” and that was il- 412 lustrated by Gould and Woodruff (1978, fig. 2) and captioned “holotype.” Gould and Woodruff’s caption cannot be accept- ed as a lectotype designation, as Article 74.5 (ICZN, 1999: 83) states, “When the original work reveals that the taxon had been based on more than one specimen, a subsequent use of the term ‘holotype’ does not constitute a valid lectotype designation unless the author, when wrongly using that term, explicitly indicated that he or she was selecting from the type series that par- ticular specimen to serve as the name- bearing type.” This specimen is here des- ignated as the lectotype to provide an ob- jective standard of reference for this spe- cies-group taxon that is consistent with Maynard's original concept of it. The re- maining 499 specimens, including MCZ 76090 and USNM 420055 (15 specimens), become paralectotypes. Lectotype Measurements. Length 30.5 mm, diameter (excluding lip) 11.9 mm; ap- erture height (including lip) 10.2 mm, ap- erture width (including lip and peristome) 5.4 mm. Type Locality. Living all over Salt Key [New Providence Island, Bahamas], and thousands dead in the sand cliff at the east end. Remarks. Maynard (1921b: 144, 1925, pl. 40, figs. 1, 2) published larger dimen- sions of 1.45 by .45 [inches; 36.8 by 11.4 mm] and illustrations of this taxon. Gould and Woodruff (1986: 475) regarded this taxon to be a synonym of Cerion glans (Ktister, 1844). Taxon 66. Strophiops larga Maynard, 1913b [February 1]: 184; Maynard, 1921b: 145; Maynard, 1925, pl. 41, figs. 3, 4. Examined 20 specimens [not illus- trated in original publication]. No mea- surements included in original descrip- tion. Type Material. The original description did not distinguish among the 20 speci- mens examined, and there was no mention of type material in the other parts of this Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 publication. Thus, all 20 specimens are syntypes. Lot number MCZ 76139 con- tains a single specimen labeled “holotype.” This lot also contains a label in Maynard’s hand with the term “type,” which includes references to illustrations (Maynard, 1925, pl. 41, figs. 3, 4) that were published 12 years after the species was described. This specimen approximates the subsequently published measurements and is the spec- imen figured in 1925 [a repaired break on the apertural surface is present in both fig- ure and specimen]. It is here designated as the lectotype to provide an objective — standard of reference for this species- group taxon that is consistent with May- nard’s concept of this taxon. The remain- ing 19 syntypes, including MCZ 76140, become paralectotypes. Lectotype Measurements. Length 32.1 mm, diameter (excluding lip) 12.8 mm; ap- erture height (including lip) 12.0 mm, ap- erture width (including lip and peristome) 10.4 mm. Type Locality. Rose Island opposite Green Key [New Providence Island, Ba- hamas |. Remarks. Maynard (1921b: 145) provid- ed measurements 1.35 by .52 inches; 34.3 by 13.2 mm] and references to subse- quently published illustrations but listed a slightly different locality, “West shore of Hog Island opposite Green Key,” and not- ed 25 specimens, some perhaps subse- quently collected. Gould and Woodruff (1986: 475) regarded this taxon to be a synonym of Cerion glans (Kiister, 1844). Taxon 67. Strophiops affinis Maynard, 1913b [February 1]: 184-185 [not il- lustrated in original publication]. Ex- amined 300 specimens. Size given as 1.15 by 60 [sic .60] [inches; 29.2 by 15.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 300 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 300 specimens are THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. syntypes. Lot number MCZ 76091 con- tains a single specimen labeled “holotype.” This lot also contains a label in Maynard’s hand with the term “Type” that includes references to illustrations (pl. 44, figs. 7, 8), which were never published. This spec- imen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consis- tent with Maynard’s concept of this taxon. The remaining 229 syntypes, including MCZ 76092, become paralectotypes. Lectotype Measurements. Length 30.9 mm, diameter (excluding lip) 12.3 mm; ap- erture height (including lip) 11.4 mm, ap- erture width (including lip and peristome) 9.4 mm. Type Locality. The original description listed this taxon as occurring on Sandy and Green Keys |New Providence Island, Ba- hamas], living under different environ- ments on each. Because the lectotype is from Sandy Key, the type locality is re- stricted to Sandy Key (Article 76.2, ICZN, 1999: 87). Remarks. Maynard (1921b: 144) provid- ed slightly different measurements [1.20 by .57 inches; 30.5 by 14.4 mm] and a ref- erence to figures [pl. 44, figs. 5, 6; these figure numbers differ from those on the label accompanying the lectotype] that were never published. Gould and Wood- ruff (1986: 474) had examined the “Ho- lotype” and synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 68. Strophiops acceptoria May- nard, 1913b [February 1]: 185 [not illustrated in original publication]. Ex- amined 200 specimens. Size given as .94 by .36 [inches; 23.9 by 9.1 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 200 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 200 specimens are syntypes. Lot number MCZ 76095 was la- beled “Holotype,” but contained two spec- 413 imens. One of the specimens approximates the measurements and subsequent figure [Maynard, 1925, pl. 41, figs. 5, 6] and is here selected as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's concept of this taxon. The re- maining 199 syntypes became co-mingled with additional specimens (see Remarks), so it is unclear which of the specimens in MCZ 76096 (223 specimens) and USNM 420075 (110 specimens), are paralecto- types. Lectotype Measurements. Length YE As mm, diameter (excluding lip) 10.6 mm; ap- erture height (including lip) 9.4 mm, ap- erture width (including lip and peristome) Tins) aman Type Locality. Low Bay Key east of Rose Island [New Providence Island, Ba- hamas]. Remarks. Maynard (1921b: 145) noted a larger size (1.00 by .40 [inches]), a larger number (528) of specimens, likely the re- sult of subsequent collections, and provid- ed references to figures (pl. 41, figs. 5, 6) that were issued in 1925. Gould and Woodruff (1986: 474) had examined the “Holotype” and synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 69. Strophiops cinereavaria May- nard, 1913b [February 1]: 185 [not illustrated in original publication]. Ex- amined 614 specimens. No measure- ments included in description. Type Material. The original description did not distinguish among the 614 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 614 specimens are syntypes. Lot number MCZ 76395 con- tained five specimens labeled “paratype.” Examination of the MCZ catalog revealed that no single specimen identified as a type was present in Maynard's collection when it was acquired from his daughter. Because neither measurements nor illustrations were ever published for this taxon, one of 414 the five specimens, the best preserved un- damaged adult, is selected to be the lec- totype to provide an objective standard of reference for this species-group taxon. The remaining specimens, recatalogued as MCZ 356476, become paralectotypes. Lectotype Measurements. Length YS). I mm, diameter (excluding lip) 11.9 mm; ap- erture height (including lip) 11.0 mm, ap- erture width (including lip and peristome) 9.9 mm. Type Locality. East end of Hog Island [Paradise Island] | New Providence Island, Bahamas]. Remarks. The taxon name originally ap- peared as S. cinerea-vara, but was used as S. cinerea-varia on the following page. A correction was published (Maynard, 1913c: 199) in which Maynard acted as the First Revisor of spellings [Article 24.2.4, ICZN, 1999: 30-31], selecting cineria-var- ia to be the correct spelling. Following Ar- ticle 32.5.2.4 (ICZN, 1999: 40), the hy- phen is removed and the taxon becomes cinereavaria. Gould and Woodruff (1986: 474) reported examining “paratypes” and synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 70. Strophiops variathorndikei Maynard, 1913b [February 1]: 186 [not illustrated in original publication]. Number of specimens examined not specified. No measurements included in description. Type Material. The original description did not specify the number of specimens examined, which are all syntypes. This tax- on was not represented in the collections of MCZ, nor was it present in Maynard’s collection at the time it was acquired from his daughter, as revealed by an examina- tion of the MCZ catalog. The USNM cat- alog revealed that USNM 359502 con- tained four syntypes that Bartsch had pur- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 ed to serve as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining three specimens, recatalogued as USNM 1093789, become paralectotypes. Lectotype Measurements. Length 19.1 mm, diameter (excluding lip) 8.4 mm; ap- erture height (including lip) 7.4 mm, ap- erture width (including lip and peristome) 6.4 mm. Type Locality. Cemetery east of Nassau [New Providence Island, Bahamas]. Remarks. Following Article 32.5.2.4 ~ (ICZN, 1999: 40) the hyphen is removed from varia-thorndikei and the taxon be- comes variathorndikei. A label accompa- nying the lectotype stated, “on an isolated grass plot, now extinct. 1915.” Gould and Woodruff (1986: 478) regarded this taxon to be a synonym of Cerion gubernatorium (Crosse, 1869). Taxon 71. Strophiops varianivia Maynard 1913b [February 1]: 186 [not illus- trated in original publication]. Number of specimens examined not specified. No measurements included in descrip- tion. Type Material. The original description did not specify the number of specimens examined, which are all syntypes. No spec- imen labeled holotype could be found at the MCZ. Lot MCZ 76392 contained 22 mostly juvenile specimens labeled “para- es.” One of the adult specimens in this lot is selected as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 21 specimens, recatalogued as MCZ 356675, become paralectotypes. Lectotype Measurements. Length 19.6 mm, diameter (excluding lip) 9.0 mm; ap- erture height (including lip) 8.0 mm, ap- erture width (including lip and peristome) chased from Maynard. Because neither measurements nor illustration were pub- lished, one of the specimens in this lot, a well-preserved undamaged adult, is select- 6.6 mm. Type Locality. About Banyan tree in Eastern Cemetery west along Shirley St. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. to St. Paul Quarry |Nassau, New Provi- dence Island, Bahamas]. Remarks. The first use of this name was in the form of a citation, as if it were pre- viously published. See remarks under Strophia curtissii nivia [Taxon 26]. Follow- ing Article 32.5.2.4 (ICZN, 1999: 40) the hyphen is removed from varia-nivia and the taxon becomes varianivia. Gould and Woodruff (1986: 478) regarded this taxon to be a synonym of Cerion gubernatorium (Crosse, 1869). Taxon 72. Strophiops nivia Maynard, 1913b [February 1]: 186 [not illus- trated in original publication]. Exam- ined 829 specimens. No measurements included in description. See remarks under Strophia curtissii nivia Maynard, 1894 [Taxon 26]. Nomenclatural Remarks. This taxon was described as new, appearing as “S. nivia new, whereas the preceding taxon, S. var- ia-nivia, was incorrectly listed as being previously described, appearing as “S. var- ia-nivia Mayn.” This is almost certainly an error in typesetting. Strophiops nivia May- nard, 1913, is a junior homonym of Stro- phia curtissti nivia Maynard, 1894. Type Material. The number of speci- mens listed in Maynard (1913b: 186, 1919b: 47) was substantially higher than the 40 listed in the original description of S. curtissti nivia (Maynard, 1894: 112). It is likely that Maynard added specimens in subsequent field work. The type localities for both uses of the name are identical ex- cept for phrasing, and only one series of specimens bearing the name nivia was present in Maynard’s collection at the time it was acquired from his daughter. The ho- lotype of Strophia curtissii nivia Maynard, 1894 (MCZ 10279), is here designated as the lectotype of Strophiops nivia Maynard, 1913, making Strophiops nivia Maynard, 1913, both a junior secondary homonym and an objective junior synonym of Stro- phia curtissii nivia Maynard, 1894. Remarks. Clench (1957: 155) listed S. 415 nivia as an error for S. nivea Maynard, 1894, rather than as a separately described taxon. Taxon 73. Strophiops rediviva Maynard, 1913c [February 9]: 187; Maynard, 1921b: 144; Maynard, 1925, pl. 40, figs. 5, 6 [not illustrated in original publica- tion]. Examined 500 specimens. No measurements included in description. Type Material. The original description did not distinguish among the 500 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 500 specimens are syntypes. Lot MCZ 76093 was labeled “Holotype” but contained two specimens and a label written by Maynard that con- tains the name of the taxon, the word “type,” and “Plate 40 figs. 5 6.” The spec- imen that closely resembles the published illustration is here selected as the lectotype to provide an objective standard of refer- ence for this species-group taxon that is consistent with Maynard’s concept of this taxon. The other specimen, recatalogued as MCZ 356478, and MCZ 76094 are among the 499 paralectotypes. Lectotype Measurements. Length 29.8 mm, diameter (excluding lip) 11.1 mm; ap- erture height (including lip) 10.5 mm, ap- erture width (including lip and peristome) 8.3 mm. Type Locality. In field just west of St. Paul Quarry, East Nassau [New Provi- dence Island, Bahamas]. Remarks. Maynard (1921b: 144) noted that the taxon was collected in 1897 but was extinct in 1921. Maynard (1924c: 6) listed S. rediviva as extinct in 1913 from the burning of vegetation prior to cultiva- tion. Maynard (1925, pl. 40, figs. 5, 6) pro- vided colored illustrations for this taxon. Gould and Woodruff (1986: 476) regarded this taxon to be “intermediate” between Cerion glans (Kiister, 1844) and C. gub- ernatorium (Crosse, 1869). Taxon 74. Strophiops variapurpurea May- nard, 1913c [February 9]: 188 [not 416 illustrated in original publication]. Ex- amined 59 specimens. No measure- ments included in description. Type Material. The original description did not distinguish among the 59 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 59 specimens are syntypes. No specimen labeled “Holotype” could be found at the MCZ. Lot MCZ 76286 contained 15 specimens labeled “paratypes.” One of these specimens, a well-preserved undamaged adult, was se- lected as the lectotype to provide an ob- jective standard of reference for this spe- cies-group taxon that is consistent with Maynard's original concept of it. The re- maining 58 specimens, including the 14 specimens recatalogued as MCZ 356676, become paralectotypes. Lectotype Measurements. Length 2oA mm, diameter (excluding lip) 10.1 mm; ap- erture height (including lip) 8.9 mm, ap- erture width (including lip and peristome) (Seman Type Locality. From the termination of Bay Street east to the Creek Settlement [Nassau, New Providence Island, Baha- mas]. Remarks. Following Article 32.5.2.4 (ICZN, 1999: 40), the hyphen is removed from varia-purpurea and the taxon be- comes variapurpurea. Gould and Wood- ruff (1986: 478) regarded this taxon to be a synonym of Cerion gubernatorium (Crosse, 1869). Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Taxon 75. Strophiops purpurea Maynard, 1913c [February 9|: 188-189; May- nard, 1921b: 150 [not illustrated in orig- inal publication]. Examined 50 speci- mens. Size given as 1.05 by .40 [inches; 26.7 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 50 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 50 specimens are syntypes. Lot MCZ 76108 contains a single specimen labeled “Holotype.” This lot also contains a label in Maynard’s hand with the term “type” that includes references to illustrations (pl. 46, figs. 2, 3) that were never published. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's concept of this taxon. The re- maining 49 syntypes become paralecto- types. Lot MCZ 76107 contains 92 speci- mens, indicating that at least 43 were col- lected after the taxon was described. Lectotype Measurements. Length 26.1 mm, diameter (excluding lip) 10.3 mm; ap- erture height (including lip) 10.1 mm, ap- erture width (including lip and peristome) 8.4 mm. Type Locality. Creek Settlement, east about 1 mile along the shore [Nassau, New Providence Island, Bahamas]. Remarks. Maynard (1919a: 24) mis- spelled the name as S. purpura. Maynard — Figures 74-86. Taxon 74. Strophiops variapurpurea Maynard, 1913. Lectotype 76286. Found from the termination of Bay Street east to the Creek Settlement, Nassau, New Providence Island, Bahamas. Taxon 75. Strophiops purpurea Maynard, 1913. Lectotype MCZ 76108. Creek Settlement, east about 1 mile along the shore, Nassau, New Providence Island, Bahamas. Taxon 76. Strophiops macularia Maynard, 1913. Lectotype 76127. South shore New Providence Island, Bahamas, from west side of first sound west 2 miles to a salina and northward along an old disused road running from the beach to Fox Hill, 1 mile. Taxon 77. Strophiops rufimaculata Maynard, 1913. Lectotype MCZ 76126. South shore of New Providence Island, Bahamas, from west side of salina to Sound Point, 2 miles. Taxon 78. Strophiops delicata Maynard, 1913. Lectotype MCZ 76099. South Key off Sound Point, New Providence Island, Bahamas. Taxon 79. Strophiops ultima Maynard, 1913. Lectotype 76154. Southwest Key, New Providence Island, Bahamas. Taxon 80. Strophiops avita Maynard, 1913. Lectotype MCZ 76394. Silver Key just west of Nassau Bar, New Providence Island, Bahamas, under a bed of limestone a foot thick, embedded in an ancient red soil. Taxon 81. Strophiops vetusta Maynard, 1913. Lectotype MCZ 117930. Fossil in the limestone of the three Silver Keys off Nassau Bar and of the Pimlico Keys off Eleuthera, Bahamas. Taxon 82. Strophiops argentia Maynard, 1913. Lectotype MCZ 76120. Living THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. A417 on the three Silver Keys, Nassau, New Providence Island, Bahamas. Taxon 83. Strophiops agavaneglecta Maynard, 1913. [Not illustrated.] The holotype (MCZ 76103) of Strophia neglecta agava Maynard, 1894 [Taxon 41], is designated as the neotype of Strophiops agavaneglecta Maynard, 1913, making Strophiops agavaneglecta Maynard, 1913, an objective junior synonym of Strophia neglecta agava Maynard, 1894. Taxon 84. Strophiops oberholseri Maynard, 1913. Lectotype MCZ 76124. Southwest Bay, New Providence Island, Bahamas. Taxon 85. Strophiops inconsueta Maynard, 1913. Lectotype MCZ 76123. Southern portion of Great Pimlico Island, Eleuthera, Bahamas. Taxon 86. Strophiops transitoria Maynard, 1913. Lectotype MCZ 117848. Northern portion of Great Pimlico Island, Eleuthera, Bahamas. 418 (1921b: 150) reported 705 mostly dead specimens collected from the back of a sandy beach, just southwest of East Point Light on New Providence Island and cited illustrations (pl. 46, figs. 2, 3) that were never issued. Maynard (1924?) listed this taxon as S. purpurea in the catalog, but in the supplement to the catalog (1924c: 6) reported S. “purpura” as extinct in 1924 from the burning of vegetation prior to cultivation. Gould and Woodruff (1986: 478) regarded this taxon to be a synonym of Cerion gubernatorium (Crosse, 1869). Taxon 76. Strophiops macularia Maynard, 1913c [February 9]: 189; Maynard, 1921b: 150 [not illustrated in original publication]. Examined 325 specimens. Size given as 1.18 by .45 [inches; 30.0 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 325 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 325 specimens are syntypes. Lot MCZ 76127 contains a single specimen labeled “holotype.” This lot also contains a label in Maynard’s hand with the term “type,” which includes references to illustrations (pl. 46, figs. 4, 5) that were never published. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of this taxon. The re- maining 324 syntypes, including MCZ 76128, become paralectotypes. Lectotype Measurements. Length LT mim, diameter (excluding lip) 11.0 mm; ap- erture height (including lip) 10.5 mm, ap- erture width (including lip and peristome) 8.4 mm. Type Locality. South shore New Provi- dence |Island, Bahamas] from west side of first sound west 2 miles to a salina and northward along an old disused road run- ning from the beach to Fox Hill, 1 mile. Remarks. Maynard (1921b: 150) provid- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 ed slightly smaller measurements (1.10 by A5 |inches; 27.9 by 11.4 mm]) and re- ported a larger number (2,041) of speci- mens, presumably from subsequent collec- tions. The illustrations cited (pl. 46, figs. 5, 6) were never issued. Gould and Woodruff (1986: 478) regarded this taxon to be a synonym of Cerion gubernatorium (Crosse, 1869). Taxon 77. Strophiops rufimaculata May- nard, 1913c [February 9]: 189-190; Maynard, 1921b: 150 [not illustrated in original publication]. Examined 325 | specimens. Size given as 1.25 by .50 inches; 32.8 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 325 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 325 specimens are syntypes. No specimen labeled “Holotype” could be found at the MCZ. Lot MCZ 76126 contained 136 specimens labeled “Paratypes.” Because this taxon was never illustrated, one of the syntypes that most closely approximates the published dimen- sions was selected as the lectotype to pro- vide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The remaining 135 specimens were _ recata- logued as MCZ 356474 and are among the remaining 324 syntypes that become par- alectotypes. Lectotype Measurements. Length Bone mm, diameter (excluding lip) 12.6 mm; ap- erture height (including lip) 12.6 mm, ap- erture width (including lip and peristome) WIL saonan Type Locality. South shore of New Providence [Island, Bahamas] from west side of salina to Sound Point, 2 miles. Remarks. Maynard (1921b: 150) cited il- lustrations (pl. 46, figs. 7, 8) that were nev- er issued. Gould and Woodruff (1986: 478) regarded this taxon to be a synonym of Cerion gubernatorium (Crosse, 1869). THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. Taxon 78. Strophiops delicata Maynard, 1913c [February 9]: 190; Maynard, 1921b: 151 [not illustrated in original publication]. Examined 107 specimens. Size given as 1.10 by .35 [inches; 27.9 by 8.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 107 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 107 specimens are syntypes. Lot MCZ 76099 was labeled “holotype” but contains two specimens ac- companied by a note in Maynard's hand- writing identifying the taxon and contain- ing the term “types (2),” as well as the no- tation “Plate 46 figs. 9-10.” Both speci- mens are nearly identical in measurements. One specimen (MCZ 76099) is here selected as the lectotype to provide an objective standard of reference for this species-group taxon that is consis- tent with Maynard's original concept of it. The other specimen was recatalogued as MCZ 356477 and is among the remaining 106 syntypes, including MCZ 76100, which become paralectotypes. Lectotype Measurements. Length 27.6 mm, diameter (excluding lip) 9.8 mm; ap- erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) 8.5 mm. Type Locality. South Key off Sound Point |New Providence Island, Bahamas]. Remarks. Maynard (1921b: 150) cited il- lustrations (pl. 46, figs. 9, 10) that were never issued. Gould and Woodruff (1986: 478) regarded this taxon to be a synonym of Cerion gubernatorium (Crosse, 1869). Taxon 79. Strophiops ultima Maynard, 1913c [February 9]: 190; Maynard, 1921b: 151 [not illustrated]. Number of specimens examined not specified [Maynard, 1921b: 151, mentioned 200 specimens]. Size given as .95 by .40 linches; 24.1 by 10.2 mm], but the term “type” was not used. 419 Type Material. The original description did not specify the number of specimens examined, which are all syntypes. Lot MCZ 76154 was labeled “Holotype” and contains a single specimen accompanied by a note in Maynard’s handwriting iden- tifying the taxon and containing the term “type” as well as the notation “Plate 45 figs. 1-2.” The illustrations were never published, nor were illustrations men- tioned in the subsequent treatment of the taxon (Maynard, 1921b: 151). This speci- men (MCZ 76154) approximates the pub- lished measurements and is here designat- ed as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. It is not clear how many of the subsequently mentioned 200 specimens were available to Maynard at the time he described this taxon. Lot MCZ 76153 contains 523 specimens, most of which could not have been part of the type series. Lectotype Measurements. Length DS, JL mim, diameter (excluding lip) 10.2 mm; ap- erture height (including lip) 9.8 mm, ap- erture width (including lip and peristome) Ouleeraninar Type Locality. No locality was men- tioned in the original publication. Maynard (1921b: 151) reported the locality as Southwest Key, New Providence Island, Bahamas. This is also the data accompa- nying the lectotype. Remarks. Maynard commented that this is “another fine example of recession with four species intervening between it and the species it mimics, two of them white!” Gould and Woodruff (1986: 474) had ex- amined “paratypes” and synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 80. Strophiops avita Maynard, 1913c [February 9]: 190 [not illus- trated in original publication]; Maynard, 1914a: front cover; Maynard, 1914b: front cover; Maynard, 1921a: 138; May- nard, !921\b, pl. 35, figs. 1, 2. Examined 90 specimens. Size given as 1.10 by .45 420 [inches; 27.9 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 50 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 50 specimens are syntypes. Lot MCZ 76394 was labeled “Holotype” and contains a single specimen accompanied by a note in Maynard’s hand- writing identifying the taxon and contain- ing the term “type.” It is within 7% of the measurements included in the original de- scription and approximates the subsequent illustration of this species on the front cov- ers of Maynard (1914a,b), more so than the later illustration (Maynard, 1921b, pl. 35, figs. 1, 2), which has a more conical spire. This specimen (MCZ 76394) is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The orig- inal description states that 50 specimens were examined; thus, the remaining 49, in- cluding MCZ 76141, become paralectoty- pes. However, Maynard, 1921a (p. 138), lists 55 specimens, some possibly subse- quently collected. Lectotype Measurements. Length 28.6 mm, diameter (excluding lip) 12.2 mm; ap- erture height (including lip) 12.2 mm, ap- erture width (including lip and peristome) 10.1 mm. Type Locality. Silver Key just west of Nassau Bar |New Providence Island, Ba- hamas]|. Found under a bed of limestone a foot thick embedded in an ancient red soil. Remarks. Maynard (1913c: 190) opined that this was “undoubtedly the oldest fossil Strophia yet found in the Bahamas.” Gould and Woodruff (1986: 474) synony- mized this taxon with Cerion agassizii Dall, 1894, but noted that the “Holotype” resembled C. glans (Kiister, 1844). Taxon 81. Strophiops vetusta Maynard, 1913c [February 9]: 191 [not illus- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 trated in original publication]; Maynard, 1921a: 138; Maynard, 1921b, pl. 35, figs. 3, 4. Examined 200 specimens. Size giv- en as 1.25 by .50 [inches; 31.8 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 200 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 200 specimens are syntypes. Lot MCZ 117930 was labeled “co-type” and contains a single specimen accompanied by a note in Maynard’s hand- ° writing dated “10/9/14,” identifying the taxon and containing the term “co-type.” This specimen is significantly (14%) short- er than the measurements included in the original description but approximates the subsequent illustration of this species (Maynard, 1921b, pl. 35, figs. 3, 4). This specimen (MCZ 117930) is here designat- ed as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. The remain- ing 199 syntypes, including MCZ 76129, become paralectotypes. Maynard (1921a: 138) listed 265 specimens, some possibly subsequently collected. Lectotype Measurements. Length 26.8 mm, diameter (excluding lip) 12.2 mm; ap- erture height (including lip) 11.2 mm, ap- erture width (including lip and peristome) 9.6 mm. Type Locality. Fossil in the limestone of the three Silver Keys off Nassau Bar |New Providence Island], and of the Pimlico Keys off Eleuthera [Bahamas]. The label accompanying the lectotype states Silver Key, thus restricting the type locality. Remarks. Clench (1952: 110) listed this taxon as a junior synonym of Cerion in- consuetum (Maynard, 1913). Gould and Woodruff (1986: 482) listed this taxon as a synonym of C. glans (Kiister, 1844). Taxon 82. Strophiops argentia Maynard, 1913c [February 9]: 191 [not illus- trated in original publication]. S. ar- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. entea Maynard, 1921a: 138; Maynard, 1921b: 139, pl. 35, figs. 5, 6 [S. argentea in plate caption]. Examined 400 speci- mens. Size given as 1.20 by .45 [inches; 30.5 by 11.43 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 400 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, there is no holotype for this taxon, and the species is based on 400 syntypes. Lot MCZ 76120 was labeled “Holotype” and contained a single speci- men accompanied by a note in Maynard's handwriting that identified the taxon and contained a reference to subsequently published illustrations (Maynard, 1921b, pl. 35, figs. 5, 6) but did not include any form of the word “type.” This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 399 syntypes (including MCZ 76121, MCZ 118064, and USNM 419999) become paralectotypes. Lectotype Measurements. Length 30.1 mm, diameter (excluding lip) 11.5 mm; ap- erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) 9.5 mm. Type Locality. Living on the three Silver Keys [New Providence Island, Bahamas]. Remarks. Maynard (1913c: 150) noted “Silver Key, the most western of the group, is remarkable as having been the home of three, quite closely allied and evidently successive species of Strophias, represen- tatives of the three distinct geological pe- riods.” This taxon name was misspelled as S. argntea in Maynard, 1921a (p. 138), and as S. argentea in captions (Maynard, 1919c, fig. 26, 1921\b, pl. 35; figs. 5, 6) and the sales catalog (Maynard, 1924?: [4]). Gould and Woodruff (1986: 474) listed this taxon as a synonym of Cerion glans (Kiister, 1844). 421 Taxon 83. Strophiops agavaneglecta May- nard, 1913c [February 9]: 192 [not illustrated]. Examined 150 specimens. No measurements provided. Type Material. The original description did not distinguish among the 150 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 150 specimens are syntypes. This taxon was not represented in the collections of MCZ, nor was it pre- sent in Maynard’s collection at the time it was acquired from his daughter, as re- vealed by an examination of the MCZ cat- alog. This taxon was not among the ma- terial purchased for the USNM collections either during Maynard’s lifetime or from his estate. Searches of the collections and/ or databases of The Academy of Natural Sciences of Philadelphia, The Field Mu- seum of Natural History, The Florida Mu- seum of Natural History, and The Muse- um of Biological Diversity, The Ohio State University, failed to uncover a single spec- imen catalogued under this name. This taxon was never illustrated, nor did it ap- pear in any of Maynard’s subsequent pub- lications or catalogs. The description is limited to the statement “Intermediate be- tween agava and neglecta.” |Strophia aga- va was originally proposed as a subspecies of S. neglecta.| No specimens bearing the name Strophiops agavaneglecta could be located in museums, nor could this taxon be identified on the basis of its limited de- scription. The holotype of Strophia neglec- ta agava Maynard, 1894 (MCZ 76103), is here designated as the neotype of Stro- phiops agavaneglecta Maynard, 1913, to provide an objective standard of reference for this species-group taxon. Strophiops agavaneglecta Maynard, 1913, becomes an objective junior synonym of Strophia neg- lecta agava Maynard, 1894. Type Locality. Maynard (1913c: 192) defined the type locality of S. agavane- glecta to be the eastern portion of the range of S. agava [fields west of Nassau, New Providence Island, Bahamas]. With 422 the neotype designation, the type locality of S. agava becomes the type locality of S. agavaneglecta (Article 76.3, ICZN, 1999: 87). Remarks. Following Article 32.5.2.4 (ICZN, 1999: 40), the hyphen is removed from agava-neglecta and the taxon be- comes agavaneglecta. Gould and Woodruff (1986: 474) synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 84. Strophiops oberholseri May- nard, 1913c [February 9]: 193 [not illustrated in original publication]; May- nard, 1920c, fig. 86; Maynard, 1921b: 140, pl. 37, figs. 1, 2. Examined 220 specimens. No measurements provided in original description. Type Material. The original description did not distinguish among the 220 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 220 specimens are syntypes. Lot MCZ 76124 contains a single specimen labeled “holotype.” This lot also contains a label in Maynard’s hand with the term “Type” that includes references to subsequently published illustrations (Maynard, 1921b, pl. 37, figs. 1, 2), which the specimen approximates. This specimen is here designated as the lectotype to pro- vide an objective standard of reference for this species-group taxon that is consistent with Maynard's concept of this taxon. The remaining 219 syntypes, including MCZ 76125 and MCZ 117655, become paralec- totypes. Lectotype Measurements. Length 26.4 mm, diameter (excluding lip) 10.6 mm; ap- erture height (including lip) 9.7 mm, ap- erture width (including lip and peristome) 8.6 mm. Type Locality. Southwest Bay, New Providence |Island, Bahamas], near the landing. Remarks. This taxon was first illustrated (Maynard, 1920c, fig. 86) as an example of Maynard’s “Law of Specific Reincarna- tion.” Maynard (1921b: 140, pl. 37, figs. 1, Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 2.) noted that this taxon was the same size as S. coryi (i.e., 1.00 by .40 linches; 25.4 by 10.2 mm]) and provided colored illus- trations. Gould and Woodruff (1986: 474) synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 85. Strophiops inconsueta May- nard, 1913c [February 9]: 193-194 [not illustrated in original publication]; Maynard, 1920a, pl. 11, fig. 1; Maynard, 1921b: 142; Maynard, 1924a, pl. 39, figs. 1, 2. Examined 675 specimens. Size giv- en as 1.27 by .50 [inches; 32.3 by 12.7. mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 675 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 675 specimens are syntypes. Lot number MCZ 76123 con- tains a single specimen labeled “Holotype” and a label in Maynard’s hand with the term “type,” which includes references to subsequently published figures (Maynard, 1924a, pl. 39, figs. 1, 2). This specimen, which approximates the measurements in the original description and the subse- quently published illustrations, is here des- ignated as the lectotype to provide an ob- jective standard of reference for this spe- cies-group taxon that is consistent with Maynard’s concept of this taxon. The re- maining 674 syntypes, including MCZ 76122, MCZ 177994 (one specimen), and USNM 420051 (75 specimens), become paralectotypes. Lectotype Measurements. Length 31.9 mm, diameter (excluding lip) 13.7 mm; ap- erture height (including lip) 11.6 mm, ap- erture width (including lip and peristome) 11.0 mm. Type Locality. Southern portion of Great Pimlico [Island, Eleuthera, Baha- mas |. Remarks. Clench (1952: 110) consid- ered this taxon to be a valid species and the senior synonym of S. vetusta Maynard, 1913 (see Taxon 81). THE CERION TAXA OF CHARLES JOHNSON MAYNARD ° Harasewych et al. Taxon 86. Strophiops transitoria May- nard, 1913c [February 9]: 194; May- nard, 1921b: 141, pl. 37, figs. 9, 10 [not illustrated in original publication]. Ex- amined 250 specimens. Size given as .90 by .35 [inches; 22.9 by 8.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 250 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 250 specimens are syntypes. Lot MCZ 117848 contains a sin- gle specimen labeled “syntype,” whereas MCZ 76119 contained 116 specimens la- beled “paratype.” All are syntypes, and most appear to be dead collected. Of these, the sequestered specimen (MCZ 117848), which roughly approximates the measurements and subsequent illustration, is here designated as the lectotype to pro- vide an objective standard of reference for this species-group taxon that is consistent with Maynard's concept of this taxon. The remaining 249 syntypes, including the 116 specimens in MCZ 76119, become para- lectotypes. Lectotype Measurements. Length Lid mm, diameter (excluding lip) 9.6 mm; ap- erture height (including lip) 9.1 mm, ap- erture width (including lip and peristome) 7.8 mm. Type Locality. Northern portion of Great Pimlico [Island, Eleuthera, Baha- mas], ranging almost into the area occu- pied by the foregoing species [S. incon- sueta |. Remarks. Maynard considered this tax- on to be “In the last stage of its specific life” with “only 15 of 250 specimens living, 10 of these young. Both species [S. incon- sueta and S. transitoria] living upon the same food and under the same environ- ment.” Maynard (1921b: 141, pl. 37, figs. 9, 10) subsequently added descriptive notes and illustrated this taxon. Clench (1952: 110) listed this taxon as a synonym of Cerion uniforme (Maynard, 1913) (see below). 423 Taxon 87. Strophiops uniformis Maynard, 1913c [February 9]: 194 [not illus- trated in original publication]; Maynard, 1921b: 141; Maynard, 1924a, pl. 38, figs. 1, 2. Examined 760 specimens. Size giv- en as 1.15 by .45 [inches; 29.2 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 760 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 760 specimens are syntypes. Lot number MCZ 76117 con- tains a single specimen labeled “Holotype” that approximates the measurements in the original description and the subse- quent illustration in Maynard (1924a, pl. 38, figs. 1, 2). This lot also contains a label in Maynard’s hand with the term “Type” that includes references to these figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of this taxon. The remaining 759 syntypes, including MCZ 76118, MCZ 117851, and USNM 420052 (30 specimens), become paralectotypes. Lectotype Measurements. Length 29.5 mm, diameter (excluding lip) 11.1 mm; ap- erture height (including lip) 10.9 mm, ap- erture width (including lip and peristome) 9.9 mm. Type Locality. Little Pimlico [Island, Eleuthera, Bahamas]. Remarks. Maynard (1921b: 141, 1924a, pl. 38, figs. 1, 2) subsequently added de- scriptive notes and illustrated this taxon. Clench (1952: 110) listed this as a valid species but did not mention type speci- mens. Taxon 88. Strophiops hyattii Maynard, 1913c [February 9]: 194-195 [not il- lustrated in original publication]; May- nard, 1921b: 142; Maynard, 1926, pl. 42, figs. 1, 2. Examined 600 specimens. Measurements not provided. Type Material. The original description 424 did not distinguish among the 600 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 600 specimens are syntypes. Lot number MCZ 76133 con- tains a single specimen labeled “holotype.” No measurements were provided in the original description, but this specimen ap- proximates the measurements (1.10 by .45 [inches; 27.9 by 14.0 mm]) and illustration in Maynard (1921b: 142, 1926, pl. 42, figs. 1, 2) [as S. hyatiii]. This lot also contains a label in Maynard’s hand with the term “type,” which includes references to these figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s concept of this taxon. The remain- ing 599 syntypes, including MCZ 76134, MCZ 118036 (one specimen), and USNM 419973 (75 specimens), become paralec- totypes. Lectotype Measurements. Length 26.3 mm, diameter (excluding lip) 11.0 mm; ap- erture height (including lip) 9.8 mm, ap- erture width (including lip and peristome) 9 )8) IAMTON, Type Locality. Bar Bay Settlement, Cur- rent Island |[Eleuthera, Bahamas]. Remarks. Clench (1952: 109) listed this taxon as a valid species. Taxon 89. Strophiops vetustapraedevina Maynard, 1913c [February 9]: 195 [not illustrated]. Examined 50 speci- mens. Size given as 1.15 by .48 [inches; 29.2, by 12.2 mm, but the term “type” was not used. Type Material. The original description Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 did not distinguish among the 50 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 50 specimens are syntypes. This taxon was not represented in the collections of the MCZ, nor was it present in Maynard’s collection at the time it was acquired from his daughter, as re- vealed by an examination of the MCZ cat- alog. Three syntypes of this taxon, which Bartsch purchased from Maynard in 1923, were located in the USNM collection. Of these, the best preserved specimen (USNM 359439) is selected as the lecto-— type. The 49 syntypes, including the two specimens in USNM 359438, become par- alectotypes. Lectotype Measurements. Length 31.7 mm, diameter (excluding lip) 13.6 mm; ap- erture height (including lip) 13.5 mm, ap- erture width (including lip and peristome) Wibod aoarany. Type Locality. Fossil on Great Pimlico [Island, Eleuthera, Bahamas]. Remarks. Following Article 32.5.2.4 (ICZN, 1999: 40), the hyphen is removed from vetusta-praedevina and the taxon be- comes vetustapraedevina. Also reported from Upper Fleming and the Silver Keys. Clench (1952: 110, 1957: 167) listed this taxon as a synonym of Cerion inconsuetum (Maynard, 1913). Taxon 90. Strophiops praedivina May- nard, 1913c [February 9]: 196 [not illustrated in original publication]; May- nard, 1921b: 141, pl. 37, figs. 5, 6. Ex- amined 100 specimens. Size given as 1.05 by .45 [inches; 26.7 by 11.4 mm], but the term “type” was not used. —-> Figures 87-99. Taxon 87. Strophiops uniformis Maynard, 1913. Lectotype MCZ 76117. Little Pimlico Island, Eleuthera, Baha- mas. Taxon 88. Strophiops hyattii Maynard, 1913. Lectotype MCZ 76133. Bar Bay Settlement, Current Island, Eleuthera, Ba- hamas. Taxon 89. Strophiops vetustapraedevina Maynard, 1913. Lectotype USNM 359439. Fossil on Great Pimlico Island, Eleuthera, Bahamas. Taxon 90. Strophiops praedivina Maynard, 1913. Lectotype MCZ 76346. Upper Fleming Key, 20 miles NE of New Providence Island, Bahamas. Taxon 91. Strophiops praedivinauniversa Maynard, 1913. [Not illustrated.] The lectotype (MCZ 76287) of Strophiops universa Maynard, 1913 [Taxon 92], is designated as the neotype of Strophiops praedivinauniversa Maynard, 1913, making Strophiops praedivinauniversa Maynard, 1913, an objective junior synonym of Strophia universa May- nard, 1913. Taxon 92. Strophiops universa Maynard, 1913. Lectotype MCZ 76287. Fossil on Green Key, Rose Island, New Providence Island, Bahamas. Taxon 93. Strophiops exigua Maynard, 1913. Lectotype MCZ 76357. Southern portion of Fleming THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 425 Key, 20 miles NE of New Providence Island, Bahamas. Taxon 94. Strophiops multa Maynard, 1913. Lectotype MCZ 76354. Northern portion of Fleming Key, 20 miles NE of New Providence Island, Bahamas. Taxon 95. Strophiops obliterata Maynard, 1913. Lectotype MCZ 76334. Booby Rock, 16 miles NE of New Providence Island, Bahamas. Taxon 96. Strophiops ornatula Maynard, 1913. Lectotype MCZ 76343. Current Settlement, Eleuthera, Bahamas. Taxon 97. Strophiops ornatulaclappiiMaynard, 1913. Lectotype MCZ 117738. North of the Current Settlement, Eleuthera, Bahamas. Taxon 98. Strophiops clappii Maynard, 1913. Lectotype MCZ 76344. Salina, north of the Current Settlement, Eleuthera, Bahamas. Taxon 99. Strophiops ornatularufina Maynard, 1913. Lectotype MCZ 118095. South of the Current Settlement, Eleuthera, Bahamas. 426 Type Material. The original description did not distinguish among the 100 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 100 specimens are syntypes. Lot number MCZ 76346 con- tains a single specimen labeled “holotype.” This lot also contains a label in Maynard’s handwriting with the name of the taxon and the term “type.” This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of this taxon. The re- maining 99 syntypes, including MCZ 76347 (62 specimens), become paralecto- es. Lectotype Measurements. Length Dilek mm, diameter (excluding lip) 11.2 mm; ap- erture height (including lip) 10.4 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. Upper Fleming |Key, 20 miles NE of New Providence Island, Ba- hamas |]. Remarks. A fossil taxon also reported from Lower Fleming and Great Pimlico. Maynard (1921b: 141, pl. 37, figs. 5, 6) subsequently provided illustrations and a different measurement (1.08 by .55 [inch- es; 27.4 by 14.0 mm]) and reported only 59 specimens. The taxon is spelled prae- divina in the original description and sub- sequent plate caption, but as predivina in Maynard (1921b: 141) and in the sales cat- alog (Maynard, 1924?: [6]). Clench (1952: 109) listed this taxon as a synonym of Cer- ion exiguum (Maynard, 1913). Taxon 91. Strophiops praedivinauniversa Maynard, 1913c [February 9]: 196 [not illustrated]. Examined 25. speci- mens. Measurements not provided. Type Material. The original description did not distinguish among the 25 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 25 specimens are syntypes. This taxon was not represented Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 in the collections of the MCZ, nor was it present in Maynard’s collection at the time it was acquired from his daughter, as re- vealed by an examination of the MCZ cat- alog. This taxon was not among the ma- terial purchased for the USNM collections either during Maynard’s lifetime or from his estate. Searches of the collections and/ or databases of The Academy of Natural Sciences of Philadelphia, The Field Mu- seum of Natural History, The Florida Mu- seum of Natural History, and The Muse- um of Biological Diversity, The Ohio State University, failed to uncover a single spec- imen catalogued under this name. This taxon was never illustrated, nor did it ap- pear in any of Maynard’s subsequent pub- lications or catalogs. The description is limited to the statement “Intermediate be- tween pdreivina [sic] and universa, but inclined to be more like the latter.” No specimens bearing the name Strophiops praedivinauniversa could be located in museums, nor could this taxon be identi- fied on the basis of its limited description. The lectotype of Strophiops universa May- nard, 1913 (MCZ 76287), is here desig- nated as the neotype of Strophiops prae- divinauniversa Maynard, 1913, making Strophiops praedivinauniversa Maynard, 1913, an objective synonym of Strophia universa Maynard, 1913. Type Locality. Maynard (1913c: 194) noted that S. praedivinauniversa occurred on Upper Fleming [Key, 20 miles NE of New Providence Island, Bahamas], both living and fossil. Although S. wniversa was reported from Upper Fleming, Great Pim- lico, and the northern end of Rose Island, the type locality of S. wniversa, and there- fore of S. praedivinauniversa, is Green Key, New Providence Island, Bahamas, per Article 76.3 (ICZN, 1999: 87). Remarks. Following Article 32.5.2.4 (ICZN, 1999: 40), the hyphen is removed from praedivina-universa and the taxon becomes praedivinauniversa. Clench (1952: 109) listed this taxon as a synonym of Cerion exiguum (Maynard, 1913). THE CERION TAXA OF CHARLES JOHNSON MAYNARD ° Harasewych et al. Taxon 92. Strophiops universa Maynard, 1913c [February 9]: 196 [not illus- trated in original publication]; Maynard, 1921b: 140, pl. 37, figs. 3, 4. Examined 30 specimens. Size given as 5) by , linches; 14.0 by 8.1 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 30 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 30 specimens are syntypes. MCZ 76287, labeled “Paratype,” con- tained 26 specimens (many fragmentary), of which one was sequestered, together with a label in Maynard’s handwriting with the taxon name and “Plate 37, figs. 3-4.” The sequestered specimen closely approx- imates the subsequently published illustra- tion and published width measurement. The published length measurement is ei- ther in error, or incorporates length mea- surements of fragmentary specimens. The sequestered specimen is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s concept of this taxon. The remain- ing 25 specimens, recatalogued as MCZ 356475, are among the 29 syntypes that become paralectotypes. Lectotype Measurements. Length 18.9 mm, diameter (excluding lip) 8.2 mm; ap- erture height (including lip) 8.2 mm, ap- erture width (including lip and peristome) Oo mM: A427 ditional specimens or whether the number is a typographical error. Clench (1952: 110) listed this taxon as a synonym of Cer- ion uniforme (Maynard, 1913). Taxon 93. Strophiops exigua Maynard, 1913c [February 9]: 196 [not illus- trated in original publication]; Maynard, 1921b: 141; Maynard, 1924a, pl. 38, figs. 3, 4. Examined 1,000 specimens. Size given as 1.30 by 50 [sic .50] [inches; 33.0 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,000 spec- imens examined, and there was no men- tion of type material in the other parts of this publication. Thus, all 1,000 specimens are syntypes. Lot MCZ 76357 contains a single specimen labeled “Holotype.” This specimen approximates the measurements in the original description and the illustra- tion in Maynard (1921b: 141, 1924a, pl. 38, figs. 3, 4). This lot also contains a label in Maynard’s hand with the term “type,” which includes references to these figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of this taxon. The remaining 999 syntypes, including MCZ 76356 (171 specimens), MCZ 117837 (1 specimen), and USNM 420011 (250 specimens), become paralec- totypes. Lectotype Measurements. Length Sil mm, diameter (excluding lip) 11.6 mm; ap- Type Locality. Fossil on Green Key [New Providence Island, Bahamas]. Remarks. Also reported from Upper Fleming, Great Pimlico, and northern end of Rose Island. Maynard (1913c: 196) sur- mised that “The wide range of this little species suggests that the islands on which it is now found were once connected.” Maynard (1921b: 140, pl. 37, figs. 3, 4) subsequently provided illustrations of this taxon and listed 50 specimens examined. It is unclear whether he later collected ad- erture height (including lip) 10.7 mm, ap- erture width (including lip and peristome) 9.1 mm. Type Locality. Southern portion of Fleming Key [20 miles NE of New Prov- idence Island, Bahamas]. Remarks. This taxon was reported from living specimens, which were abundant, and from fossils in the limestone, which were rare. Maynard (1921b: 141, 1924a, pl. 38, figs. 3, 4) provided descriptive notes and illustrations of this taxon. Clench 428 (1952: 109) listed this taxon as a valid spe- cies. Taxon 94. Strophiops multa Maynard, 1913c [February 9]: 197 [not illus- trated in original publication]; Maynard, 1921b: 141; Maynard, 1924a, pl. 38, figs. 5, 6. Examined 964 specimens. Size giv- en as 1.10 by .40 [inches; 27.9 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 964 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 964 specimens are syntypes. Lot number MCZ 76354 con- tains a single specimen labeled “Holo- type.” This specimen approximates the measurements in the original description (within 5%) and the illustration in May- nard (1924a, pl. 38, figs. 5, 6). This lot also contains a label in Maynard’s hand with the term “type,” which includes references to these figures (as pl. 38, figs. 7, 8). This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of this taxon. The remaining 963 syntypes, in- cluding MCZ 76355 and MCZ 118069, be- come paralectotypes. Lectotype Measurements. Length 28.9 mm, diameter (excluding lip) 10.8 mm; ap- erture height (including lip) 10.6 mm, ap- erture width (including lip and peristome) 9.0 mm. Type Locality. Northern portion of Fleming Key [20 miles NE of New Prov- idence Island, Bahamas]. Remarks. Maynard (1921b: 141, 1924a, pl. 38, figs. 5, 6) provided additional de- scriptive notes and illustrations of this tax- on. Clench (1934: 214) initially considered this to be a valid species, but later (1952: 109) listed this taxon as a synonym of Cer- ion exiguum (Maynard, 1913). Gould and Woodruff (1986: 474) synonymized this taxon with C. glans (Kiister, 1844). Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Taxon 95. Strophiops obliterata Maynard, 1913c [February 9]: 197 [not illus- trated in original publication]. Exam- ined 812 specimens. Size given as 1.15 by .30 [inches; 29.2 by 7.6 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 812 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 812 specimens are syntypes. There were no specimens of S. obliterata Maynard, 1913, in the collec-. tions of the MCZ, nor were there any en- tries corresponding to this taxon in the MCZ catalog either during Maynard’s life or among the specimens purchased from his estate. Maynard and Clapp (1915: 191) recognized that S. obliterata Maynard, 1913, was a homonym of Strophia scripta obliterata Maynard, 1896, and proposed the replacement name Strophiops sula [Taxon 105]. Lot number MCZ 76334 con- tains a single specimen labeled as the “Ho- lotype” of S. sula. Because Strophiops ob- literata Maynard, 1913, was based on 812 syntypes, this specimen is here designated as the lectotype of Strophiops obliterata Maynard, 1913, and consequently [Article 72.7, ICZN, 1999: 78] its replacement name, Strophiops sula Maynard and Clapp, 1915. Lots MCZ 76335 (170 spec- imens) and USNM 420096 (125. speci- mens) contain numerous specimens la- beled as paralectotypes of S. sula. These specimens are paralectotypes of S. obliter- ata Maynard, 1913, as well as of its re- placement name S. sula. Lectotype Measurements. Length 31.8 mm, diameter (excluding lip) 12.4 mm; ap- erture height (including lip) 12.3 mm, ap- erture width (including lip and peristome) 10.1 mm. Type Locality. Booby Rock [16 miles NE of New Providence Island, Bahamas]. Remarks. Although the taxon was never illustrated as S. obliterata Maynard, 1913, Maynard (1921b: 142, 1924a, pl. 38, figs. 7, 8) subsequently published additional THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. notes and illustrations of S. sula. Gould and Woodruff (1986: 474) listed both S. obliterata) Maynard, 1913, and S. sula Maynard and Clapp, 1915, as synonyms of Cerion glans (Kiister, 1844). Taxon 96. Strophiops ornatula Maynard, 1913c [February 9]: 197-198 [not il- lustrated in original publication]; May- nard, 1919a, pl. 6, fig. 4; Maynard, 1919b, fig. 11. Examined 3,528 speci- mens. Size given as 1.15 by .30 [inches; 29.2 by 7.6 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 3,528 spec- imens examined, and there was no men- tion of type material in the other parts of this publication. Thus, all 3,528 specimens are syntypes. Clench (1934: 217) men- tioned comparing the types of Cerion or- natula (Maynard) and C. laeve Plate, but did not specify the sort of type, nor the repository or catalog number of the e lot. Later, Clench (1952: 112) listed MCZ 76343 as the holotype of C. ornatula. This lot contains a single specimen labeled “Holotype” that approximates the pub- lished length but not the width and is sim- ilar to the subsequently published illustra- tion of this taxon (Maynard, 1919a, pl. 6, fig. 4). This specimen, which is accompa- nied by a label in Maynard’s hand that in- cludes the term “type” and references to these illustrations, cannot be a holotype, nor can Clench’s listing be accepted as a lectotype designation (Article 74.5, ICZN, 1999: 83). This specimen is here designat- ed as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original, or at least subsequent con- cept of it. The remaining 3,527 syntypes, including MCZ 76342 and USNM 420137 (400 specimens), become paralectotypes. Lectotype Measurements. Length 22.4 mm, diameter (excluding lip) 9.0 mm; ap- erture height (including lip) 8.9 mm, ap- erture width (including lip and peristome) 7.1 mm. 429 Type Locality. Current Settlement [Eleuthera, Bahamas]. Remarks. This taxon was later illustrated (Maynard, 1919a, pl. 6, fig. 3, 1919b, fig. 11). Clench (1934: 217) reported that Cer- ion ornatula “is an absolute synonym” of C. laeve (Plate, 1907) on the basis of com- parisons of type material. Taxon 97. Strophiops ornatulaclappii Maynard, 1913c [February 9]: 198 [not illustrated in original publication]; Maynard, 1921b: 153. Examined 50 specimens. Measurements not provided. Type Material. The original description did not distinguish among the 50 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 50 specimens are syntypes. There were no entries for this taxon in the MCZ catalog, indicating that this taxon was not represented in May- nard’s collection at the time of his death. Lot number MCZ 117738 contained nine specimens from the collection of N. A. Clapp, Maynard's collaborator and co-au- thor. Because neither measurements nor illustration were published, an adult spec- imen is selected to serve the lectotype to provide an objective standard of reference for this species-group taxon that is consis- tent with Maynard’s original concept of it. The remaining eight specimens were re- catalogued as MCZ 356993 and are among the 49 syntypes that become paralecto- types. Lectotype Measurements. Length 20.3 mm, diameter (excluding lip) 8.3 mm; ap- erture height (including lip) 7.9 mm, ap- erture width (including lip and peristome) 6.5 mm. Type Locality. Living just north of the Current Settlement | Eleuthera, Bahamas]. Remarks. Following Article 32.5.2.4 (ICZN, 1999: 40), the hyphen is removed from ornatula-clappii and the taxon be- comes ornatulaclappii. Clench (1952: 111) listed this taxon in the synonymy of Cerion laeve Plate, 1907, citing it as a nomen nu- dum. 430 Taxon 98. Strophiops clappii Maynard, 1913c [February 9]: 198 [not illus- trated in original publication]; Maynard, 1919a, fig. 10, pl. 6, fig. 5; Maynard, 1921b: 153, pl. 48, fig. 6 [plate never issued]. Number of specimens exam- ined not given, but Maynard (1913c: 198) stated the fields contained “abso- lutely millions of dead shells of this [species].” Size given as .50 by .20 [inch- es; 12.7 by 5.1 mm], but the term “type” was not used. Maynard (1921b: 153) listed 2,000 specimens. Type Material. The original description did not provide the number of specimens examined, nor is there mention of type material in the other parts of this publi- cation. Thus, there are an unspecified number of syntypes, but not more than the 2,000 specimens mentioned by Maynard (1921b: 153). Clench (1952: 112) listed MCZ 76344 as the “Holotype.” Lot MCZ 76344 was labeled “holotype” and accom- panied by a note in Maynard's handwriting identifying the taxon and containing the term “Type” as well as references to both published and unpublished illustrations. This specimen is nearly identical to the published illustration and measurements. It is here selected as the lectotype to pro- vide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 1,999 syntypes, including MCZ 76345 and MCZ 21128, become paralec- totypes. Lectotype Measurements. Length 12.6 mm, diameter (excluding lip) 5.1 mm; ap- erture height (including lip) 4.4 mm, ap- erture width (including lip and peristome) 3.8 mm. Type Locality. Living on a salina north of the Current Settlement | Eleuthera, Ba- hamas |. Remarks. This taxon was illustrated by Maynard (1919a, fig. 10, pl. 6, fig. 5 [as clappi|). Clench (1952: 111) listed this tax- on in the synonymy of Cerion laeve Plate, 1907. Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Taxon 99. Strophiops ornatularufina May- nard, 1913c [February 9]: 198 [not illustrated in original publication]. Ex- amined 50 specimens. Measurements not provided. Type Material. The original description did not distinguish among the 50 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 50 specimens are syntypes. There were no entries for this taxon in the MCZ catalog, indicating that this taxon was not represented in May- nard’s collection at the time of his death. Lot number MCZ 118095 contains a sin- gle specimen from the collection of N. A. Clapp, Maynard’s collaborator and co-au- thor. This specimen, one of the 50 syn- types, is here designated as the lectotype to provide an objective standard of refer- ence for this species-group taxon that is consistent with Maynard's original concept of it. The remaining 49 syntypes become paralectotypes. Lectotype Measurements. Length 19.5 mm, diameter (excluding lip) 8.2 mm; ap- erture height (including lip) 7.5 mm, ap- erture width (including lip and peristome) 6.9 mm. Type Locality. South of the Current Set- tlement [Eleuthera, Bahamas]. Remarks. Following Article 32.5.2.4 (ICZN, 1999: 40), the hyphen is removed from ornatula-rufina and the taxon be- comes ornatularufina. Taxon 100. Strophiops rufina Maynard, 1913c [February 9]: 198 [not illus- trated]; Maynard, 1924a: 153, pl. 48, figs. 9, 10 [plate never published]. Ex- amined 20 specimens. Size given as .75 by .40 [inches; 19.0 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 20 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 20 specimens are syntypes. Lot MCZ 76304 contains a single THE CERION TAXA OF CHARLES JOHNSON MAYNARD °* Harasewych et al. specimen labeled “Holotype” that roughly approximates the measurements of this taxon. It is accompanied by a label in May- nard’ss hand with the term “type,” which includes references to figures that were never published. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 19 syntypes, including MCZ 76303, become paralectotypes. Lectotype Measurements. Length WS) mm, diameter (excluding lip) 8.5 mm; ap- erture height (including lip) 7.9 mm, ap- erture width (including lip and peristome) 6.9 mm. Type Locality. Rare south of the Cur- rent Settlement | Eleuthera, Bahamas]. Remarks. Clench (1952: 109) listed this taxon as a synonym of Cerion hyattii (May- nard, 1913). Taxon 101. Strophiops tabida Maynard, 1913c [February 9]: 199 [not illus- trated in original publication]; Maynard, 1921b: 152 [as tibida], pl. 44, figs. 9, 10 [illustrations never published]. Type Material. Because the original de- scription did not mention the number of specimens examined, this taxon is based on an unspecified number of syntypes. Maynard (1921b: 152) subsequently re- ported 100 specimens examined and gave the size as 1.00 by .40 [inches; 25.4 by 10.2 mm]. Lot MCZ 76024 contains a single specimen labeled “Holotype” that closely approximates the subsequently published measurements for this taxon. This lot also contains a label in Maynard’s hand with the term “Type” that includes references to illustrations that were never published. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept 431 Lectotype Measurements. Length DLO mm, diameter (excluding lip) 10.3 mm; ap- erture height (including lip) 9.3 mm, ap- erture width (including lip and peristome) 8.1 mm. Type Locality. West shore of Eleuthera [Bahamas] near the Current Settlement, living on rocks close by the sea. Remarks. Maynard (1913c: 199) noted, “Some individuals appear to mimic species unknown to me. The most remarkable re- cessional species that I have thus far found as, all things considered, it is the most re- mote phylogenetically from the species it mimics.” Clench (1952: 109) listed this tax- on as a synonym of Cerion hyattii (May- nard, 1913). Taxon 102. Strophiops helena Maynard, 1914 [August 18]: 177-179, figs. SA, B, C, D. Examined 1,000 specimens. Size given as .92 by .43 [inches; 23.4 by 10.9 mm], but the term “type” was not used in connection with the measure- ments. Caption to text fig. 8, “STRO- PHIOPS HELENA: A, front. B, side view of type; c, front of aperture of type enlarged X 2; D, margin of senescent specimen X 2.” Type Material. The original description identifies a single type specimen, the ho- lotype, in the figure captions. Lot MCZ 76216 contains a single specimen labeled “Holotype.” This specimen is accompa- nied by a label in Maynard’s hand contain- ing the term “Type” and also giving the correct species number and plate number but incorrect volume and page for this tax- on in Maynard (1921b: 131, pl. 7, figs. 5— 8). Specimen MCZ 76216 is the originally designated holotype of Strophiops helena Maynard, 1914. The remaining 999 spec- imens, including MCZ 76203 (231 speci- mens) and USNM 419990 (100. speci- mens), are paratypes. Holotype Measurements. Length 22.6 mi, diameter (excluding lip) 10.1 mm; ap- of this taxon. The remaining syntypes (=99), including MCZ 76025, become paralectotypes. erture height (including lip) 8.4 mm, ap- erture width (including lip and peristome) 7.8 mm. 432 Type Locality. Fields directly south of Fresh Creek, Andros [Bahamas]. Remarks. Maynard (1919b, pl. 7, figs. 5— 8) re-illustrated this taxon. Taxon 103. Strophiops thompsoni May- nard and Clapp, 1915 [February 8]: 179 bis [not illustrated in original pub- lication]; Maynard, 1921b: 139, pl. 35, figs. 9, 10. Examined 20 specimens. Size given as 1.05 by .45 [inches; 26.7 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 20 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 20 specimens are syntypes. Lot MCZ 76186 contains a single specimen labeled “Holotype.” This speci- men closely approximates the measure- ments in the original description and the subsequent illustrations (Maynard, 1921b, pl. 35, figs. 9, 10) and is accompanied by a label in Maynard’s hand containing the term “Type” that refers to the published figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. Lot MCZ 76187 contains 264 specimens (most dam- aged, incomplete, or juvenile) and was la- beled “paratype.” Because Maynard added specimens after publishing the descrip- tion, it is not clear which 19 of the 264 specimens are paralectotypes. Lectotype Measurements. Length 25.7 mm, diameter (excluding lip) 11.3 mm; ap- erture height (including lip) 10.0 mm, ap- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 erture width (including lip and peristome) 8.5 mm. Type Locality. Occurs as a fossil in the rock on the south shore of Hog Island [ = Paradise Island, Nassau, New Providence Island, Bahamas]. Remarks. Maynard (1921b: 139, pl. 35, figs. 9, 10) provided additional descriptive information and illustrations, noting 370 specimens in his collection. Gould and Woodruff (1986: 482) provisionally attri- buted this taxon to Cerion glans (Kiister, 1844). Taxon 104. Strophiops morula Maynard and Clapp, 1915 [February 8]: 179 bis-180 bis [not illustrated in original publication]; Maynard, 1921b: 139, pl. 36, figs. 1, 2. Neither the number of specimens examined nor measurements are provided in the original description. Type Material. The original description did not specify the number of specimens examined. In the subsequent monograph, Maynard (1921b: 139, pl. 36, figs. 1, 2) list- ed 60 specimens. The size of the syntype lot is uncertain, but does not exceed 60 syntypes. The original description provid- ed neither measurements nor illustration but stated that there were 10 whorls, with 23 costae on the final whorl. Lot number MCZ 76017 contains a single specimen la- beled “Holotype.” This specimen consists of nine whorls but has 23 costae on the final whorl. This specimen is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the subsequently published figures. However, it does not closely match either the figures — Figures 100-113. Taxon 100. Strophiops rufina Maynard, 1913. Lectotype MCZ 76304. South of the Current Settlement Eleuth- era, Bahamas. Taxon 101. Strophiops tabida Maynard, 1913. Lectotype MCZ 76024. Near the Current Settlement, west shore of Eleuthera, Bahamas. Taxon 102. Strophiops helena Maynard, 1914. Holotype MCZ 76216. Fields directly south of Fresh Creek, Andros, Bahamas. Taxon 103. Strophiops thompsoni Maynard and Clapp, 1915. Lectotype MCZ 76186. Fossil, south shore of Hog Island [Paradise Island], Nassau, New Providence Island, Bahamas. Taxon 104. Strophiops morula Maynard and Clapp, 1915. Lectotype MCZ 76017. Spruce Key, 4 miles E of Nassau, New Providence Island, Bahamas. Taxon 105. Strophiops sula Maynard and Clapp, 1915. [Not illustrated.] Replacement name for Strophiops obliterata Maynard, 1913 [Taxon 95], a junior homonym of S. scripta obliterata Maynard, 1896 [Taxon 53]. The type specimens and type locality of S. obliterata Maynard, 1913 are also the type specimens and type locality of S. sula Maynard and Clapp, 1915. Taxon 106. Strophiops hesternia Maynard and Clapp, 1915. Lectotype MCZ 76157. West end of Booby Rock, 16 miles NE of New Providence Island, Bahamas. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 433 Taxon 107. Strophiops caerulea Maynard and Clapp, 1915. Lectotype MCZ 76308. Field north of Ft. Charlotte, Nassau, New Providence Island, Bahamas. Taxon 108. Strophiops devereuxi Maynard and Clapp, 1915. Lectotype MCZ 76137. Devereux Estate west end of New Providence Island, Bahamas. Taxon 109. Strophiops praedicta Maynard and Clapp, 1915. Lectotype MCZ 76393. Great Pimlico Island, Eleuthera, Bahamas. Taxon 110. Strophiops prisca Maynard and Clapp, 1915. Lectotype 76353. Fossil in the rocks on east end of Salt Key, New Providence Island, Bahamas. Taxon 111. Strophiops tenui Maynard and Clapp, 1915. Lectotype MCZ 76350. East end of New Providence Island, Bahamas. Taxon 112. Strophiops fulmineaMaynard and Clapp, 1915. Lectotype MCZ 76276. Rocky shore east of Ft. Winton, New Providence Island, Bahamas. Taxon 113. Stro- phiops longidentata Maynard, 1919. Nomen nudum. [Not illustrated. ] 434 or the measurements in the monograph. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining syntypes (<59), including MCZ 76018, become paralectotypes. Lectotype Measurements. Length 21.6 mm, diameter (excluding lip) 8.8 mm; ap- erture height (including lip) 7.7 mm, ap- erture width (including lip and peristome) 5. mone Type Locality. Spruce Key [4 miles E of Nassau, New Providence Island, Baha- mas]. Remarks. Semi-fossil. Maynard (1921b: 139, pl. 36, figs. 1, 2) provided additional descriptive information, including mea- surements [1.10 by .50 inches; 27.9 by 12.7 mm] as well as illustrations. Gould and Woodruff (1986: 474) synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 105. Strophiops sula Maynard and Clapp, 1915 [February 8]: 180 bis. Replacement name for Strophiops obli- terata Maynard, 1913 [Taxon 95], a ju- nior secondary homonym of Strophia scripta obliterata Maynard, 1896 [Taxon Doe Type Material. The type specimens and type locality of S. obliterata Maynard, 1913 [Taxon 95], are also the type specimens of S. sula Maynard and Clapp, 1915. Remarks. Strophiops obliterata May- nard, 1913, was not illustrated, nor was S. sula Maynard and Clapp, 1915, at the time of its publication. Maynard (1921b: 142, 1924a, pl. 38, figs. 7, 8) subsequently pub- lished additional notes and illustrations of S. sula. Gould and Woodruff (1986: 474) synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 106. Strophiops hesternia Maynard and Clapp, 1915 [February 8]: 180 bis [not illustrated in original publica- tion]; Maynard, 1921b: 142; Maynard, Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 1924a, pl. 38, figs. 9, 10. Examined 200 specimens. Measurements not provided. Type Material. The original description did not distinguish among the 200 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 200 specimens are syntypes. No specimen labeled “Holotype” was located at the MCZ. An examination of the MCZ catalog revealed that no spec- imen identified as “type” was present in Maynard's collection at the time of its sale. Lot number MCZ 76157 contained 27 specimens labeled “paratypes” together with a note in Maynard’s handwriting list- ing the taxon name and locality, but lack- ing the word “type” or references to the illustrations. Gould and Woodruff (1986: 474) reported examining “paratypes” but did not see a “holotype.” One of these specimens, which most closely matches Maynard’s only illustration of this taxon in size and proportions, is here designated as the lectotype to provide an objective stan- dard of reference for this species-group taxon that is consistent with Maynard's concept of it. The remaining 26 specimens were recatalogued as MCZ 356994 and are among the 199 syntypes that become par- alectotypes. Lectotype Measurements. Length 27.9 mm, diameter (excluding lip) 11.6 mm; ap- erture height (including lip) 11.2 mm, ap- erture width (including lip and peristome) 9.6 mm. Type Locality. West end of Booby Rock [16 miles NE of New Providence Island, Bahamas] by a low place over which the sea frequently breaks. Remarks. Gould and Woodruff (1986: 474) synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 107. Strophiops caerulea Maynard and Clapp, 1915 [February 8]: 181 [not illustrated in original publication]; Maynard, 1921b: 140, pl. 36, figs. 7, 8. Examined 50 specimens. Size given as .92 by .40 [inches; 23.4 by 10.2 mm], but the term “type” was not used. THE CERION TAXA OF CHARLES JOHNSON MAYNARD °* Harasewych et al. Type Material. The original description did not distinguish among the 50 speci- mens examined, and there was no mention of type material in the other parts of this publication. Thus, all 50 specimens are syntypes. Lot number MCZ 76308 con- tains a single specimen labeled “Holotype” that approximates the illustration and the measurements of this taxon and is accom- panied by a label in Maynard’s hand with the term “type,” which includes references to subsequently published figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original con- cept of it. The remaining 49 syntypes be- come paralectotypes. Lectotype Measurements. Length 22.8 mm, diameter (excluding lip) 10.1 mm; ap- erture height (including lip) 9.0 mm, ap- erture width (including lip and peristome) ToD) maven, Type Locality. In field N of Ft. Char- lotte, Nassau [New Providence Island, Ba- hamas]. Remarks. Maynard considered this tax- on to be extinct at the time it was de- scribed. This taxon was subsequently illus- trated (Maynard, 1921b, pl. 36, figs. 7, 8). Gould and Woodruff (1986: 474) synony- mized this taxon with Cerion glans (Kiis- ter, 1844). Taxon 108. Strophiops devereuxi May- nard and Clapp, 1915 [February 8]: 181 [not illustrated in original publica- tion]; Maynard, 1921b: 140, pl. 36, figs. 9, 10. Unspecified number of specimens examined. Size given as .85 by .40 [inch- es; 21.6 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not specify the number of specimens examined. In the subsequent monograph, Maynard (1921b: 140) lists 164 specimens. The size of the syntype lot is uncertain but does not exceed 164 syntypes. Lot number MCZ 76137 contains a single specimen la- 435 beled “Holotype” that approximates the measurements and subsequently pub- lished illustration of this taxon and is ac- companied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining syntypes (=163), including MCZ 76138 and MCZ 117730 (one specimen), become paralectotypes. Lectotype Measurements. Length 20.9 mm, diameter (excluding lip) 9.7 mm; ap- erture height (including lip) 8.4 mm, ap- erture width (including lip and peristome) 6.9 mm. Type Locality. Devereux Estate west end of New Providence [Island, Bahamas]. Remarks. This taxon was later listed as S. deveruxi (Maynard, 1921b: 140) and S. devereuaxi (Maynard, 1924?: [7]) and il- lustrated under its correct original spelling (Maynard, 1921b, pl. 36, figs. 9, 10). Gould and Woodruff (1986: 475) synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 109. Strophiops praedicta Maynard and Clapp, 1915 [February 8]: 181 [not illustrated]. Number of specimens not specified; measurements not provid- ed. Type Material. The original description did not specify the number of specimens examined. This taxon does not appear in any of Maynard’s subsequent publications or catalogs. The size of the type series is uncertain. An examination of the MCZ catalog revealed that no specimen identi- fied as “type” was present in Maynard's collection at the time of its sale. Lot MCZ 76393 contains 91 specimens (many dam- aged, incomplete, or juvenile) and was la- beled “paratype.” Because no characteris- tics were provided by which an individual specimen could be identified, a large, ma- ture, complete specimen is selected from 436 the syntype series to serve as lectotype to provide an objective standard of reference for this species-group taxon. The remain- ing 90 specimens were recatalogued as MCZ 356995 and are paralectotypes. Lectotype Measurements. Length 29.5 mm, diameter (excluding lip) 13.4 mm; ap- erture height (including lip) 11.3 mm, ap- erture width (including lip and peristome) 10.0 mm. Type Locality. Great Pimlico [Island, Eleuthera, Bahamas]. Remarks. Fossil in the rocks. Clench (1952: 110) listed this taxon as a synonym of Cerion uniforme (Maynard, 1913). Taxon 110. Strophiops prisca Maynard and Clapp, 1915 [February 8]: 182 [not illustrated in original publication]; Maynard, 1921b: 143; Maynard, 1924a, pl. 39, figs. 5, 6. Unspecified number of specimens examined. Size given as 1.15 by .45 [inches; 29.2 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not specify the number of specimens examined. In the subsequent monograph, Maynard (1921b: 143) lists 15 specimens. The size of the syntype lot is uncertain but does not exceed 15 syntypes. Lot MCZ 76353 contains a single specimen labeled “Holotype” that approximates the mea- surements of this taxon as well as the sub- sequently published illustrations. This specimen is accompanied by a label in Maynard’s hand with the term “type,” which includes references to these illustra- tions. This specimen is here designated as the lectotype to provide an objective stan- dard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining syn- types (=14), including MCZ 76352, be- come paralectotypes. Lectotype Measurements. Length 29.3 mm, diameter (excluding lip) 11.7 mm; ap- erture height (including lip) 10.7 mm, ap- erture width (including lip and peristome) 7.9 mm. Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Type Locality. Fossil in the rocks on east end of Salt Key [New Providence Island, Bahamas]. Remarks. Maynard (1921b: 143, 1924a, pl. 39, figs. 5, 6) noted 15 specimens and provided illustrations for this taxon. Gould and Woodruff (1986: 482) synonymized this taxon with Cerion glans (Kiister, 1844). Taxon 111. Strophiops tenui Maynard and Clapp, 1915 [February 8]: 182 [not illustrated in original publication]; Maynard, 1921b: 149-150, pl. 46, figs. - 1, 2 [pl. 46 was never issued]. Unspec- ified number of specimens examined. Size given as .70 by .33 [inches; 17.8 by 8.4 mm], but the term “type” was not used. Type Material. The original description did not specify the number of specimens examined. In the subsequent monograph, Maynard (1921b: 150) lists 600 specimens. The size of the syntype lot is uncertain, but does not exceed 600 syntypes. Lot MCZ 76350 contains a single specimen la- beled “Holotype” that approximates the measurements of this taxon. This speci- men is accompanied by a label in May- nard’s hand with the term “type,” which includes references to illustrations that were never published. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining syntypes (=599), including MCZ 76351, become paralectotypes. Lectotype Measurements. Length 17.6 mm, diameter (excluding lip) 8.5 mm; ap- erture height (including lip) 7.8 mm, ap- erture width (including lip and peristome) 6.8 mm. Type Locality. Isolated on a portion of shore on east end of New Providence [Is- land, Bahamas]. Remarks. Maynard and Clapp (1915: 182) considered this taxon to be extinct at the time it was described, but later, May- THE CERION TAXA OF CHARLES JOHNSON MAYNARD ° Harasewych et al. nard (1921b: 149-150) wrote that they were common along a short reach of shore just west of East Point Light, New Provi- dence, and that 600 specimens were col- lected, chiefly dead. Gould and Woodruff (1986: 477-478) synonymized this taxon with Cerion gubernatorium (Crosse, 1869). Taxon 112. Strophiops fulminea Maynard and Clapp, 1915 [February 8]: 182 [not illustrated in original publication]; Maynard, 1921b: 149. Examined 1,000 specimens. Measurements not provided. Type Material. The original description did not distinguish among the 1,000 spec- imens examined, and there was no men- tion of type material in the other parts of this publication. Thus, all 1,000 specimens are syntypes. Lot MCZ 76276 was labeled “holotype,” but contains three specimens and a label in Maynard’s hand containing the term “Type” as well as plate and figure numbers (pl. 45, figs. 9, 10) for illustra- tions of this taxon that were never pub- lished. The specimen that most closely ap- proximates the subsequently published measurements (Maynard, 1921b: 149) is here designated as the lectotype. The re- maining two specimens, recatalogued as MCZ 356996, MCZ 76277 (262 speci- mens), and USNM 420088 (50. speci- mens), are among the 999 syntypes that become paralectotypes. Lectotype Measurements. Length 25.2 mm, diameter (excluding lip) 9.7 mm; ap- erture height (including lip) 9.4 mm, ap- erture width (including lip and peristome) $.2 mm. Type Locality. Very common on rocky shore east of Ft. Winton, New Providence [Island, Bahamas]. Remarks. In the original description, Maynard noted that nearly all were dead in 1913. Later, Maynard (1921b: 149) pro- vided measurements (1.00 by .40 [inches; 25.4 by 10.2 mm]); listed 2,000 specimens, likely by subsequent collection; and ref- erenced illustrations (pl. 45, figs. 5, 6) that were never issued. 437 Taxon 113. Strophiops longidentata May- nard, 1919a [October 16]: 24 [nomen nudum]. Remarks. Not listed in Clench (1957). Taxon 114. Strophiops peravita Maynard, 1919b [November 6]: 47; Maynard, 1O20dsSeptemben 350i ple etiess IF 2 [captioned “S. perantiqua M. & C.”]. Nomenclatural Remarks. The binomen Strophiops peravita was used by Maynard (1919b: 47) as a example of a species with a conically tapering shell and was accom- anied by a reference to subsequently published figures (Maynard, 1920d, pl. 1, figs. 1, 2) that were captioned “S. peran- Higua Win Ga) Cr eeAnticle 1Osleie EZN: 1999: 9) states that if publication of the data relating to a new nominal taxon or a nomenclatural act is interrupted and con- tinued at a later date, the name or act be- comes available only when the require- ments of the relevant Articles have been met. Strophiops peravita became available on November 6, 1919, on the basis of the publication of the meager description, which was adequate to meet the require- ments of the relevant article [Article 12.1, ICZN, 1999: 16], rather than from the lat- er publication of the captioned figure. Au- thorship is attributed to Maynard, the sole author of the description, rather than to Maynard and Clapp, as specified in the caption to the subsequently published il- lustrations. Ironically, this is the only time that Maynard used the binomen Stro- phiops peravita. It is a lapsus calami be- cause Maynard used the name Strophiops perantiqua in two prior references (May- nard, 1919a [October 16]: 15, 1919b [| No- vember 6]: 31; in both instances as a no- men nudum) and one subsequent refer- ence (Maynard and Clapp in Maynard, 1920d [September 30]: 115, map 1, 2, pl. 1, figs. 1, 2) to this taxon. The figure re- ferred to in the inadvertent description of S. peravita is the identical figure refer- enced in the intended description of this taxon [as Strophiops perantiqua], in which 438 it was noted that it was “contemporary with S. avita Mayn.” Type Material. The description of S. peravita did not specify the number of specimens examined but included a ref- erence to a subsequently published figure of a single specimen. The 25 specimens mentioned in the intended description of this taxon (as S. perantiqua Maynard and Clapp in Maynard, 1920d: 115) were avail- able to Maynard in 1919 and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops peravita is thus based on 25 syntypes. Lot number MCZ 187537 contains a single specimen that closely matches the illustration mentioned in the taxon description. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of this taxon. The remaining 24 syntypes become para- lectotypes. Lectotype Measurements. Length SLY) mm, diameter (excluding lip) 14.7 mm; ap- erture height (including lip) 11.1 mm, ap- erture width (including lip and peristome) 9.5 mm. Type Locality. Found fossil in red earth near west coast of Great Guana Key |Ex- umas, Bahamas], about 1 mile from south- ern end (see Maynard, 1920d: 117, map 1, location 2 [from the intended taxon de- scription of Strophiops perantiqua]). May- nard (1924a: 164) described in detail the locality in which these fossils were collect- ed. Remarks. Although it seems clear that Maynard intended the taxon to be known as perantiqua, it is not possible to consider peravita as an “incorrect original spelling” Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 within the confines of Article 32.5.1 (ICZN, 1999: 39) because there is no clear evidence to suggest an alternative spelling within the original publication itself. The name S. peravita did not appear in the Batchelder (1951) compilation of May- nard’s taxa, nor in Clench’s (1957) catalog of Cerion taxa. The lectotype of S. peravita is also the lectotype of Strophiops peran- tiqua Maynard and Clapp in Maynard, 1920, which is a junior objective synonym of S. peravita [see Taxon 139, below]. Taxon 115. Strophiops scalariformis May- nard, 1919b [November 6]: 47-48: Maynard and Clapp in Maynard, 1920d [September 30]; 116, map 1; 3, pl. 1, figs. 5, 6 [captioned “S. scalariformis M. & C.”]. Examined 50 specimens. Size given as 1.20 by .45 [inches; 38.1 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. Maynard (1919b: 48) first used the binomen Stro- phiops scalariformis as a example of a spe- cies with a turreted shell and included a reference to subsequently published fig- ures (Maynard, 1920d, pl. 1, figs. 5, 6) that were captioned “S. scalariformis M. & C.” Article 10.1.1 (ICZN, 1999: 9) states that if publication of the data relating to a new nominal taxon or a nomenclatural act is in- terrupted and continued at a later date, the name or act becomes available only when the requirements of the relevant Ar- ticle have been met. Strophiops scalarifor- mis became available on November 6, 1919, on the basis of the publication of the meager description, which was adequate to meet the requirements of the relevant articles [Article 12.1, ICZN, 1999: 16], —-= Figures 114-125. Taxon 114. Strophiops peravita Maynard, 1919. Lectotype MCZ 187537. Fossil in red earth near west coast of Great Guana Key, Exuma Group, Bahamas, about 1 mile from southern end. Taxon 115. Strophiops scalariformis Maynard, 1919. Lectotype MCZ 76180. Limited area near beach Great Guana Key, Exuma Group, Bahamas. Taxon 116. Strophiops inornata Maynard, 1919. Lectotype MCZ 76049. North side of Bell Key, Exuma Group, Bahamas. Taxon 117. Strophiops caprana Maynard, 1919. Lectotype MCZ 76386. North Goat Key off Fresh Creek, Andros, Bahamas. Taxon 118. Strophiops lobata Maynard, 1919. Lectotype MCZ 76207. Eastern Soldier Key, Berry Islands, Bahamas. Taxon 119. Strophiops variata Maynard and Clapp in Maynard, 1919. Lectotype MCZ 76270. Crab Key, Berry Islands, Bahamas. Taxon 120. Strophiops recessa Maynard and Clapp in Maynard, 1919. Lectotype MCZ 76135. Southern portion of Stocking Island, Exuma Group, Bahamas. Taxon 121. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 439 Ler AUR AS ‘ ; Ww Ny) \ Ps " penned “A : - Strophiops palmata Maynard and Clapp in Maynard, 1919. Lectotype MCZ 76179. East and west sides of Wax Key, Exuma Group, Bahamas. Taxon 122. Strophiops agricola Maynard and Clapp in Maynard, 1919. Lectotype MCZ 76004. Farmer’s Key which lies about 1 mile southwest of the southern end of Great Guana Key, Exuma Group, Bahamas. Taxon 123. Strophiops elegantissima Maynard, 1919. Lectotype MCZ 76375. Norman’s Pond Key, Exuma Group, Bahamas. Taxon 124. Strophiops arbusta Maynard and Clapp in Maynard, 1919. Lectotype MCZ 76273. Guana Key, Berry Islands, Bahamas. Taxon 125. Stro- phiops aspera Maynard, 1920. Lectotype MCZ 76176. Among dwarf palms in an area about 300 yards long by some 150 wide along the third beach north of south end of Great Guana Key, Exuma Group, Bahamas. 440 rather than from the later publication of the captioned figure. Authorship is attri- buted to Maynard, the sole author of the description, rather than to Maynard and Clapp, as specified in the caption to the subsequently published illustrations. Type Material. The description of S. scalariformis did not specify the number of specimens examined, but included a reference to a subsequently published fig- ure of a single specimen. The 50 speci- mens mentioned in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 1920d: 116) were likely available to Maynard in 1919 and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops scalariformis is thus based on 50 syntypes. In a subsequent and more detailed description of this taxon, Maynard (1924a: 165) included the state- ment “dimensions of type .45 by 1.20 [inches; 11.4 by 30.5 mm], aperture .34 linches; 8.6 mm].” This is regarded as a lectotype designation, and the remaining 49 specimens became paralectotypes. Clench and Aguayo (1952: 430, pl. 56, fig. 2) listed and illustrated MCZ 76180 as the “holotype,” presumably on the basis of Maynard’s handwritten label using the term “type” that accompanied the speci- men. This specimen is the lectotype des- ignated by Maynard. Lectotype Measurements. Length 30.0 mim, diameter (excluding lip) 11.3 mm; ap- erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) 9.2 mm. Type Locality. Found in a limited area near beach [Great Guana Key, Exuma Group, Bahamas] (see Maynard, 1920d: 117, map 1, location 3). Maynard (1924a: 165) described in detail the locality in which these fossils were collected. Remarks. Clench and Aguayo (1952) considered Cerion scalariforme to be a synonym of Cerion asperum (Maynard and Clapp, 1920). Taxon 116. Strophiops inornata Maynard, 1919b [November 6]: 49; Maynard, Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 1920b [April 8]: 87; Maynard, 1920c [July 10], fig. 73, pl. 17, fig. 2; Maynard, 1920d [September 30]: 112; Maynard and Clapp in Maynard, 1920d [Septem- ber 30]: 126, pl. 3, figs. 11-13. Exam- ined 1,060 specimens. Size given as 1.12 by .45 [inches; 28.4 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. Maynard (1919b: 49) first used the binomen Stro- phiops inornata as a example of a taxon in which the ultimate whorl and aperture are contracted, and included a reference to. subsequently published figures (Maynard, 1920d, pl. 3, figs. 11-13) that were cap- tioned “S. inornata.” Article 10.1.1 (ICZN, 1999: 9) states that if publication of the data relating to a new nominal taxon or a nomenclatural act is interrupted and con- tinued at a later date, the name or act be- comes available only when the require- ments of the relevant Articles have been met. Strophiops inornata became available on November 6, 1919, on the basis of the publication of the meager description, which was adequate to meet the require- ments of the relevant articles | Article 12.1, ICZN, 1999: 16], rather than from the lat- er publication of the captioned figure. Au- thorship is attributed to Maynard, the sole author of the description. Before the publication of plate 3, May- nard (1920b [April 8]: 87) again used the name S. inornata as an example of mor- phological reversion following shell dam- age and cited a figure of a different spec- imen (Maynard, 1920c [July 10], fig. 73) that was reproduced in color (Maynard, 1920c, pl. 17, fig. 2). In the next issue, Maynard (1920d [September 30]: 112) published a correction stating that figure numbers 72 and 73, as well as plate 17, figures 1 and 2, were transposed and that the figures 73 and plate 17, figure 2, are actually S. salinaria, whereas figures 72 and plate 17, figure 1, represent S. inor- nata. Type Material. The description of Stro- phiops inornata did not specify the num- THE CERION TAXA OF CHARLES JOHNSON MAYNARD ° Harasewych et al. ber of specimens examined but included a reference to a subsequently published fig- ure of a single specimen. The 1,060 spec- imens mentioned in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 1920d: 126) were likely available to Maynard in 1919 and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops inornata is thus based on 1,060 syntypes. Lot MCZ 76049 was labeled “Holotype” but contained three specimens. Of these, one has an off- set final whorl and clearly matches the fig- ures cited in the inadvertent as well as the intended taxon description (Maynard and Clapp in Maynard, 1920d [September 30]: 126, pl. 3, figs. 11-13). This specimen is designated to be the lectotype of Stro- phiops inornata Maynard, 1919, to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining two specimens were recatalogued as MCZ 357001 and together with MCZ 76050, MCZ 118190, and USNM 420056, become paralectotypes. Lectotype Measurements. Length 31.9 mm, diameter (excluding lip) 13.7 mm; ap- erture height (including lip) 11.6 mm, ap- erture width (including lip and peristome) 11.0 mm. Type Locality. North side of Bell Key [Exuma Group, Bahamas]. Taxon 117. Strophiops caprana Maynard, 1919b [November 6]: 52. Strophiops capraia Maynard and Clapp in May- mard, 1919c [December 31], fig. 19; Maynard, 1920c, pl. 17, fig. 3; Maynard and Clapp in Maynard, 1921a: 131, pl. 28, figs. 7, 8. Examined 1,620 speci- mens. Size given as 1.12 by .47 [inches; 28.4 by 17.8 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen Strophiops caprana was first used by May- nard (1919b: 52) as an example of a spe- cies with a thick margin, and accompanied by a reference to a subsequently published 44] figure (Maynard, 1919c, fig. 19) that was captioned “S. capraia M. & C.” A colored version of this figure, also labeled S. ca- praia, was published (Maynard, 1920c, pl. Le, fig. 3). Article 10.1.1 (ICZN, 1999: 9) states that if publication of the data relat- ing to a new nominal taxon or a nomen- clatural act is interrupted and continued at a later date, the name or act becomes available only when the requirements of the relevant Articles have been met. Stro- phiops caprana became available on No- vember 6, 1919, on the basis of the pub- lication of the meager description, which was adequate to meet the requirements of the relevant articles [Article 12.1, ICZN, 1999: 16], rather than from the later pub- lication of the captioned figure. Author- ship is attributed to Maynard, the sole au- thor of the description, rather than to Maynard and Clapp as specified in the caption to the subsequently published il- lustrations. Type Material. The description of Stro- phiops caprana did not specify the num- ber of specimens examined but included a reference to a subsequently published fig- ure of a single specimen. The 1,620 spec- imens mentioned in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 1921a: 131) were likely available to Maynard in 1919 and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops caprana is thus based on 1,620 syntypes. The specimen il- lustrated in the originally referenced figure (Maynard, 1919c, fig. 19) could not be identified at the MCZ. Maynard clearly re- garded a different specimen to be the “Type” of Strophiops capraia. Lot MCZ 76386 was labeled “Holotype” but con- tains two specimens. One of the specimens approximates the measurements and fig- ures in the intended taxon description and is here selected as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of this taxon. The re- maining specimen, recatalogued as MCZ 356997, MCZ 76215, and USNM 420060, 442 are among the 1,619 syntypes that become paralectotypes. Lectotype Measurements. Length 28.9 mm, diameter (excluding lip) 12.2 mm; ap- erture height (including lip) 10.4 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. North Goat Key off Fresh Creek [Andros, Bahamas]. Remarks. This taxon has been the victim of multiple lapsus calami, having been spelled caprana in the unintended original description (Maynard, 1919b: 52), capraia in text and figure captions (Maynard, 1919¢, fig. 19, 1920b: 81, 1920e, pl. 17, fig: 3) and in the intended taxon description (Maynard and Clapp in Maynard, 1921a: 131, pl. 28, figs. 7, 8), and capria (May- nard, 1924bP: [3]) in the sales catalog. Al- though it seems clear that Maynard in- tended the taxon to be known as capraia, it is not possible to consider caprana as an “incorrect original spelling” within the confines of Article 32.5.1 (ICZN, 1999: 39) because there is no clear evidence to sug- gest an alternative spelling within the orig- inal publication itself. This taxon was also listed as S. capraia by Clench (1957: 139). Taxon 118. Strophiops lobata Maynard, 1919b [November 6]: 52; Maynard and Clapp in Maynard, 1919c |Decem- ber 31], fig. 20; Maynard, 1920b: 81; Maynard, 1920c, fig. 64, pl. 14, fig. 4; Maynard and Clapp in Maynard, 192 1a: 134, pl. 31 [erroneously listed as pl. 30], figs. 7, 8. Examined 925 specimens. Size given as 1.11 by .45 [inches; 30.5 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen Strophiops lobata was used by Maynard (1919b: 527) as an example of a species with a thickened and lobed margin and in- cluded a reference to a subsequently pub- lished figure (Maynard, 1919c, fig. 20) that was labeled “S. lobata M. & C.” Article 10.1.1 (ICZN, 1999: 9) states that if pub- lication of the data relating to a new nom- inal taxon or a nomenclatural act is inter- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 rupted and continued at a later date, the name or act becomes available only when the requirements of the relevant Articles have been met. Strophiops lobata became available on November 6, 1919, on the ba- sis of the publication of the meager de- scription, which was adequate to meet the requirements of the relevant articles [Ar- ticle 12.1, ICZN, 1999: 16], rather than from the later publication of the captioned figure. Authorship is attributed to May- nard, the sole author of the description, rather than to Maynard and Clapp, as specified in the caption to the subsequent- ly published illustrations. Type Material. The unintended taxon description of Strophiops lobata did not specify the number of specimens exam- ined but included a reference to a subse- quently published figure of a single spec- imen. The 925 specimens mentioned in the intended description of this taxon (Maynard and Clapp in Maynard, 1921a: 134) were likely available to Maynard in 1919 and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Stro- phiops lobata is thus based on 925 syn- types. The specimen illustrated in figure 20 could not be identified at the MCZ. Maynard clearly regarded a different spec- imen to be the “Type” of Strophiops lob- ata. Lot MCZ 76207 contains a single specimen labeled “Holotype” that approx- imates the illustration and the measure- ments of this taxon in the intended taxon description and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to these figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining specimens, including MCZ 76208 and USNM 420147, are paralectotypes. Lectotype Measurements. Length 29.1 mm, diameter (excluding lip) 11.9 mm; ap- erture height (including lip) 11.4 mm, ap- erture width (including lip and per one) 8.9 mm. THE CERION TAXA OF CHARLES JOHNSON MAYNARD ¢ Harasewych et al. Type Locality. Eastern Soldier Key, Ber- ry Islands, Bahamas. Remarks. Maynard (1920b: 81) again used the name S. lobata prior to the in- tended description of the taxon, referenc- ing a subsequently published figure (May- nard, 1920c, fig. 64) that was identical to figure 20 but captioned “S. caduca M. & C.” A colored version of figures 20 and 64 (Maynard, 1920c, pl. 14, fig. 4) is cap- tioned “S. caduca.” Maynard (1920d: 112) published corrections, stating that the cap- tions to figures 64 and pl. 14, fig. 4, should read lobata not caduca. Taxon 119. Strophiops variata Maynard and Clapp in Maynard, 1919c [De- cember 31]: 55, figs. 27, 28; Maynard, 1920a: 77, 78, fig. 54; Maynard, 1920c, pl. 14, fig. 5; Maynard and Clapp in Maynard, 1921a: 134, pl. 32, figs. 1, 2. Examined 1,132 specimens. Measure- ments not provided. Nomenclatural Remarks. The binomen Strophiops variata was used by Maynard (1919c: 55, figs. 27, 28) as an example of a taxon with margins malformed after in- corporation of a limestone granule in the peristome and accompanied by figures of three specimens labeled “S. variata M. & C.” The association of a new species group name with an illustration of the taxon be- ing named before 1931 makes the name Strophiops variata Maynard and Clapp in Maynard, 1919c [December 31]: 55, figs. 27, 28. available (Article 12.2.7, ICZN, 1999: 17) prior to the intended description of this species (Maynard and Clapp in Maynard, 1921a [May 2]: 134, pl. 32, figs. ee 2), Type Material. Maynard’s (1919c: 55— 56) first use of the name S. variata was as an example of a deformation that could oc- cur in some specimens after incorporation of a limestone granule in the peristome. He mentioned that this produced a pro- tuberance in a total of six specimens of S. variata and illustrated portions of three of the malformed specimens. It is likely that the six deformed specimens were selected 443 from among the 1,132 specimens men- tioned in the intended description of this taxon (Maynard and Clapp in Maynard, 1921a: 134) that were available to him at the time. All 1,132 specimens are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76], and Strophiops lobata is based on 1,132 syntypes. None of the figured specimens could be identified at the MCZ. Maynard clearly regarded a different spec- imen to be the “Type” of Strophiops var- iata. Lot MCZ 76270 contained two spec- imens in a lot labeled “Holotype,” one smooth, with weak costae limited to the final whorl, the other with strong costae over the entire shell. The smooth speci- men, which more closely agrees with the intended taxon description and figures, is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The strongly costate specimen was recata- logued as MCZ 356998. It and the re- maining specimens, including MCZ 76271, MCZ 118258, and USNM 420123, are paralectotypes. Lectotype Measurements. Length Ses mm, diameter (excluding lip) 10.7 mm; ap- erture height (including lip) 10.7 mm, ap- erture width (including lip and peristome) 8.8 mm. Type Locality. Crab Key, Berry Islands, Bahamas. Remarks. Other uses of the binomen prior to its intended original description include Maynard (1920a: 77, 78, fig. 54, 1920c, caption to pl. 14, fig. 5). Taxon 120. Strophiops recessa Maynard and Clapp in Maynard, 1919c [De- cember 31]: 57, figs. 29A, B; May- nard, 1920c, pl. 17, fig. 6; Maynard and Clapp in Maynard, 1920d: 122, pl. 24, figs. 2, 3. Examined 1,147 specimens. Size given as 1.15 by .45 [inches; 29.2 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. Maynard (1919¢: 57, figs. 29A, B) first used the bi- +4 nomen Strophiops recessa as an example of a species that has an enlarged, ear-like frontal bar encroaching on the umbilicus and included illustrations (figs. 29A, B) captioned “S. recessa M. & C.” The asso- ciation of a new species group name with an illustration of the taxon being named before 1931 makes the name Strophiops recessa Maynard and Clapp in Maynard, 1919c (p. 57, figs. 29A, B), available on December 31, 1919 (Article 12.2.7, ICZN, 1999: 17), prior to the intended descrip- tion of this species (Maynard and Clapp in Maynard, 1920d: 122, pl. 24, figs. 2, 3). Type Material. The description of Stro- phiops recessa did not specify the number of specimens examined but included fig- ures of a single specimen. The 1,147 spec- imens mentioned in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 1920d: 122) were likely available to Maynard at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops recessa is based on 1,147 syntypes. The originally illustrated specimen (Maynard, 1919c, figs. 29A, B) could not be identified at the MCZ. Maynard clearly regarded a different specimen to be the ily peerol Strophiops recessa. Lot MCZ 76135 contained a single specimen labeled “Holotype” that approximates the mea- surements and illustration of this taxon in the intended description (Maynard and Clapp in Maynard, 1920d: 122, pl. 24, figs. 2, 3) and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen (MCZ 76135) is here des- ignated as the lectotype of Strophiops re- cessa Maynard and Clapp in Maynard, 1919, to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original con- cept of it. The remaining specimens, in- cluding MCZ 76136, MCZ 118346, and USNM 420106, are paralectotypes. Lectotype Measurements. Length 29.9 mm, diameter (excluding lip) 11.3 mm; ap- erture height (including lip) 10.3 mm, ap- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 erture width (including lip and peristome) 8.1 mm. Type Locality. Southern portion of Stocking Island, Exuma Cays, Bahamas. Remarks. It is not surprising that, after Maynard's death, the specimens labeled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens were dispersed among MCZ, USNM, and other institu- tions (Turner, 1957: 151). Given the qual- ity of Maynard’s illustrations, it is unlikely that the initially illustrated specimen could ever be discerned unambiguously from among the large number of remaining specimens, even if it were present among them. Taxon 121. Strophiops palmata Maynard and Clapp in Maynard, 1919c [De- cember 31]: 59-60, 64, figs. 32, 32*, 33, 34A_D, 35, [35]B, pl. 10, figs. 1, 2, 8; Maynard, 1920a: 76, 77; Maynard, 1920b, fig. 58, pl. 12, fig. 2; Maynard, 1920e: 86, fig. 67, pl. 14, fig. 2, fig. 71, pl. 16, figs. 4, 5; Maynard and Clapp in Maynard, 1920d: 120, pl. 21, figs. 6, 7; Maynard, 1926: 203-212. Examined 3,422 specimens. Size given as 1.25 by 45 l[inches; 31.8 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen Strophiops palmata was first used by May- nard (1919c: 59-60, 64, figs. 32, 32*, 33, 34A—D, 35, [35]B, pl. 10, figs. 1, 2, 8) to illustrate pathological duplication and/or migration of teeth within the shell aper- ture after shell damage. Portions of at least six different specimens are illustrated in these figures, of which figures 32, 32*, and 33 and pl. 10, figs. 1, 2, and 8, are cap- tioned “S. palmata M & C.,” whereas fig- ures 34A—D, 35, and 35B are captioned “S. palmata.” The association of a new species group name with an illustration of the taxon being named before 1931 makes the name Strophiops palmata Maynard and Clapp in Maynard 1919c (pp. 59-60, 64, figs. 32, 32*, 33, 34A_D, 35, [35]B, pl. THE CERION TAXA OF CHARLES JOHNSON MAYNARD °* Harasewych et al. 10, figs. 1, 2, 8), available (Article 12.2.7, ICZN, 1999: 17) prior to the intended de- scription of this species (Maynard and Clapp in Maynard, 1920d: 120, pl. 21, figs. O77). Maynard (1920a: 76, 77) again used S. palmata as an example of a taxon that ag- glutinates sand, citing a subsequently pub- lished illustration of another specimen (Maynard, 1920b, fig. 58, pl. 12, fig. 2). Additional damaged specimens are illus- trated (Maynard, 1920c: 86, fig. 67, re- peated on pl. 14, fig. 2, fig. 71, repeated on pl. 16, figs. 4, 5). All of these uses of the name preceded the intended original description of Strophiops palmata. Type Material. The description of Stro- phiops palmata did not specify the num- ber of specimens examined, but included figures of portions of six specimens. The 3,422 specimens mentioned in the intend- ed description of this taxon (Maynard and Clapp in Maynard, 1920d: 120) were likely available to Maynard at the time, and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops re- cessa is based on 3,422 syntypes. None of the six initially figured specimens could be identified at the MCZ. Maynard clearly re- garded a different specimen to be the “Type” of Strophiops palmata. Lot MCZ 76179 contains a single specimen labeled “Holotype” that approximates the mea- surements and illustration of this taxon in the intended description (Maynard and Clapp in Maynard, 1920d: 122, pl. 24, figs. 2, 3) and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen (MCZ 76179) is here des- ignated as the lectotype of Strophiops pal- mata Maynard and Clapp in Maynard, 1919, to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 3,415 specimens, including MCZ 76178, MCZ 118352, and USNM 420024, are paralectotypes. Lectotype Measurements. Length 27.6 mm, diameter (excluding lip) 11.0 mm; ap- 445 erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) Tail saan Type Locality. East and west sides of Wax Key [Exuma Group, Bahamas]. Remarks. It is not surprising that, fol- lowing Maynard's death, the specimens la- beled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens were dis- persed among MCZ, USNM, and other in- stitutions (Turner, 1957: 151). The anom- alies in dentition make the six illustrated specimens distinctive, yet they were not found in the collections of either MCZ or USNM. Maynard (1920d: 120) reported this taxon to be very abundant on palm fronds and under them. Maynard (1926: 203-212) provided additional, detailed in- formation on the animals and anatomy, noting that 2,683 specimens were collect- ed in 2 or 3 hours. Taxon 122. Strophiops agricola Maynard and Clapp in Maynard, 1919c [De- cember 31]: 64, figs. 35C, D; May- nard, 1920d, pl. 22, figs. 6, 7; Maynard, 1924c: 1. Examined 80 specimens. Size given as 1.15 by .40 [inches; 29.2 by 10.2 mm], but the term “type” was not used. Nomenclatural Remarks. The first use of the binomen Strophiops agricola appeared in a discussion of the occasional appear- ance of a tooth-like concretion along the shell margin (Maynard, 1919c, figs. 35C, D, p. 64). The caption to the figures, which illustrate the malformed aperture of a specimen (fig. 35C) as well as an enlarge- ment of the concretion (fig. 35D), reads “S. agricola M & C..,” establishing the au- thorship of the taxon as Maynard and Clapp in Maynard. The association of a new species group name with an illustra- tion of the taxon being named before 1931 makes the name Strophiops agricola May- nard and Clapp in Maynard, 1919c (figs. 30C, D), available (Article 12.2.7, ICZN, 1999: 17) prior to the intended description of this species (Maynard, 1924c: 1). The 446 name was next used in the caption to il- lustrations (Maynard, 1920d, pl. 22, figs. 6, 7) published 4 years before the intended description. Type Material. Only a single specimen was referred to in the inadvertent descrip- tion of Strophiops agricola. The intended taxon description (Maynard, 1924c: 1) mentioned 80 specimens. It is likely that at least some if not all of these were avail- able to Maynard in 1919. Strophiops agri- cola is thus based on =80 syntypes. Neither of the figures on which the tax- on is based shows an entire specimen; rather, they provide details of a concretion on a portion of the outer lip. An exami- nation of the 49 specimens in MCZ 76004 that were labeled “paratypes” uncovered the originally illustrated specimen (May- nard, 1919c, fig. 35). This specimen is here designated as the lectotype. The remaining specimens were recatalogued as MCZ 356999 and together with MCZ 76003 and MCZ 118252 are among the =79 speci- mens that become paralectotypes. Lectotype Measurements. Length 28.0 mm, diameter (excluding lip) 10.7 mm; ap- erture height (including lip) 9.1 mm, ap- erture width (including lip and peristome) Tt) Tamia, Type Locality. Farmer's Key which lies about 1 mile southwest of the southern end of Great Guana Key, Exuma Group, Bahamas. Remarks. Maynard clearly regarded a different specimen to be the “Type” of Strophiops agricola. Lot MCZ 76003 con- tains a single specimen labeled “Holotype” that approximates the subsequent illustra- tions (Maynard, 1920d, pl. 22, figs. 6, 7) and the measurements in the intended de- scription (Maynard, 1924c: 1). This speci- men, a paralectotype, is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the previously published plates. Taxon 123. Strophiops elegantissima May- nard, 1919c [December 31]: 64, figs. 36A-—D; Maynard, 1920a: 80; Maynard, Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 1920b, fig. 60, pl. 13, fig. 2; Multistro- phia elegantissima Maynard and Clapp in Maynard, 1920d: 126, pl. 25, figs. 1, 2. Unspecified number of specimens ex- amined. Size given as 1.25 by .50 [inch- es; 31.8 by 12.7 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen Strophiops elegantissima was first pub- lished by Maynard (1919c: 64, figs. 36A— D) together with illustrations labeled “S. elegantissima.” The text provided detailed descriptions of dentition stated to be a specific character in S. elegantissima. The name Strophia elegantissima Maynard be- comes available on December 31, 1919 (Articles 12.1 and 12.2.7, ICZN, 1999: 17), prior to the intended description of the taxon as Multicostata elegantissima May- nard and Clapp in Maynard, 1920d (p. 126, pl. 25, figs. 1, 2). Because figures 36A—D are captioned “S. elegantissima,” the taxon must be attributed to Maynard rather than to Maynard and Clapp, as cit- ed by Clench (1957: 144). The name S. elegantissima was again used by Maynard (1920a: 80) as an example of a taxon with color banding, citing a subsequently pub- lished illustration (Maynard, 1920b, fig. 60, identical image colored on pl. 13, fig. 2, both captioned “S. elegantissima M. & Cx Type Material. The description of Stro- phiops elegantissima did not specify the number of specimens examined but in- cluded figures of portions of a single spec- imen. The number of specimens examined was not specified in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 1920d: 126). Although these were likely available to Maynard in 1919 and are part of the type series [Article 72.4.1.1, ICZN, 1999: 76], the number of syntypes is unclear. The initially illustrated specimen could not be located at the MCZ. Maynard clearly regarded a differ- ent specimen to be the “Type” of Stro- phiops elegantissima. Lot MCZ 76375 contains a single specimen labeled “Ho- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. lotype” that approximates the measure- ments and illustration of this taxon in the intended description (Maynard and Clapp in Maynard, 1920d: 126, pl. 25, figs. 1, 2) and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen (MCZ 76375) is here designated as the lectotype of Strophiops elegantissi- ma Maynard, 1919, to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. The remain- ing specimens, including MCZ 76183, MCZ 118239, and USNM 420105, are paratypes. Lectotype Measurements. Length OP) mm, diameter (excluding lip) 11.1 mm; ap- erture height (including lip) 10.8 mm, ap- erture width (including lip and peristome) 8.7 mm. Type Locality. Norman’s Pond Key [Ex- uma Group, Bahamas]. Remarks. It is not surprising that after Maynard's death the specimens labeled as “types” by Maynard were the ones to be segregated at the MCZ while the many re- maining specimens were dispersed among MCZ, USNM, and other institutions (Turner, 1957: 151). Although Maynard’s illustrations (Maynard, 1919c, figs. 36A—D, 1920b, fig. 60, pl. ISS. fig. 2, 1920d: 126, pl. 25, figs. 1, 2) are of sufficient quality to conclude that the shells figured are differ- ent specimens, we have not been able to match the earliest figure with any of the specimens available to us. Taxon 124. Strophiops arbusta Maynard and Clapp in Maynard, 1919c [De- cember 31]: 65, fig. 38; Maynard, 1920a: 74, fig. 49; Maynard and Clapp in Maynard, 1921a: 133, pl. 30, figs. 5, 6. Examined 2,454 specimens. Size giv- en as 1.24 by .48 [inches; 30.5 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen Strophiops arbusta was first published by Maynard (1919c: 65, fig. 38) as an example 447 of a species that lacked an upper [colu- mellar] tooth. The figure was captioned “S. arbusta M. & C.” This illustration was lat- er reproduced in color (Maynard, 1920c, pl. 16, fig. 6). The association of a new species group name with an illustration of the taxon being named before 1931 makes the name Strophiops arbusta Maynard and Clapp in Maynard (1919c: 65, fig. 38) available (Article 12.2.7, ICZN, 1999: 17) prior to the intended description of this species (Maynard and Clapp in Maynard, 1921a: 133, pl. 30, figs. 5, 6). Maynard (1920a: 74, figs. 49A, B) again used the name in conjunction with figures before the intended introduction of the taxon. These illustrations were later reproduced in color (Maynard, 1920b, pl. 12, figs. 4, ey) Type Material. The description of Stro- phiops arbusta did not specify the number of specimens examined but included a fig- ure of a single specimen. The 2,454 spec- imens mentioned in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 1921a: 133) were likely available to Maynard at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops arbusta is based on 2,454 syntypes. The originally illustrated specimen (Maynard, 1919c, fig. 38) could not be identified at the MCZ. Maynard clearly regarded a different specimen to be the “Type” of Strophiops arbusta. Lot MCZ 76273 contains a single specimen la- beled “Holotype” that approximates the il- lustration more than the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “Holo- type” added later. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining specimens, including MCZ 76272, USNM 419968, and USNM 420027, are paralectotypes. Lectotype Measurements. Length 29.8 mim, diameter (excluding lip) 11.0 mm; ap- erture height (including lip) 10.4 mm, ap- 448 erture width (including lip and peristome) 8.6 mm. Type Locality. Guana Key, Berry Is- lands, Bahamas. Remarks. It is not surprising that, after Maynard's death, the specimens labeled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens were dispersed among MCZ, USNM, and other institu- tions (Turner, 1957: 151). Given the qual- ity of Maynard’s illustrations, it is unlikely that the initially figured specimen could ever be discerned unambiguously from among the large number of remaining specimens, even if it were present among them. Taxon 125. Strophiops aspera Maynard, 1919a [October 16]: 14 [nomen nu- dum]; Maynard, 1920a [February 29]: 77; Maynard and Clapp in May- nard, 1920b [April 8]: 89, fig. 57, pl. 12, fig. 1; Maynard and Clapp in Maynard, 1920 ANC sma sel oll eenicis OhelO: Examined 1,500 specimens. Size given as 1.06 by .30 [inches; 26.9 by 7.6 mm], but the term “type” was not used. Nomenclatural Remarks. Maynard (1919a: 14) first introduced the binomen Strophiops aspera as a nomen nudum. He again (Maynard, 1920a: 77) used this taxon name as an example of a species in which the animal gathered grains of carbonate sand and fastened them to outer and inner surfaces of the shell, citing a subsequently published illustration (Maynard, 1920b, fig. 57) captioned “S. aspera M & C.” Ar- ticle 10.1.1 (ICZN, 1999: 9) states that if publication of the data relating to a new nominal taxon or a nomenclatural act is in- terrupted and continued at a later date, the name or act becomes available only when the requirements of the relevant Ar- ticles have been met. Strophiops aspera became available on February 29, 1920, on the basis of the publication of the descrip- tion, which was adequate to meet the re- quirements of the relevant articles [Article Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 12.1, ICZN, 1999: 16], rather than from the later publication of the captioned fig- ure. Authorship is attributed to Maynard, the sole author of the description. Type Material. The description of S. as- pera did not specify the number of speci- mens examined, but included a reference to a subsequently published figure of a sin- gle specimen. The 1,500 specimens men- tioned in the intended description of this taxon (Maynard and Clapp in Maynard, 1920d: 116) were likely available to May- nard at the time he wrote the unintended species description and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops aspera is therefore based on 1,500 syntypes. The figured specimen (Maynard, 1920b, fig. 57) could not be recognized at the MCZ. Maynard clearly regarded a different specimen to be the “Type” of Strophiops aspera. Lot MCZ 76176 contains a single specimen labeled “Holotype” that, at best, approximates the measurements and illustrations of this tax- on in the intended description. This spec- imen is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures on plate 1. Clench and Aguayo (1952: 430, pl. 56, fig. 1) illustrated this specimen as the “Ho- lotype” of Cerion asperum. Article 74.5 (ICZN, 1999: 82) states, “When the orig- inal work reveals that the taxon had been based on more than one specimen, a sub- sequent use of the term ‘holotype’ does not constitute a valid lectotype designation unless the author, when wrongly using that term, explicitly indicated that he or she was selecting from the type series that par- ticular specimen to serve as the name- bearing type.” Thus, Clench and Aguayo’s treatment of MCZ 76176 can not be con- sidered a lectotype designation. This spec- imen (MCZ 76176) is here designated as the lectotype of Strophiops aspera May- nard, 1920, to provide an objective stan- dard of reference for this species-group taxon that is consistent with Maynard's original, and Clench and Aguayo’s subse- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. quent, concepts of it. The remaining spec- imens are paralectotypes. Lectotype Measurements. Length 24.8 mm, diameter (excluding lip) 9.6 mm; ap- erture height (including lip) 7.6 mm, ap- erture width (including lip and peristome) 7.0 mm. Type Locality. Among dwarf palms in an area about 300 yards long by some 150 wide along the third beach N of the S end of [Great Guana] key [Exuma Group, Ba- hamas |]. Remarks. It is not surprising that, after Maynard’s death, the specimens labeled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens were dispersed among MCZ, USNM, and other institu- tions (Turner, 1957: 151). Given the qual- ity of Maynard’s illustrations of this spe- cies, it is unlikely that the initially figured specimen, shown only in right lateral view, could ever be discerned unambiguously from among the large number of speci- mens. Maynard (1920d: 116) reported this taxon to be abundant, but mostly dead. Out of 1,500 specimens collected, only 70 were living. Maynard (1924a: 167) contra- dicted this number, stating that of 953 col- lected, 70 were living, and provided more detailed descriptions of the shell, ecology, and anatomy of this taxon. Clench and Aguayo (1952) considered Cerion asperum to be a valid species and the senior syno- nym of Strophiops scalariformis. Taxon 126. Strophiops fruticosa Maynard, 1920a [February 29]: 77 [nomen nu- dum]; Maynard and Clapp in May- nard, 1920b [April 8], fig. 55, pl. 12, fig. 3; Maynard and Clapp in Maynard, 1920d: 125, pl. 20, figs. 3, 4. Examined 1,040 specimens. Size given as 1.25 by 45 [inches; 31.8 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. Maynard (1920a: 77) first introduced the binomen Strophiops fruticosa as one of two exam- ples of a taxon that occasionally has de- 449 pressions in their surface sculpture but noted explicitly that this was not a specific character. This use was a nomen nudum, but included a reference to a subsequently published illustration (Maynard, 1920b, fig. 55, identical illustration colored on pl. 12, fig. 3, both captioned “S. fruticosa M. & C.”). The association of a new species group name with an illustration of the tax- on being named prior to 1931 makes the name Strophiops fruticosa Maynard and Clapp in Maynard, 1920b (fig. 55, pl. 12, HieanS) peavailable) (Articles 2:22. KEZNp 1999: 17) prior to the intended description of this species (Maynard and Clapp in Maynard, 1920d: 125, pl. 20, figs. 3, 4). Type Material. Although the unintended description of Strophiops fruticosa makes reference only to a single specimen shown in figure 55, it is likely that the 1,040 spec- imens mentioned in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 1920d: 125) were available to Maynard at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops fruticosa is based on 1,040 syntypes. The specimen illustrated in Maynard (1920b, fig. 55, pl. 12, fig. 3) could not be located at the MCZ. Maynard clearly re- garded a different specimen to be the “Type” of Strophiops fruticosa. Lot MCZ 76239 contains a single specimen labeled “Holotype” that approximates the mea- surements and illustration of this taxon in the intended description (Maynard and Clapp in Maynard, 1920d: 125, pl. 20, figs. 3, 4) and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen (MCZ 76239) is here des- ignated as the lectotype of Strophiops fru- ticosa Maynard, 1920, to provide an objec- tive standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. The remaining specimens, including MCZ 76240, MCZ 118290, and USNM 420099, are paralectotypes. Lectotype Measurements. Length 29.3 450 mm, diameter (excluding lip) 11.5 mm; ap- erture height (including lip) 10.5 mm, ap- erture width (including lip and peristome) 8.8 mm. Type Locality. Group, Bahamas]. Remarks. It is not surprising that, fol- lowing Maynard’s death, the specimens la- beled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens were dis- persed among MCZ, USNM, and other in- stitutions (Turner, 1957: 151). Given the quality of Maynard's illustration of a dorsal view of the single shell, it is unlikely that this specimen could ever be discerned un- ambiguously from among the large num- ber of remaining specimens, even if it were present among them. Maynard (1920d: 125) reported this tax- on to be common on the ground or about the bases of bushes. Bush Key [Exuma Taxon 127. Strophiops caduca Maynard, 1919b [November 6]: 35 [nomen nu- dum]; Maynard, 1920a [February 29]: 77 [nomen nudum]; Maynard and Clapp in Maynard, 1920b [April 8], fig. 56, 82; Maynard, 1920c, figs. 61A, 64, pl. 14, fig. 4 [see Nomenclatural Re- marks], pl. 15, fig. 4; Maynard and Clapp in Maynard, 1921a: 133, pl. 30, figs. 9, 10. Examined 1,278 specimens. Size given as 1.00 by .45 [inches; 25.4 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen Strophiops caduca first appeared as a no- men nudum (Maynard, 1919b: 35). May- nard (1920a: 77) again used this binomen as one of two examples of a taxon that oc- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 casionally had depressions in its surface sculpture, but noted explicitly that this was not a specific character. This use, also a nomen nudum, included a reference to a subsequently published illustration (May- nard, 1920b, fig. 56, captioned “S. caduca M. & C.”). The association of a new spe- cies group name with an illustration of the taxon being named before 1931 makes the name Strophiops caduca Maynard and Clapp in Maynard, 1920b (fig. 56), avail- able (Article 12.2.7, ICZN, 1999: 17) prior to the intended description of this species (Maynard and Clapp in Maynard, 1921a: 133, pl. 30, figs. 9, 10). Maynard (1920b: 82) again used the binomen citing figures (Maynard, 1920c, fig. 61A) of a different specimen, captioned “S. caduca.” The bi- nomen “S. caduca M. & C.” appears in the caption to figure 64 (Maynard, 1920c), which is identical to figure 20 (Maynard, 1919c), captioned “S. lobata M. & C.” A colored version of figures 20 and 64 (May- nard, 1920c, pl. 14, fig. 4) is captioned “S. caduca.” Maynard (1920d: 112) published corrections, stating that the captions to fig- ures 64 and pl. 14, fig. 4, should read lob- ata not caduca. Type Material. Although the unintended description of Strophiops caduca makes reference only to a single specimen shown in figure 56, it is likely that the 1,278 spec- imens mentioned in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 1921a: 133) were available to Maynard at the time, and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops caduca is based on 1,278 syntypes. The specimen illustrated in Maynard, 1920b (fig. 56), could not be identified at —_ Figures 126-137. Taxon 126. Strophiops fruticosa Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76239. Bush Key, Exuma Group, Bahamas. Taxon 127. Strophiops caduca Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76366. West side of Cabbage Key, Berry Islands, Bahamas. Taxon 128. Strophiops persuasa Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76189. Along shore north of Fresh Creek, Andros, Bahamas. Taxon 129. Strophiops santesoni Maynard, 1920. Lectotype MCZ 358073. Along northern shore of New Providence Island, west of Nassau from second mile post to beyond the fourth. Taxon 130. Strophiops inconstans Maynard, 1920. Lectotype MCZ 76341. West side of Bird Key, Exuma Group, Bahamas. Taxon 131. Strophiops plebeia Maynard, 1920. Lectotype USNM 420125. Lignumvitae Key, Berry Islands, Bahamas. Taxon 132. 451 THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. Strophiops cana Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76374. Fortune Key, Berry Islands, Bahamas. Taxon 133. Strophiops territa Maynard, 1920. Lectotype MCZ 76396. Methodist churchyard and vicinity, Sherley Street, Nassau, New Providence Island, Bahamas. Taxon 134. Strophiops albolabra Maynard, 1920. Lectotype MCZ 76255. Great Harbor Key, Berry Islands, Bahamas. Taxon 135. Strophiops fragilis Maynard, 1920. Lectotype MCZ 76166. Second westernmost Brigadier Key, Great Exuma, Exuma Group, Bahamas. Taxon 136. Strophiops latonia Maynard, 1920. Lectotype MCZ 76314. St. Paul’s Quarry off West Sherley St., Nassau, New Providence Island, Bahamas. Taxon 137. Strophiops mariae Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76204. Maria Key near Little Exuma, Exuma Group, Bahamas. 452 the MCZ. Maynard clearly regarded a dif- ferent specimen to be the “Type” of Stro- phiops caduca. Lot MCZ 76366 contains a single specimen labeled “Holotype” that approximates the illustration more than the measurements of this taxon in the in- tended taxonomic description (Maynard and Clapp in Maynard, 1921a: 133, pl. 30, figs. 9, 10) and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures on plate 30. This specimen is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. Maynard’s re- maining specimens, including MCZ 76365 and USNM 420005, are paralectotypes. Lectotype Measurements. Length 27.4 mm, diameter (excluding lip) 11.8 mm; ap- erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) Same Type Locality. West side of Cabbage Key [Berry Islands], Bahamas. Remarks. It is not surprising that, after Maynard’s death, the specimens labeled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens were dispersed among MCZ, USNM, and other institu- tions (Turner, 1957: 151). Given the qual- ity of Maynard’s illustrations of the dorsal view of the shell, it is unlikely that this specimen could ever be discerned unam- biguously from among the large number of remaining specimens, even if it were present among them. Taxon 128. Strophiops persuasa Maynard and Clapp in Maynard, 1920b [April 8], fig. 59*, pl. 13, fig. 1; Maynard, 1920d: 112; Maynard and Clapp in Maynard, 1921a: 131, pl. 28, figs. 9, 10. Examined 2,428 specimens. Size given as .98 by .40 [inches; 24.9 by 10.2 mm], but the term “type” was not used. Nomenclatural Remarks. This taxon be- came available when an illustration (May- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 nard, 1920b, fig. 59*, pl. 13, fig. 1) cap- tioned “S. persuasa M. & C.” was pub- lished. The association of a new species group name with an illustration of the tax- on being named before 1931 makes the name Strophiops persuasa Maynard and Clapp in Maynard, 1920, available (Article 12.2.7, ICZN, 1999: 17) prior to the in= tended description of this species (May- nard and Clapp in Maynard, 1921a: 131, pl. 28, figs. 7, 8). The name next appeared (Maynard, 1920d: 112) in a text correction [see nomenclatural remarks under S. san- tesoni, Taxon 129]. Type Material. Although the unintended description of Strophiops persuasa makes reference only to a single specimen illus- trated in figure 59*, it is likely that the 2,428 specimens mentioned in the intend- ed description of this taxon (Maynard and Clapp in Maynard, 1921a: 131) were avail- able to Maynard at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops persuasa is based on 2,428 syntypes. The specimen figured both in figure 59* and on plate 13, figure 1, could not be located at the MCZ. Maynard clearly regarded a different spec- imen to be the “Type” of Strophiops per- suasa. Lot MCZ 76189 was labeled “Ho- lotype” but contained two specimens. The larger of these specimens approximates the subsequently published measurements and figures and is here selected as the lec- totype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s concept of this taxon. The remaining specimen was recatalogued as MCZ 357000 and, togeth- er with MCZ 76188 and USNM 420124, are among the 2,427 specimens that be- come paralectotypes. Lectotype Measurements. Length 25.8 mm, diameter (excluding lip) 10.8 mm; ap- erture height (including lip) 8.9 mm, ap- erture width (including lip and peristome) 7.9 mm. Type Locality. Along shore north of Fresh Creek, Andros, Bahamas. Remarks. It is not surprising that, after THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. Maynard's death, the specimens labeled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens were dispersed among MCZ, USNM, and other institu- tions (Turner, 1957: 151). Given the qual- ity of Maynard’s illustration of an apertural view, it is unlikely that this specimen could ever be discerned unambiguously from among the large number of specimens, even if it were present among them. Taxon 129. Strophiops santesoni Maynard, 1920a [February 29]: 80, 81 [nomen nu- dum]; Maynard, 1920b [April 8], fig. 59; Maynard, 1920c [July 10], fig. 62, pl. 15, fig. 5 [both of sinistral speci- men]; Maynard and Clapp in Maynard, 1921b: 139, pl. 36, figs. 3, 4, pl. 15, fig. 5. Examined 1,964 specimens. Size giv- en as 1.20 by .47 [inches; 30.5 by 11.9 mm], but the term “type” was not used. Nomenclatural Remarks. Maynard (1920a: 80) first used the binomen Stro- phiops santesoni in error as one of three taxa that, in rare cases, can have a shell encircled by a band of color. Because this character is not diagnostic of S. santesoni, this passage cannot be considered a de- scription of the taxon and the use is a no- men nudum. This use also includes a clearly incorrect reference to a subse- quently published figure (Maynard, 1920b, fig. 59) that was captioned “S. nuda Mayn.” Earlier on the same page is anoth- er reference to figure 59 that correctly re- fers to it as S. nuda. Maynard (1920d: 112) published a correction to the first use on page 80, stating “for santesoni read per- suasa.” This statement makes it clear that the intended figure reference was figure 59* (Maynard, 1920b), which is captioned “S. persuasa M. & C. X2, banded below suture.” The next use of the name S. san- tesoni appeared in a discussion of sinis- trality in Cerions (Maynard, 1920a: 81). Again, the text notes that this rare char- acter is present in both S. santesoni and S. inconstans, and the use is a nomen nu- 453 dum. However, it does include a reference to a subsequently published illustration (Maynard, 1920c, fig. 62) that was cap- tioned “S. santesoni.” Because the publi- cation of the association of a new species group name (Maynard, 1920a: 81) with an illustration (Maynard, 1920b, fig. 62) of the taxon being named before 1931 was interrupted, the name Strophiops sante- soni became available only when the re- quirements of the relevant Articles (Arti- cles 10.1.1 and 12.2.7, ICZN, 1999: 9, 17) have been met. The name Strophiops san- tesoni Maynard, 1920, became available July 10, 1920, on publication of figure 62, prior to the intended description of this species (Maynard and Clapp in Maynard, 1921b: 139, pl. 36, figs. 3, 4, pl. 15, fig. 5). Because of Maynard’s unintended taxo- nomic act, both the date of publication and the authorship of Strophiops santesoni were altered. Type Material. The unintended descrip- tion linked the binomen Strophiops san- tesoni with a figure of a single sinistral specimen illustrated in Maynard, 1920c (fig. 62, pl. 15, fig. 5). Maynard (1920b: 81) noted that sinistrality is very rare in Cer- ion, and only two have been found, “one in S. santesoni.” This clearly indicates that he had multiple specimens of C. santesoni, of which one was sinistral, and suggests that the 1,964 specimens mentioned in the intended description of this taxon (May- nard and Clapp in Maynard, 1921a: 139) were available to him at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops san- tesoni is thus based on 1,963 syntypes. Lot MCZ 76237 contained 216 specimens la- beled “paratypes,” among them the illus- trated sinistral specimen, which is here designated as the lectotype and has been recatalogued as MCZ 358073. Maynard clearly regarded a different specimen to be the “Type” of Strophiops santesoni. Lot MCZ 76238 contains a single specimen la- beled “Holotype” that resembles the illus- tration in the intended description, but differs substantially from the measure- 454 ments. This specimen is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures on plate 36. Maynard’s remaining 1,962 specimens of S. santesoni, includin MCZ 76237, MCZ 76238, USNM 420028, USNM 419959, and USNM 419964, are paralectotypes. Lectotype Measurements. Length 23.5 mm, diameter (excluding lip) 11.2 mm; ap- erture height (including lip) 9.7 mm, ap- erture width (including lip and peristome) 7.9 mm. Type Locality. Along northern shore of New Providence [Island] west of Nassau from second mile post to beyond the fourth. Remarks. It is not surprising that, after Maynard’s death, the specimens labeled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens were dispersed among MCZ, USNM, and other institu- tions (Turner, 1957: 151). Fortunately, the sinistral specimen originally illustrated by Maynard was segregated in a vial and la- beled as the specimen appearing in figure 62. Surprisingly, the large lot collected by Maynard actually contained two sinistral specimens, both illustrated by Gould et al. (1985: 1368, fig. 1) as Cerion glans (Kiis- ter, 1844), New Providence sample. Gould and Woodruff (1986: 475) considered S. santesoni to be a synonym of C. glans. Taxon 130. Strophiops inconstans May- nard, 1920b [April 8]: 81 [nomen nu- dum]; Maynard, 1920c [July 10], fig. 63, pl. 15, fig. 6; Maynard and Clapp in Maynard, 1920d: 119, pl. 3, figs. 7, 8. Examined 584 specimens. Size given as 1.10 by .40 [inches; 27.9 by 10.2 mm], but the term “type” was not used. Also 1920c [July 10], pl. 15, fig. 6. Nomenclatural Remarks. The first use of the binomen Strophiops inconstans ap- peared in Maynard (1920b: 81) in a dis- cussion of sinistrality in Cerionidae and referenced an illustration of a_ sinistral Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 specimen published in the following issue (Maynard, 1920c, fig. 63). The identical image appeared in color in the later issue (Maynard, 1920c, pl. 15, fig. 6). Both il- lustrations were captioned “S. inconstans.” The use of S. inconstans in Maynard (1920b: 81) is anomen nudum. For names published before 1931, the association of a new species group name with an illus- tration is sufficient to make the species group name available [Article 12.2.2.7, ICZN, 1999: 17]. Because of Maynard’s unintended taxonomic act, the binomen Strophiops inconstans became available on July 10, 1920, with Maynard as the author. The intended description of this taxon was published in the subsequent issue (May- nard and Clapp in Maynard, 1920d: 119, pl. 3, figs. 7, 8) and was based on a dextral specimen. Type Material. Although inadvertently, this taxon is based on a sinistral specimen illustrated in Maynard, 1920c (fig. 63, pl. 15, fig. 6). It is likely that the 584 speci- mens mentioned in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 192la: 131) were available to Maynard at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops inconstans is based on 584 syntypes. Lot MCZ 76341 con- tained multiple specimens labeled “para- types,” among them the illustrated sinistral specimen, which is here designated as the lectotype of Strophiops inconstans and has been recatalogued as MCZ 356677. May- nard clearly regarded a different specimen to be the “Type” of Strophiops inconstans. Lot MCZ 76340 contained a single speci- men labeled “Holotype” that approximates the measurements and illustrations of this taxon in the intended description. This specimen is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures in the intended description. It becomes a paralectotype, as do the specimens in MCZ 76340, MCZ 76341, and USNM 4920121. Lectotype Measurements. Length 26.4 THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. mm, diameter (excluding lip) 11.2 mm; ap- erture height (including lip) 7.6 mm, ap- erture width (including lip and peristome) 7.9 mm. Type Locality. West side of Bird Key [Exuma Group, Bahamas]. Remarks. It is not surprising that, after Maynard’s death, the specimens labeled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens were dispersed among MCZ, USNM, and other institu- tions (Turner, 1957: 151). Fortunately, the sinistral specimen was retained among the material at the MCZ and was easily rec- ognized. Maynard (1920d: 119) reported Strophiops inconstans to be common on cacti. Maynard (1925: 185-194) provided additional information on the anatomy, habitat, and morphology of this taxon. Taxon 131. Strophiops plebeia Maynard, 1920b [April 8]: 85 [nomen nudum]. Strophiops plebea Maynard 1920c [July 10], fig. 66, pl. 14, fig. 1. Stro- phiops plebia Maynard and Clapp in Maynard, 1921a [May 2]: 130, pl. 28, figs. 3, 4. Examined 1,500 specimens. Size given as 1.20 by .40 [inches; 30.5 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. This binomen was first used by Maynard (1920b: 85) [as plebeia| as an example of a fairly common- ly decollated species [80 of 1,500 speci- mens| and included a reference to an il- lustration of an apertural view of a shell published in the following issue (Maynard 1920c, fig. 66, pl. 14, fig. 1). Both the text illustration and the identical colored figure were captioned “S. plebea.” The April 8, 1920, use as plebeia is a nomen nudum. The association of a new species group name with an illustration of the taxon be- ing named before 1931 makes the name Strophiops plebea Maynard (1920c, fig. 66, pl. 14, fig. 1) available (Article 12.2.7, ICZN, 1999: 17) prior to the intended de- scription of this species (Maynard and 455 Clapp in Maynard, 1921a: 131, pl. 28, figs. 7, &). Type Material. Although the unintended description of Strophiops plebea makes reference only to a single decollated spec- imen illustrated in figure 66, it is likely that the 1,500 specimens mentioned in the in- tended description of this taxon (Maynard and Clapp in Maynard, 1921a: 130) were available to Maynard at the time and are thus spar Ormbhe typemsemess || Article 72.4.1.1, ICZN, 1999: 76]. Strophiops ple- bea is based on 1,500 syntypes. In a search for the specimen illustrated in figure 66, examination of large series of this taxon from the Maynard collection now at the MCZ and USNM revealed over 25 decollated specimens. About a third of these specimens could be excluded from consideration on the basis of the degree of decollation or the presence of other dam- age undocumented in the illustration. The majority of decollated specimens, however, all resembled the illustration to a similar degree, and we are not able to match any one of them to the illustration with any degree of certainty. Maynard clearly regarded a different specimen to serve as the “Type” of Stro- phiops plebea. Lot MCZ 76401 was la- beled “Holotype,” but contained two spec- imens, neither a close match to the figures or measurements in the intended taxon description. Nor were any labels in May- nard’s handwriting present. Lot USNM 420125 contained numerous specimens from among the 1,500 syntypes. A speci- men that closely approximates the illustra- tions and measurements published togeth- er with the intended taxon description was selected as a lectotype (USNM 420125) to provide an objective standard of reference for this species-group taxon that is consis- tent with Maynard’s original concept of it. The remaining specimens were recata- logued as USNM 1093795, and together with MCZ 76401 and MCZ 76169, are paralectotypes. Lectotype Measurements. Length 30.9 mim, diameter (excluding lip) 11.3 mm; ap- 456 erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) Sei aman Type Locality. Lignumvitae Key, Berry Islands, Bahamas. Remarks. This taxon has been the victim of multiple lapsus calami, having been spelled plebeia when first used as a nomen nudum, plebea in the unintended, but val- id description, and plebia in the intended taxonomic description, as well as in a sub- sequent listing in the sales catalogue (May- nard, 1924?P: 3). Article 32.5.1 (ICZN, 1999: 39) dictates that the spelling plebea be used for this taxon because there is no clear evidence to suggest an alternative spelling within the original publication it- self. Strophiops plebea Maynard, 1920, has a different spelling, authorship, and date of publication from that cited by Clench GIS Si 5s8)r Taxon 132. Strophiops cana Maynard, 1920b [April 8]: 86, 88 [nomen nudum]; Maynard and Clapp in Maynard, 1920c [July 10], figs. 69, 76, pl. 15, fig. 1, pl. 17, fig. 5; Maynard and Clapp in Maynard, 1921a [May 2]: 137; 1921b [July 15], pl. 34, figs. 3, 4. Ex- amined 760 specimens. Size given as 1.10 by .45 [inches; 27.9 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen Strophiops cana was first used by Maynard (1920b: 86, 88) in a discussion of shell re- pair after severe damage and referred to subsequently published illustrations (May- nard, 1920c, figs. 69, 76, pl. 15, fig. 1, pl. 17, fig. 5) of damaged specimens of S. cana as examples of snails surviving crush- ing predation, noting that color change oc- casionally accompanied shell repair. The appearances of S. cana in the text (May- nard, 1920b: 86, 88) are as a nomen nu- dum. The association of a new species group name with an illustration of the tax- on being named before 1931 made the name Strophiops cana Maynard and Clapp in Maynard, 1920, available (Article 12.2.7, Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 ICZN, 1999: 17) on July 10, with the pub- lication of figures 69 and 76, the first cap- tioned “S. cana M & C” and the second “S. cana,” prior to the intended descrip- tion of this species (Maynard and Clapp in Maynard, 1921a: 137). Type Material. In addition to the two damaged specimens of Strophiops cana that were illustrated in figures 69 and 76 (as well as the colored versions of these figures, pl. ls) aie, phi figs) a its likely that the 760 specimens mentioned in the intended description of this taxon (Maynard and Clapp in Maynard, 1921a: 137) were available to Maynard at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops cana is based on 760 syntypes. Neither of the specimens illustrated in figures 69 and 76 could be located at the MCZ. Maynard clearly regarded a different specimen to be the “Type” of Strophiops cana. Lot MCZ 76374 contains a single specimen labeled “Holotype” that approximates the mea- surements of this taxon in the intended de- scription (Maynard and Clapp in Maynard, 1921a: 137) and the subsequently pub- lished illustrations (Maynard, 1921b, pl. 34, figs. 3, 4) and is accompanied by a la- bel in Maynard’s hand with the term “type” that includes references to the fig- ures. This specimen (MCZ 76374) is here designated as the lectotype of Strophiops cana Maynard and Clapp in Maynard, 1920, to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining specimens, in- cluding MCZ 76373, MCZ 118217, and USNM 420091, are paralectotypes. Lectotype Measurements. Length 28.3 mm, diameter (excluding lip) 11.5 mm; ap- erture height (including lip) 10.1 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. Fortune Key, Berry Is- lands, Bahamas. Remarks. It is not surprising that, after Maynard's death, the specimens labeled as “types” by Maynard were the ones to be THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. segregated at the MCZ, while the many remaining specimens, possibly including those in figures 69 and 76, were dispersed among MCZ, USNM, and other institu- tions (Turner, 1957: 151). Taxon 133. Strophiops territa Maynard, 1920c [July 10], fig. 70*, pl. 15, fig. 2; Maynard, 1920d: 112; Maynard and Clapp in Maynard, 1921b: 147; May- nard, 1926, pl. 43, figs. 5, 6. Examined 393 specimens. Size given as .94 by .40 [inches; 23.9 by 10.2 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen Strophiops territa was first used by May- mard (1920c, fig. 70*, pl. 15, fig. 2) in the caption to a figure of an apertural view of a crushed and repaired shell. Figure 70* and the identical colored image on plate 15, figure 2, are both captioned “S. territa M.”; the authorship of the taxon is restrict- ed to Maynard. The association of a new species group name with an illustration of the taxon being named before 1931 makes the name Strophiops territa Maynard, 1920c (fig. 70*, pl. 15, fig. 2), available (Ar- Hele 12.2.7, ICZN, 1999: 17) on July 10, 1920, prior to the intended description of this species (Maynard and Clapp in May- nard, 1921b: 147). Type Material. Although the unintended description of Strophiops territa makes reference only to a single damaged speci- men shown in figure 70*, it is likely that the 393 specimens mentioned in the in- tended description of this taxon (Maynard and Clapp in Maynard, 1921b: 147) were available to Maynard at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops ter- rita is based on 393 syntypes. The speci- men illustrated in figure 70* could not be located at the MCZ. Maynard clearly re- garded a different specimen to be the “Type” of Strophiops territa. Lot MCZ 76396 contains a single specimen labeled “Holotype” that approximates the mea- surements of this taxon in the intended de- 457 scription and the subsequently published illustrations. It is accompanied by a label in Maynard’s hand with the term “type,” that includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 392 speci- mens, including MCZ 76316 and USNM 419995, are paralectotypes. Lectotype Measurements. Length 23.4 mim, diameter (excluding lip) 11.3 mm; ap- erture height (including lip) 9.3 mm, ap- erture width (including lip and peristome) 8.1 mm. Type Locality. Methodist Churchyard and vicinity |Sherley Street, Nassau, New Providence Island, Bahamas]. Remarks. It is not surprising that, after Maynard’s death, the specimens labeled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens, possibly including the damaged specimen in figure 70*, were dispersed among MCZ, USNM, and other institutions (Turner, 1957: 151). Gould and Woodruff (1986: 476) regarded this taxon to be an “anomalously located species at- tributable to Cerion glans.” Taxon 134. Strophiops albolabra Maynard, 1919b [November 6]: 25 [nomen nu- dum]; Maynard, 1920b [April 8]: 87, 88 [nomina nuda]; Maynard, 1920c [July 10], figs. 74a, b, 75, 77, pl. 16, figs. 1-3, pl. 17, fig. 4; Maynard and Clapp in Maynard, 192la [May 2]: 129-130, pl. 27, figs. 3, 4. Examined 4,560 spec- imens. Size given as 1.30 by .50 [inches; 33.0 by 12.7 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen Strophiops albolabra first appeared as a nomen nudum (Maynard, 1919b: 25). Maynard (1920b: 87, 88) discussed shell repair in this species prior to its intended description and included citations to illus- trations of four broken specimens that 458 were published in the following issue (Maynard, 1920c [July 10], figs. 74a, b, 75, 77). These same figures were reproduced in color in the same issue (Maynard, 1920c, pl. 16, figs. 1-3, pl. 17, fig. 4). The appearances of the taxon name in the text are as a nomen nudum. The association of a new species group name with an illustration of the taxon be- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 1921a, pl. 27, figs. 3, 4). This specimen is here designated as the lectotype of Stro- phiops albolabra to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. The remain- ing 4,559 specimens are paralectotypes. Lectotype Measurements. Length DO mim, diameter (excluding lip) 12.4 mm; ap- ing named before 1931 makes the name Strophiops albolabra Maynard, 1920c (figs. 74a, b, 75, 77, pl. 16, figs. 1-3, pl. 17, fig. 4), available (Article 12.2.7, ICZN, 1999: 17) on July 10, 1920, prior to the intended description of this species (May- nard and Clapp in Maynard, 1921a: 129- 130). Because the illustrations were cap- tioned “S. albolabra,” the taxon must be attributed to Maynard rather than May- nard and Clapp as cited by Clench (1957: 137). Type Material. Although the unintended description of Strophiops albolabra illus- trated four broken specimens, it is likely that these were selected from among the 4,560 specimens mentioned in the intend- ed description of this taxon (Maynard and Clapp in Maynard, 1921a: 130) that were available to Maynard and are part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops albolabra is based on 4,560 syntypes. Over a thousand speci- mens contained in MCZ 76241, MCZ 118211, USNM 420023, and USNM 420098 were examined. Only two exhibit- ed significant shell damage but did not correspond to any of the four figured syn- types. These four syntypes are lost because it is unlikely that badly damaged speci- mens would have been selected for sale or exchange by Maynard. Maynard clearly re- garded a different specimen to be the “Type” of Strophiops albolabra. Lot MCZ 76255 contains a single specimen labeled “Holotype” that approximates the illustra- tion more than the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures in the intended taxon description (Maynard, erture height (including lip) 11.0 mm, ap- erture width (including lip and peristome) 9.1 mm. Type Locality. Great Harbor Key Weng Islands, Bahamas]. Romane It is not surprising that, after Maynard's death, the specimens labeled as “types” by Maynard were the ones to be segregated at the MCZ, while the many remaining specimens were dispersed among MCZ, USNM, and other institu- tions (Turner, 1957: 151). The four severe- ly damaged syntypes would be easily rec- ognizable, yet could not be found in the collections of the MCZ and might have been discarded, especially since in excess of 4,000 specimens were available. Taxon 135. Strophiops fragilis Maynard, 1920b [April 8]: 91, 92 [nomen nudum]; Maynard, 1920c [July 10], figs. 81, 82, pl. 18, figs. 4, 5; Maynard and Clapp in Maynard, 1921a [May 2]: 128, pl. 26, figs. 3, 4. Examined 200 speci- mens. Size given as 1.00 by .40 |inches; 25.4 by 10.2 mm], but the term “type” was not used. Nomenclatural Remarks. Maynard (1920b: 91-92) used the binomen Stro- phiops fragilis prior to its intended de- scription as an example of his “Law of In- dividual Reincarnation” and referenced subsequently published illustrations (May- nard, 1920c, figs. 81, 82) representing ap- ertural views of two different specimens that were reproduced in color (Maynard, 1920c, pl. 18, figs. 4, 5). Article 10.1.1 (ICZN, 1999: 9) states that if publication of the data relating to a new nominal taxon or a nomenclatural act is interrupted and THE CERION TAXA OF CHARLES JOHNSON MAYNARD °* Harasewych et al. continued at a later date, the name or act becomes available only when the require- ments of the relevant Articles have been met. Thus, the first uses of the name (Maynard, 1920b: 91-92) were nomina nuda. The association of a new species group name with an illustration of the tax- on being named before 1931 makes the name Strophiops fragilis Maynard, 1920c (figs. 81, 82, pl. 18, figs. 4, 5), available (Article 12.2.7, ICZN, 1999: 17) on July 10, 1920, prior to the intended description of this species (Maynard and Clapp in Maynard, 1921a: 128). As the illustrations were captioned “S. fragilis,” the taxon must be attributed to Maynard rather than Maynard and Clapp as cited by Clench (1957: 146). Type Material. Two different specimens of S. fragilis were illustrated (Maynard, IO20c) figs) Ole 32) pl lis) figs. 455) One (fig. 81, pl. 18, fig. 4) was reported to closely resemble S. agrestina (figs. 80, pl. 18, fig. 3) the other (fig. 82, pl. 18, fig. 5) was described as indistinguishable from S. nuda (fig. 83, pl. 18, fig. 6). These two specimens were selected from the 200 specimens mentioned in the intended de- scription of this taxon (Maynard and Clapp in Maynard, 1921a: 128) that were avail- able to Maynard and are part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops territa is based on 200 syn- types. Lot MCZ 76166 contains a single specimen labeled “Holotype” that is ac- companied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures in the intended de- scription (pl. 26, figs. 3, 4). Although this specimen closely matches the measure- ments provided in the subsequent descrip- tion, it more closely resembles one of the two illustrations (Maynard, 1920c, fig. 81, pl. 18, fig. 4) on which the taxon is based than it does the subsequent illustration in the intended description. In our view, it is probable that this specimen served as the basis for both illustrations. This specimen is here designated as the lectotype to pro- vide an objective standard of reference for 459 this species-group taxon that is consistent with Maynard’s original concept of it. The remaining syntypes become paralecto- types. Lectotype Measurements. Length 25.5 mm, diameter (excluding lip) 10.2 mm; ap- erture height (including lip) 9.4 mm, ap- erture width (including lip and peristome) 8.1 mm. Type Locality. Second westernmost Brigadier Key, Great Exuma |[Exuma Group, Bahamas]. Remarks. Maynard (1920c: 92) com- mented that five specimens of S. fragilis were indistinguishable from S. nuda [from Clarence Harbor, Long Island, Bahamas]. Taxon 136. Strophiops latonia Maynard, 1920c [July 10]: 95, fig. 84, pl. 18, fig. 1; Maynard and Clapp in Maynard, 1921b [July 15]: 147-148; Maynard and Clapp in Maynard, 1926 [March 24], pl. 43, figs. 7, 8. Examined 600 specimens. Size given as 1.10 by .42 [inches; 27.9 by 10.7 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen Strophiops latonia was used prior to its in- tended description (Maynard, 1920c: 95, fig. 84) as an example of the “Law of Spe- cific Reincarnation.” The figure was repro- duced in color (Maynard, 1920c, pl. 18, fig. 1). The association of a new species group name with an illustration of the tax- on being named before 1931 makes the name Strophiops latonia available (Article 12.2.7, ICZN, 1999: 17) at the time of publication of the illustrations (July 10, 1920) rather than on July 15, 1921, when the intended original description was pub- lished. As these illustrations were cap- tioned “S. latonia,” the taxon is attributed to Maynard, rather than Maynard and Clapp, as cited by Clench (1957: 150). Type Material. Although the unintended description of Strophiops latonia is based on the illustration of a single specimen (fig. 84), it is likely that the 600 specimens mentioned in the intended description of 460 this taxon (Maynard and Clapp in May- nard, 1921b: 148) were available to May- nard at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops latonia is therefore based on 600 syntypes. The specimen shown in figure 84 could not be located at the MCZ. Maynard clearly regarded a different spec- imen to be the “Type” of Strophiops la- tonia. Lot MCZ 76314 contains a single specimen labeled “Holotype” that approx- imates the measurements of this taxon in the intended description (Maynard, 1921b: 147-148) and the subsequently published illustrations (Maynard, 1926, pl. 43, figs. 7, 8) and is accompanied by a la- bel in Maynard’s hand with the term “type,” which includes references to the figures. This specimen (MCZ 76314) is here designated as the lectotype of Stro- phiops latonia Maynard, 1920, to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining specimens, including MCZ 76315 and USNM 420146, are paralectotypes. Lectotype Measurements. Length 28.1 mm, diameter (excluding lip) 10.8 mm; ap- erture height (including lip) 9.6 mm, ap- erture width (including lip and peristome) 8.0 mm. Type Locality. St. Paul’s Quarry off West Sherley Street, Nassau [New Providence Island, Bahamas]. Remarks. Maynard (1924c: 6) listed S. latonia as extinct in 1924 from the burning of vegetation prior to cultivation. Gould and Woodruff (1986: 476) regarded this taxon to be “intermediate” between Cer- ion glans (Kiister, 1844) and C. guberna- torium (Crosse, 1869). Taxon 137. Strophiops mariae Maynard and Clapp in Maynard, 1920c [July 10]: 96, fig. 88; Maynard, 1920d [Sep- tember 30], pl. 24, fig. 10 [apertural view of costate specimen], pl. 25, fig. 5 [left lateral view of smooth specimen]; Maynard and Clapp in Maynard, 1921a [May 2]: 128. Examined 216 specimens Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 [96 smooth and 120 costate]. Size given as 1.10 by .40 [inches; 27.9 by 10.2 mm], but the term “type” was not used. Nomenclatural Remarks. The taxon name Strophiops mariae was first used and figured by Maynard (1920c: 96, fig. 88) as an example of his “Law of Specific Rein- carnation” earlier than the intended de- scription of the species. The association of a new species group name with an illus- tration of the taxon being named before 1931 makes the name Strophiops mariae available (Article 12.2.7, ICZN, 1999: 17) prior to the intended description of this species (Maynard and Clapp in Maynard, 1921la: 128). The illustrations referenced in the intended description were pub- lished in the previous issue (Maynard, 1920d, pl. 24, fig. 10, pl. 25, fig. 5) and depict two different specimens. Because figure 88 is captioned “S. mariae M & C.,” the taxon is attributed to Maynard and Clapp in Maynard and dates from July 10, 1920. Type Material. Although the unintended description of Strophiops mariae is based on the illustration of a single specimen (fig. 88), it is likely that the 216 specimens mentioned in the intended description of this taxon (Maynard and Clapp in May- nard, 1921a: 128) were available to May- nard at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops mariae is therefore based on 216 syntypes. The specimen shown in figure 84 could not be located at the MCZ. Lot MCZ 76204 contains 120 specimens and was labeled “Paratypes.” Of these 120 specimens, two were sequestered in glass vials and labeled as figured specimens. One of the labels in Maynard’s handwrit- ing associated with the smooth specimen illustrated on plate 25, figure 5, states “Fig. 88.” This specimen, which retains the number MCZ 76204, is here designated as the lectotype of Strophiops mariae May- nard and Clapp in Maynard, 1920, to pro- vide an objective standard of reference for this species-group taxon that is consistent THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. with Maynard’s original concept of it. The remaining 119 specimens were recata- logued as MCZ 357002 and, together with MCZ 118193, are among the 215 speci- mens that become paralectotypes. Lectotype Measurements. Length 27.8 mm, diameter (excluding lip) 10.1 mm; ap- erture height (including lip) 9.7 mm, ap- erture width (including lip and peristome) 8.0 mm. Type Locality. Maria Key near Little Ex- uma [Exuma Group, Bahamas]. Remarks. Maynard (1920c: 96) noted that the species S. mariae closely resem- bles S. repetita from Nassau, New Provi- dence Island. Taxon 138. Strophiops repitita Maynard, 1919b: 36 [nomen nudum]; S. repetita Maynard and Clapp in Maynard, 1920c [July 10]: 96, fig. 89; Maynard and Capp in Maynard, 1921b [July 15]: 149, pl. 45, figs. 5, 6 [pl. 45 was never issued]. Examined 1,052 specimens. Size given as 1.10 by .40 [inches; 27.9 by 10.2 mm], but the term “type” was not used. Nomenclatural Remarks. The binomen re repitita first appeared as a nomen nu- dum (Maynard, 1919b: 36). The name was used as S. repetita (Maynard, 1920c: 96, fig. 89) prior to the intended taxon de- scription as an example of Maynard’s “Law of Specific Reincarnation.” The association of a new species group name with an il- lustration of the taxon being named before 1931 makes the name Strophiops repetita Maynard and Clapp in Maynard, 1920, available (Article 12.2.7, ICZN, 1999: 17) prior to the intended description of this species (Maynard and Clapp in Maynard, 1921b: 149). The caption to figure 89 reads “S. repetita M & C.,” establishing the authorship as Maynard and Clapp in Maynard, 1920. Type Material. The unintended descrip- tion of Strophiops repetita is based on the 46] illustration of a single specimen (fig. 89). It is likely that this was one of 1,052 spec- imens mentioned in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 1921b: 149) that were available to Maynard at the time and are thus part of the type series [Article 72.4.1.1, ICZN, 1999: 76]. Strophiops repetita is therefore based on 1,052 syntypes. No specimen la- beled as the “Holotype” of Strophiops re- petita was catalogued at the MCZ. Lot MCZ 76336 contained 198 specimens and was labeled “Paratypes.” A single speci- men was sequestered in a vial and labeled “figured.” This specimen closely approxi- mates both the illustration in figure 89 and the measurements in the intended de- scription. Also present was a label in May- nard’s handwriting referring to “Fig 89” and “Plate 45 figs 5-6.” This label indi- cates that the sequestered specimen is that in figure 89, as well as the specimen that Maynard regarded as the “type” because he planned to illustrate it as part of the intended taxon description. This speci- men, which retains the number MCZ 76336 is here designated as the lectotype of Strophiops repetita Maynard and Clapp in Maynard, 1920, to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. The remain- ing 197 specimens were recatalogued as MCZ 357003 and are among the remain- ing 1,051 specimens that become paralec- totypes. Lectotype Measurements. Length 27.3 mm, diameter (excluding lip) 10.4 mm; ap- erture height (including lip) 9.3 mm, ap- erture width (including lip and peristome) 8.0 mm. Type Locality. Old fields off Kemp’s Road, East Nassau [New Providence Is- land, Bahamas]. Remarks. The spelling Strophiops repi- tita appeared only as a nomen nudum (Maynard, 1919b: 36). Both the unintend- ed and the intended taxon descriptions used the spelling S. repetita. The sales cat- alog (Maynard, 1924bp?: [4]) used the spell- 462 ing repitita, but its supplement (Maynard, 1924c: 6) listed S. repetita as nearly extinct (only three specimens) in 1924 from the burning of vegetation prior to cultivation. Gould and Woodruff (1986: 477) regarded this taxon to be a synonym of Cerion gub- ernatorium (Crosse, 1869). Taxon 139. Strophiops perantiqua May- nard, 1919a [October 16]: 15 [nomen nudum]; Maynard, 1919b [November 6]: 31 [nomen nudum]; Maynard and Clapp in Maynard, 1920d [Septem- ber 30]: 115, pl. 1, figs. 1, 2, map 1, 2 [published as map 1, 1]. Examined 25 specimens. Size given as 1.25 by .60 linches; 31.8 by 15.2 mm], but the term “type” was not used. Nomenclatural Remarks. Maynard’s (1919a: 15) first use of this binomen was a nomen nudum. Maynard (1919b: 31) again used this name, this time along with a ref- erence to illustrations that were issued lat- er (Maynard, 1920d, pl. 1, figs. 1, 2), to- gether with the intended taxon descrip- tion. The second use of the name in 1919 was again a nomen nudum. Article 10.1.1 (ICZN, 1999: 9) states that if publication of the data relating to a new nominal taxon or a nomenclatural act is interrupted and continued at a later date, the name or act becomes available only when the require- ments of the relevant Articles have been met. Thus, the name S. perantiqua be- comes available on September 30, 1920, upon publication of the illustrations but does not have priority over the simulta- neously published intended taxon descrip- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 tion, which references the same illustra- tions. The taxon S. perantiqua dates from the intended species description (Maynard and Clapp in Maynard, 1920d: 115, pl. 1, figs. 1, 2), as cited by Clench and Aguayo (1952). Strophiops perantiqua is an junior objective synonym of Strophiops peravita Maynard, 1919, because they share the same type series, including the same lec- totype [see Taxon 114]. Type Material. The taxon description by Maynard and Clapp in Maynard, 1920d (p. 115) did not distinguish among the 25 specimens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in the original description or in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Clench and Aguayo (1952: 440, pl. 55, fig. 6) listed and illustrated MCZ 187537 as the “holotype,” but the speci- men is not accompanied by Maynard’s la- bel. Article 74.5 (ICZN, 1999: 82) states, “When the original work reveals that the taxon had been based on more than one specimen, a subsequent use of the term ‘holotype’ does not constitute a valid lec- totype designation unless the author, when wrongly using that term, explicitly indicat- ed that he or she was selecting from the type series that particular specimen to serve as the name-bearing type.” Thus, there is no holotype for this taxon, nor can Clench and Aguayo’s treatment of MCZ 187537 be considered a lectotype desig- nation. The species is based on 25 syn- —- Figures 138-150. Taxon 138. Strophiops repetita Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76336. Old fields off Kemp’s Road, East Nassau, New Providence Island, Bahamas. Taxon 139. Strophiops perantiqua Maynard and Clapp in May- nard, 1920. [Not illustrated.] The binomen Strophiops perantiqua is an objective junior synonym of Strophiops peravita Maynard, 1919 [Taxon 114], as the lectotype of Strophiops peravita is also the lectotype of S. perantiqua. Taxon 140. Strophiops processa Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76148. In area just south of the third beach north of the south end of Great Guana Key Exumas, Bahamas. Taxon 141. Strophiops mutatoria Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76234. Northwest of small pond near south end of Great Guana Key, Exuma Group, Bahamas. Taxon 142. Strophiops mitra Maynard and Clapp in Maynard, 1920. Lectotype MCZ 187538. Back of fourth beach from south end of Great Guana Key, Exuma Group, Bahamas. Taxon 143. Strophiops intentata Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76305. Small area east or tract 5 just south of the third beach north of the south end of Great Guana Key, Exumas, Bahamas. Taxon 144. Strophiops prognata Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76163. Between small pond and beach, southern end of Great Guana Key, Exuma Group, Bahamas. Taxon 145. Strophiops extrema Maynard and Clapp in Maynard, 1920. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 463 Lectotype MCZ 76146. Near well, at extreme south end of Great Guana Key, Exuma Group, Bahamas. Taxon 146. Strophiops imperfecta Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76313. Near well on south end of Great Guana Key, Exuma Group, Bahamas. Taxon 147. Strophiops progressa Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76197. Near Black Point, north end of Great Guana Key, Exuma Group, Bahamas. Taxon 148. Strophiops cyclura Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76045. Bitter Guana Key, Exuma Group, Bahamas. Taxon 149. Strophiops dissimila Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76057. East Cistern Key, Exuma Group, Bahamas. Taxon 150. Strophiops veta Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76221. Strout’s [Stroud’s] Key, Exuma Group, Bahamas. 464 types. Of these, MCZ 187537 (the lecto- type of Strophiops peravita [Taxon 114)]), which closely approximates the published measurements and clearly matches the published illustrations, is here designated as the lectotype of Strophiops perantiqua to provide an objective standard of refer- ence for this species-group taxon that is consistent with Maynard’s original and Clench and Aguayo’s subsequent concepts of this taxon. The remaining 24 syntypes become paralectotypes. Lectotype Measurements. Length 31.9 mm, diameter (excluding lip) 14.7 mm; ap- erture height (including lip) 11.1 mm, ap- erture width (including lip and peristome) 9.5 mm. Type Locality. Found fossil in red earth near west coast of Great Guana Key |Ex- umas, Bahamas], about 1 mile from south- ern end. Remarks. Maynard (1924a: 164) de- scribed in detail the locality in which these fossils were collected. Clench and Aguayo (1952) considered Cerion perantiquum to be a hybrid between the “scalarinum ele- ment and a member of the glans com- plex.” Taxon 140. Strophiops processa Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 116, pl. 1, figs. 7, 8, map 1, 5. Examined 300 specimens. Size giv- en as .85 by .35 [inches; 26.9 by 7.6 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 300 speci- mens examined. Later, Maynard (1924a: 168) wrote that only 122 were collected. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43— 44) specified that types were selected. However, there was no indication in the original description or in any of Maynard’s publications to associate a single specimen with the term “type.” Clench and Aguayo (1952: 431, pl. 56, fig. 3) listed and illus- trated MCZ 76148 as the “holotype,” pre- sumably on the basis of a handwritten la- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 bel accompanying the specimen with the term “typical.” Article 74.5 (ICZN, 1999: 82) states, “When the original work reveals that the taxon had been based on more than one specimen, a subsequent use of the term ‘holotype’ does not constitute a valid lectotype designation unless the au- thor, when wrongly using that term, ex- plicitly indicated that he or she was se- lecting from the type series that particular specimen to serve as the name-bearing type.” Thus, there is no holotype for this taxon, nor can Clench and Aguayo’s treat- ment of MCZ 76148 be considered a lec- totype designation. The species is based on either 300 or 122 syntypes. Of these, MCZ 76148 is here designated as the lectotype to provide an objective standard of refer- ence for this species-group taxon that is consistent with Maynard’s original and Clench and Aguayo’s subsequent concepts of this taxon. The remaining 121 or 299 syntypes become paralectotypes. Lectotype Measurements. Length 24.4 mm, diameter (excluding lip) 8.8 mm; ap- erture height (including lip) 8.5 mm, ap- erture width (including lip and peristome) Too) Wi Type Locality. In area just S of the third beach north of the south end of Great Guana Key [Exumas, Bahamas]. Remarks. Maynard (1920d: 116) report- ed that this taxon was rather common, but all 300 specimens collected were dead. Maynard (1924a: 168) later contradicted this number, stating that 122 specimens were collected. Clench and Aguayo (1952: 431) considered this taxon to be a subspe- cies of Cerion asperum (Maynard, 1920). Taxon 141. Strophiops mutatoria May- nard and Clapp in Maynard, 1920d [September 30]: 116, pl. 1, figs. 3, 4, map 1, 6; Maynard, 1924a: 170. Ex- amined 500 specimens. Size given as 1.00 by .26 [inches; 25.4 by 6.6 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 500 speci- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. mens examined. Later, Maynard (1924a: 170) wrote that 627 were collected. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. How- ever, there was no indication in the origi- nal description or in any of Maynard's pub- lications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 500 or 627 syntypes. Of these, MCZ 76234 was labeled “Holotype- command.” and contains a single specimen accompanied by a note in Maynard's hand- writing with the term “type.” This speci- men is here designated as the lectotype to 465 types were selected. However, there was no indication in the original description or in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 75 syn- types. Of these, MCZ 187538, which was labeled “Holotype” and contains a single specimen, is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon. The remaining 74 syntypes become paralecto- types. Lectotype Measurements. Length oleZ mm, diameter (excluding lip) 13.7 mm; ap- erture height (including lip) 12.2 mm, ap- provide an objective standard of reference for this species-group taxon that is consis- tent with Maynard’s original concept of it. The remaining 499 or 626 syntypes be- come paralectotypes. Lectotype Measurements. Length 25.9 mm, diameter (excluding lip) 7.4 mm; ap- erture height (including lip) 7.6 mm, ap- erture width (including lip and peristome) 6.4 mm. Type Locality. In area south of last [Strophiops processa = area just south of the third beach north of the south end of Great Guana Key, Exuma Group, Baha- mas |. Remarks. Maynard (1920d: 116) report- ed this taxon to be common, but all 500 specimens examined were collected dead. Later, Maynard (1924a: 170) reported that 623 specimens were collected dead, of which two were fossilized. Taxon 142. Strophiops mitra Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 118, pl. 2, figs. 4, 5, map 1, 1. Examined 75 specimens. Size giv- en as 1.25 by .50 [inches; 31.8 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 75 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that erture width (including lip and peristome) 9.9 mm. Type Locality. Back of fourth beach from south end of Great Guana Key [Ex- uma Group, Bahamas], under fallen palm fronds. Remarks. Maynard (1924a: 170) provid- ed supplemental information on the shell, animal, anatomy, and habitat. Taxon 143. Strophiops intentata Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 118, pl. 2, figs. 6, 7, map 1, 7; Maynard, 1925: 171. Examined 256 specimens. Size given as .75 by .30 [inches; 19.1 by 7.6 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 256 speci- mens examined. Later, Maynard (1925: 171) wrote that 239 were collected. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. How- ever, there was no indication in the origi- nal description or in any of Maynard’s pub- lications to associate a single specimen with the term “type.” Clench and Aguayo, (1952: 431, pl. 56, fig. 4) identified and illustrated MCZ 76305 as “holotype,” pre- sumably on the basis of a handwritten la- bel identifying a type specimen. Article 4.5 (ICZN, 1999: 82) states, “When the orig- 466 inal work reveals that the taxon had been based on more than one specimen, a sub- sequent use of the term ‘holotype’ does not constitute a valid lectotype designation unless the author, when wrongly using that term, explicitly indicated that he or she was selecting from the type series that par- ticular specimen to serve as the name- bearing type.” Thus, there is no holotype for this taxon, nor can Clench and Aguayo’s treatment of MCZ 76305 be con- sidered a lectotype designation. The spe- cies is based on 256 or 239 syntypes. Of these, MCZ 76305 is here designated as the lectotype to provide an objective stan- dard of reference for this species-group taxon that is consistent with Maynard’s original and Clench and Aguayo’s subse- quent concepts of it. The remaining 255 or 238 syntypes become paralectotypes. Lectotype Measurements. Length 20.3 mm, diameter (excluding lip) 8.4 mm; ap- erture height (including lip) 7.3 mm, ap- erture width (including lip and peristome) 6.8 mm. Type Locality. Rather common in a small area east of tract 5 | = type locality of Strophiops processa = area just south of the third beach north of the south end of Great Guana Key, Exumas, Bahamas]. Remarks. Maynard (1920d: 118) report- ed that all of the 256 specimens were col- lected dead and some were very old. May- nard (1925: 170) reported that 239 speci- mens were collected. Clench and Aguayo (1952: 431) listed this taxon in the synon- ymy of Cerion asperum processum (May- nard and Clapp, 1920) [Taxon 140]. Taxon 144. Strophiops prognata Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 118, pl. 2, figs. 8, 9, map 1, 8; Maynard, 1924: 171-172. Exam- ined 300 specimens. Size given as .92 by 40 [inches; 23.4 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 300 speci- mens examined. In the preamble to the Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in the original description or in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 300 syn- types. Of these, MCZ 76163, which was labeled “Holotype” and contains a single specimen accompanied by a note in May- nard’s handwriting with the term “type,” is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 299 syntypes become paralecto- types. Lectotype Measurements. Length LOS) mm, diameter (excluding lip) 9.8 mm; ap- erture height (including lip) 9.0 mm, ap- erture width (including lip and peristome) 7.6 mm. Type Locality. Common in area be- tween a small pond P [map 1, 8, p. 117] and beach. [Southern end of Great Guana Key, Exuma Group, Bahamas. | Remarks. Maynard (1920d: 118) report- ed that all 300 specimens were collected dead, but were more recent than Stro- phiops intentata. Maynard (1924: 171— 172) added some additional information, but did not mention the number collected. Taxon 145. Strophiops extrema Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 118, pl. 3, figs. 1, 2 [these figures are cited in the original descrip- tion but captioned “S. imperfecta”, pl. 2, figs. 10, 11 [these figures are not cit- ed in the original description but are captioned “S. extrema M. & C.”], map 1, 9; Maynard, 1924: 173. Examined 150 specimens. Size given as 1.05 by .49 [inches; 26.7 by 12.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 150 speci- mens examined. In the preamble to the THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in the original description or in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 150 syn- types. Of these, MCZ 76146, which was labeled “Holotype” and contains a single specimen accompanied by a note in May- nard’s handwriting with the term “type,” is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 149 syntypes become paralecto- types. Lectotype Measurements. Length 25.9 mm, diameter (excluding lip) 10.3 mm; ap- erture height (including lip) 9.6 mm, ap- erture width (including lip and peristome) Tot Tamir, Type Locality. Found in vicinity of well at extreme south end of [Great Guana] Key |Exuma Group, Bahamas]. Remarks. Maynard and Clapp in May- Mand lLOZ0d: (ells) ered, an! citing the wrong illustrations in their original de- scription. The illustrations were correctly captioned and published at the same time as the description. Clench (1957: 145) cor- rected this error. Maynard (1920d: 118) reported that all 150 specimens were col- lected dead and were of about the same age as Strophiops prognata, with some specimens showing reddish markings. Maynard (1924: 173) added ecological in- formation and confirmed that 150 speci- mens were collected. Taxon 146. Strophiops imperfecta May- nard and Clapp in Maynard, 1920d [September 30]: 118-119, pl. 1 [sic, actually 3], figs. 1, 2, map 1, 10. Ex- amined two specimens. Size given as 1.05 by .40 [inches; 26.7 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the two speci- 467 mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in the original description or in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on two syn- es. Of these, MCZ 76313, which was labeled “Holotype” and contains a single specimen accompanied by a note in May- nard’s handwriting with the term “type” is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynards original concept of it. The other syntype become a paralectotype. Lectotype Measurements. Length 27.7 mm, diameter (excluding lip) 11.6 mm; ap- erture height (including lip) 10.4 mm, ap- erture width (including lip and peristome) 9.8 mm. Type Locality. Near well, on south end of Great Guana Key [Exuma Group, Ba- hamas |]. Remarks. Maynard and Clapp in May- nard (1920d: 118) erred in citing the wrong plate number but the correct figure numbers in their description of this taxon. The illustrations were correctly captioned and published at the same time as the de- scription. Clench (1957: 148) corrected this error. Maynard (1920d: 119) reported this taxon to be fossil, embedded in rock. Taxon 147. Strophiops progressd Maynard, 1919b [November 6]: 32, 34, 35 [nonem nudum]; Maynard and Clapp in May- nard, 1920d [September 30]: 119, pl. 2, figs. 1-3. Examined 1,738 spec- imens. Size given as 1.15 by .45 [inches; 29.2 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,738 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that 468 types were selected. However, there was no indication in the original description or in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 1,738 syntypes. Lot MCZ 76197 contains a single specimen labeled “Holotype” that approx- imates the measurements and illustration of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 1,737 syn- types, including MCZ 118179 and USNM 419947 (100 specimens), become paralec- totypes. Lectotype Measurements. Length 29.6 mim, diameter (excluding lip) 12.1 mm; ap- erture height (including lip) 11.4 mm, ap- erture width (including lip and peristome) 10.0 mm. Type Locality. Near Black Point, north end of Great Guana Key |Exuma Group, Bahamas]. Remarks. This binomen appeared as a nomen nudum (Maynard, 1919b: 32, 34, 35) before its description. Taxon 148. Strophiops cyclura Maynard, 1919a [October 16]: 24 [nomen nu- dum]; Maynard, 1919b [November 6]: 25 [nomen nudum]; Maynard and Clapp in Maynard, 1920d [Septem- ber 30]: 119, pl. 3, figs. 3-5; Maynard, 1925: 181-185. Examined 4,422 speci- mens. Size given as 1.10 by .40 [inches; 27.9 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 4,422 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in the original description or Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Thus, there is no holotype for this taxon and the species is based on 4,422 syntypes. Lot MCZ 76045 contains a single specimen labeled “Holotype” that approx- imates the measurements and illustration of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 4,421 syn- types, including MCZ 76046, MCZ 118235, and USNM 419949 (500 speci- mens), become paralectotypes. Lectotype Measurements. Length 28.8 mm, diameter (excluding lip) 11.6 mm; ap- erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. Bitter Guana [Key, Exu- ma Group, Bahamas]. Remarks. This binomen first appeared as a nomen nudum (Maynard, 1919a: 24, 1919b: 25). Maynard (1925: 181-185) pro- vided additional anatomical information on this taxon and noted that he collected 82 specimens from a single lily plant. Taxon 149. Strophiops dissimila Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 119, pl. 24, figs. 8, 9. Ex- amined 86 specimens. Measurements not provided. Type Material. The original description did not distinguish among the 86 speci- mens examined. Maynard (1926: 195) re- ported that 275 specimens were collected. Presumably 189 of these specimens were collected after the taxon was described. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. How- ever, there was no indication in the origi- nal description or in any of Maynard's pub- lications to associate a single specimen THE CERION TAXA OF CHARLES JOHNSON MAYNARD *¢ Harasewych et al. with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 86 syntypes. Of these, MCZ 76057, which was labeled “Holotype” and contains a single specimen accompanied by a note in Maynard's handwriting iden- tifying the taxon but not using the term “type,” is here designated as the lectotype to provide an objective standard of refer- ence for this species-group taxon that is consistent with Maynard's original concept of it. The remaining 85 syntypes, which might have been co-mingled with 189 sub- sequently collected non-type specimens, become paralectotypes. Lectotype Measurements. Length 29.4 mm, diameter (excluding lip) 11.7 mm; ap- erture height (including lip) 11.2 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. East Cistern Key [Exuma Group, Bahamas]. Remarks. Maynard (1920d: 119) report- ed this taxon to occur on palm fronds and bushes. Taxon 150. Strophiops veta Maynard and Clapp in Maynard, 1920d [Septem- ber 30]: 120, pl. 21, figs. 3, 4. Ex- amined 80 specimens. Size given as 1.25 by .50 [inches; 31.8 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 80 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in the original description or in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 80 syn- types. Lot MCZ 76221 contains a single specimen labeled “Holotype” that approx- imates the illustration and the measure- ments of this taxon and is accompanied by a label in Maynard’s hand with the term “type” that includes references to the fig- 469 ures. This specimen is here designated as the lectotype to provide an objective stan- dard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 79 syntypes become paralectotypes. Lectotype Measurements. Length 31.4 mm, diameter (excluding lip) 12.2 mm; ap- erture height (including lip) 11.5 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. Strout’s [Stroud’s|] Key [Exuma Group, Bahamas]. Fossil in cliff on west side. Remarks. Maynard (1920d: 120) report- ed this taxon to be the immediate ancestor of S. stroutii. Taxon 151. Strophiops stroutii Maynard, 1919a [October 16]: 15 [nomen nu- dum]; Maynard, 1919b [November 6]: 35 [nomen nudum]; Maynard and Clapp in Maynard, 1920d [Septem- ber 30]: 120, pl. 21, figs. 1, 2; May- nard, 1926: 198-203. Examined 1,951 specimens. Measurements not provided. Type Material. The original description did not distinguish among the 1,951 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in the original description or in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 1,951 syntypes. Lot MCZ 76130 contains a single specimen labeled “Holotype” that approx- imates the illustration of this taxon [no measurements were published] and is ac- companied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 1,951 syntypes, including MCZ 470 76131, MCZ 76132, MCZ 118195, USNM 420000 (200 specimens), and USNM 42122 (200 specimens), become paralec- totypes. Lectotype Measurements. Length 30.3 mim, diameter (excluding lip) 12.2 mm; ap- erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) 8.3 mm. Type Locality. Maynard (1920d: 120) and Clench (1957: 164) listed this taxon as occurring on Little Strout [Stroud], Strout [Stroud], East and West Hawksbill and Cistern Keys |[Exuma Group, Bahamas]. The lectotype is from Strout [Stroud] Key. Remarks. Maynard (1919a: 15, 1919b: 35) twice introduced this binomen as a no- men nudum. Maynard (1926: 198-203) provided additional data on the basis of dissections and noted that this was one of a very few taxa that occurred on more than a single key. Taxon 152. Strophiops palmata minuta Maynard and Clapp in Maynard, 1920d [September 30]: 120, pl. 21, fig. 5. Examined 600 specimens. Size given as .95 by .40 [inches; 24.13 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 600 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in the original description or in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 600 syn- types. Because no specimen labeled “Ho- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 lotype” or accompanied by a label in May- nard’s hand could be found, a specimen, USNM 420093, that most closely approx- imates the published measurements and figure is here designated as lectotype from a large syntype lot at USNM to provide an objective standard of reference for this species-group taxon. The remaining 599 syntypes, including MCZ 76214, MCZ 118243, and USNM_ 1093792, become paralectotypes. Lectotype Measurements. Length 24.2, mm, diameter (excluding lip) 9.9 mm; ap- erture height (including lip) 8.6 mm, ap- erture width (including lip and peristome) 7.4 mm. Type Locality. Wax Key [Great Exuma Island, Exuma Cays, Bahamas]. Remarks. Maynard (1926: 213-218) provided additional data on shell mor- phology and anatomy. Taxon 153. Strophiops degenis Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 120-121, pl. 4, figs. 6, 7. Examined 34 specimens. Size given as 1.10 by .45 linches; 27.9 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 34 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in the original description or in any of Maynard’s publications to asso- ciate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 34 syn- types. Lot MCZ 76404 contains a single specimen labeled “Holotype” that approx- => Figures 151-162. Taxon 151. Strophiops stroutii Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76130. Strout [Stroud], Key, Exuma Group, Bahamas. Taxon 152. Strophiops palmata minuta Maynard and Clapp in Maynard, 1920. Lectotype USNM 420093. Wax Key, Great Exuma Island, Exuma Cays, Bahamas. Taxon 153. Strophiops degenis Maynard and Clapp jn Maynard, 1920. Lectotype MCZ 76404. Harvest Key, Harvey Cay, Exuma Cays, Bahamas. Taxon 154. Strophiops sampsoni Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76235. North beach of Sampson’s Key, Exuma Cays, Bahamas. Taxon 155. Strophiops tenucostata Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76194. South Beach of Sampson’s Key, Exuma Cays, Bahamas. Taxon 156. Strophiops fulvia Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76159. East side of Pipe Key, Exuma Group, Bahamas. Taxon 157. Strophiops inquita Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76182. Fowl THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 471 Key, Exuma Group, Bahamas. Taxon 158. Strophiops cervina Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76047. Refuge Key, east of Norman’s Pond Key, Exuma Cays, Bahamas. Taxon 159. Strophiops pulla Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76217. Key opposite Roseville, Great Exuma, Exuma Cays, Bahamas. Taxon 160. Strophiops caerules- cens Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76198. Key north of key opposite Roseville, Great Exuma, Bahamas. Taxon 161. Strophiops nebula Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76039. First key north of Stocking Island, Exuma Cays, Bahamas. Taxon 162. Strophiops exorta Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76405. Interior of Refuge Key, Exuma Group, Bahamas. A472 imates the illustration and the measure- ments of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. The remain- ing 33 syntypes become paralectotypes. Lectotype Measurements. Length 28.0 mm, diameter (excluding lip) 11.1 mm; ap- erture height (including lip) 9.4 mm, ap- erture width (including lip and peristome) 7.6 mm. Type Locality. Harvest Key [Harvey Cay, Exuma Cays, Bahamas]. Remarks. Maynard (1920d: 121) report- ed this taxon as occurring very scattered on palms, bushes, and rocks. Taxon 154. Strophiops sampsoni Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 121, pl. 4, figs. 1, 10. Ex- amined 1,540 specimens. Size given as 1.20 by .50 [inches; 30.5 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,540 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,540 syntypes. Lot MCZ 76235 was labeled “Holotype” but contains two syntypes, one of which matches closely the published dimensions and illustration of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The other specimen has Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 been recatalogued as MCZ 357004. The remaining 1,539 syntypes, including MCZ 357004, MCZ 76236, MCZ 118237, and USNM 420090 (100 specimens), become paralectotypes. Lectotype Measurements. Length 30.4 mim, diameter (excluding lip) 11.6 mm; ap- erture height (including lip) 10.4 mm, ap- erture width (including lip and peristome) 8.4 mm. Type Locality. North beach of Samp- son's Key |Exuma Cays, Bahamas]. Remarks. Maynard (1920d: 121) report- ed this taxon was common on palms. j Taxon 155. Strophiops tenucostata May- nard and Clapp in Maynard, 1920d [September 30]: 121, pl. 4, figs. 3 [sic, actually 8], 9. Examined 112 spec- imens. Measurements not provided. Type Material. The original description did not distinguish among the 112 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 112 syntypes. Lot MCZ 76194 contains a single specimen labeled “Ho- lotype” that matches the illustrations of this taxon and is accompanied by a label in Maynard’s hand with the term “type.” This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The remaining 111 syntypes, including MCZ 76195, become paralecto- types. Lectotype Measurements. Length 28.5 mm, diameter (excluding lip) 11.0 mm; ap- erture height (including lip) 9.9 mm, ap- erture width (including lip and peristome) 7.9 mm. Type Locality. South Beach of Samp- son’s Key [Exuma Cays, Bahamas]. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. Remarks. Maynard (1920d: 121) report- ed this occurred on palms but was not very common. Taxon 156. Strophiops fulvia Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 121, pl. 4, figs. 4, 5. Number of specimens examined not given. Size given as 1.15 by .40 [inches; 29.2 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not specify the number of specimens examined but noted that they were com- mon. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on an un- determined number of syntypes. Lot MCZ 76159 contains a single specimen labeled “Holotype” that matches closely the pub- lished dimensions and illustration of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining syntypes, in- cluding MCZ 76158, MCZ 118223, and USNM 420039 (100 specimens), become paralectotypes. Lectotype Measurements. Length 29.5 mm, diameter (excluding lip) 11.4 mm; ap- erture height (including lip) 9.9 mm, ap- erture width (including lip and peristome) 7.6 mm. Type Locality. On the east side of Pipe Key [Exuma Group, Bahamas]. Remarks. Maynard (1920d: 121) report- ed this taxon to be common beneath palm fronds. Taxon 157. Strophiops inquita Maynard and Clapp in Maynard, 1920d [Sep- 473 tember 30]: 121, pl. 4, figs. 2, 3 |fig- ure caption as S. inquieta]. Neither the number of specimens nor the measure- ments are provided. Type Material. The original description did not distinguish among the unspecified number of specimens examined but noted that this taxon was found in 11 small, iso- lated groups, indicating, at a minimum, 11 syntypes. In the preamble to the taxonom- ic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 11 or more syntypes. Lot MCZ 76182 contains 14 adult specimens and a juvenile and was labeled “Paratype.” No labels by Maynard accompany this lot. The specimen that most closely resembles the illustrations in size and proportion and in the visibility of a weak columellar tooth is selected as lec- totype. The remaining 13 adults and ju- venile, recatalogued as MCZ 357005, are paralectotypes. Lectotype Measurements. Length 30.6 mim, diameter (excluding lip) 12.4 mm; ap- erture height (including lip) 10.8 mm, ap- erture width (including lip and peristome) Sain, Type Locality. Fowl Key [Exuma Group, Bahamas]. Remarks. The taxon name originally ap- peared as Strophiops inquita in the head- ing of the original description, but as S. inquieta in the caption to the figures in the plate. The spellings inquita and inquieta are thus different original spellings. May- nard never referred to this taxon again, and it was not listed in his catalog. Clench (1957: 149) listed this taxon as inquita, but cannot be considered the First Revisor [Article 24.2.3, ICZN, 1999: 30] because he did not list both original spellings. Stro- phiops inquita is here formally selected as the correct original spelling. Maynard (1920d: 121) reported that this taxon was A74 nearly exterminated by cultivation when he encountered it in 1915. Taxon 158. Strophiops cervina Maynard, 1919a [October 16]: 21 [nomen nu- dum]; Maynard and Clapp in May- nard, 1920d [September 30]: 121, pl. 22, figs. 8, 9. Examined 3,480 spec- imens. Size given as 1.30 by .47 [inches; 33.0 by 11.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 3,480 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 3,480 syntypes. Perusal of the MCZ catalog indicated that no specimen identified by Maynard as “type” could be recognized in his collection when it reached MCZ. Lot MCZ 76047 comprised 158 specimens that were labeled “para- types.” Segregated in a vial were two spec- imens, a juvenile and a transversely broken adult shell, together with a label referring to “fig. 124a,” which was never published. A specimen matching the published mea- surements and approximating the illustra- tions was selected as lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 157 specimens were recatalogued as MCZ 357006. These and MCZ 118251 and USNM 419938 are among the 3,479 syntypes that become paralectotypes. Lectotype Measurements. Length 33.0 mm, diameter (excluding lip) 11.9 mm; ap- erture height (including lip) 11.1 mm, ap- erture width (including lip and peristome) 8.6 mm. Type Locality. Refuge Key (east of Nor- man’s Pond Key) [Exuma Cays, Bahamas]. Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Remarks. Maynard's (1919a: 21) first use of this binomen was a nomen nudum. Lat- er (1920d: 121), he reported this taxon to be exceedingly abundant on palms, bush- es, and small trees. Taxon 159. Strophiops pulla Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 122, pl. 23, figs. 3, 4. Ex- amined 1,572 specimens. Size given as 1.05 by .45 linches; 26.7 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,572 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,572 syntypes. Lot MCZ 76217 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and illustration of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 1,571 syntypes, including MCZ 76218 and USNM 419948 (100 speci- mens), become paralectotypes. Lectotype Measurements. Length 26.6 mm, diameter (excluding lip) 12.0 mm; ap- erture height (including lip) 9.8 mm, ap- erture width (including lip and peristome) 8.2 mm. Type Locality. Key opposite Roseville, Great Exuma, Exuma Cays, Bahamas. Remarks. Maynard (1920d: 122) report- ed this taxon to be common on bushes. Taxon 160. Strophiops caerulescens May- nard and Clapp in Maynard, 1920d [September 30]: 122, pl. 22, fig. 10, pl. 23, fig. 5. Examined 2,000 speci- THE CERION TAXA OF CHARLES JOHNSON MAYNARD ° Harasewych et al. mens. Size given as 1.30 by .50 [inches; 33.0% by 127 rom, but the term type™ was not used. Type Material. The original description did not distinguish among the 2,000 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 2,000 syntypes. Lot MCZ 76198 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and illustration of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 1,999 syntypes, including MCZ 76199, MCZ 118268, MCZ 357007 and USNM 419965 (300 specimens), become paralectotypes. Lectotype Measurements. Length 33.0 mim, diameter (excluding lip) 13.3 mm; ap- erture height (including lip) 11.4 mm, ap- erture width (including lip and peristome) 10.0 mm. Type Locality. Key north of key opposite Roseville, Great Exuma, Bahamas. Remarks. Maynard (1920d: 122) report- ed this taxon to be common, chiefly on palms. Clench (1934: 214) noted that he compared specimens from Strachan, Salt- water Pond, and Clem Cays off Clarence- town, Long Island, with the type specimen of Cerion caerulescens from Exuma and thought the differences to be only colonial. Taxon 161. Strophiops nebula Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 122, pl. 23, fig. 10, pl. 24, fig. 1. Examined 250 specimens. Size given as 1.20 by .44 [inches; 30.5 A475 by 11.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 250 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 250 syntypes. Of these, MCZ 76039, which was labeled “Holotype” and contains a single specimen accompanied by a note in Maynard’s handwriting with the term “type,” is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 249 syntypes become paralectotypes. Lectotype Measurements. Length DST mm, diameter (excluding lip) 11.5 mm; ap- erture height (including lip) 10.8 mm, ap- erture width (including lip and peristome) 9.3 mm. Type Locality. First key north of Stock- ing Island, Exuma Cays, Bahamas. Remarks. Maynard (1920d: 122) report- ed this taxon to be rather uncommon on palms. Taxon 162. Strophiops exorta Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 122, pl. 24, figs. 4, 5. Ex- amined nine specimens. Size given as .90 by .35 [inches; 22.9 by 8.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the nine speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is 476 based on nine syntypes. Lot MCZ 76405 contains a single specimen labeled “Ho- lotype” that approximates the illustration and the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining eight syntypes become paralecto- types. Lectotype Measurements. Length Quik mm, diameter (excluding lip) 10.0 mm; ap- erture height (including lip) 8.8 mm, ap- erture width (including lip and peristome) TAS) TANI, Type Locality. Interior of Refuge Key, Exuma Group, Bahamas. Remarks. Maynard (1920d: 122) report- ed this taxon to be limited to a small col- ony of nine specimens. Taxon 163. Strophiops cylindriata May- nard and Clapp in Maynard, 1920d [September 30]: 123, pl. 23, figs. 8, 9. Examined 132 specimens. Size given as 1.35 by .50 [inches; 34.3 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 132 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 based on 132 syntypes. Of these, MCZ 76031, which was labeled “Holotype” and contains a single specimen accompanied by a note in Maynard’s handwriting with the term “type,” is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 131 syntypes become paralectotypes. Lectotype Measurements. Length 33.4 mm, diameter (excluding lip) 13.3 mm; ap- erture height (including lip) 11.4 mm, ap-. erture width (including lip and peristome) 923 mmar Type Locality. First key north of Leward Stocking Key, Exuma Group, Bahamas. Remarks. Maynard (1920d: 123) report- ed this taxon to occur beneath palm fronds. Taxon 164. Strophiops exasperata May- nard and Clapp in Maynard, 1920d [September 30]: 123, pl. 23, figs. 1, 2. Examined 572 specimens. Size given as 1.27 by .47 [inches; 32.3 by 11.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 572 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 572 syntypes. Lot MCZ 76041 contains a single specimen labeled “Ho- lotype” that approximates the illustration —= Figures 163-174. Taxon 163. Strophiops cylindriata Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76031. First key north of Leward Stocking Key, Exuma Group, Bahamas. Taxon 164. Strophiops exasperata Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76041. Northeast point, Key east of Anna’s Tract, Great Exuma, Bahamas. Taxon 165. Strophiops crassa Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76220. East Hill, Little Norman’s Key, Exuma Cays, Bahamas. Taxon 166. Strophiops normanii Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76248. West Hill, Little Norman’s Key, Exuma Group, Bahamas. Taxon 167. Strophiops semipolita Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76033. Third key southeast of Roseville, Great Exuma, Bahamas. Taxon 168. Strophiops accuminator Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76349. Long Key, Long Cay, Exuma Group, Bahamas. Taxon 169. Strophiops navalis Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76035. South end of Ship Channel Key, Exuma Group, Bahamas. Taxon 170. Strophiops valida THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. AT7 Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76043. Near well on west coast of Ship Channel Key, Exuma Group, Bahamas. Taxon 171. Strophiops genetiva Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76037. On south end of Ship Channel Key, Exuma Group, Bahamas. Taxon 172. Strophiops marmorosa Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76185. Well Key, a little north of Leward Stocking Key, Exuma Group, Bahamas. Taxon 173. Strophiops pusilla Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76053. Key near Long Rock, Great Exuma, Exuma Group, Bahamas. Taxon 174. Strophiops inexpecta Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76307. Fossil in soft rock next to westernmost Brigadier Key, Exuma Group, Bahamas. 478 and the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 571 syntypes, including MCZ 76042, MCZ 118234, and USNM 420006 (75 specimens), become paralectotypes. Lectotype Measurements. Length BT mm, diameter (excluding lip) 12.5 mm; ap- erture height (including lip) 11.3 mm, ap- erture width (including lip and peristome) 8.6 mm. Type Locality. Northeast point, key east of Anna’s Tract, Great Exuma, Bahamas. Remarks. Maynard (1920d: 123) report- ed this taxon to occur in a small area on shrubs. Taxon 165. Strophiops crassa Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 123, pl. 21, figs. 9, 10. Examined 400 specimens. Size given as 1.37 by .50 [inches; 34.8 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 400 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 400 syntypes. Lot MCZ 76220 contains a single specimen labeled “Ho- lotype” that approximates the illustration and the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Maynard's original concept of it. The re- maining 399 syntypes, including MCZ 76219 and USNM 420007 (100 speci- mens), become paralectotypes. Lectotype Measurements. Length OES mim, diameter (excluding lip) 12.8 mm; ap- erture height (including lip) 11.4 mm, ap- erture width (including lip and peristome) 9.6 mm. Type Locality. East Hill, Little Nor- man’s Key, Exuma Cays, Bahamas. Remarks. Maynard (1920d: 123) report- ed this taxon to occur not very commonly at the base of shrubbery. Taxon 166. Strophiops normanii Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 123, pl. 21, fig. 8, pl. 20, fig. 5 [as normandi]|. Examined 400 specimens. Size given as 1.25 by .38 inches; 31.8 by 9.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 400 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 400 syntypes. Lot MCZ 76248 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and illustration of this taxon and is accompanied by a label in Maynard's hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 399 syntypes, including MCZ 76249 and USNM 420094 (100 speci- mens), become paralectotypes. Lectotype Measurements. Length oles mm, diameter (excluding lip) 12.7 mm; ap- erture height (including lip) 11.5 mm, ap- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. erture width (including lip and peristome) 9.1 mm. Type Locality. West Hill, Little Nor- mans Key, Exuma Group, Bahamas. Remarks. The taxon name originally ap- peared as Strophiops normanii in the heading of the original description, but as S. normandi in the caption to the figures in the plate. The spellings normanii and normandi are thus different original spell- ings. Maynard never referred to this taxon again, and it was not listed in his catalog. Clench (1957: 155) listed this taxon as nor- manii, but cannot be considered the First Revisor [Article 24.2.3, ICZN, 1999: 30] because he did not list both original spell- ings. Strophiops normanii is here formally selected as the correct original spelling. Maynard (1920d: 123) reported this taxon to occur on the ground, with few living and many dead specimens. Taxon 167. Strophiops semipolita May- nard and Clapp in Maynard, 1920d [September 30]: 123, pl. 23, figs. 6, 7. Examined 385 specimens. Size given asp es Ombya a Omiiimnenes. 35.0 mln. Ln? mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 385 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 385 syntypes. Lot MCZ 76033 contains a single specimen labeled “Ho- lotype” that approximates the illustration and the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to subsequently published figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original A479 concept of it. The remaining 384 syntypes, including MCZ 76034 and MCZ 118254, become paralectotypes. Lectotype Measurements. Length 34.2 mm, diameter (excluding lip) 14.0 mm; ap- erture height (including lip) 11.7 mm, ap- erture width (including lip and peristome) 9.7 mm. Type Locality. Third key southeast of Roseville, Great Exuma, Bahamas. Taxon 168. Strophiops acuminata May- nard, 1920b [April 8]: 84 [nomen nu- dum]. Strophiops accuminator May- nard and Clapp in Maynard, 1920d [September 30]: 124, pl. 3, fig. 6 [ju- venile specimens], pl. 20, figs. 1, 2 [S. accumulata on pl. 20]. Examined 1,500 specimens. Size given as 1.15 by .47 inches; 29.2 by 11.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,500 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,500 syntypes. Lot MCZ 76349 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and illustration of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 1,499 syntypes, including MCZ 76348 and USNM 420062 (100 speci- mens), become paralectotypes. Lectotype Measurements. Length 29.0 mm, diameter (excluding lip) 11.5 mm; ap- erture height (including lip) 9.7 mm, ap- 450 erture width (including lip and peristome) 7 Al iamiam, Type Locality. Long Key [Long Cay, Ex- uma Group, Bahamas]. Remarks. The taxon name Strophiops acuminata first appeared as a nomen nu- dum (Maynard, 1920b [April 8]: 84). The next to appear was S. accuminator in the heading of the original description, but as S. accumulata in the caption to the figures in one of two plates. The spellings accu- minator and accumulata are thus different original spellings. Maynard (1924bp: [3]) listed the taxon as acuminator [one c] in his catalog. Clench (1957: 136) listed this taxon as accuminator, but cannot be con- sidered the First Revisor [Article 24.2.3, ICZN, 1999: 30] because he did not list both original spellings. Strophiops accu- minator is here formally selected as the correct original spelling. Taxon 169. Strophiops navalia Maynard, 1919b [November 6]: 32 [nomen nu- dum]; Maynard, 1919b [November 6]: 35 [nomen nudum]. Strophiops navalis Maynard and Clapp in Maynard, 1920d [September 30]: 124, pl. 20, figs. 8-10. Examined 1,070 specimens. Size given as 1.25 by .50 [inches; 31.8 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,070 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,070 syntypes. Lot MCZ 76035 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and illustration of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 1,069 syntypes, including MCZ 76036, MCZ 118226, and USNM 420015 (50 specimens), become paralectotypes. Lectotype Measurements. Length 30.9 mm, diameter (excluding lip) 12.3 mm; ap- erture height (including lip) 11.4 mm, ap- erture width (including lip and peristome) 9.2 mm. Type Locality. South end of Ship Chan- nel Key [Exuma Group, Bahamas]. Remarks. The binomen S. navalia ap- peared twice as a nomen nudum (May- nard, 1919b [November 6]: 32, 35). Taxon 170. Strophiops valida Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 124, pl. 22, figs. 1, 2. Ex- amined 400 specimens. Size given as 1.20 by .55 [inches; 30.5 by 14.0 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 400 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 400 syntypes. MCZ 76043 con- tains a single specimen labeled “Holotype” that approximates the measurements and the illustration for this taxon and is accom- panied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here des- ignated as the lectotype to provide an ob- jective standard of reference for this spe- cies-group taxon that is consistent with Maynard's original concept of it. The re- maining 399 syntypes, including MCZ 76044, become paralectotypes. Lectotype Measurements. Length 30.3 THE CERION TAXA OF CHARLES JOHNSON MAYNARD ¢ Harasewych et al. mm, diameter (excluding lip) 14.1 mm; ap- erture height (including lip) 11.9 mm, ap- erture width (including lip and peristome) 10.4 mm. Type Locality. Near well on west coast of Ship Channel Key [Exuma Group, Ba- hamas]. Remarks. Because Strophiops is consid- ered to be a subgenus of Cerion in current classifications, Strophiops valida Maynard and Clapp in Maynard, 1920, is a junior secondary homonym of Cerion (Maynar- dia) columna var. valida Pilsbry and Van- atta, 1895. In our view, it is likely that fu- ture research will identify a synonym for Strophiops valida Maynard and Clapp in Maynard, 1920 [Article 60.2, ICZN, 1999: 62], and that establishing a replacement name would only create an additional syn- onym. Taxon 171. Strophiops genitiva Maynard, 1919b [November 6]: 32, 35 [nomen nudum]. Strophiops genetiva Maynard and Clapp in Maynard, 1920d [Sep- tember 30]: 124 [genitiva in text], pl. 22, figs. 3, 4 [genetiva on plate]. Ex- amined 400 specimens. Size given as 1.50 by .60 [inches; 38.1 by 15.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 400 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s _ publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 400 syntypes. Lot MCZ 76037 contains a single specimen labeled “Ho- lotype” that approximates the illustrations and measurements of this taxon. This specimen is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard 481 of reference for this species-group taxon that is consistent with Maynard's original concept of it. The remaining 399 syntypes become paralectotypes. Lectotype Measurements. Length 38.6 mm, diameter (excluding lip) 14.5 mm; ap- erture height (including lip) 12.3 mm, ap- erture width (including lip and peristome) 10.0 mm. Type Locality. On south end of Ship Channel Key [Exuma Group, Bahamas]. Remarks. The binomen S. genitiva first appeared as a nomen nudum (Maynard, 1919b [November 6]: 32, 35). The taxon name was spelled genitiva in the original description and genetiva on the plate cap- tion, which was published together with the text; thus, both names are different original spellings. Maynard (1924b?: [7]) listed the taxon as genetiva in his catalog, thus serving as the First Revisor of spell- ings [Article 24.2.4, ICZN, 1999: 30-31] and fixing the spelling as genetiva. Clench (1957: 146) listed this species as genitiva. Taxowms 02. Strophiops marmorosa May- nard and Clapp in Maynard, 1920d [September 30]: 125, pl. 24, figs. 6, 7. Unspecified number of specimens ex- amined. Size given as 1.30 by .47 [inch- es; 33.0 by 11.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the unspecified number of specimens examined, but noted that this taxon was not common. The taxon is listed for sale in Maynard’s (1924b?: [6]) catalog, indicating that more than one specimen was at hand. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on an uncertain number of syntypes. Lot MCZ 76185 contains a single speci- men labeled “Holotype” that approximates 482 the measurements and the illustration for this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining syntypes, in- cluding MCZ 76184, MCZ 118242, and USNM 419944 (25 specimens), become paralectotypes. Lectotype Measurements. Length 31.8 mm, diameter (excluding lip) 12.4 mm; ap- erture height (including lip) 10.6 mm, ap- erture width (including lip and peristome) 8.1 mm. Type Locality. Well Key, a little north of Leward Stocking Key |Exuma Group, Ba- hamas |. Taxon 173. Strophiops pusilla Maynard and Clapp in Maynard, 1920d [Sep- tember 30], pl. 22, fig. 5; Maynard and Clapp in Maynard, 1921a [May 2]: 127. Examined 700 specimens. Size giv- en asim (0by2 30) inches) ives byaMi6 mm], but the term “type” was not used. Nomenclatural Remarks. The captioned illustration (Maynard 1920, pl. 25, fig. 5) was published before the text. The taxon thus dates from the publication of the name in the caption accompanying the il- lustrations, not from the subsequently published text (Article 12.2.7, ICZN, 1999: 17). The date of publication is September 30, 1920. Plate 22 was issued as part of “Part Two” of the Contributions to the History of Cerionidae; therefore, author- ship of this taxon is attributed to Maynard and Clapp. Type Material. Although the figure cap- tion refers only to the single specimen il- lustrated, it is likely that the 700 speci- mens mentioned in the intended descrip- tion of this taxon (Maynard and Clapp in Maynard, 1921la: 127) were available to Maynard at the time and are thus part of the ‘typelsenes |Articles 7214 iGZiNG Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 1999: 76]. Strophiops pusilla is based on 700 syntypes. Lot MCZ 76053 contains a single specimen labeled “Holotype” that approximates the illustration more than the subsequently published measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype. The remaining 699 specimens, including MCZ 76054 and USNM 420133, are paralectotypes. Lectotype Measurements. Length 26.6 mm, diameter (excluding lip) 12.0 mm; ap- erture height (including lip) 9.8 mm, ap- erture width (including lip and peristome) 8.2. mm. Type Locality. Key near Long Rock, Great Exuma [Exuma Group, Bahamas]. Remarks. Maynard (1921a: 127) report- ed that of 700 specimens examined, only 12 were living. Taxon 174. Strophiops inexpecta Maynard and Clapp in Maynard, 1920d [Sep- tember 30], pl. 25, figs. 3, 4; Maynard and Clapp in Maynard, 1921a [May 2]: 127. Unspecified number of specimens examined. Size given as 1.06 by .45 linches; 29.9 by 11.4 mm], but the term “type” was not used. Nomenclatural Remarks. The captioned illustrations (Maynard 1920, pl. 25, figs. 3, 4) were published before the text. The tax- on thus dates from the publication of the name in the caption accompanying the il- lustrations, not from the subsequently published text (Article 12.2.7, ICZN, 1999: 17). The date of publication is September 30, 1920. Plate 25 was issued as part of “Part Two” of the Contributions to the History of Cerionidae; therefore, author- ship of this taxon is attributed to Maynard and Clapp. Type Material. There was no mention of other specimens in the plate caption, and the text portion published 7 months later did not specify the number of specimens that might have been available to Maynard THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. at the time the plate was prepared. The type series of Strophiops inexpecta likely consists of multiple specimens [Article 72.4.1.1, ICZN, 1999: 76], but the number is not known. There was no specimen la- beled “Holotype” at the MCZ. Lot MCZ 76307 contains 9 specimens and was la- beled “Paratypes.” This lot contains a label in Maynard’s hand that does not include the term “type” but does include refer- ences to the figures. Only one of these nine specimens is complete enough to match the illustration, although it does have some minor damage along the colu- mella that is not represented in the figure. This specimen is the figured specimen and is here designated as the lectotype. The remaining specimens were recatalogued as MCZ 357008. These and MCZ 118187 are among, the unspecified number of paralec- totypes. Lectotype Measurements. Length Dll mim, diameter (excluding lip) 12.0 mm; ap- erture height (including lip) 10.1 mm, ap- erture width (including lip and peristome) 5.9 mm. Type Locality. Fossil in soft rock next to westernmost Brigadier Key [Exuma Group, Bahamas]. Taxon 175. Strophiops fitzgeraldi May- nard and Clapp in Maynard, 1920d [September 30], pl. 25, figs. 6, 7; Maynard and Clapp in Maynard, 1921a [May 2]: 127. Unspecified number of specimens examined. Size given as .85 by .37 [inches; 21.6 by 9.4 mm], but the term “type” was not used. Nomenclatural Remarks. The captioned illustrations (Maynard 1920, pl. 25, figs. 6, 7) were published before the text. The tax- on thus dates from the publication of the name in the caption accompanying the il- lustrations, not from the subsequently published text (Article 12.2.7, ICZN, 1999: 17). The date of publication is September 30, 1920. Plate 25 was issued as a part of “Part Two” of the Contributions to the History of Cerionidae; therefore, author- 483 ship of this taxon is attributed to Maynard and Clapp in Maynard. Type Material. There was no mention of other specimens in the plate caption, and the text portion published 7 months later did not specify the number of specimens that might have been available to Maynard at the time the plate was prepared. The type series of Strophiops fitzgeraldi likely consists of multiple specimens [Article 72.4.1.1, ICZN, 1999: 76], but the number is not known. Lot MCZ 76051 contains a single specimen labeled “Holotype” that approximates the illustrations and is within a half millimeter of the subsequently pub- lished measurements of this taxon. This specimen, which is accompanied by a label in Maynard’s hand with the term “type” that includes references to the figures, is here designated as the lectotype. Lectotype Measurements. Length YD, JA mm, diameter (excluding lip) 9.9 mm; ap- erture height (including lip) 8.2 mm, ap- erture width (including lip and peristome) 7. Te ana as Type Locality. North end of Little Ex- uma |[Exuma Group, Bahamas], on the Fitzgerald Estate. Taxon 176. Strophiops adumbra Maynard and Clapp in Maynard, 1920d [Sep- tember 30], pl. 25, figs. 10, 11; May- nard, 1924c [10 December]: 2. Exam- ined 79 specimens. Size given as .75 by .30 [inches; 19.1 by 7.6 mm], but the term “type” was not used. Nomenclatural Remarks. The captioned illustrations (Maynard, 1920d, pl. 25, figs. 10-11) were published before the text. The taxon thus dates from the publication of the name in the caption accompanying the illustrations, not from the subsequent- ly published text (Article 12.2.7, ICZN, 1999: 17). Plate 25 was published in “Part Two” of the Contributions to the History of the Cerionidae; therefore, the author- ship of this taxon is attributed to Maynard and Clapp in Maynard. Type Material. There was no mention of 454 other specimens in the plate caption, but it is likely that the 79 specimens referred to in the text portion published 7 months later were available to Maynard at the time the plate was prepared. The type series of Strophiops adumbra thus consists of 79 specimens [Article 72.4.1.1, ICZN, 1999: 76] that are syntypes. Lot MCZ 76029 contains a single specimen labeled “Ho- lotype” that closely matches the illustra- tions and the subsequently published mea- surements of this taxon. This specimen is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is the figured specimen and is here desig- nated as the lectotype. The remaining 78 specimens are paralectotypes. Lectotype Measurements. Length 19.1 mm, diameter (excluding lip) 7.6 mm; ap- erture height (including lip) 7.2 mm, ap- erture width (including lip and peristome) 5.7 mm. Type Locality. North side of a small key just south of Green Turtle Cut and north of Great Exuma. Remarks. Maynard (1924c: 2) reported that all of the specimens were dead col- lected but that some were fresh. Taxon 177. Strophiops transmutata May- nard and Clapp in Maynard, 1921a [May 2]: 127, pl. 26, figs. 1, 2 [figures labeled S. transimutata]. Examined 150 specimens. Size given as .95 by .35 inches; 24.1 by 8.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 150 speci- mens examined. In the preamble to the taxonomic portion to this publication, Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 150 syntypes. Lot MCZ 76298 contains a single specimen labeled “Ho- lotype” that approximates the illustration and the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 149 syntypes, including MCZ 76297 and MCZ 118183 (one specimen), become paralectotypes. Lectotype Measurements. Length 22.6 mm, diameter (excluding lip) 8.9 mm; ap- erture height (including lip) 8.1 mm, ap- erture width (including lip and peristome) 6.4 mm. Type Locality. Muddy Point Key, Great Exuma [Exuma Group, Bahamas]. Remarks. The taxon name appears as transmutata in the text and transimutata on the plate, both names were published on the same date and are different original spellings. Maynard (1924b?: [6]) listed this taxon as transmutata, acting as the First Revisor of spellings [Article 24.2.4, ICZN, 1999: 30-31]. Maynard (1921a: 127) re- ported that most of the specimens collect- ed were dead. Taxon 178. Strophiops similaria Maynard and Clapp in Maynard, 1921a [May 2]: 128, pl. 25, figs. 5, 6. Examined — Figures 175-186. Taxon 175. Strophiops fitzgeraldi Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76051. North end of Little Exuma, Exuma Group, Bahamas, on the Fitzgerald Estate. Taxon 176. Strophiops adumbra Maynard and Clapp in Maynard, 1920. Lectotype MCZ 76029. North side of a small key just south of Green Turtle Cut and north of Great Exuma. Taxon 177. Strophiops transmutata Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76298. Muddy Point Key, Great Exuma, Exuma Group, Bahamas. Taxon 178. Strophiops similaria Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76055. Westernmost Brigadier Key, Great Exuma, Exuma Group, Bahamas. Taxon 179. Strophiops flamea Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76155. Southwest Point of Great Exuma, Exuma Group, Bahamas. Taxon 180. Strophiops proavita Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76143. Fossil in cliff at north end of Goat Key, Great Harbor, THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 485 Berry Islands, Bahamas. Taxon 181. Strophiops intercalaria Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76160. Fossil in cliff at north end of Goat Key, Great Harbor, Berry Islands, Bahamas. Taxon 182. Strophiops profunda Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76145. Fossil in cliff at north end of Goat Key, Great Harbor, Berry Islands, Bahamas. Taxon 183. Strophiops mixta Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76175. Bonds Key, Berry Islands, Bahamas. Taxon 184. Strophiops reliqua Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76206. Holmes’ Key, Bahamas. Taxon 185. Strophiops aviaria Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76171. Bird Key, Berry Islands, Bahamas. Taxon 186. Strophiops confusa Maynard and Clapp in Maynard, 1921. Lectotype USNM 420009. Little Harbor Key, Bahamas. 486 210 specimens. Original description states “size of last” | = S. fragilis = 1.00 by .40 (inches; 25.4 by 10.2 mm)]. Type Material. The original description did not distinguish among the 210 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 210 syntypes. Lot MCZ 76055 contains a single specimen labeled “Ho- lotype” that approximates the illustration more than the measurements of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 209 syntypes, in- cluding MCZ 76056, become paralectoty- es. fb Lectotype Measurements. Length DDS) mm, diameter (excluding lip) 9.8 mm; ap- erture height (including lip) 9.2 mm, ap- erture width (including lip and peristome) 1) WHIM, Type Locality. Westernmost Brigadier Key, Great Exuma [Exuma Group, Baha- mas]. Remarks. Maynard (1921a: 128) report- ed that most of the specimens collected were dead. Taxon 179. Strophiops flamea Maynard and Clapp in Maynard, 1921a [May 2]: 128, pl. 25, figs. 8, 9. Examined 340 specimens. Size given as 1.00 by .40 inches; 25.4 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 340 speci- mens examined. In the preamble to the taxonomic portion to this publication, Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 340 syntypes. Lot MCZ 76155 contains a single specimen labeled “Ho- lotype” that closely approximates the illus- tration and the measurements of this tax- on. This specimen is accompanied by a la- bel in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. The remain- ing 339 syntypes, including MCZ 76156 and MCZ 118184 (one specimen), become paralectotypes. Lectotype Measurements. Length Weyl mm, diameter (excluding lip) 10.1 mm; ap- erture height (including lip) 9.3 mm, ap- erture width (including lip and peristome) 8.1 mm. Type Locality. Southwest Point of Great Exuma [Exuma Group, Bahamas]. Remarks. Maynard (1921a: 128) report- ed that of 340 specimens, only 38 were living. Taxon 180. Strophiops proavita Maynard and Clapp in Maynard, 1921a [May 2]: 129, pl. 26, figs. 9, 10. Examined 40 specimens. Size given as 1.40 by .55 [inches; 35.6 by 14.0 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 40 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 40 syntypes. Lot MCZ 76143 THE CERION TAXA OF CHARLES JOHNSON MAYNARD ° Harasewych et al. contains a single specimen labeled “Ho- lotype” that does not closely approximate either the illustrations or the measure- ments of this taxon. This specimen is ac- companied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 39 syntypes, including MCZ 76144 and MCZ 118257, become paralec- totypes. Lectotype Measurements. Length OLEO mm, diameter (excluding lip) 12.9 mm; ap- erture height (including lip) 12.1 mm, ap- erture width (including lip and peristome) 10.3 mm. Type Locality. Fossil in cliff at north end of Goat Key, Great Harbor [Berry Islands, Bahamas]. axon 1S 1. Strophiops intercalaria May- nard and Clapp in Maynard, 1921a [May 2]: 129, pl. 26, figs. 7, 9. Ex- amined 60 specimens. Size given as 1.30 by .50 [inches; 33.0 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 60 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 60 syntypes. Lot MCZ 76160 contains a single specimen labeled “Ho- lotype” that approximates the illustration more than the measurements of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that 487 is consistent with Maynard’s original con- cept of it. The remaining 59 syntypes, in- cluding MCZ 76161, become paralectoty- es. , Lectotype Measurements. Length 359.8 mm, diameter (excluding lip) 14.5 mm; ap- erture height (including lip) 13.7 mm, ap- erture width (including lip and peristome) JUL} Taman, Type Locality. Fossil in cliff at north end of Goat Key, Great Harbor [Berry Islands, Bahamas]. Taxon 182. Strophiops profunda Maynard and Clapp in Maynard, 1921a [May 2]: 129, pl. 27, figs. 1, 2. Examined 115 specimens. Size given as 1.15 by .45 linches; 29.2 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 115 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 115 syntypes. Lot number MCZ 76145 was labeled “paratype,” contained two specimens, and was accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. Neither specimen matches closely the published measurements. The speci- men that more closely approximates the il- lustrations is here designated as the lec- totype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original con- cept of it. The other specimen was reca- talogued as MCZ 357009. It is one of the remaining 114 syntypes that become par- alectotypes. Lectotype Measurements. Length 32.0 mim, diameter (excluding lip) 11.7 mm; ap- erture height (including lip) 11.2 mm, ap- erture width (including lip and peristome) 9.4 mm. 488 Type Locality. Fossil in cliff at north end of Goat Key, Great Harbor [Berry Islands, Bahamas]. Taxon 183. Strophiops mixta Maynard and Clapp in Maynard, 1921a [May 2]: 130, pl. 27, figs. 5, 6. Examined 1,005 specimens. Size given as 1.26 by .45 [inches; 32.0 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,005 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,005 syntypes. Lot MCZ 76175 contains a single specimen labeled “Ho- lotype” that closely approximates the mea- surements and illustration of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 1,004 syntypes, including MCZ 76174, MCZ 118215, and USNM 420115 (100 specimens), become paralectotypes. Lectotype Measurements. Length 32.0 mm, diameter (excluding lip) 12.2 mm; ap- erture height (including lip) 11.3 mm, ap- erture width (including lip and peristome) LOOP mim: Type Locality. Bonds Key [Berry Is- lands, Bahamas]. Remarks. Maynard (1921a: 130) report- ed that this taxon was common on palms. Taxon 184. Strophiops reliqua Maynard and Clapp in Maynard, 1921a [May 2]: 130 [as relequa in text], pl. 27, figs. 7, 8 [as S. reliqua on plate]. Examined Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 192 specimens. Size given as 1.16 by .40 inches; 31.8 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 192 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho-- lotype for this taxon, and the species is based on 192 syntypes. Lot MCZ 76206 contains a single specimen labeled “Ho- lotype” that approximates the illustration and the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 191 syntypes, including MCZ 76205, become paralectotypes. Lectotype Measurements. Length 31.4 mm, diameter (excluding lip) 12.0 mm; ap- erture height (including lip) 11.4 mm, ap- erture width (including lip and peristome) 9.4 mm. Type Locality. Holmes’ Key [SW Grand Bahama Island, Bahamas]. Remarks. The taxon name appears as re- lequa in the text and reliqua on the plate, both published on the same date and both different original spellings. Maynard (1924b?: [7]) listed this taxon as S. reliqua, acting as the First Revisor of spellings [Ar- ticle 24.2.4, ICZN, 1999: 30-31]. Clench (1957: 160) listed this taxon as S. relequa. Taxon 185. Strophiops aviaria Maynard and Clapp in.Maynard, 1921a [May 2]: 130, pl. 27, figs. 9, 10. Examined 534 specimens. Size given as 1.16 by .40 [inches; 31.8 by 12.7 mm], but the term “type” was not used. re THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. Type Material. The original description did not distinguish among the 534 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 534 syntypes. Lot MCZ 76171 contains a single specimen labeled “Ho- lotype” that closely approximates the mea- surements and illustration of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 533 syntypes, in- cluding MCZ 118206, MCZ 76170, and USNM 419942, become paralectotypes. Lectotype Measurements. Length 28.2 mm, diameter 12.0 (excluding lip) mm; ap- erture height (including lip) 10.5 mm, ap- erture width (including lip and peristome) 8.6 mm. Type Locality. Bird Key, Berry Islands, Bahamas. Taxon 186. Strophiops confusa Maynard and Clapp in Maynard, 1921a [May 2]: 130, pl. 28, figs. 1, 2, and 136, pl. 34, figs. 1, 2 [see Remarks]. Examined 1,272 specimens. Size given as 1.20 by A5 |inches; 30.5 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,272 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is 489 based on 1,272 syntypes. Lot MCZ 76324 contains three specimens and was labeled “Holotype.” MCZ 118205 contains a single specimen and was labeled “paratype.” None of these closely approximates the measurements or illustration of this taxon. A syntype, USNM 420009, that more closely matches the measurements in the original descriptions and conforms to the first illustration [pl. 28, figs. 1, 2] is here designated as the lectotype to provide an objective standard of reference for this species-group taxon. The remaining 1,271 syntypes, including MCZ 76168, MCZ 76324, MCZ 118205, and USNM 1093790, become paralectotypes. Lectotype Measurements. Length Sly mm, diameter (excluding lip) 11.7 mm; ap- erture height (including lip) 11.3 mm, ap- erture width (including lip and peristome) 8.6 mm. Type Locality. Little Harbor Key | Berry Islands, Bahamas]. Remarks. This taxon appears twice in the same publication: once on page 130 (pl. 28, figs. 1, 2), then again on page 136 (pl. 34, fig. 1, 2) [the second illustration appeared in the following issue dated July 15, 1921]. The descriptions differ (size, number of costae) but have the same type locality and list the same number of spec- imens examined. The figures are similar but do not represent the same specimen. The taxon originates from the first descrip- tion (p. 130, pl. 28, figs. 1, 2). The second appearance of this taxon in the same issue is likely a lapsus on the part of Maynard. It is listed twice in the sales catalog (May- nard, 1924?: [3] nos. 63 [as coufusa] and 94, both from Little Harbor). Taxon 187. Strophiops candida Maynard and Clapp in Maynard, 192la [May 2]: 131, pl. 28, figs. 5, 6. Examined 68 specimens. Size given as 1.25 by 0.50 linches; 31.8 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 68 speci- 490 mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 68 syntypes. Lot number MCZ 76151 contains a single specimen labeled “Holotype” that approximates the illustra- tion and the measurements of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 67 syntypes, in- cluding MCZ 76152 and MCZ 118200, be- come paralectotypes. Lectotype Measurements. Length 30.5 mm, diameter (excluding lip) 12.5 mm; ap- erture height (including lip) 11.6 mm, ap- erture width (including lip and peristome) 10.2 mm. Type Locality. East Marketfish Key [Berry Islands, Bahamas]. Taxon 188. Strophiops primordia May- nard and Clapp in Maynard, 1921a [May 2]: 132, pl. 29, figs. 1, 2. Ex- amined 2,008 specimens. Size given as 1.35 by .50 l[inches; 34.3 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 2,008 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 2,008 syntypes. Lot MCZ 76173 contains 179 syntypes [labeled “para- types” |, including a segregated specimen that approximates the figures and mea- surements listed in the original descrip- tion. The segregated specimen is here se- lected as the lectotype to provide an ob- jective standard of reference for this spe- cies-group taxon that is consistent with Maynard’s concept of this taxon. The re- maining 178 specimens were recatalogued as MCZ 357010 and are among the 2,007 syntypes, including USNM 420126 (150 specimens), that become paralectotypes. Lectotype Measurements. Length Sond mm, diameter (excluding lip) 12.9 mm; ap- erture height (including lip) 11.9 mm, ap- erture width (including lip and peristome) 10.2 mm. Type Locality. Great Srirrup Key [Ba- hamas]. Taxon 189. Strophiops balaena Maynard and Clapp in Maynard, 1921a [May 2]: 132 [as balaene], pl. 29, figs. 3, 4 [as S. balaena|. Examined 765 speci- mens. Size given as 1.20 by .40 [inches; 30.5 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 765 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's => Figures 187-198. Taxon 187. Strophiops candida Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76151. East Marketfish Key, Bahamas. Taxon 188. Strophiops primordia Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76173. Great Srirrup Key, Bahamas. Taxon 189. Strophiops balaena Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76190. Whale Key, Bahamas. Taxon 190. Strophiops porcina Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76193 Frazar’s Hog Key, Bahamas. Taxon 191. Strophiops obtusa Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76213. Cat Key, Berry Islands, Bahamas. Taxon 192. Strophiops albata Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76256. Hog Cay, 4 miles NW of Morgan’s Bluff, Andros, Bahamas. Taxon 193. Strophiops litorea Maynard and Clapp in Maynard, 1921. Lectotype USNM THE CERION TAXA OF CHARLES JOHNSON MAYNARD *¢ Harasewych et al. 491 oy iA AYA \, ot 420058. South border of Guana Key, Berry Islands, Bahamas. Taxon 194. Strophiops rara Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76023. West side of Cabbage Key, Bahamas. Taxon 195. Strophiops scutata Maynard and Clapp in Maynard, 1921. Lectotype USNM 420100. Petit Key, Berry Islands, Bahamas. Taxon 196. Strophiops procliva Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76210. Goat Key, Great Harbor, Berry Islands, Bahamas. Taxon 197. Strophiops jenneyi Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76012. Anderson’s Key, Berry Islands, Bahamas. Taxon 198. Strophiops travellii Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76363. Bridgewater Key, Berry Islands, Bahamas. 492 publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 765 syntypes. Lot number MCZ 76190 contains a single specimen labeled “Holotype” that approximates the illustra- tion more than the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 764 syntypes, including MCZ 76191, MCZ 118213, and USNM 419943 (200 specimens), become paralectotypes. Lectotype Measurements. Length 30.2 mm, diameter (excluding lip) 11.38 mm; aperture height (including lip) 11.4 mm, aperture width (including lip and _ peri- stome) 9.1 mm. Type Locality. Whale Key [Berry Is- lands, Bahamas]. Remarks. The name of this taxon is spelled as balaene in the original descrip- tion, balaena in the plate caption, and bal- anae in Maynard's (1924b?: [4]) sales cat- alog. The spellings balaene and balaena are different original spellings, whereas balanae is a subsequent lapsus calami. Clench (1957: 138) listed this taxon as ba- laena but cannot be considered the First Revisor [Article 24.2.3, ICZN, 1999: 30] because he did not list both original spell- ings. Strophiops balaena is here formally selected as the correct original spelling. Taxon 190. Strophiops porcina Maynard and Clapp in Maynard, 1921a [May 2]: 132, pl. 29, figs. 7, 8. Examined 2,305 specimens. Size given as 1.15 by A5 [inches; 29.2 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 2,305 spec- imens examined. In the preamble to the taxonomic portion to this publication, Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 2,305 syntypes. Lot number MCZ 76193 contains a single specimen la- beled “Holotype” that approximates the il- lustration more than the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 2,304 syn- types, including MCZ 76192 and USNM 420012 (150 specimens), become paralec- totypes. Lectotype Measurements. Length 29.3 mm, diameter (excluding lip) 11.9 mm; ap- erture height (including lip) 11.4 mm, ap- erture width (including lip and peristome) 9.5 mm. Type Locality. Frazar’s Hog Key |Berry Islands, Bahamas]. Taxon 191. Strophiops obtusa Maynard and Clapp in Maynard, 1921a [May 2]: 132, pl. 29, figs. 9, 10. Examined 1,500 specimens. Size given as 1.10 by 45 [inches; 27.9 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,500 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,500 syntypes. Lot MCZ 76213 contains a single specimen labeled “Ho- lotype” that approximates the illustration more than the measurements of this taxon THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 1,499 syntypes, including MCZ 76212, MCZ 118216, and USNM 420102 (500 specimens), become paralectotypes. Lectotype Measurements. Length 28.1 mim, diameter (excluding lip) 11.2 mm; ap- erture height (including lip) 10.5 mm, ap- erture width (including lip and peristome) 9.1 mm. Type Locality. Cat Key, Berry Islands, Bahamas. Taxon 192. Strophiops albata Maynard and Clapp in Maynard, 1921a [May 2]: 132-133, pl. 30, figs. 3, 4. Exam- ined 60 specimens. Size given as 1.08 by 45 [inches; 30.5 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 60 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 60 syntypes. Lot number MCZ 76256 contains a single specimen labeled “Holotype” that approximates the illustra- tion and less so the measurements of this taxon. It is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original con- cept of it. The remaining 59 syntypes, in- cluding MCZ 76257, become paralecto- types. 493 Lectotype Measurements. Length 27.8 mm, diameter (excluding lip) 11.1 mm; ap- erture height (including lip) 10.8 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. “Hog Key near Morgan's Bluff.” Clench (1957: 136) clarified the type locality as Hog Cay [4 miles NW of] Morgan’s Bluff |Andros, Bahamas]. Remarks. See also Strophiops albata Maynard and Clapp in Maynard, 1921b [Taxon 215], and Strophiops vagabunda Maynard and Clapp in Maynard, 1925 [Taxon 248]. Taxon 193. Strophiops litorea Maynard and Clapp in Maynard, 1921a [May 2]: 133, pl. 30, figs. 7, 8. Examined 300 specimens. Measurements not pro- vided. Type Material. The original description did not distinguish among the 500 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 500 syntypes. Lot number MCZ 76370 was labeled “holotype,” but contains two specimens, neither of which closely match the published figure. A lectotype, USNM 420058, that more closely approx- imates the figure is here selected from a large syntype lot at USNM to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 499 syntypes, including MCZ 76369, MCZ 76370, and USNM 1093791, become paralectotypes. Lectotype Measurements. Length 34.0 mm, diameter (excluding lip) 12.4 mm; ap- erture height (including lip) 11.1 mm, ap- erture width (including lip and peristome) 9.9 mm. Type Locality. South border of Guana Key, Berry Islands, Bahamas. 494 Taxon 194. Strophiops rara Maynard and Clapp in Maynard, 1921a [May 2]: 133, pl. 31, figs. 1, 2. Examined 10 specimens. Size given as 2.25 [sic, 1.25] by .52 [inches; 31.8 by 13.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 10 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 10 syntypes. Lot MCZ 76023 contains a single specimen labeled “Ho- lotype” that approximates the illustration and the inferred measurements (see Re- marks) of this taxon and is accompanied by a label in Maynard's hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. The remain- ing nine syntypes, including MCZ 76402 (one specimen), become paralectotypes. Lectotype Measurements. Length 31.6 mm, diameter (excluding lip) 13.5 mm; ap- erture height (including lip) 11.6 mm, ap- erture width (including lip and peristome) 10.6 mm. Type Locality. West side of Cabbage Key [Berry Islands], Bahamas. Remarks. Fossil, embedded in rock. The length measurement provided, 2.25 inch- es, is likely a typographical error. The il- lustrated lectotype corresponds to a mea- surement of 1.25 inches. Taxon 195. Strophiops scutata Maynard and Clapp in Maynard, 1921a [May 2|: 133-134, pl. 31, figs. 3, 4. Exam- ined 1,690 specimens. Size given as larger, 1.17 by .50 [inches; 30.5 by 12.7 mm], but the term “type” was not used. Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Type Material. The original description did not distinguish among the 1,690 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,690 syntypes. Of the specimens at the MCZ, none were found labeled “ho- lotype,” and only a single specimen (MCZ 118208) labeled “paratype” was located. A lectotype, USNM 420100, selected to most closely approximate the illustration and published measurements of this taxon, is here designated to provide an objective standard of reference for this species- group. The remaining 1,689 syntypes, in- cluding MCZ 76211, MCZ 118208, and USNM 1093793, become paralectotypes. Lectotype Measurements. Length 29.4 mim, diameter (excluding lip) 12.7 mm; ap- erture height (including lip) 10.8 mm, ap- erture width (including lip and peristome) 9.5 mm. Type Locality. Petit Key | Berry Islands], Bahamas. Taxon 196. Strophiops procliva Maynard and Clapp in Maynard, 1921a [May 2]: 134, pl. 31, figs. 5, 6. Examined 145 specimens. Size given as 1.10 by .45 [inches; 27.9 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 145 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 145 syntypes. Lot number MCZ 76210 contains a single specimen labeled “Holotype” that approximates the illustra- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. tion and the measurements of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 144 syntypes, in- cluding MCZ 76209, become paralecto- types. Lectotype Measurements. Length 27.6 mm, diameter (excluding lip) 10.9 mm; ap- erture height (including lip) 9.5 mm, ap- erture width (including lip and peristome) 7.9 mm. Type Locality. Goat Key, Great Harbor, Berry Islands, Bahamas. Taxon 197. Strophiops jenneyi Maynard and Clapp in Maynard, 1921a |May 2]: 134, pl. 31, figs. 9, 10. Examined 1,131 specimens. Size given as 1.15 by 40 [inches; 30.5 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,131 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,131 syntypes. Lot number MCZ 76012 contains a single specimen la- beled “Holotype” that approximates the il- lustrations and measurements of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original con- cept of it. The remaining 1,130 syntypes, including MCZ 76103 and USNM 419985 (75 specimens), become paralectotypes. 495 Lectotype Measurements. Length 31:1 mm, diameter (excluding lip) 11.6 mm; ap- erture height (including lip) 11.1 mm, ap- erture width (including lip and peristome) 9.3 mm. Type Locality. Anderson’s Key, Berry Is- lands, Bahamas. Taxon 198. Strophiops travellii Maynard and Clapp in Maynard, 1921a [May 2]: 135 [as travelii|, pl. 32, figs. 3, 4 [as travellii]. Examined 3,416 speci- mens. Size given as 1.35 by .45 [inches; 34.3 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 3,416 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 3,416 syntypes. Lot number MCZ 76363 contains a single specimen la- beled “Holotype” that approximates the il- lustration and the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 3,416 syn- types, including MCZ 76364, MCZ 118198, and USNM 419946 (200 speci- mens), become paralectotypes. Lectotype Measurements. Length 33.9 mm, diameter (excluding lip) 11.9 mm; ap- erture height (including lip) 11.3 mm, ap- erture width (including lip and peristome) 9.1 mm. Type Locality. Bridgewater Key, Berry Islands, Bahamas. Remarks. The taxon name appeared as S. travelii in the heading of the original 496 description and S. travellii in the figure caption. By listing this taxon as S. travellii in the sales catalog, Maynard (1924bp: [3]) acted as the First Revisor of spellings [Ar- ticle 24.2.4, ICZN, 1999: 30-31], fixing the spelling as travellii. Clench (1957: 165) listed this taxon as S. travelii in his catalog. Taxon 199. Strophiops picturata Maynard and Clapp in Maynard, 1921a [May 2]: 135, pl. 32, figs. 7, 8. Examined eight specimens. Size given as 1.25 by 59 [inches; 31.8 by 14.0 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the eight spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on eight syntypes. Lot number MCZ 76019 contains a single specimen labeled “Holotype” that approximates the illustra- tion and the measurements of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining seven syntypes become paralectotypes. Lectotype Measurements. Length 31.9 mm, diameter (excluding lip) 13.6 mm; ap- erture height (including lip) 11.4 mm, ap- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 erture width (including lip and peristome) 9.8 mm. Type Locality. Cliffs on Cabbage Key, Berry Islands, Bahamas. Taxon 200. Strophiops lenticularia May- nard and Clapp in Maynard, 1921la [May 2]: 135, pl. 32, figs. 9, 10. Ex- amined 325 specimens. Size given as 1.19 by .40 [inches; 30.2 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 325 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 325 syntypes. Lot number MCZ 76362 contains a single specimen labeled “Holotype” that closely approximates the illustration and less so the measurements of this taxon. It is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 324 syntypes, including MCZ 76361, become paralecto- types. Lectotype Measurements. Length 28.0 mm, diameter (excluding lip) 10.6 mm; ap- erture height (including lip) 9.7 mm, ap- erture width (including lip and peristome) 8.3 mm. —-> Figures 199-210. Taxon 199. Strophiops picturata Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76019. Cliffs on Cabbage Key, Berry Islands, Bahamas. Taxon 200. Strophiops lenticularia Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76362. Staniard Creek, Andros, Bahamas. Taxon 201. Strophiops stupida Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76281. North Key, Staniard Creek, Andros, Bahamas. Taxon 202. Strophiops columbiana Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76339. Pigeon Key, Staniard Creek, Andros, Bahamas. Taxon 203. Sfrophiops carnale Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76299. West of Morgan’s Bluff, Staniard Creek, Andros, Bahamas. Taxon 204. Stro- phiops panda Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76269. Saddle-Back Key, Andros, Bahamas. Taxon 205. THE CERION TAXA OF CHARLES JOHNSON MAYNARD ° Harasewych et al. 497 Strophiops crescentia Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76385. Calabash Key, Andros, Bahamas. Taxon 206. Sirophiops thayeri Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76367. East end of Thompson’s Key, Berry Islands, Bahamas. Taxon 207. Strophiops sylvatica Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76371. Chub Point Key, Berry Islands, Bahamas. Taxon 208. Strophiops ralla Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76301. Joulter Keys, Andros, Bahamas. Taxon 209. Strophiops rosacea Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76246. West Silver Key, Nassau, New Providence Island, Bahamas. Taxon 210. Strophiops angustocostata Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76295. Fossil on Lower Fleming, Eleuthera, Bahamas. 498 Type Locality. Staniard Creek, Andros, Bahamas. Taxon 201. Strophiops stupida Maynard and Clapp in Maynard, 1921a [May 2]: 135, pl. 33, figs. 1, 2. Examined 266 specimens. Size given as 1.00 by .45 [inches; 25.4 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 266 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 266 syntypes. Lot MCZ 76281 contained 45 specimens and was labeled “paratype.” The only slip of paper in May- nard’s handwriting gave the locality data but made no mention of types. Of these, a single specimen that most closely ap- proximated the published illustrations and measurements is here designated as the lectotype (MCZ 76281) to provide an ob- jective standard of reference for this spe- cies-group taxon that is consistent with Maynard's original concept of it. The re- maining 44 specimens were recatalogued as MCZ 357011 and are among the 265 syntypes that become paralectotypes. Lectotype Measurements. Length 24.9 mim, diameter (excluding lip) 10.7 mm; ap- erture height (including lip) 9.3 mm, ap- erture width (including lip and peristome) 8.1 mm. Type Locality. North Key, Staniard Creek, Andros, Bahamas. Taxon 202. Strophiops columbiana May- nard and Clapp in Maynard, 1921a [May 2]: 136, pl. 33, figs. 3, 4. Ex- amined 900 specimens. Size given as .92 by .40 [inches; 23.4 by 10.2 mm], but the term “type” was not used. Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Type Material. The original description did not distinguish among the 900 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 900 syntypes. Lot MCZ 76339 contains a single specimen labeled “Ho- lotype” that approximates the illustration and the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 899 syntypes, including MCZ 76338 and USNM 420054 (75 specimens), become paralectotypes. Lectotype Measurements. Length DSO) mm, diameter (excluding lip) 10.2 mm; ap- erture height (including lip) 9.0 mm, ap- erture width (including lip and peristome) TS) vaqany, Type Locality. Pigeon Key, Staniard Creek, Andros, Bahamas. Taxon 203. Strophiops carnale Maynard and Clapp in Maynard, 1921a [May 2]: 136, pl. 33, figs. 5, 6. Examined 266 specimens. Size given as .90 by OO inches; 22.9 by 8.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 266 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 266 syntypes. Lot number MCZ THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 76299 contains a single specimen labeled “Holotype” that approximates the illustra- tion and the measurements of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original con- cept of it. The remaining 265 syntypes, in- cluding MCZ 76300, become paralecto- types. Lectotype Measurements. Length LED) mm, diameter (excluding lip) 8.9 mm; ap- erture height (including lip) 8.8 mm, ap- erture width (including lip and peristome) 7. mm. Type Locality. West of Morgan's Bluff, Staniard Creek, Andros, Bahamas. Remarks. Clench (1957: 140) lists the type locality as 1 mile west of Morgan’s Bluff. Taxon 204. Strophiops panda Maynard and Clapp in Maynard, 1921a [May 2|: 136, pl. 33, figs. 7, 8. Examined 72 specimens. Size given as .98 by .45 inches; 24.9 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 72 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 72 syntypes. Lot number MCZ 76269 contains a single specimen labeled “Holotype” that approximates the illustra- tion and the measurements of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of 499 reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 71 syntypes, in- cluding MCZ 76268, become paralecto- types. Lectotype Measurements. Length 26.0 mim, diameter (excluding lip) 11.2 mm; ap- erture height (including lip) 9.8 mm, ap- erture width (including lip and peristome) CO) OMINN, Type Locality. Saddle-Back Key, An- dros, Bahamas. Taxon 205. Strophiops crescentia May- nard and Clapp in Maynard, 1921la [May 2]: 136, pl. 33, figs. 9, 10. Ex- amined 240 specimens. Size given as .95 by .40 [inches; 24.1 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 240 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 240 syntypes. Lot number MCZ 76385 contains a single specimen labeled “Holotype” that approximates the illustra- tion and the measurements of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original con- cept of it. The remaining 239 syntypes, in- cluding MCZ 76260, become paralecto- types. Lectotype Measurements. Length DAS mim, diameter (excluding lip) 10.2 mm; ap- erture height (including lip) 9.4 mm, ap- erture width (including lip and peristome) 7.9 mm. Type Locality. Calabash Key, Andros, Bahamas. 500 Taxon 206. Strophiops thayeri Maynard and Clapp in Maynard, 1921a [May 2]: 137; Maynard, 1921b [July 14], pl. 34, figs. 5, 6 [as thayerii in figure cap- tion]. Examined 2,876 specimens. Size given as 1.35 by .40 [inches; 34.3 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 2,876 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 2,876 syntypes. Lot MCZ 76367 contains a single specimen labeled “Ho- lotype” that approximates the illustration and the measurements of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 2,875 syntypes, including MCZ 76368, USNM 420010 (50 specimens), and USNM 420092 (400 specimens), be- come paralectotypes. Lectotype Measurements. Length 30.0 mm, diameter (excluding lip) 9.8 mm; ap- erture height (including lip) 9.7 mm, ap- erture width (including lip and peristome) 8.1 mm. Type Locality. East end of Thompson’s Key, Berry Islands, Bahamas. Remarks. The taxon name appeared as S. thayeri in the heading of the original description and S. thayerii in the figure captions, which were published in a later issue than the text. This taxon was listed as S. thayerii in the sales catalog (May- nard, 1924b?: [3]). Clench (1957: 165) list- ed this taxon as S. thayeri in his catalog. Lot MCZ 118209 has a single specimen labeled “Paratype,” from Cabbage Key, Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Berry Islands. This specimen was not the Same species, nor is there any mention of Cabbage Key in the description of Cerion thayeri. This specimen is likely mislabeled and/or miscatalogued. Taxon 207. Strophiops sylvatica Maynard and Clapp in Maynard, 1921a [May 2]: 137; Maynard, 1921b [July 15], pl. 34, figs. 7, 8. Examined 1,500 speci- mens. Size given as 1.12 by .47 [inches; 28.4 by 11.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,500 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,500 syntypes. Lot MCZ 76371 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and the subsequently published il- lustration and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 1,499 syn- types, including MCZ 76372, MCZ 118119, and USNM 420004 (100 speci- mens), become paralectotypes. Lectotype Measurements. Length 27.4 mm, diameter (excluding lip) 10.7 mm; ap- erture height (including lip) 9.8 mm, ap- erture width (including lip and peristome) 8.8 mm. Type Locality. Chub Point Key, Berry Islands, Bahamas. Taxon 208. Strophiops ralla Maynard and Clapp in Maynard, 1921la [May 2]: 137; Maynard, 1921b [July 15], pl. 34, figs. 8, 9. Examined 65 specimens. Size THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. given as 1.20 by .47 [inches; 30.5 by 11.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 65 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 65 syntypes. Lot MCZ 76301 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and subsequently published illustra- tions of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to these figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. The remain- ing 64 syntypes, including MCZ 76302 and MCZ 118356, become paralectotypes. Lectotype Measurements. Length 29.4 mm, diameter (excluding lip) 11.9 mm; ap- erture height (including lip) 10.8 mm, ap- erture width (including lip and peristome) So Wit. Type Locality. Joulter Keys, Andros, Ba- hamas. Taxon 209. Strophiops rosacea Maynard and Clapp in Maynard, 1921b [July 15]: 139, pl. 35, figs. 7, 8. Examined 1,759 specimens. No measurements provided. Type Material. The original description did not distinguish among the 1,759 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- 501 lotype for this taxon, and the species is based on 1,759 syntypes. No specimen la- beled “Holotype” was located at the MCZ, nor had any been catalogued as such in the material from Maynard's collection. MCZ 76246 contains 159 paratypes, of which one was segregated in a vial and la- beled “figured.” This specimen approxi- mates the published figure and measure- ments and is here designated as the lec- totype to provide an objective standard of reference for this species-group taxon. The remaining 158 specimens were recata- logued as MCZ 356678, and, together with MCZ 76246, USNM 419966 (100 speci- mens), and USNM 420026 (100 speci- mens), are among the 1,758 syntypes that become paralectotypes. Lectotype Measurements. Length 29:1 mm, diameter (excluding lip) 11.7 mm; ap- erture height (including lip) 10.0 mm, ap- erture width (including lip and peristome) 9.1 mm. Type Locality. West Silver Key, Nassau, New Providence Island, Bahamas. Taxon 210. Strophiops angustocostata Maynard and Clapp in Maynard, 1921b [July 15]: 141, pl. 37, figs. 7, 8. Examined eight specimens. Size giv- en as 1.05 by .55 [inches; 26.7 by 14.0 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the eight spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on eight syntypes. Lot MCZ 76295 contains a single specimen labeled “Para- type” that approximates the length mea- surement and illustrations of this taxon and is accompanied by a label in May- nard’s hand that includes references to the figures but does not contain the term 502 “type.” This specimen is here designated as the lectotype to provide an objective standard of reference for this species- group taxon that is consistent with May- nard’s original concept of it. The remain- ing seven syntypes become paralectotypes. Lectotype Measurements. Length 26.5 mm, diameter (excluding lip) 11.6 mm; ap- erture height (including lip) 10.5 mm, ap- erture width (including lip and peristome) 9.5 mm. Type Locality. Fossil on Lower Fleming [Eleuthera, Bahamas]. Remarks. Clench (1952: 109) listed this taxon as a synonym of Cerion exiguum (Maynard, 1913). Taxon 211. Strophiops crassalabra May- nard and Clapp in Maynard, 1921b [July 15]: 143; Maynard, 1924a |Octo- ber 25], pl. 39, figs. 7, 8 [figures pub- lished with the label S. crassamarga}. Examined 300 specimens. Size given as 1.15 by .40 linches; 29.2 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 300 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 300 syntypes. Lot MCZ 76359 contains a single specimen labeled “Ho- lotype” that approximates the measure- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 ments and later illustrations of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 299 syntypes, in- cluding MCZ 76360 and MCZ 118274, be- come paralectotypes. Lectotype Measurements. Length 28.7 mm, diameter (excluding lip) 11.3 mm; ap-- erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) 8.1 mm. Type Locality. Cliffs on east side of Rose Island, opposite Low Bay Key, New Prov- idence Island, Bahamas. Remarks. Misspelled as S. crassilabra in the sales catalog (Maynard, 1924c?: [6]). See also Strophiops crassamarga |Taxon 233]. Gould and Woodruff (1986: 481) re- ported that they had seen the “Holotype” and synonymized Cerion crassalabra with C. glans (Kiister, 1844). Taxon 212. Strophiops angustalabra May- nard and Clapp in Maynard, 1921b [July 15]: 143-144; Maynard, 1924a [October 25], pl. 39, figs. 9, 10. Exam- ined 500 specimens. Size given as .95 by 45 [inches; 24.1 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 500 speci- mens examined. In the preamble to the taxonomic portion to this publication, —- Figures 211-222. Taxon 211. Strophiops crassalabra Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76359. Cliffs on east side of Rose Island, opposite Low Bay Key, New Providence Island, Bahamas. Taxon 212. Strophiops angustalabra May- nard and Clapp in Maynard, 1921. Lectotype MCZ 76329. Low cliffs on west side of Rose Island, opposite Green Key, New Providence Island, Bahamas. Taxon 213. Strophiops palidula Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76274. Extreme east end of Hog Island [Paradise Island], New Providence Island, Bahamas. Taxon 214. Strophiops saxitina Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76326. Hog Island [Paradise Island], east of Three Bays on rock of shore, New Providence Island, Bahamas. Taxon 215. Strophiops albata Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76258. Southern end of Rose Island, New Providence Island, Bahamas. Strophiops vagabunda Maynard and Clapp in Maynard, 1925 [Taxon 248], is a new name for S. albata Maynard and Clapp in Maynard, 1921, not S. albata Maynard and Clapp ijn Maynard, 1921 [Taxon 192]. Taxon 216. Strophiops mobile Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76309. Extreme west end of Rose Island, New Providence Island, Bahamas. Taxon 217. Strophiops hartbennetii Maynard and Clapp in Maynard, THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. 503 1921. Lectotype MCZ 76001. Poters Key, New Providence Island, Bahamas. Taxon 218. Strophiops oscula Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76333. Old Thompson Place, off East Bay Street, Nassau, New Providence Island, Bahamas. Taxon 219. Strophiops eratica Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76387. Near Fox Hill Village, New Prov- idence Island, Bahamas. Taxon 220. Strophiops castra Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76388. Field west of Williams Street, Nassau, New Providence Island, Bahamas. Taxon 221. Strophiops rubiginosa Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76006. Field east of Methodist Church on Sherley Street, Nassau, New Providence Island, Bahamas. Taxon 222. Strophiops migratoria Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76312. Grounds directly about Methodist Sunday School building, Sherley Street, Nassau, New Providence Island, Bahamas. 504 Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 500 syntypes. Lot number MCZ 76329 contains a single specimen labeled “Holotype” that approximates the mea- surements and subsequently published il- lustrations of this taxon and is accompa- nied by a label in Maynard’s hand with the term “type,” which includes references to these figures. This specimen is here des- ignated as the lectotype to provide an ob- jective standard of reference for this spe- cies-group taxon that is consistent with Maynard's original concept of it. The re- maining 499 syntypes, including MCZ 76328 and MCZ 118269, become paralec- totypes. Lectotype Measurements. Length DAL? mm, diameter (excluding lip) 11.0 mm; ap- erture height (including lip) 10.0 mm, ap- erture width (including lip and peristome) 8.1 mm. Type Locality. Low cliffs on west side of Rose Island, opposite Green Key, New Providence Island, Bahamas. Remarks. Gould and Woodruff (1986: 481) reported that they had seen the “Ho- lotype” and synonymized Cerion angustal- abra with C. glans (Kiister, 1844). Taxon 213. Strophiops palidula Maynard and Clapp in Maynard, 1921b [July 15]: 145; Maynard, 1925 [July 18], pl. 40, figs. 9, 10. Examined 1,054 speci- mens. Size given as 1.10 by .40 [inches; 27.9 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,054 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,054 syntypes. Lot number MCZ 76274 contains a single specimen la- beled “Holotype” that approximates the measurements and later illustrations of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 1,053 syn-— types, including MCZ 76275 and USNM 419991 (250 specimens), become paralec- totypes. Lectotype Measurements. Length 27.8 mm, diameter (excluding lip) 10.7 mm; ap- erture height (including lip) 9.6 mm, ap- erture width (including lip and peristome) 7.4 mm. Type Locality. Extreme east end of Hog Island [Paradise Island, New Providence Island, Bahamas]. Remarks. Gould and Woodruff (1986: 475) reported that they had seen the “Ho- lotype.” These authors synonymized Stro- phiops palidula with Cerion glans (Kiister, 1844). Taxon 214. Strophiops saxitina Maynard and Clapp in Maynard, 1921b [July 15]: 145; 1925 [July 18], pl. 41, figs. 1, 2. Examined 100 specimens. Size given as .76 by .35 [inches; 19.3 by 8.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 100 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 100 syntypes. Lot number MCZ 76326 was labeled “Holotype” but con- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. tains two specimens and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. The specimen that more closely approximates the measurements and later illustrations of this taxon is here designat- ed as the lectotype (MCZ 76326) to pro- vide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The other specimen was recatalogued as MCZ 357012. It and MCZ 76327 are among the remaining 99 syntypes that become para- lectotypes. Lectotype Measurements. Length 19.4 mm, diameter (excluding lip) 9.8 mm; ap- erture height (including lip) 7.6 mm, ap- erture width (including lip and peristome) 6.3 mm. Type Locality. Hog Island, east of Three Bays on rock of shore, New Providence Is- land, Bahamas. Remarks. The plate containing the illus- trations of this taxon was issued 4 years after the publication of the description. Gould and Woodruff (1986: 481) consid- ered this taxon to be a synonym of Cerion glans (Kiister, 1844). Taxon 215. Strophiops albata Maynard and Clapp in Maynard, 1921b [July 15]: 145; Maynard and Clapp in May- nard, 1925 [July 18], pl. 41, figs. 7, 8 [the figures are labeled S. vagabunda] [junior primary homonym of S. albata Maynard and Clapp in Maynard, 1921a; S. vagabunda Maynard and Clapp in Maynard, 1925 (Taxon 248), is the next available name]. Examined 72. speci- mens. Size given as 1.10 by .45 [inches; 27.9 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 72 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's 505 publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 72 syntypes. Lot MCZ 76258 contains a single specimen accompanied by two MCZ labels that refer to this spec- imen as the holotype of Strophiops vaga- bunda, a replacement name for the pre- occupied S. albata (see Remarks). A label in Maynard’s hand contains the name S. vagabunda and the term “type” and in- cludes references to the figures. This spec- imen is here designated as the lectotype of S. albata Maynard and Clapp in Maynard, 1921b, and also [Article 72: 7, ICZN, 1999: 78] its replacement name Strophiops vagabunda Maynard and Clapp in May- nard, 1925, to provide an objective stan- dard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The remaining 71 syntypes, including MCZ 76256 and MCZ 76257, become paralectotypes of S. albata Maynard and Clapp in Maynard, 1921b, and consequently [Article 72: 7, ICZN, 1999: 78] its replacement name Strophiops vagabunda Maynard and Clapp in May- nard, 1925. Lectotype Measurements. Length 28.0 mm, diameter (excluding lip) 11.2 mm; ap- erture height (including lip) 10.3 mm, ap- erture width (including lip and peristome) 8.8 mm. Type Locality. Southern end of Rose Is- land, New Providence Island, Bahamas. Remarks. The binomen Strophiops al- bata had been previously proposed by Maynard and Clapp in Maynard, 1921 ([May 2]: 132-133, pl. 30, figs. 3, 4) (see Taxon 192). The second description of S. albata cites different measurements, a dif- ferent number of specimens examined, and a different type locality and refers to an illustration of a different specimen in- tended to serve as the type. Strophiops al- bata Maynard and Clapp in Maynard, 1921b, is a junior primary homonym of Strophiops albata Maynard and Clapp in Maynard, 192la. Maynard (1925, pl. 41, figs. 7, 8) apparently recognized the hom- 506 onymy and captioned the subsequently published illustrations S. vagabunda. As noted by Clench (1957: 166), Strophiops vagabunda Maynard, 1925, is a next avail- able name for S. albata Maynard and Clapp in Maynard, 1921b, not S. albata Maynard and Clapp in Maynard, 1921a. Gould and Woodruff (1986: 474) errone- ously reported S. albata Maynard and Clapp in Maynard, 1921b, to be a nomen nudum and considered S. vagabunda to be a synonym of Cerion glans (Kiister, 1844). Taxon 216. Strophiops mobile Maynard and Clapp in Maynard, 1921b [July 15]: 146; Maynard and Clapp in May- nard, 1925 [July 18], pl. 41, figs. 9, 10. Examined 61 specimens. Size given as 1.00 by .40 linches; 25.4 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 61 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 61 syntypes. Lot number MCZ 76309 contains a single specimen labeled “Holotype” that approximates the mea- surements and later illustrations of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 60 syntypes, including MCZ 76310, become paralecto- types. Lectotype Measurements. Length 25.8 mim, diameter (excluding lip) 10.6 mm; ap- erture height (including lip) 9.7 mm, ap- erture width (including lip and peristome) 7.8 mm. Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Type Locality. Extreme west end of Rose Island, New Providence Island, Ba- hamas. Remarks. The figures cited in the orig- inal description were issued 4 years later. Taxon 217. Strophiops hart-bennetii May- nard, 1919b |November 6]: 36 [nomen nudum]. Strophiops hartbennetii May- nard and Clapp in Maynard, 1921b [July 15]: 146 [as hart-bennetii—see Remarks]; Maynard and Clapp in May- nard, 1926 |March 24], pl. 42, figs. 3, 4 _ [as hart-benettii]. Examined 500 speci- mens. Size given as .80 by .32 [inches; 20.3 by 8.1 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 500 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 500 syntypes. Lot number MCZ 76001 contains a single specimen labeled “Holotype” that approximates the mea- surements and later illustrations of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 499 syntypes, including MCZ 76002, MCZ 118271, and USNM 420022 (50 specimens), become paralectotypes. Lectotype Measurements. Length 20.9 mm, diameter (excluding lip) 8.4 mm; ap- erture height (including lip) 7.4 mm, ap- erture width (including lip and peristome) 5.7 mm. Type Locality. Poters Key |New Provi- dence Island, Bahamas]. THE CERION TAXA OF CHARLES JOHNSON MAYNARD ¢ Harasewych et al. Remarks. The binomen first appeared as a nomen nudum (Maynard, 1919b: 36). The taxon name is spelled hart-bennetii in the original description, on the handwrit- ten label, and in the sales catalog (May- nard, 1924?: [6]). The plate, with the spell- ing hart-benettii, was issued 6 years later and is considered a subsequent misspell- ing. According to Article 32.5.2.4 (ICZN, 1999: 40), the taxon name is hartbennetii because hyphens are to be removed. Taxon 218. Strophiops oscula Maynard and Clapp in Maynard, 1921b [July 15]: 146; Maynard and Clapp in May- nard, 1926 [March 24], pl. 42, figs. 5, 6. Examined 442 specimens. Size given as 95 by 35 [sic = .95 by .35] [inches; 24.1 by 8.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 442 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 442 syntypes. Lot MCZ 76333 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and later illustrations of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 44] syntypes, in- cluding MCZ 76332 and USNM 420116 (100 specimens), become paralectotypes. Lectotype Measurements. Length 23.8 mm, diameter (excluding lip) 10.0 mm; ap- erture height (including lip) 8.3 mm, ap- erture width (including lip and peristome) 7.1 mm. 507 Type Locality. Old Thompson Place off East Bay Street, Nassau [ New Providence Island, Bahamas]. Remarks. The illustrations were issued nearly 5 years after the original description was published. Gould and Woodruff (1986: 476) regarded this taxon to be “in- termediate” between Cerion glans (Kiister, SAAD) anid G- gubernatorium (Crosse, 1869). Taxon 219. Strophiops eratica Maynard and Clapp in Maynard, 1921b [July 15]: 147; Maynard and Clapp in May- nard, 1926 |March 24], pl. 42, figs. 7, 8. Examined 14 specimens. Size given as .90 by .35 [inches; 22.9 by 8.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 14 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 14 syntypes. Lot number MCZ 76387 contains a single specimen labeled “Holotype” that approximates the mea- surements and later illustrations of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 13 syntypes become paralectotypes. Lectotype Measurements. Length KS mm, diameter (excluding lip) 9.5 mm; ap- erture height (including lip) 9.1 mm, ap- erture width (including lip and peristome) 7.3 mm. Type Locality. Near Fox Hill Village, New Providence Island, Bahamas. Remarks. The illustrations were issued 508 nearly 5 years after the original description was published. Gould and Woodruff (1986: 477) regarded this taxon to be typ- ical of coastal Cerion gubernatorium (Crosse, 1869). Taxon 220. Strophiops castra Maynard and Clapp in Maynard, 1921b [July 15]: 147; Maynard and Clapp in May- nard, 1926 [March 24], pl. 42, figs. 9, 10. Examined 14 specimens. Size given as 1.12 by .42 [inches; 28.4 by 10.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 14 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 14 syntypes. Lot number MCZ 76388 contains a single specimen labeled “Holotype” that approximates the mea- surements and later illustrations of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 13 syntypes become paralectotypes. Lectotype Measurements. Length el mm, diameter (excluding lip) 9.9 mm; ap- erture height (including lip) 8.7 mm, ap- erture width (including lip and peristome) oman Type Locality. Field west of Williams Street, Nassau [New Providence Island, Bahamas]. Taxon 221. Strophiops rubiginosa May- nard and Clapp in Maynard, 1921b [July 15]: 147; Maynard and Clapp in Maynard, 1926 [March 24], pl. 43, figs. 1, 2. Examined 385 specimens. Size giv- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 en as .85 by .28 [inches; 21.6 by 7.1 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 385 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 385 syntypes. Lot MCZ 76006 - contains a single specimen labeled “Ho- lotype” that differs substantially from the published measurements for this taxon but approximates the later illustrations and is accompanied by a label in Maynard’s hand with the term “type,” which includes ref- erences to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 384 syntypes, including MCZ 76007, MCZ 356679, and USNM 420089 (50 specimens), become paralectotypes. Lectotype Measurements. Length 27.7 mm, diameter (excluding lip) 11.3 mm; ap- erture height (including lip) 9.5 mm, ap- erture width (including lip and peristome) 7.8 mm. Type Locality. Field east of Methodist Church on Sherley Street, Nassau |New Providence Island, Bahamas]. Remarks. The illustrations were issued nearly 5 years after the taxon description was published. Gould and Woodruff (1986: 476) regarded this taxon to be “in- termediate” between Cerion glans (Kiister, 1844) and C. gubernatorium (Crosse, 1869). Taxon 222. Strophiops migratoria May- nard and Clapp in Maynard, 1921b [July 15]: 147; Maynard and Clapp in Maynard, 1926 [March 24], pl. 43, figs. 3, 4. Examined 75 specimens. Size given as 1.10 by .40 [inches; 27.9 by 10.2 mm], but the term “type” was not used. THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. Type Material. The original description did not distinguish among the 75 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 75 syntypes. Lot number MCZ 76312 contains a single specimen labeled “Holotype” that approximates the mea- surements and later illustrations of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 74 syntypes, including MCZ 76311, become paralecto- types. Lectotype Measurements. Length 27.8 mm, diameter (excluding lip) 11.2 mm; ap- erture height (including lip) 9.9 mm, ap- erture width (including lip and peristome) 8.2 mm. Type Locality. Grounds directly about Methodist Sunday School building, Sher- ley Street, Nassau [New Providence Is- land, Bahamas]. Remarks. The illustrations were issued nearly 5 years after the original description was published. Gould and Woodruff (1986: 476) regarded this taxon to be “in- termediate” between Cerion glans (Kiister, 1844) and C. gubernatorium (Crosse, 1869). Taxon 223. Strophiops mayoi Maynard and Clapp in Maynard, 1921b [July 15]: 148; Maynard and Clapp in May- nard, 1926 [March 24], pl. 43, figs. 9, 10. Examined 575 specimens. Size given as 1.03 by .40 [inches; 26.2 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 575 speci- 509 mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 575 syntypes. Lot MCZ 76318 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and later illustrations of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original con- cept of it. The remaining 574 syntypes, in- cluding MCZ 76317 and USNM 420025 (200 specimens), become paralectotypes. Lectotype Measurements. Length 26.3 mm, diameter (excluding lip) 10.6 mm; ap- erture height (including lip) 9.3 mm, ap- erture width (including lip and peristome) OS. bamime: Type Locality. In field east of Mackey Street, Nassau [New Providence Island, Bahamas]. Remarks. The illustrations of this taxon were issued nearly 5 years after the origi- nal description was published. Gould and Woodruff (1986: 476) regarded this taxon to be “intermediate” between Cerion glans (Ktister, 1844) and C. gubernatorium (Crosse, 1869). Taxon 224. Strophiops reincarnata May- nard and Clapp in Maynard, 1921b [July 15]: 148, pl. 44, figs. 1, 2 [pl. 44 was never issued]. Examined 176 spec- imens. Size given as 1.12 by .40 [inches; 28.4 by 10.2 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 176 speci- mens examined. In the preamble to the taxonomic portion to this publication, 510 Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 176 syntypes. Lot number MCZ 76322 contains a single specimen labeled “Holotype” that approximates the mea- surements and later illustrations of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The remaining 175 syntypes, including MCZ 76321, become paralecto- types. Lectotype Measurements. Length 28.5 mm, diameter (excluding lip) 10.9 mm; ap- erture height (including lip) 9.9 mm, ap- erture width (including lip and peristome) Ona) mann: Type Locality. Near Ocean Hole E of Mackey Street, Nassau [New Providence Island, Bahamas]. Remarks. The illustrations of this taxon were never issued. Gould and Woodruff (1986: 476) regarded this taxon to be an “anomalously located species attributable to Cerion glans.” Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Taxon 225. Strophiops fincastlei Maynard and Clapp in Maynard, 1921b [July 15]: 148, pl. 44, figs. 5, 6 [pl 44 was never issued]. Examined 150 specimens. Size given as .85 by .47 [inches; 21.6 by 11.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 150 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that — types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 150 syntypes. Lot number MCZ 76015 contains a single specimen labeled “Holotype” that approximates the mea- surements of this taxon and is accompa- nied by a label in Maynard’s hand with the term “type,” which includes references to the unpublished figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 149 syntypes, including MCZ 76016 and MCZ 118102, become paralec- totypes. Lectotype Measurements. Length 21.7 mm, diameter (excluding lip) 9.5 mm; ap- = Figures 223-236. Taxon 223. Strophiops mayoi Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76318. Field east of Mackey Street, Nassau, New Providence Island, Bahamas. Taxon 224. Strophiops reincarnata Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76322. Near Ocean Hole east of Mackey Street, Nassau, New Providence Island, Bahamas. Taxon 225. Strophiops fincastlei Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76015. Vicinity of Fort Fincastle, West Nassau, New Providence Island, Bahamas. Taxon 226. Strophiops novita Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76319. In park west of Fort Montague, East Nassau, New Providence Island, Bahamas. Taxon 227. Strophiops phoenecia Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76330. Waterloo, East Nassau, New Providence Island, Bahamas. Taxon 228. Stro- phiops muralia Maynard and Clapp in Maynard, 1921. [Not illustrated.] Lectotype MCZ 76389. In walls at East Nassau, New Providence Island, Bahamas. Taxon 229. Strophiops rosea Maynard and Clapp in Maynard, 1921. The type of Cerion (Maynardia) agassizii Dall, 1894, is designated as the neotype of Strophiops rosea Maynard and Clapp in Maynard, 1921, making Strophiops rosea Maynard and Clapp in Maynard, 1921, an objective junior synonym of Cerion (Maynardia) agassizii Dall, 1894. Taxon 230. Strophiops gigantea Maynard and Clapp in Maynard, 1921. Holotype MCZ 76244. In wall on Village Road, near Sherley Street, Nassau, New Providence Island, Bahamas. Strophiops ajax Maynard, 1924 [Taxon 246], is a replacement name for this taxon, a homonym of Strophia grayi gigantea Maynard, 1894 [Taxon 36]. Taxon 231. Strophiops leva Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76337. In rock above Queen’s Staircase, Nassau, New Providence Island, Bahamas. Taxon 232. Strophiops flacida Maynard and Clapp in Maynard, 1921. Lectotype MCZ 76397. In rock above Queen’s Staircase, Nassau, -THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. oll New Providence Island, Bahamas. Taxon 233. Strophiops crassamarga Maynard and Clapp in Maynard, 1924. [Not illustrated. ] Strophiops crassamarga has the same type specimens and type locality and is an objective junior synonym of Strophiops crassalabra Maynard and Clapp in Maynard, 1921 [Taxon 210]. Taxon 234. Strophiops extranea Maynard, 1924. Holotype MCZ 76142. Roseville Key, Exuma Group, Bahamas. Taxon 235. Strophiops albicostata Maynard and Clapp in Maynard, 1924. Lectotype MCZ 76109. Long Key southeast of Highburn Key, Exuma Group, Bahamas. Taxon 236. Strophiops extensa Maynard, 1924. Lectotype MCZ 76008. Churchyard of the Baptist Chapel on St. James St., East Nassau, New Providence Island, Ba- hamas. 512 erture height (including lip) 8.0 mm, ap- erture width (including lip and peristome) 6.3 mm. Type Locality. Vicinity of Fort Fincastle, West Nassau [New Providence Island, Ba- hamas |]. Remarks. This taxon was never illustrat- ed. In the original description, Maynard noted that many of the 150 specimens were dead and later (1924c: 6) listed S. fincastlei as extinct in 1924 because of the burning of vegetation prior to cultivation. Gould and Woodruff (1986: 476) regarded this taxon to be “intermediate” between Cerion glans (Kiister, 1844) and C. gub- ernatorium (Crosse, 1869). Taxon 226. Strophiops novita Maynard, 1919b [November 6]: 36 [nomen nu- dum]; Maynard and Clapp in May- nard, 1921b [July 15]: 148, pl. 45, figs. 1, 2 [pl. 45 was never issued]. Ex- amined 251 specimens. Size given as .90 by .35 [inches; 22.9 by 8.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 251 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 251 syntypes. Lot MCZ 76319 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments of this taxon and is accompanied by a label in Maynard’s hand with the term “type,” which includes references to the unpublished figures. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original concept of it. The re- maining 250 syntypes, including MCZ 118104, become paralectotypes. Lectotype Measurements. Length 23.0 Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 mm, diameter (excluding lip) 9.2 mm; ap- erture height (including lip) 8.6 mm, ap- erture width (including lip and peristome) 6.9 mm. Type Locality. In park west of Fort Montague, East Nassau [New Providence Island, Bahamas]. Remarks. This taxon, which first ap- peared as a nomen nudum (Maynard, 1919b: 36), was never illustrated. Maynard (1924c: 6) listed S. novita as nearly extinct (only one specimen) in 1924 from the burning of vegetation prior to cultivation. . Gould and Woodruff (1986: 476) regarded this taxon to be “intermediate” between Cerion glans (Kiister, 1844) and C. gub- ernatorium (Crosse, 1869). Taxon 227. Strophiops phoenicia Maynard, 1919b [November 6]: 36 [nomen nu- dum]. S. phoenecia Maynard and Clapp in Maynard, 1921b [July 15]: 149, pl. 45, figs. 3, 4 [pl. 45 was never issued]; Maynard, 1924b?: [4]; Maynard, 1924c: 6. Examined 1,550 specimens. Size given as .85 by .33 [inches; 21.6 by 8.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 1,550 spec- imens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard's publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 1,550 syntypes. Lot MCZ 76330 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and later illustrations of this taxon and is accompanied by a label in May- nard’s hand with the term “type,” which includes references to the figures. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. cept of it. The remaining 1,549 syntypes, including MCZ 76331, MCZ 118156, and USNM 420059 (50 specimens), become paralectotypes. Lectotype Measurements. Length DD mm, diameter (excluding lip) 8.6 mm; ap- erture height (including lip) 7.9 mm, ap- erture width (including lip and peristome) 6.2 mm. Type Locality. Waterloo, East Nassau, New Providence Island, Bahamas. Remarks. The binomen S. phoenicea first appeared as a nomen nudum (May- nard, 1919b: 36). The taxon description used the spelling S. phoenecia, as did the sales catalog (Maynard, 1924b?: [4]). May- nard (1924c: 6) listed S. phoenecia as ex- tinct in 1924 from the burning of vegeta- tion prior to cultivation. Gould and Wood- ruff (1986: 477) regarded this taxon to be a synonym of Cerion gubernatorium (Crosse, 1869). Taxon 228. Strophiops muralia Maynard and Clapp in Maynard, 1921b [July 15]: 151, pl. 47, figs. 5, 6 [pl. 47 was never issued]. Examined 4 specimens. Size given as 1.05 by .45 [inches; 26.7 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the four speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on four syntypes. Lot MCZ 76389 contains a single specimen labeled “Ho- lotype” that approximates the measure- ments and later illustrations of this taxon and is accompanied by a label in May- nard’s hand with the term “type.” This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that 513 is consistent with Maynard’s original con- cept of it. The remaining three syntypes become paralectotypes. Lectotype Measurements. Length 26.4 mm, diameter (excluding lip) 12.1 mm; ap- erture height (including lip) 10.5 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. In walls at East Nassau, New Providence Island, Bahamas. Remarks. Gould and Woodruff (1986: 481) examined the “Holotype” of this tax- on and attributed it to the synonymy of Cerion agassizii Dall, 1894. Taxon 229. Strophiops rosea Maynard and Clapp in Maynard, 1921b [July 15]: 151, pl. 48, figs. 9, 10 [pl. 48 was never issued]. Examined one specimen. Size given as .73 by .36 [inches; 18.5 by 9.1 mm], but the term “type” was not used. Type Material. The original description was based on a single specimen, the unique holotype, for which measurements were provided but for which the illustra- tions were never published. The specimen was not at the MCZ, nor was there any catalog entry for this taxon at the MCZ at the time Maynard’s collection was cata- logued. Searches of the collections and/or databases of the USNM, the Academy of Natural Sciences of Philadelphia, the Field Museum of Natural History, the Florida Museum of Natural History, and the Mu- seum of Biological Diversity, Ohio State University, failed to uncover a single spec- imen catalogued under this name. May- nard’s original brief description—*“Similar to last [S. agassizii Dall] but smaller, .73 by .36, with 8 instead of 11 whorls, and is tinged with dull orange”—is inadequate to differentiate this taxon from S. agassizii unambiguously. The holotype of Cerion (Maynardia) agassizii Dall, 1894 (MCZ IP 113595, Invertebrate Paleontology Collec- tion), is here designated as the neotype of Strophiops rosea Maynard and Clapp in Maynard, 1921, to provide an objective 514 standard of reference for this species- group taxon. Strophiops rosea Maynard and Clapp in Maynard, 1921, becomes a junior objective synonym of Cerion (May- nardia) agassizii Dall, 1894. Type Locality. Strophiops rosea was ini- tially described as being from the north- east end of Rose Island, New Providence Island, Bahamas (Maynard and Clapp in Maynard, 1921: 151). According to Article 76.3 (ICZN, 1999: 87), the type locality of Cerion agassizii Dall, 1894, “The calcare- ous sand-rock at the W. quarry, top of Nas- sau Ridge” |New Providence Island, Ba- hamas], becomes the type locality of Stro- phiops rosea. Remarks. The illustrations of this taxon were never published. Gould and Wood- ruff (1986: 481) mentioned that they had not seen the unique holotype of this taxon but considered it to be a “fairly unambig- uous” synonym of Cerion agassizii Dall, 1894, on the basis of the limited descrip- tion. Taxon 230. Strophiops gigantea Maynard and Clapp in Maynard, 1921b [July 15]: 152, pl. 47, figs. 9, 10 [pl. 47 never issued]. Examined one specimen. Size given as 1.75 by .65 [inches; 44.5 by 16.5 mm], but the term “type” was not used. Type Material. The original description is based on a single specimen, the unique holotype, for which measurements were provided, but for which the illustrations were never issued. Lot MCZ 76244 con- tains a single specimen accompanied by two MCZ labels that refer to this specimen as the holotype of Strophiops ajax, a re- placement name for the preoccupied S. gi- gantea (see Remarks). A handwritten label with the specimen contains the name Stro- phiops gigantea, with “gigantea” partially erased and overwritten “ajax.” This speci- men is the holotype both of S. gigantea and of S. ajax. Holotype Measurements. Length 43.9 mim, diameter (excluding lip) 17.4 mm; ap- erture height (including lip) 17.1 mm, ap- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 erture width (including lip and peristome) 36) 1aanam Type Locality. In wall on Village Road, near Sherley Street, Nassau, New Provi- dence Island, Bahamas. Remarks. Strophiops gigantea Maynard and Clapp in Maynard, 1921, is a hom- onym of Strophia grayi gigantea Maynard, 1894 |Taxon 36]. Strophiops ajax Maynard, 1924 |Taxon 246], was proposed as a re- placement name. Gould and Woodruff (1986: 480) attributed this fossil taxon to Cerion agassizti Dall, 1894. Taxon 231. Strophiops leoa Maynard and Clapp in Maynard, 1921b [July 15]: 152, pl. 48, figs. 1, 2 [pl. 48 was never issued]. Examined 12 specimens. Size given as 1.10 by .50 [inches; 27.9 by 12.7 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 12 speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on 12 syntypes. Lot number MCZ 76337 was labeled “Paratype” and contains 93 specimens, most damaged or incom- plete, of which one was sequestered. The labels make no mention of the term “type,” nor is there a reference to plates or figures. The sequestered specimen, which is complete and well preserved and had been carefully cleaned of matrix, is likely the specimen used by Maynard to prepare the species description. This spec- imen is here selected as the lectotype (MCZ 76337). The remaining specimens were recatalogued as MCZ 357013. The original description referred to 12 speci- mens, indicating that Maynard had added at least 81 specimens since the species was described. It is unclear which 11 of the remaining 92 specimens are paralecto- types. THE CERION TAXA OF CHARLES JOHNSON MAYNARD ¢ Harasewych et al. Lectotype Measurements. Length 29.6 mm, diameter (excluding lip) 12.7 mm; ap- erture height (including lip) 11.5 mm, ap- erture width (including lip and peristome) 9.0 mm. Type Locality. In rock above Queen's Staircase, Nassau, New Providence Island, Bahamas. Taxon 232. Strophiops flacida Maynard and Clapp in Maynard, 1921b [July 15]: 152, pl. 48, figs. 3, 4 [pl. 48 was never issued]. Examined two specimens. Size given as .98 by .34 [inches; 24.9 by 8.6 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the two speci- mens examined. In the preamble to the taxonomic portion to this publication, Maynard (1919b: 43-44) specified that types were selected. However, there was no indication in this or any of Maynard’s publications to associate a single specimen with the term “type.” Thus, there is no ho- lotype for this taxon, and the species is based on two syntypes. Lot number MCZ 76397 contains a single specimen labeled “Holotype” that roughly approximates the published length of this taxon and is ac- companied by a label in Maynard’s hand with the term “type” that refers to the un- published illustrations. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The other syntype (MCZ 76398) becomes a paralec- totype. Lectotype Measurements. Length 25.4 mm, diameter (excluding lip) 10.9 mm; ap- erture height (including lip) 10.1 mm, ap- erture width (including lip and peristome) 8.7 mm. Type Locality. In rock above Queen’s Staircase, Nassau, New Providence Island, Bahamas. Remarks. Gould and Woodruff (1986: 481) regarded S. flacida to be a “ribby ver- sion” of Cerion agassizii Dall, 1894. 515 Maxonme2 oor Strophiops crassamarga May- nard and Clapp in Maynard, 1924a [October 25], pl. 39, figs. 7, 8. Nomenclatural Remarks. The binomen S. crassamarga was published in the cap- tion beneath the figures (pl. 39, figs. 7, 8) cited in the original description of Stro- phiops crassalabra Maynard and Clapp in Maynard, 1921b (p. 143), and was not used subsequently by Maynard, who listed the taxon as S. crassilabra in the sales cat- alog (Maynard, 1924b?: [6]). The speci- men labeled Strophiops crassamarga on plate 39 (figs. 7, 8) is the lectotype of Stro- phiops crassalabra |Taxon 211], making Strophiops crassamarga Maynard and Clapp in Maynard, 1924, a junior objective synonym of Strophiops crassalabra May- nard and Clapp in Maynard, 1921. Taxon 234. Strophiops extranea Maynard, 1924c [10 December]: 2 [not illus- trated]. Examined one specimen. Size given as 1.17 by .50 [inches; 29.7 by 12.7 mm], but the term “type” was not used. Type Material. The original description stated that this taxon was based on a single fossil found embedded in rock and provid- ed its measurements. This taxon is thus based on a unique holotype. Lot MCZ 76142 contains a single specimen labeled “Holotype” that corresponds closely to the published measurements and is here con- sidered to be the holotype. Holotype Measurements. Length 29.9 mm, diameter (excluding lip) 12.7 mm; ap- erture height (including lip) 10.8 mm, ap- erture width (including lip and peristome) 9.2 mm. Type Locality. Roseville Key, Exuma Group [Bahamas]. Taxon 235. Strophiops albicostata May- nard and Clapp in Maynard, 1924bpP: [3] [nomen nudum]; Maynard, 1924c [10 December]: 2 [not illustrated]. Exam- ined 1,000 specimens. Size given as 1.36 by .48 linches; 34.5 by 12.2 mm], but the term “type” was not used. 516 Nomenclatural Remarks. Strophiops al- bicostata was listed for sale in Maynard’s (1924bP: [3]) catalog, whereas the descrip- tion of this taxon was published in the sup- plement to this catalog. On page [2] of the catalog, Maynard noted that, “Unless oth- erwise designated, all species should be accredited to Maynard and Clapp.” Thus, Strophiops albicostata Maynard and Clapp in Maynard (1924b?) is a nomen nudum. The taxon became available as Strophiops albicostata’ Maynard on December 10, 1924. Type Material. The original description did not distinguish among the 1,000 spec- imens examined. There was no indication in this publication to associate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 1,000 syntypes. Lot MCZ 76109 contains a single specimen la- beled “Holotype.” This specimen approx- imates the measurements in the original description and is here designated as the lectotype to provide an objective standard of reference for this species-group taxon. All other syntypes, including, MCZ 76110 and USNM 420061 (100 specimens), be- come paralectotypes. Lectotype Measurements. Length 34.1 mm, diameter (excluding lip) 12.9 mm; ap- erture height (including lip) 11.64 mm, ap- erture width (including lip and peristome) 9.4 mm. Type Locality. Long Key southeast of Highburn Key [Exuma Group, Bahamas]. Taxon 236. Strophiops extensa Maynard, 1924c [10 December]: 2, 3 [not illus- trated]. Examined 23,000 specimens. No measurements provided. Type Material. The original description did not distinguish among the 23,000 spec- imens examined. There was no indication in this publication to associate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 23,000 syntypes. An ex- amination of the MCZ catalog revealed Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 that there was no specimen segregated as type for this species in Maynard’s collec- tion at the time of his death. Lot MCZ 76008 contains 278 specimens labeled “paratypes.” Because this species had not been figured, nor have any measurements been published to help select the lecto- type, an intact, adult, live-collected speci- men that conforms to the brief description is here designated as the lectotype (MCZ 76008) to provide an objective standard of reference for this species-group taxon. The remaining 277 specimens were recata- - logued as MCZ 352331. These and all oth- er syntypes, including USNM 419996 (1,500 specimens), become paralectotypes. Lectotype Measurements. Length 31.5 mm, diameter (excluding lip) 10.1 mm; ap- erture height (including lip) 9.8 mm, ap- erture width (including lip and peristome) 7.9 mm. Type Locality. Churchyard of the Bap- tist Chapel on St. James Street, East Nas- sau, New Providence Island, Bahamas. Remarks. Maynard (1924c: 2—3) report- ed this taxon to be “abundant living in the church-yard of the Baptist Chapel on St. James St. East Nassau, on the west and south side of this street to Kemp’s Road, but, except about the chapel, many were dead. 23,000 were collected. But out of this number only about 3000 were living.” Because the lectotype was a live-collected specimen, the type locality is here restrict- ed to the church-yard. Gould and Wood- ruff (1986: 476) regarded this taxon to be “intermediate” between Cerion glans (Ktister, 1844) and C. gubernatorium (Crosse, 1869). Taxon 237. Strophiops sparsa Maynard, 1924c [10 December]: 3 [not illus- trated]. Examined “about 280” speci- mens. Size given as 1.16 by .45 [inches; 29.5 by 11.4 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the “about 280” THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. oly, Figures 237-248. Taxon 237. Strophiops sparsa Maynard, 1924. Lectotype MCZ 76399. Field at St. James Corner, East Nassau, New Providence Island, Bahamas. Taxon 238. Strophiops rufula Maynard, 1924. Lectotype USNM 420135. West side of Kemp’s Road near the terminus of St. James St. and south of that street to St. James Corner, Nassau, New Providence Island, Bahamas. Taxon 239. Strophiops gracila Maynard, 1924. Lectotype MCZ 76285. Found in and near a cutting through rocks on Soldiers Road about a mile and a half from South Shore, New Providence Island, Bahamas. Taxon 240. Strophiops montana Maynard, 1924. Lectotype USNM 420103. Sunnyside Estate East Bay St. near the house, Nassau, New Providence Island, Bahamas. Taxon 241. Strophiops clara Maynard, 1924. Lectotype MCZ 76111. Church on the road to Fox Hill from East Bay St., Nassau, New Providence Island, Bahamas. Taxon 242. Strophiops livida Maynard, 1924. Lectotype MCZ 76101. On West Bay Street, Nassau, about opposite North Silver Key, New Providence Island, Bahamas. Taxon 243. Strophiops minima Maynard, 1924. Lectotype MCZ 76022. St. James Corner, East Nassau, New Providence Island, Bahamas. Taxon 244. Stro- phiops concina Maynard, 1924. Holotype MCZ 76020. St. James Corner, East Nassau, New Providence Island, Bahamas. Taxon 245. Strophiops pygmea Maynard, 1924. Holotype MCZ 76021. St. James Corner, East Nassau, New Providence Island, Ba- hamas. Taxon 246. Strophiops ajax Maynard, 1924. [Not illustrated.] Replacement name for Strophiops gigantea Maynard and Clapp in Maynard, 1921 [Taxon 230], not Strophia grayi gigantea Maynard, 1894 [Taxon 36]. The type specimens and type locality of S. gigantea Maynard and Clapp in Maynard, 1921 are also the type specimens of S. ajax Maynard, 1924. Taxon 247. Strophiops leucophera Maynard and Clapp in Maynard, 1925. Lectotype MCZ 76283. Gray hill at the extreme northern end of Great Guana Key, Exuma Group, Bahamas. Taxon 248. Strophiops vagabunda Maynard, 1925. [Not illustrated.] Replacement name for S. albata Maynard and Clapp in Maynard, 1921b [Taxon 215], not S. albata Maynard and Clapp in Maynard, 1921 [Taxon 192]. The type specimens and type locality of Strophiops albata Maynard and Clapp in Maynard, 1921 (see Taxon 214), are also the type specimens of S. vagabunda Maynard, 1925. 518 specimens examined. There was no indi- cation in this publication to associate a sin- gle specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on “about 280” syntypes. An examination of the MCZ catalog re- vealed that there was no specimen segre- gated as type for this species in Maynard's collection at the time of his death. Lot MCZ 76399 was labeled “Paratype” and contained 174 specimens. As this species had not been figured, an intact, adult, live- collected specimen from this lot that most closely approximates the published mea- surements and conforms to the brief de- scription is here designated as the lecto- type (MCZ 76399) to provide an objective standard of reference for this species- group taxon. The remaining 173 speci- mens, which were recatalogued as MCZ 357014, are among the “about 280” syn- types that become paralectotypes. Lectotype Measurements. Length 29.3 mm, diameter (excluding lip) 11.4 mm; ap- erture height (including lip) 10.4 mm, ap- erture width (including lip and peristome) 8.9 mm. Type Locality. Field at St. James Cor- ner, East Nassau |New Providence Island, Bahamas]. Remarks. Maynard (1924c: 3) reported 20 specimens found living in a field at St. James Corner, East Nassau, and about 260 found dead scattered in fields from this point east to the estate called Sunnyside, but chiefly as shells occupied by hermit crabs. Because the lectotype was collected living (as evidenced by vestiges of an epi- phragm around the aperture), the type lo- cality is restricted to the field at St. James Corner, East Nassau, New Providence Is- land, Bahamas. Gould and Woodruff (1986: 476) re- garded this taxon to be an “anomalously located species attributable to Cerion glans.” Taxon 238. Strophiops rufula Maynard, 1924c [10 December]: 3 [not illus- Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 trated]. Examined 3,439 specimens. Measurements not provided. Type Material. The original description did not distinguish among the 3,439 spec- imens examined. There was no indication in this publication to associate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 3,439 syntypes. An ex- amination of the MCZ catalog revealed that there was no specimen segregated as type for this species in Maynard’s collec- tion at the time of his death. Large syntype ~ lots are present in the USNM and MCZ collections. Because this species was never figured, nor had any measurements been published to help select the lectotype, an intact, adult, live-collected specimen that conforms to the brief description is here designated as the lectotype (USNM 420135) to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of this taxon. The remaining syn- types, including MCZ 76098 and USNM 1093796, become paralectotypes. Lectotype Measurements. Lot USNM 420135, length 28.3 mm, diameter (ex- cluding lip) 10.7 mm; aperture height (in- cluding lip) 10.9 mm, aperture width (in- cluding lip and peristome) 8.8 mm. Type Locality. On the west side of Kemp’s Road, near the terminus of St. James St. [Street] and south of that street to St. James Corner, Nassau, New Provi- dence Island, Bahamas. Remarks. Maynard (1924c: 3) noted that of the 3,439 specimens collected, only 649 were living. Taxon 239. Strophiops gracila Maynard, 1924c [10 December]: 3 [not illus- trated]. Examined 61 specimens. Only diameter given as .40 [inches; 10.2 mm]. Type Material. The original description did not distinguish among the 61 speci- mens examined. There was no indication in this publication to associate a single specimen with the term “type.” Thus, THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. there is no holotype for this taxon, and the species is based on 61 syntypes. An ex- amination of the MCZ catalog revealed that there was no specimen segregated as type for this species in Maynard's collec- tion at the time of his death. Lot MCZ 76285 was labeled “Paratypes” and con- tained 33 specimens. Because this species was never illustrated and only a measure- ment for the diameter was published, an intact, adult, live-collected specimen from this lot that most closely approximates the published diameter and conforms to the brief description is here designated as the lectotype to provide an objective standard of reference for this species-group taxon. The remaining 32 specimens, which were recatalogued as MCZ 357015, are among the 60 syntypes that become paralectoty- es. : Lectotype Measurements. Length 26.5 mim, diameter (excluding lip) 10.2 mm; ap- erture height (including lip) 10.4 mm, ap- erture width (including lip and peristome) 8.5 mm. Type Locality. Found in and near a cut- ting through rocks on Soldier's Road about a mile and a half from South Shore, New Providence Island, Bahamas. Remarks. Gould and Woodruff (1986: 478) regarded this taxon to be a synonym of Cerion gubernatorium (Crosse, 1869). Taxon 240. Strophiops montana Maynard, 1924c [10 December]: 3, 4 [not illus- trated]. Examined 1,219 specimens. Only diameter given as .35 [inches; 8.9 mm]. Type Material. The original description did not distinguish among the 1,219 spec- imens examined. There was no indication in this publication to associate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 1,219 syntypes. An ex- amination of the MCZ catalog revealed that there was no specimen segregated as type for this species in Maynard’s collec- tion at the time of his death. Large syntype O19 lots are present in the USNM and MCZ collections. Because this species had not been figured and only the diameter was published, an intact, adult, live-collected specimen that conforms to the brief de- scription is here designated as the lecto- type (USNM 420103) to provide an objec- tive standard of reference for this species- group taxon that is consistent with May- nard’s original concept of this taxon. The remaining syntypes, including MCZ 76113 and USNM 1093794, become paralecto- types. Lectotype Measurements. Number USNM 420103, length 26.4 mm, diameter (excluding lip) 9.8 mm; aperture height (including lip) 9.4 mm, aperture width (in- cluding lip and peristome) 7.4 mm. Type Locality. Occurs on Sunnyside Es- tate East Bay St. near the house, Nassau, New Providence Island, Bahamas. Remarks. Maynard (1924c: 3-4) noted that of the 1,219 specimens collected, only 33 were living. Gould and Woodruff (1986: 478) reported that they did not examine specimens of this taxon but regarded it to be a synonym of Cerion gubernatorium (Crosse, 1869). Taxon 241. Strophiops clara Maynard, 1924c [10 December]: 4 [not illus- trated]. Examined four specimens. Size given as .80 by .35 [inches; 20.3 by 8.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the four speci- mens examined. There was no indication in this publication to associate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on four syntypes. Lot MCZ 76111 contains a single specimen la- beled “Holotype” that roughly approxi- mates the published measurements for this taxon. This specimen is here designat- ed as the lectotype to provide an objective standard of reference for this species- group taxon. The remaining three speci- mens become paralectotypes. 520 Lectotype Measurements. Length 19.2 mm, diameter (excluding lip) 8.3 mm; ap- erture height (including lip) 7.8 mm, ap- erture width (including lip and peristome) 6.5 mm. Type Locality. About a church on the road to Fox Hill from East Bay St., Nas- sau, New Providence Island, Bahamas. Remarks. Gould and Woodruff (1986: 477) considered this taxon to be a syno- nym of Cerion gubernatorium (Crosse, 1869). Taxon 242. Strophiops livida Maynard, 1924c [10 December]: 4 [not illus- trated]. Examined 795 specimens. Mea- surements not provided. Type Material. The original description did not distinguish among the 795 speci- mens examined. There was no indication in this publication to associate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 795 syntypes. Neither measurements nor illustration are includ- ed in the original description. Lot MCZ 76101 contains a single specimen labeled “Holotype” that is accompanied by a label in Maynard’s hand with the term “type,” which is dated July 27, 1924, presumably the date it was collected. This specimen is here designated as the lectotype to provide an objective standard of reference for this species-group taxon that is consistent with Maynard's original concept of it. The re- maining 794 specimens, including, MCZ 76102 and USNM 420104 (200 specimens including one sinistral), become paralec- totypes. Lectotype Measurements. Length 28.5 mim, diameter (excluding lip) 11.5 mm; ap- erture height (including lip) 11.7 mm, ap- erture width (including lip and peristome) 9.7 mm. Type Locality. On West Bay Street, Nas- sau, about opposite North Silver Key, New Providence Island, Bahamas. Remarks. Gould and Woodruff (1986: 477) considered this taxon to be a syno- nym of Cerion glans (Kiister, 1844). Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Taxon 243. Strophiops minima Maynard, 1924c [10 December]: 4 [not illus- trated]. Examined 4 specimens. Size given as .75 by .31 [inches; 19.1 by 7.9 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the four speci- mens examined. There was no indication in this publication to associate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on four syntypes. An ex- amination of the MCZ catalog revealed - that there was no specimen segregated as type for this species in Maynard's collec- tion at the time of his death. Lot MCZ 76022 was labeled “Paratype” and con- tained three specimens |MCZ catalog lists five], all damaged to some degree. The best preserved specimen (lacking a portion of the protoconch) is here selected as the lectotype (MCZ 76022) to provide an ob- jective standard of reference for this spe- cies-group taxon. The remaining two spec- imens, recatalogued as MCZ 357016, be- come paralectotypes. Lectotype Measurements. Length 21.0 mm, diameter (excluding lip) 9.6 mm; ap- erture height (including lip) 8.9 mm, ap- erture width (including lip and peristome) 7.4 mm. Type Locality. St. James Corner, East Nassau [New Providence Island, Baha- mas]. Remarks. From crab hole, with S. con- cina and S. pygmea. Gould and Woodruff (1986: 482) considered this taxon to be a synonym of Cerion gubernatorium (Crosse, 1869), but questioned whether it were truly “fossil” or just long dead. Taxon 244. Strophiops concina Maynard, 1924c [10 December]: 4 [not illus- trated]. Examined one specimen. Size given as .65 by .30 [inches; 16.5 by 7.6 mm], but the term “type” was not used. Type Material. The original description states that this taxon is known from one specimen, the measurements provided ap- THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. plying to the unique holotype (MCZ 76020). Holotype Measurements. Length 16.0 mm, diameter (excluding lip) 7.7 mm; ap- erture height (including lip) 6.6 mm, ap- erture width (including lip and peristome) 6.1 mm. Type Locality. St. James Corner, East Nassau [New Providence Island, Baha- mas]. Remarks. From crab hole, with S. min- ima and S. pygmea. Gould and Woodruff (1986: 481) considered this taxon to be a synonym of Cerion universum (Maynard, 1913). Taxon 245. Strophiops pygmea Maynard, 1924c [10 December]: 4, 5 [not illus- trated]. Examined one specimen. Size given as .50 by .25 [inches; 12.7 by 6.4 mm], but the term “type” was not used. Type Material. The original description states that this taxon is known from one specimen, the measurements provided ap- plying to the unique holotype (MCZ 76021). Holotype Measurements. Length 12.2 mm, diameter (excluding lip) 6.1 mm; ap- erture height (including lip) 4.5 mm, ap- erture width (including lip and peristome) 4.6 mm. Type Locality. St. James Corner, East Nassau [New Providence Island, Baha- mas]. Remarks. From crab hole, with S. min- ima and S. concina. Maynard (1924c: 4) noted that this was the smallest fossil Cer- ion known to him. Gould and Woodruff (1985: 481) considered this a synonym of Cerion universum (Maynard, 1913). Taxon 246. Strophiops ajax Maynard, 1924c [10 December]: 5. Replace- ment name for Strophiops gigantea Maynard and Clapp in Maynard, 1921 [Taxon 230], not Strophia grayi gigantea Maynard, 1894 [Taxon 36]. Type Material. The e specimen (Ho- lotype, MCZ 76244) and type locality of S. 521 gigantea Maynard and Clapp, 1921 (see Taxon 230), are also the type specimen and type locality of S. ajax Maynard, 1924. Taxon 247. Strophiops leucophera May- nard, 1925 [July 18]: 181, pl. 52, figs. 1, 2 (pl. 52 never published). Examined 50 specimens. Size given as .50 by 1.50 [inches; 12.7 by 38.1 mm], but the term “type” was not used. Type Material. The original description did not distinguish among the 50 speci- mens examined. There was no indication in this publication to associate a single specimen with the term “type.” Thus, there is no holotype for this taxon, and the species is based on 50 syntypes. Lot num- ber MCZ 76283 contains a single speci- men labeled “Holotype” that is accompa- nied by a label in Maynard’s hand with the term “type,” which includes references to figures that were never published. This specimen is here designated as the lecto- type to provide an objective standard of reference for this species-group taxon that is consistent with Maynard’s original con- cept of it. The remaining 49 specimens be- come paralectotypes. Lectotype Measurements. Length 32.5 mm, diameter (excluding lip) 13.1 mm; ap- erture height (including lip) 12.0 mm, ap- erture width (including lip and peristome) 10.2 mm. Type Locality. Gray hill at the extreme northern end of Great Guana Key, Exuma Group, Bahamas. Remarks. The published measurements are transposed. Because this taxon was published in “Part Three” of Contribu- tions to the History of the Cerionidae, the authorship is attributed to Maynard, rather than Maynard and Clapp, as listed in Clench (1957: 151). Taxon 248. Strophiops vagabunda May- nard, 1925 [July 18], pl. 41, figs. 7, 8 [appears in figure caption only]. Nomenclatural Remarks. The binomen Strophiops vagabunda appeared under fig- 522 ures cited in the original description of Strophiops albata Maynard and Clapp in Maynard, 1921b (see Taxon 215), but pub- lished 4 years later. Strophiops albata Maynard and Clapp in Maynard, 1921b, is a junior homonym of S. albata Maynard and Clapp in Maynard, 1921a (see Taxon 192). Clench (1957: 166) regarded Stro- phiops vagabunda to be a replacement name for S. albata Maynard and Clapp in Maynard, 1921b, not S. albata Maynard and Clapp in Maynard, 1921a, but attri- buted authorship to Maynard and Clapp. As the figures are captioned “S. vagabun- da” authorship of the taxon is confined to Maynard, 1925. Type Material. The type specimens (Lectotype, MCZ 76258) and type locality of Strophiops albata Maynard and Clapp in Maynard, 1921 (see Taxon 215), are also the type specimens and type locality of S. vagabunda Maynard, 1925. LITERATURE CITED ABELE, S. D. 2002. Discovering Charles Johnson Maynard, Naturalist and Teacher. Newton, Mas- sachusetts: Jackson Homestead Museum. 30 pp. BATCHELDER, C. F. 1951. A bibliography of the pub- lished writings of Charles Johnson Maynard [1845-1929]. Journal of the Society for the Bib- liography of Natural History, 2(7): 227—260. CLENCH, W. J. 1933. Notes and descriptions of land mollusks from the Bahama Islands, based mainly upon collections obtained during the Utowana Expedition of 1932 and 1933. Proceedings of the New England Zoélogical Club, 13: 77—100, pl. 1. . 1934. Notes and descriptions of new Cerions from Hispaniola and the Bahama Islands, based mainly upon collections obtained during the Uto- wana Expedition of 1934. Proceedings of the Boston Society of Natural History, 40(2): 205— 218, pls. 1-2. 1952. Land and freshwater mollusks of Eleuthera Island, Bahama Islands. Revista de la Sociedad Malacologia “Carlos de la Torre,” 8: 97-116. . 1957. A catalog of the Cerionidae (Mollusca: Pulmonata). Bulletin of the Museum of Com- parative Zoology, 116: 121—169. . 1959. Land and freshwater mollusks of Great and Little Inagua, Bahama Islands. Bulletin of the Museum of Comparative Zoology, 121(2): 29-53, pl. 1. . 1963. Land and freshwater Mollusca of the Crooked Island Group, Bahamas. Bulletin of the Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 Museum of Comparative Zoology, 128(8): 393— 414. . 1964. Land and freshwater Mollusca of the Cayman Islands, West Indies. Occasional Papers on Mollusks Museum of Comparative Zoology, 2(31): 345-380, pls. OOS) CLENCH, W. J., AND C. G. AGUAYO. 1952. The sca- larinum species complex (Umbonis) in the genus Cerion. Occasional Papers on Mollusks, Harvard University, 1: 413-440. FRENCH, M. V. 1930. Charles J. Maynard. The Flor- ida Naturalist, 3: 58. GOULD, S. J., AND D. S. WOODRUFF. 1986. Evolution and systematics of Cerion (Mollusca: Pulmonata) on New Providence Island: a radical revision. Bulletin of the American Museum of Natural History, 182(4): 389-490. GOULD, S. J., N. D. YOUNG, AND B. KASSON. 1985. The consequences of being different: sinistral coiling in Cerion. Evolution, 39(6): 1364-1379. HUMMELINCK, P. W. 1980. Caribbean land molluscs, Cerion in the Cayman Islands. Uitgaven Natu- urwetenschappelijke Studiekring voor Suriname en de Nederlandse Antillen, 1980: 1—67. ICZN. 1999. International Code of Zoological No- menclature. Fourth ed. London: The Interna- tional Trust for Zoological Nomenclature. xxix + 306 pp. JOHNSON, C. W. 1930. Charles Johnson Maynard. The Nautilus, 43(3): 101-103, 1 pl. MAYNARD, C. J. 1889a [April]. Monograph of the ge- nus Strophia, a group of tropical and sub-tropical land shells. Contributions to Science, 1(1): 1—29, pls. 1-2. . 1889b [July]. Monograph of the genus Stro- phia. [Continued]. Contributions to Science, 1(2): 68-79, pl. 7. . 1889c [October]. Monograph of the genus Strophia. [Continued]. Contributions to Science, 1(3): 125-135, pl. 16. . 1890 [January (dated “January, 1889” on page 1)]. Monograph of the genus Strophia. [Continued]. Contributions to Science, 1(4): 188-197. . 1894a [December]. Monograph of the genus Strophia. [Continued]. Contributions to Science, 2 (8) O72) . 1894b [December]. Monograph of the genus Strophia. [Continued]. Contributions to Science, 24) bso. . 1896 [March]. Monograph of the genus Stro- phia. [Continued]. Contributions to Science, 3(1): 1-40, pls. 1-7. . 1913a [January]. Descriptions of some spe- cies of the family Cerionidae. Records of Walks and Talks with Nature, 5(Appendix): 177—182. . 1913b [February 1]. Descriptions of some species of the family Cerionidae. Records of Walks and Talks with Nature, 5(Appendix): 183— 186. . 1913c [February 9]. Descriptions of some THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. species of the family Cerionidae. Records of Walks and Talks with Nature, 5(Appendix): 187— 200. . 1914 [August 18]. A species of land shell of the family Cerionidae redescribed. Records of Walks and Talks with Nature, 6(Appendix): 177— 179 . 1919a [October 16]. Contributions to the his- tory of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(1, Appendix): 1-24, text figs. 1-3, pls. 5, 6. . 1919b [November 6]. Contributions to the history of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(2, Appendix): 25-52, text figs. 4-18 [including 13*, 14*, 15*], pls. 7, 8. . 1919c [December 31]. Contributions to the history of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(3, Appendix): 53-68, text figs. 19-42 [including 30*, 32*], pls. 9, 10. . 1920a [February 29]. Contributions to the history of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(4, Appendix): 69-80, text figs. 43-53, pls. 10*, 11. . 1920b [April 8]. Contributions to the history of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, Volume 10(5, Appendix): 81-92, text figs. 55-60 [including 59*], pls. 12, 13. . 1920c [July 10]. Contributions to the history of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(6, Appendix): 93-110, text figs. 61-89 [in- cluding 70*], pls. 14-19. . 1920d [September 30]. Contributions to the history of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(7, Appendix): 111-126, text figs. 90-92, Map l, pls. 1-4, 20-25. . 1921a [May 2]. Contributions to the history of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(8, Appendix): T= NBS. pls. 26-33. . 1921b [July 15]. Contributions to the history of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(9, Appendix): 139-154, pls. 34-37. . 1924a [October 25]. Contributions to the his- tory of the Cerionidae with descriptions of many 923 new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(10, Appendix): 155-170, text figs. 93-94, pls. 38, 39. . 1924bP [undated]. Catalogue of Specimens of the Family Cerionidae for Sale by Charles J. Maynard Containing Many New Species. West Newton, Massachusetts: Published privately. 8 unnumbered pp. . 1924c [December 10]. Supplement to Sale Catalogue of Cerionidae with Descriptions of New Species Collected in the Bahama Islands in Summer of 1924. [Footnote on p. 6 “Issued De- cember 10, 1924”] West Newton, Massachusetts: Published privately. Pp. 1-6. . 1925 [July 18]. Contributions to the history of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(11, Appendix): 171-194, text figs. 95-103, maps 2-4, pls. 40-41. . 1926 [March 24]. Contributions to the his- tory of the Cerionidae with descriptions of many new species and notes on evolution in birds and plants. Records of Walks and Talks with Nature, 10(12, Appendix): 195-218, text figs. 104-116, maps 5-6, pls. 42, 43. MAYNARD, C. J., AND N. A. CLApp. 1914 [August 18]. Descriptions of new Strophias. Records of Walks and Talks with Nature, 6(Appendix): 180. . 1915 [February 8]. Descriptions of new Stro- phias. Records of Walks and Talks with Nature. 6(Appendix): 179 bis, 180 bis, 181, 182. [PILsBRY, H. A.] 1901. Francis C. Browne [Obituary]. The Nautilus, 14(11): 132. PiLsBry, H. A. 1901—02. Family Cerionidae. Manual of conchology. Ser. 2. Pulmonata. Academy of Natural Sciences of Philadelphia, 14: 174-286, pls. 27-47. PILSBRY, H. A., AND E. G. VANATTA. 1896. Catalogue of the species of Cerion, with descriptions of new forms. Proceedings of the Academy of Natural Sciences of Philadelphia, (1896): 315-338, pl. 11. PLATE, L. 1907. Die Variabilitét und die Artbildung nach dem Prinzip geographischer Formenketten bei den Cerion-Landschnecken der Bahamas-In- seln. Archiv fiir Rassen- und Gesselschafts-Biol- ogie, 4: 433-470, 581-614, pls. 1-5. TOWNSEND, C. W. 1930. Charles Johnson Maynard. Bulletin of the Boston Society of Natural History, 54: 1-7, 2 pls. TURNER, R. D. 1957. Charles Johnson Maynard and his work in malacology. Occasional papers on Mollusks Museum of Comparative Zoology, Har- vard, 2(21): 137-152, portrait. WoopkRUFF, D. S. 1978. Evolution and adaptive ra- diation of Cerion: a remarkably diverse group of West Indian land snails. Malacologia, 17: 223— 239. 524 INDEX OF SPECIES-LEVEL TAXA acceptoria Maynard, 1913—Taxon 68 ___......_.... 413 accuminator Maynard and Clapp in Maynard, 1920 > TaxonwiliGS he s+ 3 ee ls Sie 479 accumulata Maynard and Clapp in Maynard, 1920 (misspelling)—Taxon 168 ____. 479 acuminata Maynard, 1920 (nomen nudum)— PaxOnr liGSr 4 ss tine Cid ce sas, Mies Gade hee eee ee 479 acuta Maynard, 1889—Taxon 4 376 adumbra Maynard and Clapp in Maynard, 1920S. © ijn ae 5 toh oe GR ee ween 483 affinis Maynard, 1913—Taxon 67 ____-_--- 412 agava Maynard, 1894—Taxon 41 __.. 398 agavaneglecta Maynard, 1913—Taxon 83 ___... 421 agrestina Maynard, 1894—Taxon 51] ___.. 402 agricola Maynard and Clapp in Maynard, LOUG2 axons les, fea eee eae 445 agristina Maynard, 1894 (misspelling)—Taxon i Ub eet ae A Sarit ee ccichte cA E ONRL Sa lb ge ted ats 402 agsestina Maynard, 1919 (misspelling)—Taxon Bull» | wieteiues Ned PS CS, eae DN Dien 402 ajax Maynard, 1924—Taxon 246 Sw alba Maynard, 1889—Taxon 17 384 albata Maynard and Clapp in Maynard, LO2de=slaxon® LOZ Ss 28 ee. es Eee ee 493 albata Maynard and Clapp in Maynard, 1921 (homonym of Strophia albata Maynard and Clapp in Maynard, 1921 [Taxon 192]; Strophiops vagabunda Maynard and Clapp in Maynard, 1925 [Taxon 248], is a replacement name)—Taxon 215 _ 505 albea Maynard, 1894—Taxon 32 393 albicostata Maynard, 1924—Taxon 235 ___. 515 albolabra Maynard, 1920—Taxon 134 457 angustalabra Maynard and Clapp in Maynard, 192 Rax one ie aire ROD eo 502 angustocostata Maynard and Clapp in Maynard, 1921—Taxon 210 ___ 501 antiqua Maynard, 1913—Taxon 63 ___.... 410 arbusta Maynard and Clapp in Maynard, O19 = axon e lO es Rai ae eel a 447 argentea Maynard, 1919, 1921, 1924? (misspelling)—Taxon 82 420 argentia Maynard, 1913—Taxon 82 ____.- 420 argntea Maynard, 1921 (misspelling)—Taxon OD co: Bo cae EA Ae ee ek ee. ee 420 aspera Maynard, 1920—Taxon 125 __.. 448 aviaria Maynard and Clapp in Maynard, 92 |= Taxon US = ose ie 2a Seen Pee ee 488 avita Maynard, 1913—Taxon 80 ____._._-..---------- 419 balaena Maynard and Clapp in Maynard, 192) 1—TVaxona S9ie_ 2 we 2 ee ee 490 balaene Maynard and Clapp in Maynard, 1921 (incorrect original spelling)—Taxon 189 .. 490 balanae Maynard, 1924? (misspelling)—Taxon USO. bse kecte has oe gS MONA sl Pies See ioe eee aeee 490 bimarginata Maynard, 1894—Taxon 45 _____.... 400 brownei Maynard, 1889—Taxon 22 _____..........- 386 caduca Maynard and Clapp in Maynard, 1920! fasent 12742 A ee ae 450 Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 caerulea Maynard and Clapp, 1915—Taxon LOVcai Ane. oll emund Hey Dain ee als oy Pople etal EA ho vs! 514 levigata Maynard, 1889—Taxon 2 375 lineota Maynard, 1889—Taxon 7 379 litorea Maynard and Clapp in Maynard, LOO PAXOM LOS Mee oe elem n Li eer nt 493 livida Maynard, 1924—Taxon 242 _.. 520 lobata Maynard, 1919—Taxon 118 442 longidentata Maynard, 1919 (nomen nudum) SEPP O LGR LOL ph Sree es as yancss i SE Ae ENN ited oe 437 macularia Maynard, 1913—Taxon 76 _ 418 mariae Maynard and Clapp in Maynard, LOZ O==Waxonwe bo iene te eee oe ye 460 marmorosa Maynard and Clapp in Maynard, 9202S Axon il aes erie leet. apr a 481 mayoi Maynard and Clapp in Maynard, 1921— PP ANOS iep enw ae Rae ny ae EN Th Ae eee Be 509 media Maynard, 1896—Taxon 58 407 migratoria Maynard and Clapp in Maynard, ROE axenie 2 Oe es een. Oe ert 508 minima Maynard, 1924—Taxon 243 _ 520 minuta Maynard and Clapp in Maynard, OZ 0==Faxomelo 2. 2b ee SE ripe, eth Lb 470 mitra Maynard and Clapp in Maynard, SZ 02 faxonslAa. wi 5 ain eee eh oe) 465 526 mixta Maynard and Clapp in Maynard, 1921— Taxon: S822 3. ee a ee oes 488 mobile Maynard and Clapp in Maynard, 192 l= Taxon 2ilo-8 2 ae ee ee: 506 montana Maynard, 1924—Taxon 240 519 morula Maynard and Clapp, 1915—Taxon 104 oA. ete Ti eae 1 a | Tee eee ee 432 multa Maynard, 1913—Taxon 94 428 muralia Maynard and Clapp in Maynard, LODE Taxont 22 Sk ba te ee ee ee yks} mutata Maynard, 1894—Taxon 31 393 mutatoria Maynard and Clapp in Maynard, 192022 Taxon: Alig) cae ee Se ee, Wie 464 nana Maynard, LS SO=== Ax @ eee 380 navalia Maynard, 1919 (nomen nudum)— axont GO: 24... Lobo. aes Pale 2. a 480 navalis Maynard and Clapp in Maynard, 19205—Taxont iGO; seek oe eee 480 nebula Maynard and Clapp in Maynard, ISAO ——Pewicoyiny WOE eee ets et cee 475 neglecta Maynard, 1894—Taxon 40 _. 398 nitela Maynard, 1889—Taxon 16 _ 384 nivea Maynard, 1894 (incorrect original ove inmey) I ey COn AG) nee 390 nivia Maynard, 1894—Taxon 26 (see Taxon TD) eRe AA. Se) les ore Sele 390 nivia Maynard, 1913—Taxon 72 (see Taxon DG ope re a Noe: ke. seni ole 3 the AR A hu te ect italied ohe 415 normandi Maynard and Clapp in Maynard, 1920 (incorrect original spelling)—Taxon GO sxseee te Pe ee EES Set eh 478 normanti Maynard and Clapp in Maynard, L920 == FaxOnnliGOx ce 2A eet, et tome ads A478 novita Maynard and Clapp in Maynard, LOD Faxon 2 Ooe = ee ee oa aoe Pe, nuda Maynard, 1889—Taxon 12 __ 381 oberholseri Maynard, 1913—Taxon 84 4292, obliterata Maynard, 1896—Taxon 53 404 obliterata Maynard, 1913 (homonym of Strophia scripta obliterata Maynard, 1896 [Taxon 53]; Strophiops sula Maynard and Clapp, 1915 [Taxon 105], is a replace- mmavermne mewn) ——Iesxorm G)5) 428 obscura Maynard, lS96—laxon) 60) 2a 408 obtusa Maynard and Clapp in Maynard, LOZ = TaxonvlOll 2) we eee 4992, orbicularia Maynard, 1889—Taxon 24 _______- 388 ornatula Maynard, 1913—Taxon 96 429 ornatulaclappii Maynard, 1913—Taxon 97 ___. 429 ornatularufina Maynard, 1913—Taxon 99 ____. 430 oscula Maynard and Clapp in Maynard, LOQI=— Taxon i218) oa eh 507 palida Maynard, 1889—Taxon 14 __ 382 palidula Maynard and Clapp in Maynard, AS PA eecona SAG) 504 pallida Maynard, 1889 (misspelling)—Taxon 14 1 i 55 5 oe ieee BET ie ee ee 382 palmata Maynard and Clapp in Maynard, 191.9 Paxon2iie 22 ee cea eee 444 panda Maynard and Clapp in Maynard, 1921 -—Taxon- 204) 2) ee ee 499 Bulletin Museum of Comparative Zoology, Vol. 158, No. 7 pannosa Maynard, 1889—Taxon 1] _ parva Maynard, 1889—Taxon 9 _ perantiqua Maynard and Clapp in Maynard, 192 0—Vaxon W509. 22> 6 ee eee peravita Maynard, 1919—Taxon 114 perplexa Maynard, 1889—Taxon 15 _ persuasa Maynard and Clapp in Maynard, 1920—Taxon 128 phoenecia Maynard and Clapp in Maynard, 1921 Maxong22jjyeel = 2 Oe) See eae Taxon 227 (3 sens i Se eee picta Maynard, 1889—Taxon 6 picturata Maynard and Clapp in Maynard, 1921—Taxon 199 pilsbryi Maynard, 1894—Taxon 47 _ plebea Maynard, 1920—Taxon 13] __ plebeia Maynard, 1920 (misspelling)—Taxon plebia Maynard and Clapp in Maynard, 1921 (misspelling)—Taxon 131 polita Maynard, 1896—Taxon 57 __ porcina Maynard and Clapp in Maynard, 1921-—Taxontl 90 ies)... Seah eee praedicta Maynard and Clapp, 1915—Taxon 1092 4611 a ee ees AR) Se praedivina Maynard, 1913—Taxon 90 ________ praedivinauniversa Maynard, 1913—Taxon predivina Maynard, 1921 (misspelling)—Taxon DO)» 2072 case hy es, Nt ae ee primigenia Maynard, 1913—Taxon 64 primordia Maynard and Clapp in Maynard, 1921-—-Vaxonvl SS), 8 Sais! 2 Pe ee DO) 32 oni oats SOS NE aes Tepes Boeeaeie eee proavita Maynard and Clapp in Maynard, 192 1—Taxonv SO) = see oe eee eee processa Maynard and Clapp in Maynard, 192022 taxon e410 eee procliva Maynard and Clapp in Maynard, IC AA fey cory ICG uk a profunda Maynard and Clapp in Maynard, 1921—Taxon 182 prognata Maynard and Clapp in Maynard, 1920—Taxon 144 progressa Maynard and Clapp in Maynard, IES PAO Penouny WIZAM pulla Maynard and Clapp in Maynard, 1920— Taxon 159 pumilia Maynard, 1894—Taxon 37 pumilla Maynard, 1894 (misspelling in the index)=—TaxOmesi(i et ele purpura Maynard, 1919, 1924 (misspelling)— Taxon 75 purpurea Maynard, 1913—Taxon 75 _.... pusilla Maynard and Clapp in Maynard, 1.99.0 =Waxonsli7o Geka eer cee eee pygmea Maynard, 1924—Taxon 245 ralla Maynard and Clapp in Maynard, 1921— Taxon 208 426 424 411 490 436 486 464 494 487 466 467 A474 395 395 416 416 482 521 THE CERION TAXA OF CHARLES JOHNSON MAYNARD * Harasewych et al. rara Maynard and Clapp in Maynard, 1921— “TBS SON mail INC Ys eine ease eres ee ee Ee Oe 494 recessa Maynard and Clapp in Maynard, HOGS Taxomell2 Ome oe. 2s ee 443 rediviva Maynard, 1913—Taxon 73 __..... AMS regula Maynard, 1894—Taxon 44 ___. 400 regular Maynard, 1919 (misspelling)—Taxon ZA LA Na EIN GL WT a WS SR REINS cee Re 28 aE ee Tt 400 reincarnata Maynard and Clapp in Maynard, MO Oi AX OME Aig eee mee i a 509 relequa Maynard and Clapp in Maynard, 1921 (misspelling)—Taxon 184 _ 488 reliqua Maynard and Clapp in Maynard, ROME saxo Ares eee eo ee, eee 488 repetita Maynard and Clapp in Maynard, 2 OS Rea @ MM Gee re coe sere oe eee 461 repitita Maynard and Clapp in Maynard, 1920 (ronments mica) ——ax@mel oS) omenene nna unnene 461 restricta Maynard, 1894—Taxon 49 _. 401 ritchei Maynard, 1920 (misspelling)—Taxon 34 Je CoN ERO 9 SE ESE A De, SE EO eee te 394 ritchiei Maynard, 1894—Taxon 34 __ 394 robusta Maynard, 1894—Taxon 29 392 rosacea Maynard and Clapp in Maynard, OM——Tenorn SAVE) o01 rosea Maynard and Clapp in Maynard, 1921— Tey coni OAC) Re as cate res ae ae Ae, Sete a DIES rubiginosa Maynard and Clapp in Maynard, IS Teoin BOA a ee 508 rufumaculata Maynard, 1913—Taxon 77 418 rufina Maynard, 1913—Taxon 100 _ 430 rufula Maynard, 1924—Taxon 238 _. 518 salinaria Maynard, 1913—Taxon 65 _ All sampsoni Maynard and Clapp in Maynard, O20 = axOnmlod pee ees en le ale 472, santesoni Maynard, 1G? 0= axon yeas 453 saxitina Maynard and Clapp in Maynard, UG PAs Pep Once abe i ee 904 scalariformis Maynard, 1919—Taxon 115 ____ 438 scripta Maynard, 1896—Taxon 52 __ 404 scutata Maynard and Clapp in Maynard, NO FNS lax Ont ae ee eee ee Boer Log 494 semipolita Maynard and Clapp in Maynard, N92 O==Waxonel Ong eaee aes ele mere ey A479 similaria Maynard and Clapp in Maynard, NOD Maou Greets Wee oT te oy See 484 sparsa Maynard, 1924—Taxon 237 516 stroutii Maynard and Clapp in Maynard, L920 SSVaxonel lta eer eh ie See ee 469 stupida Maynard and Clapp in Maynard, 1921—Taxon 201 sula Maynard and Clapp, 1915 (new name for obliterata Maynard, 1913)—Taxon 105 __. sylvatica Maynard and Clapp in Maynard, OD aoe Olena a cade se a tabida Maynard, 1913—Taxon 101 tenucostata Maynard and Clapp in Maynard, 1920—Taxon 155 tenui Maynard and Clapp, 1915—Taxon 111 ~ territa Maynard, 1920—Taxon 133 __ thayeri Maynard and Clapp in Maynard, 1921—Taxon 206 thayerii Maynard and Clapp in Maynard, 1921(misspelling)—Taxon 206 thompsoni Maynard and Clapp, 1915—Taxon COS. eee 20 irae eae 0 ager 5 eee thorndikei Maynard, 1894—Taxon 27 ___. thorndikeii Maynard, 1921 (misspelling)— Taxon 27 tibida Maynard, 1921 (misspelling)—Taxon tracta Maynard, 1894—Taxon 30 __ transitoria Maynard, 1913—Taxon 86 ____. transimutata Maynard and Clapp in Maynard, 1921 (misspelling)—Taxon 177 __ transmutata Maynard and Clapp in Maynard, MS) TST Retox opines ih 0h en ses ee Pe ae a travelii Maynard and Clapp in Maynard, 1921 (incorrect original spelling)—Taxon 198 travellii Maynard and Clapp in Maynard, 1921—Taxon 198 ultima Maynard, 1913—Taxon 79 ______-.- uniformis Maynard, 1913—Taxon 87 universa Maynard, NOKS==axonn92 ee aes vagabunda Maynard and Clapp in Maynard, 1925—Taxon 248 valida Maynard and Clapp in Maynard, 92 0==Taxom WEOe se eae et ee, varianivia Maynard 1913—Taxon 71 variapurpurea Maynard, 1913—Taxon 74 ___. variata Maynard and Clapp in Maynard, OTS Saxons lle ares eae IE variathorndikei Maynard, 1913—Taxon 70 __. veta Maynard and Clapp in Maynard, 1920— Arax@ milla Oran wees eS eke h IR ee vetusta Maynard, 1913—Taxon 81 ____- vetustapraedevina Maynard, 1913—Taxon a27 wow eat we Lee ‘al rf a ‘ ? iat ee! ee Ad her ria a Pereil Me ‘ , y ie Le tay ae oy os 7 aa ae " \ at »8 eo tas moh at. " sb a A 090 co ie : Nib wit iprewae s ssa ee bss ba ties a ee ¥ . 4 t pr) y