AMERICAN MALACOLOGICAL BULLETIN THE NATURAL-1 HISTORY **MSEUM ' - 1 APR 2010 PUrtUiA>i:D | ZOOLOGY LIBRARY 1 Journal of the American Malacological Society http://www. malacological. org volume 28 26 February 2010 number 1/2 Invited Paper Dams, zebras, and settlements: The historical loss of freshwater mussels in the Ohio River mainstem. G. THOMAS WATTERS and CAROL J. MYERS FLAUTE 1 Symposium Papers Current research on land snails and land snail conservation: Leslie Hubricht Memorial Symposium on Terrestrial Gastropods. KATHRYN E. PEREZ 13 Leslie Hubricht (1908-2005), his publications and new taxa. JOCHEN GERBER 15 Pupillid land Snails of eastern North America. JEFFREY C. NEKOLA and BRIAN F. COLES 29 The lesser families of Mexican terrestrial molluscs. EDNA NARANJO-GARCIA and NEIL E. FAHY 59 Diversity and conservation of the land snail fauna of the western Pacific islands of Belau (Republic of Palau, Oceania). REBECCA J. RUNDELL 81 Analysis of museum records highlights unprotected land snail diversity in Alabama. RUSSELL L. MINTON and KATHRYN E. PEREZ 91 continued on back cover Kenneth M. Brown, Editor-in-Chief Department of Biological Sciences Louisiana State University Baton Rouge, Louisiana 70803, U.S.A. AMERICAN MALACOLOGICAL BULLETIN BOARD OF EDITORS Cynthia D. Trowbridge, Managing Editor Oregon Institute of Marine Biology P.O. Box 1995 Newport, Oregon 97365, U.S.A. Janice Voltzow Department of Biology University of Scranton Scranton, Pennsylvania 18510-4625, U.S.A. Robert H. Cowie Center for Conservation Research and Training University of Hawaii 3050 Maile Way, Gilmore 408 Honolulu, Hawaii 96822-2231, U.S.A. Carole S. Hickman University of California Berkeley Department of Integrative Biology 3060 VLSB #3140 Berkeley, California 94720, U.S.A. Paula M. Mikkelsen Paleontological Research Institution 1259 Trumansburg Road Ithaca, New York 14850-1313, U.S.A. Alan J. Kohn Department of Zoology Box 351800 University of Washington Seattle, Washington 98195, U.S.A. Dianna Padilla Department of Ecology and Evolution State University of New York Stony Brook, New York 11749-5245, U.S.A. Roland C. Anderson The Seattle Aquarium 1483 Alaskan Way Seattle, Washington 98101, U.S.A. Timothy A. Pearce Carnegie Museum of Natural History 4400 Forbes Avenue Pittsburgh, Pennsylvania 15213-4007, U.S.A. Janet Voight The Field Museum 1400 S. Lake Shore Dr. Chicago, Illinois 60605-2496, U.S.A. The American Malacological Bulletin is the scientific journal of the American Malacological Society, an international society of professional, student, and amateur malacologists. Complete information about the Society and its publications can be found on the Society’s website: h ttp://www. malacological. org AMERICAN MALACOLOGICAL SOCIETY MEMBERSHIP MEMBERSHIP INFORMATION: Individuals are invited to com- plete the membership application available at the end of this issue. SUBSCRIPTION INFORMATION: Institutional subscriptions are available at a cost of $75 plus postage for addresses outside the U.S.A. Further information on dues, postage fees (for members outside the U.S.A.), and payment options can be found on the membership application at the end of this issue. ALL MEMBERSHIP APPLICATIONS, SUBSCRIPTION ORDERS, AND PAYMENTS should be sent to the Society Treasurer: Dawn E. Dittman Tunison Laboratory of Aquatic Science 3075 Gracie Rd. Cortland, New York 13045-9357, U.S.A. CHANGE OF ADDRESS INFORMATION should be sent to the Society Secretary: Amanda Lawless Department of Malacology The Academy of Natural Sciences of Philadelphia 1900 Benjamin Franklin Parkway Philadelphia, Pennsylvania 19103-1195, U.S.A. INFORMATION FOR CONTRIBUTIONS is available on-line and appears at the end of this issue. MANUSCRIPT SUBMISSION, CLAIMS, AND PERMISSIONS TO REPRINT JOURNAL MATERIAL should be sent to the Editor-in-Chief: Kenneth M. Brown, Editor-in-Chief Department of Biological Sciences Louisiana State University Baton Rouge, Louisiana 70803, U.S.A. Voice: 225-578-1740 • Fax: 225-578-259 7 E-mail: kmbrown@lsu.edu AMERICAN MALACOLOGICAL BULLETIN 28(1/2) AMER. MALAC. BULL. ISSN 0740-2783 Copyright © 2010 by the American Malacological Society Cover photo: Sacoglossan opisthobranchs from Okinawa, Japan; clockwise from top left: Elysia rufescens, Stiliger ornatus, Caliphylla sp., and Placida sp. Sacoglossans offer a model system to investigate trophic specialization (Trowbridge et al. 167-181). HISTORY MUSEUM - 1 APR 2010 I PURCHASED j ZOOLOG Y LIBRARY AMERICAN MAJLACOLOGICAL BULLETIN CONTENTS VOLUME 2 8 | NUMBER l/2 Invited Paper Dams, zebras, and settlements: The historical loss of freshwater mussels in the Ohio River mainstem. G. THOMAS WATTERS and CAROL J. MYERS FLAUTE Symposium Papers Current research on land snails and land snail conservation: Leslie Hubricht Memorial Symposium on Terrestrial Gastropods. KATHRYN E. PEREZ 13 Leslie Hubricht (1908-2005), his publications and new taxa. JOCHEN GERBER 15 Pupillid land snails of eastern North America. JEFFREY C. NEKOLA and BRIAN F. COLES 29 The lesser families of Mexican terrestrial molluscs. EDNA NARANJO-GARCIA and NEILE.FAHY 59 Diversity and conservation of the land snail fauna of the western Pacific islands of Belau ( Republic of Palau, Oceania ) . REBECCA J. RUNDELL 81 Analysis of museum records highlights unprotected land snail diversity in Alabama. RUSSELL L. MINTON and KATHRYN E. PEREZ 91 Strategies for collecting land snails and their impact on conservation planning. MARLA L. COPPOLINO 97 Surfing snails: Population genetics of the land snail Ventridens ligera (Stylommatophora: Zonitidae) in the Potomac Gorge. COLLEEN S. SINCLAIR 105 Reproductive biology and the annual population cycle of Oxyloma retusum (Pulmonata: Succineidae). AYDIN ORSTAN 113 Independent Papers Distribution, density, and population dynamics of the Anthony Riversnail (Athearnia anthonyi) in Limestone Creek, Limestone County, Alabama. JEFFREY T. GARNER and THOMAS M. HAGGERTY 121 Epiphyton or macrophyte: Which primary producer attracts the snail Hebetancylus moricandP ROGER PAULO MORMUL, SIDINEI MAGELA THOMAZ, MARCIO JOSE DA SILVEIRA, and LILIANA RODRIGUES 127 Distribution and environmental influences on freshwater gastropods from lotic systems and springs in Pennsylvania, USA, with conservation recommendations. RYAN R. EVANS and SALLY J. RAY 135 Fish hosts of the Carolina heelsplitter ( Lasmigona decora ta), a federally endangered freshwater mussel (Bivalvia: Unionidae). CHRIS B. EADS, ROBERT B. BRINGOLF, RENAE D. GREINER, ARTHUR E. BOGAN, and JAY F. LEVINE 151 Population studies of an endemic gastropod from waterfall environments. DIEGO E. GUTIERREZ GREGORIC, VERONICA NUNEZ, and ALEJANDRA RUMI 159 Subtropical sacoglossans in Okinawa — at “special risk” or “predictably rare”? CYNTHIA D. TROWBRIDGE, YAYOI M. HIRANO, YOSHIAKI J. HIRANO, KOSUKE SUDO, YOICHI SHIMADU, TOMOHIRO WATANABE, MAKIKO YORIFUJI, TARO MAEDA, YUKI ANETAI, and KANAKO KUMAGAI 167 Research Notes Self-adhesive wire markers for bivalve tag and recapture studies. LANCE W. RILEY, SHIRLEY M. BAKER, and EDWARD J. PHLIPS 183 Occurrence of the red abalone Haliotis rufescens in British Columbia, Canada. ALAN CAMPBELL, RUTH E. WITHLER, and K. JANINE SUPERNAULT 185 Index to Vol. 28 189 Society Business Seventy-five years of molluscs: A history of the American Malacological Society on the occasion of its 75th annual meeting PAULA M. MIKKELSEN 191 James W. Nybakken: September 16, 1936 - June 20, 2009 An Appreciation ALAN J. KOHN 215 Information for Contributors 2010 219 Membership Form 2010 221 Financial Report 2007 222 AMS/WSM Meeting Announcement 223 ii Amer. Make. Bull. 28: 1-12 (2010) INVITED PAPER* Dams, zebras, and settlements: The historical loss of freshwater mussels in the Ohio River mainstem G. Thomas Watters1 and Carol J. Myers Flaute2 1 Department of Evolution, Ecology and Organismal Biology, The Ohio State University, 1315 Kinnear Road, Columbus, Ohio 43212, U.S.A. 2 College of Environment and Design, The University of Georgia, Athens, Georgia 30602, U.S.A. Corresponding author: Watters.l@osu.edu Abstract: The decline of the freshwater mussel fauna of the Ohio River, U.S.A. is compared to the dates of service of the existing dams, the arrival of the exotic zebra mussel, and the presence of urban centers on the mainstem. Based upon historical records we know that most pools supported 20-50 species of mussels; today many have fewer than ten. The results presented here show a mixed effect of the dams on the mussel fauna, ranging from marked deleterious effects in Hannibal, McAlpine, and Smithland pools to comparatively little effect in pools such as Dashields, Greenup, and Markland. In nearly all cases, the most dramatic declines in mussels were associated with the arrival of zebra mussels in the Ohio River in 1991. Pools with significant urban centers often had a loss of diversity well before the construction of dams or the arrival of zebra mussels; these losses are attributed to water quality problems associated with urban centers. Mussel diversity has thus declined in the Ohio River as the result of a three-fold problem: loss of water quality, existing dams, and zebra mussels. Key words: Unionacea, diversity, impoundments, Dreissenidae, water quality “The Ohio is the most beautiful river on earth. Its current gentle, waters clear, and bosom smooth and unbroken by rocks and rapids, a single instance only excepted ” (Thomas Jefferson 1782: 256). The Ohio River system is one of the largest in the world (1,579 km long with a watershed area of 490,603 km2). It was an important navigational route for both Native Americans and European settlers. Forts were established on the river by the early 1700s, many of which became large urban centers (Pittsburgh, Cincinnati, Louisville, etc.). Early on it was rec- ognized that the river supported a diverse assemblage of freshwater mussels. Historically the Ohio River is thus one of the most important rivers in North America to the science of freshwater malacology. It was at the Falls of the Ohio that Rafinesque, one of the most influential (and eccentric) natu- ralists of the Ohio region, had his epiphany. Thomas Lea, a citizen of Cincinnati, sent specimens to his brother Isaac Lea in Philadelphia, the most prolific freshwater malacologist in North America. Because of the long history of human use, it is not surprising that the river has suffered some indignities. It has been impounded with numerous dams, both for navi- gation and hydropower (Fig. 1). In more recent times, it has been invaded by the exotic zebra mussel. However, early cit- ies began to impact the river’s water quality and mussel fauna long before the arrival of the current dams and the zebra mussels. That impoundments are detrimental to aquatic life in general, and mussels in particular, has been demonstrated by many case studies (see review of Watters 2000). Perhaps several dozen mussel species in North America, and numer- ous more freshwater snails, have been driven to extinction by the effects of impoundments (Stansbery 1973, Layzer et al. 1993, Lydeard and Mayden 1995). Although mussel faunas change through time, with or without dams, and species become extinct as a matter of course, the interference of humanity has dramatically accelerated this process. For exam- ple, the Tennessee River’s mussel diversity decreased from 100 species to 44, mainly due to impoundment (Isom 1969). In the Fort Loudoun Reservoir on the Tennessee River, Isom (1971) found only four mussel species, whereas Ortmann (1918) reported 64 species from the same general area before impoundment. The area of the Chickamauga Reservoir of the Tennessee River supported 46 species for perhaps 2000 years This is a paper invited by the editor, based on a presentation given at the 2008 AMS meeting in Carbondale. Tom Watters has been responsible for a number of influential papers, some published in the AMB. This paper elegantly shows the effects that urbanization, impoundments, and invasive species have had on unionids in the Ohio River. AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 1. The Ohio River mainstem with locations of dams. 1, Ems- worth; 2, Dashields; 3, Montgomery; 4, New Cumberland; 5, Pike Is- land; 6, Hannibal; 7, Willow Island; 8, Belleville; 9, Racine; 10, Byrd; 1 1, Greenup; 12, Meldahl; 13, Markland; 14, McAlpine; 15, Cannel- ton; 16, Newburgh; 17, Myers; 18, Smithland; 19, Olmsted. prior to impoundment (Parmalee etal. 1982). After impound- ment, 28 species were extirpated, and several are now extinct. Similar reductions in diversity after impoundment were documented for the Cumberland River (Schmidt 1986, Layzer et al. 1993), where the construction of the Center Hill hydroelectric dam resulted in the loss of 78% of the original mussel species. In the Little Tennessee River, only six of the original 50 mussel species at Tellico Lake remained after impoundment (Parmalee and Hughes 1993). The mus- sels of other rivers have met a similar fate, such as the Kaskaskia River (Suloway et al. 1981) and the Tombigbee River (Williams et al. 1992). After impoundment, the original mussel fauna may be eliminated or greatly reduced (Holland-Bartels 1990), or changed in favor of silt-tolerant species, such as ano- dontines and species of Leptodea and Potamilus (Clark and Gillette 1911, Ellis 1931, Bates 1962, Isom 1969, Klippel and Parmalee 1979, Parmalee and Hughes 1993, Blalock and Sickel 1996). Thus while some impoundments con- tain numerous mussels, these invariably are invading, soft-substrate-adapted species that have replaced the orig- inal fauna. In addition to affecting mussels, dams may be physical barriers to mussel hosts as well. Declines in fish diversity due to dams, particularly in migrating fish, have been document- ed (Branson 1974, Hubbs and Pigg 1976, Steubner 1993, Col- lier et al. 1996). Some mussel populations may no longer be recruiting juveniles because their hosts are absent (Suloway et al. 1981, Jones 1991, Burkhead et al. 1992). Lowhead dams may be as disruptive as high-lift dams (Dean et al. 2002); Watters (1996) showed that dams as low as one meter in height affected host movements and restricted the range of some mussels. The recently introduced zebra and quagga mussels are highly detrimental to native mussels (Haag etal. 1993, Strayer 1999). Zebra mussels compete with native mussels for food, consume native mussel gametes, interfere with the closure of the native mussels’ shells, and result in beach strandings of native mussels (Parker et al. 1998). Since their introduction, zebra mussel densities have fluctuated widely in the Ohio River and elsewhere. However, where native and zebra mus- sels coexist, zebra mussels often result in the local extirpation of native mussels (Gillis and Mackie 1994, Nalepa 1994). For example, native mussels have been nearly extirpated from the western basin of Lake Erie by zebra and quagga mussels (Ecological Specialists, Inc. 1999). Before the current dams were completed and zebra mussels arrived, the Ohio River had been impacted by the rapidly growing urban centers along its length. Unchecked waste water discharge, logging, and mining all contributed to a decrease in water and habitat quality. Undoubtedly, water quality degradation, impound- ments, and zebra mussels have had a negative impact on native mussels in the Ohio River, but the degree and timing of these changes have not been gauged. Fortunately, the Ohio River was sampled extensively by early naturalists and later by professional survey crews. This fact allows us to examine the changes in the mussel fauna of this river over nearly two centuries. MATERIALS AND METHODS Historical records Mussel records were derived from the unpublished data- base assembled by H. Dunn (Ecological Specialists, Inc.), the survey of Williams and Schuster (1989), the surveys of the USFWS Ohio River Islands National Wildlife Refuge, and the collection records of the Ohio State University Museum of Biological Diversity (OSUM). The river mile, date of oper- ation, and numbers of mussel taxa in each pool are given in Table 1. The presence of mussel species, regardless of year, in each pool is given in the Appendix. Data were not available for individual Pools 52 and 53; these were combined into the Olmsted Pool in this study. The Cairo “Pool” is that area from the confluence to Olmsted and is formed by the impoundment of the Mississippi River backing up into the Ohio River. Cumulative mussel species richness was based on the assumption that species recorded from a later date had always existed in the pool prior to that survey. For example, a species first recorded in 1980 was considered to have existed in the pool back to 1800. This assumption is based on the nature of mussel surveys in the Ohio River. Prior to intensive sampling beginning in the 1980s the Ohio River had only been sampled by specimen collectors or commercial clammers. These col- lectors may not have comprehensively sampled any given area, and certainly did not have access to the deeper reaches LOSS OF MUSSELS IN OHIO RIVER 3 Table 1. Ohio river dams included in this study and their maximum recorded mussel species richness. Unlike most river systems, river miles on the Ohio River are numbered from the source (Pittsburgh) to the confluence (Cairo). The “date placed in service” may span the years of first construction of locks to completion of the present dam. Approximate dates are given for the arrival of zebra mussels in each pool. Sizeable population centers are given for each pool. *The Cairo “Pool” is that area from the confluence to Olmsted and is formed by impoundment of the Mississippi River backing up into the Ohio River. N/A, not available. Dam Ohio river mile Date placed in service # mussel species Zebra mussel arrival Population centers Emsworth 6.2 1919-1922 N/A 1994 Pittsburgh, Pennsylvania Dashields 13.2 1927-1929 48 1994 Montgomery 31.7 1932-1936 2 1994 Rochester, Pennsylvania New Cumberland 54.4 1955-1961 8 1994 Pike Island 84.2 1959-1965 5 1994 Steubenville, Ohio Hannibal 126.4 1967-1975 21 1993 Willow Island 161.7 1972-1976 24 1993 Belleville 203.9 1963-1968 46 1993 Marietta, Ohio Racine 237.5 1965-1967 41 1993 Parkersburg, West Virginia Byrd 279.2 1937 27 1993 Pt. Pleasant, West Virginia Greenup 341.0 1955-1962 37 1992 Gallipolis, Ohio Meldahl 436.2 1959-1964 48 1992 Huntington, West Virginia Portsmouth, Ohio Markland 531.5 1959-1964 63 1992 Maysville, Kentucky Cincinnati, Ohio McAlpine 606.8 1961 45 1992 Cannelton 720.7 1963-1974 40 1992 Louisville, Kentucky Newburgh 776.1 1969-1975 30 1992 Owensboro, Kentucky Myers 846.0 1969-1977 35 1992 Evansville, Indiana Smithland 918.5 1971-1980 35 1992 Olmsted 938.9-962.6 1928-1929 29 1991 Paducah, Kentucky (Dams 52, 53) Cairo* confluence N/A 33 1991 now accessible by modern scuba techniques, except through the biased sampling of the crow-foot brail, a device dragged behind a boat to snag mussels. Thus many species were missed in earlier efforts that have since been collected. Although some recently collected species may in fact be introductions or taxa increasing their ranges, these species are a small per- centage of the overall fauna. Anodonta suborbiculata Say, 1831 and Quadrula aspera (Lea, 1831) maybe the only recent arriv- als (Watters, unpubl. data) although soft-substrate-adapted species such as anodontines may be more abundant now than in the past. Because of this, individual pool species richness may be slightly over-estimated. Based upon the data presented here, the cumulative number of mussel species per pool is plotted as a function of time (Fig. 2). The dates of data for each pool range from 1800 to 2000. The dates of operation of each dam are overlain, as is the general advent of zebra mussels in the Ohio River (begin- ning in 1991). RESULTS The mussel data for several pools were insufficient to draw any conclusions because of a lack of historical records. These pools are not further considered: Byrd, Cairo, Ems- worth, Montgomery, New Cumberland, Newburgh, Olm- sted, Pike Island, Racine, and Willow Island. Results for remaining pools and averages are illustrated (Fig. 2). Pool by pool accounts (in alphabetical order) Belleville Belleville had among the best historical records of any Ohio River pool, dating back to the early 1800s. Except for a small decline in the 1880s, the diversity had remained fairly constant until the late 1970s, despite the presence of Marietta and Parkersburg in the pool. This decline may reflect the con- struction of the dam in the 1960s. However, a marked decline 4 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2 -2010 1790 1820 1850 1880 1910 1940 1970 2000 1790 1820 1850 1880 1910 1940 1970 2000 Belleville Greenup o z o z 1790 1820 1850 1880 1910 1940 1970 Smithland O 2000 z 50 1790 1820 1850 1880 1910 1940 1970 2000 1790 1820 1850 1880 1910 1940 1970 2000 Meldahl Markland 0 2 O Z 20 10 50 40 30 20 10 0 1790 1820 1850 1880 1910 1940 1970 2000 1790 1820 1850 1880 1910 1940 1970 2000 McAlpine Cannelton O z O z 50 1 “l \ 40 • 1 20 • , , , , , , , 0 ■ A 1790 1820 1850 1880 1910 1940 1970 Average O 50 ■ 40 30 ■ 20 • 10 • 0 ■ 2000 z 1790 1820 1850 1910 1940 1970 2000 1790 1820 1850 1910 1940 1970 2000 1790 1820 1850 1880 1910 1940 1970 2000 Year Figure 2. Mussel species richness (y-axis) plotted against time (x-axis) for each Ohio River Pool and averaged across all pools. O, dam placed in operation; Z, arrival of zebra mussels in river. The width of the grey bars spans the years of present dam construction and the arrival of zebra mussels. Pools are arranged from upstream to downstream. was associated with the advent of zebra mussels. Of the sev- eral endangered species recorded from the reach, only Cyprogenia stegaria (Rafinesque, 1820) apparently still existed during the last survey. Cannelton Cannelton mussel records dated to -1900 and suggested a robust original fauna of -40 species. Louisville is at the McAlpine Dam within this pool but there are insufficient data to document its effects, if any. The extinct Epioblasma torulosa (Rafinesque, 1820) and other very rare species were found as subfossil material in 2000, but when they were lost from the pool is not known. Most of the fauna persisted until the construction of the dam in 1972, then declined during the late 1970s into the 1980s. However, by comparison, mussel diversity plummeted after the arrival of zebra mussels. Dashields The Dashields Pool had one of the longest records of mussels in the river, dating to the 1800s. Data indicated that this reach once supported a thriving mussel community of perhaps 40 species, including the extinct Epioblasma torulosa, last seen in 1800. Recent surveys have found fewer than ten species. It is clear that the decline began as early as the late 1800s, undoubtedly due to water quality degradation down- stream of Pittsburgh. The dam was not placed in operation until 1929 and no further surveys were conducted until the 1970s. At that time, the pool probably still supported over 20 species. No marked decline attributable to the dam was apparent in the data superimposed as it was on the already existing decline. The fauna dramatically declined with the arrival of zebra mussels in 1991. The most recent fauna was largely comprised of mussels using freshwater drum or LOSS OF MUSSELS IN OHIO RIVER 5 catfish as their hosts. The endangered Cyprogenia stegaria probably still exists in the pool. Greenup Greenup mussel data showed an alarming decline in diversity after the arrival of zebra mussels — from nearly 40 species to fewer than five. The decline associated with the dam was relatively minor. Huntington and Portsmouth occur in the pool but records were insufficient to demonstrate any loss of diversity as a result. The endangered Lampsilis abrupta (Say, 1831) still occurred in this reach in a recent survey. Hannibal Mussel data dated from the early 1900s. The negative effects of both dam construction and zebra mussels were evi- dent. No major urban centers exist in the pool. The only endangered mussel recorded from the pool was Pleurobema plenum (Lea, 1840), last seen in 1901. Markland The Markland Pool may have had more mussel data than any other Ohio River reach. Data were available from the 1800s through 2000, revealing an original mussel diversity probably exceeding 60 species. A marked decline in mussel diversity occurred in the 1840s. The fauna slowly declined to the early 1980s until the construction of the dam. This loss of diversity is probably caused by water quality problems associ- ated with Cincinnati. An accelerated decline occurred at this time, followed by yet another decline when zebra mussels arrived. Numerous rare, endangered, and extinct species once lived in this reach. With the exception of Cyprogenia stegaria, apparently still living here, none of these species have been seen in the past 50 years. McAlpine Mussel data were available from 1800. The lack of large urban centers in the pool is apparent in the high and continuous level of diversity for over 150 years. The negative effect of dam construction on the mussel fauna was obvious. Mussel diversity had declined dramatically to -1990, the last date for which we have data. Several rare or endangered mussels were known only as weathered or subfossil shells — Cyprogenia stegaria, Lampsilis abrupta, Obovaria retusa (Lamarck, 1819), Plethoba- sus cicatricosus (Say, 1829), Pleurobema clava (Lamarck, 1819), and Pleurobema plenum. Meldahl Mussel data dated to the early 1900s. Over 50 mussel species originally lived in this river reach. Surveys revealed a slow but constant decline in mussel diversity before and after the construction of the dam; the arrival of zebra mussels led to a severe decline in diversity. The early decline may be attributable to the presence of Maysville in the pool. One extinct mussel once occurred here, Epioblasma torulosa, as well as the very rare Epioblasma obliquata obliquata (Rafinesque, 1820). Several other endangered species also were recorded from here, but none have been seen in the past 50 years. Myers (Uniontown) We had mussel data for the Myers Pool dating back to the early 1900s. Evansville is within this pool but there are insufficient data to document its effects, if any. Mussel diver- sity seemed to have remained fairly constant until the advent of zebra mussels, with a small decline after dam construction. After the arrival of zebra mussels, mussel diversity precipi- tously declined. The Myers Pool had two mussel species that are usually considered more southern taxa — Plectomerus dombeyanus (Valenciennes, 1827) and Quadrula aspera. This pool probably represented the northernmost extent of these species. Smithland Records for the Smithland Pool dated to 1800 and clearly showed the impact of impoundment in 1980. Before that time, there does not appear to have been a dramatic decline, testimony to the fact that no large urban centers exist in the pool. After impoundment, the fauna was reduced from its original 36 taxa to fewer than 15. This included the extinct Epioblasma torulosa and several endangered species. Effects from zebra mussels were not as dramatic here as in other pools. Average across all pools Mussel species richness showed a steady decline from the earliest surveys. The construction of the present dams contin- ued this decline, particularly beginning in the late 1970s when the faunal loss accelerated. This loss quickened with the arrival of zebra mussels. DISCUSSION The Ohio River has been sampled for freshwater mussels since at least the 1700s. The records used here date back to 1800 in several instances, providing a lengthy record of mus- sels in several pools. Prior to impoundment, the average depth of the upper Ohio River was -0.3 m deep. Frontier accounts clearly depict the upper Ohio River as an often wadeable, meandering series of riffles, runs, and pools (Eck- ert 1996). Mark Twain (1917: 39) remarked that “We reached Louisville - at least the neighborhood of it. We stuck hard and fast on the rocks in the middle of the river, and lay there four days” and “The Ohio river was friz to the bottom - which 6 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 warn’t no great shakes in the freezing line, considering that krick aint never got more’n forty barls of water in it, no how.” Walt Whitman (1848) reminisced “The bottom of the boat grated harshly more than once on the stones beneath, and the pilots showed plainly that they did not feel altogether as calm as a summer morning.” Mussels in relatively unaltered riverine systems occupy distinct river reaches: some species are most common in rif- fles and runs whereas others prefer pools and backwaters. This was undoubtedly the case for the Ohio River as well. The construction of dams eliminated the riffles and pools, pre- sumably driving some species (and/or their hosts) to extirpa- tion or even extinction. Most of the currently extinct North American mussel species, predominately in the genus Epio- blasma , were apparently big river riffle/run species. The results presented here show a mixed effect of current dams on the mussel fauna. In pools such as Hannibal, McAlpine, and Smithland, the effects are marked. But in oth- er pools comparatively little effect is apparent: Dashields, Greenup, and Markland, for example. In some cases it is dif- ficult to separate a dam effect from an existing water quality effect. Often the dam effect does not appear to be immediate; in some cases, nearly a decade passes before declines are apparent. This may be due to two causes. First, it may be an artifact created by the timing of the next survey after dam construction. If the next survey occurred ten years after the dam was placed in operation, then the diversity estimate would not have been updated until that time. Second, there may be a biological reason for the lag time. This may reflect the lack of recruitment in some mussel populations while their adults age and eventually die off in the pool. However, there is little evidence for the latter interpretation in the results presented here. Where data are available, there appears to have been an immediate, if often slight, decline in mussel diversity once the current dams were constructed. Once zebra mussels invaded, their impact on native mussels was also immediate. The current suite of 18 high-lift dams replaced an ear- lier system of 51 movable wicket and lock dams, although even earlier dams date to 1885. These wicket dams were placed in service in 1929. There is no evidence that they seri- ously impacted the mussel fauna. This is probably due to the fact that the dams did not permanently impound the river — during normal flow the dams were lowered. Based upon historical records, we know that most pools supported 20-50 species of mussels; today many have fewer than ten. By contrast, most pools now have 60-80 species of fish (Watters et al. 2003). Unlike mussel species, most fish spe- cies have persisted in pools for the past several decades. Com- parisons with the known potential hosts for the mussels in each pool indicate that the decline of native mussels is not obviously linked to a loss of available hosts. While it is possible that some very host-specific mussels, perhaps now extinct, may have been affected by the loss of necessary fish, this is probably the exception. Therefore, the loss of native mussels does not seem to be attributable to a loss of hosts, as most pools have a wide variety of fish that could act as hosts to these mussels. For some pools with extensive chronological data, such as Dashields and Markland, it is evident that the mussel fauna had already suffered several declines in the 1800s prior to cur- rent dam construction. These declines may be associated with the rapid growth and industrialization of the Ohio River cor- ridor in general — logging, mining, discharges, etc. The Dashields river reach is only 21 km below Pittsburgh and shows a significant decline in diversity beginning nearly 50 years prior to current dam construction. The Markland river reach contains Cincinnati and had been in decline for over 120 years prior to the current dam construction. By contrast, less affected reaches had smaller urban centers; for example, Marietta and Parkersburg are the only sizeable urban centers in the Belleville reach. The Smithland reach is devoid of large urban centers, as is the McAlpine pool, with the exception of Louisville at its extreme downstream reach. For pools having sufficient data, pools lacking urban centers do not show a pre-existing decline in diversity prior to dam construction. Thus, in some cases the completion of the present dams seems to have been part of an ongoing decline originated by other factors associated with urbanization and industrialization, the Markland and Meldahl pools being good examples. Land surface temperatures in the Ohio Valley have increased since 1880 an average of 0.05 °C/decade, largely after 1980 (National Oceanic and Atmospheric Administra- tion 2009). Although mussel reproductive cycles are believed to be tied to water temperature variation (Watters and O’Dee 2000, Watters et al. 2001), the effect of these slight changes is unknown. We do not believe there is any evidence for cli- matic warming in the Ohio Valley impacting native mussels at this time. Zebra mussels first appeared in the lowest reach of the Ohio River in 1991 and had spread upstream to Pittsburgh, Pennsylvania by 1994 (U. S. Geological Survey 2009). In nearly all cases the most dramatic declines in mussels are associated with the arrival of zebra mussels in the Ohio River. Often these declines are precipitous. Although zebra mussel densities vary among pools, with upstream pools having the lowest densities, the decline is not associated with river mile. Upstream pools such as Dashields show the same loss in diversity as do downstream pools such as Newburgh. Regard- less of river position, declines due to zebra mussels seem to work on an already decreased diversity caused by impound- ment, pollutants, and other factors. The overall loss of mussels in each pool is rarely attribut- able to any one cause. Although impoundment clearly has LOSS OF MUSSELS IN OHIO RIVER 7 negatively influenced the overall diversity, zebra mussels appear to have had a more serious and immediate effect. But the decline of Ohio River mussels began nearly 200 years ago and is the result of numerous, synergistic perturbations to the aquatic ecosystem. ACKNOWLEDGMENTS We thank Heidi Dunn (Ecological Specialist, Inc.) and Peter Dodgion (formerly U.S. Army Corps of Engineers) for contributing data. Funding for this project was provided by the U.S. Army Corps of Engineers and the Ohio Department of Natural Resources Division of Wildlife. We are further indebted to Mr. Dodgion for overseeing the project and Sar- ah Hazzard (OSU) and Jennifer Carpenter (OSU) for proof- reading the manuscript. LITERATURE CITED Bates, J. M. 1962. The impact of impoundment on the mussel fauna of Kentucky Reservoir, Tennessee River. American Midland Naturalist 68: 232-236. Blalock, H. N. and J. B. Sickel. 1996. Changes in mussel (Bivalvia: Unionidae) fauna within the Kentucky portion of Lake Barkley since impoundment of the lower Cumberland River. American Malacological Bulletin 13: 111-116. Branson, B. A. 1974. American paddlefish: Signs of distress. National Parks and Conservation Magazine 48: 21-23. Burkhead, N., J. Williams, and B. J. Freeman. 1992. 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Decline of native unionid bivalves in Lake St. Clair after infestation by the zebra mussel, Dreissena polymor- pha. Canadian Journal of Fisheries and Aquatic Sciences 51: 2227-2233. National Oceanic and Atmospheric Administration. 2009. Global climate at a glance. Available at: http://www.ncdc.noaa.gov/ gcag; accessed 1 June 2009. Ortmann, A. E. 1918. The nayades (freshwater mussels) of the upper Tennessee drainage, with notes on synonymy and distribution. Proceedings of the American Philiosophical Society 57: 521-626. Parker, B. C., M. A. Patterson, and R. J. Neves. 1998. Feeding inter- actions between native freshwater mussels (Bivalvia: Unioni- dae) and zebra mussels ( Dreissena polymorpha) in the Ohio River. American Malacological Bulletin 14: 173-179. Parmalee, P. W. and M. H. Hughes. 1993. Freshwater mussels (Mol- lusca: Pelecypoda: Unionidae) of Tellico Lake: Twelve years after impoundment of the Little Tennessee River. Annals of the Carnegie Museum 62: 81-93. Parmalee, P. 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Tankersley, D. I. Warmolts, G. T. Watters, B. J. Armitage, P. D. Johnson, and R. S. Butler, eds., Freshwater Mollusk Sympo- sia Proceedings, Special Publication of the Ohio Biological Survey. Pp. 135-140. Watters, G. T„ S. H. O’Dee, and S. Chordas. 2001. Patterns in verti- cal migration in freshwater mussels. Journal of Freshwater Ecol- ogy 16: 541-549. Watters, G. T., T. Cavender, C. J. Myers, M. Kibbey, V. Gordon, B. Pittinger, and T. Pohlman. 2003. Fish passage study, Product 3 — Distribution of mussels and fish within the Ohio River. Final Report to the US Army Corps of Engineers, Huntington Dis- trict. Huntington, West Virginia. Whitman, W. 1848. Excerpts from a traveller’s notebook. Daily Crescent [New Orleans], 10 March. Unpaginated. Williams, J. C. and G. A. Schuster. 1989. Freshwater Mussel Inves- tigations of the Ohio River. Mile 317.0 to Mile 981.0. Kentucky Department of Fish and Wildlife Resources, Division of Fisher- ies, Frankfort, Kentucky. Williams, J. D., S. L. H. Fuller, and R. Grace. 1992. Effects of im- poundments on freshwater mussels (Mollusca: Bivalvia: Union- idae) in the main channel of the Black Warrior and Tombigbee Rivers in western Alabama. Bulletin of the Alabama Museum of Natural History 13: 1-10. Submitted: 3 June 2009; accepted: 31 July 2009; final revisions received: 19 October 2009 Appendix. Mussel species presence by pool. X, species recorded from pool. LOSS OF MUSSELS IN OHIO RIVER 9 10 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 LOSS OF MUSSELS IN OHIO RIVER 11 Mussel species Belle Byrd Cairo Cann Dash Green Hann Mark McAlp Meld Mont Myers NewC Newb Olms Pike Racine Smith Willow Quadrula cylindrica XX X XXX X cylindrica (Say, 1817) 12 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 X X X X X X X X X X X X X m co X X X X X X X X X X X X LT) X X X X X X X ON X X X X X X X o cO X X X 00 X X X X X X X ro X 2 mm) and fine (0.6-2 mm) fractions are observed in the field (with magnification as necessary) to establish an estimate of species richness and abundance. With practice, this approach allows rapid and reliable field identification of preferred microhabitats. These appropriate microsites are then targeted for additional sampling, with approx. 50-500 ml of fine material (0.6-2.0 mm) being collected per site. Sievings are removed from the field, dried at room temperature, and then passed through a 0.6-mm sieve, with fractions being handpicked against a neutral background using low magni- fication as necessary. Through this process, typically 101- 103 individuals per taxon were recovered per site, with 102- 104 total individuals per taxon being observed across their entire ecological and geographic range. Over the last 15 years, we have collected >250,000 total pupillid individuals from the field, representing all but three valid eastern North American taxa. Material from the Coles collection is held at the National Museum of Wales (NMW) and the Florida Museum of Natural History (FM), while material from the Nekola collection is currently being maintained at the University of New Mexico. Species concepts Because pupillids demonstrate a high degree of aphallism and limited levels of anatomical variation (Pokryszko 1987), both species-level and supra-specific taxonomy has historically relied entirely upon conchological features. Some investigators have considered much of this variation to simply represent environmental plasticity, and have subsequently recom- mended the wholesale lumping of taxa ( e.g ., Bequaert and Miller 1973, Metcalf and Smartt 1997). However, other investigators have advocated a much more liberal approach and have suggested that even the most subtle shell differences demarcate biologically distinct species {e.g., Frest 1991). Throughout this work, we have chosen to let the observed variation in shell characters guide the determination of species level distinctions, rather than by blindly following either of these two camps. To do this we have used our extensive collections to define typical levels of variation for roughly 20 separate conchological features (Appendix 1) across all individuals in a given taxon both within and between populations across the entire known geographic and ecological range. We have then noted which features (if any) reliably Figure 3. A, Pupilla muscorum (European exotic), Crawford Quarry, Linn Co., Iowa, 41°59T2”N, 91°44’24”W, JCN 14592; B, Pupilla muscorum (native), Lake Bemidji State Park, Beltrami Co., Minne- sota, 47°31’58”N, 94°49’29”W, JCN 9054; C, Pupilla muscorum xe- robia, Folsom, Union Co., New Mexico, 36°55’00”N, 103°46’48”W, JCN 16491; D, Pupilla blandi. Las Vegas, San Miguel Co., New Mexico, 35°35’35”N, 105°12T7”W, JCN 12788; E, Pupoides horda- ceus, Duran, Torrance Co., New Mexico, 34°26’56”N, 105°25’6”W, JCN 14844; F, Pupoides inornatus, Folsom, Union Co., New Mexico, 36°54’18”N, 103°46’59”W, JCN 16521; G, Pupoides modicus, Cedar Key, Levy Co., Florida, CM 62.21320 (please note that shell has fad- ed in long-term storage); H, Pupoides albilabris, Gettle Farm, Wright Co., Missouri, 37°10’57”N, 92°35>32”W, JCN 11938. distinguish a given taxon, with a taxon being considered a distinct species when more variation in its key identifying features was noted between it and other taxa than was observed within that taxon. We have lumped taxa when we noted introgression between critical features either within a single population or between populations spread across either ecological or geographic space. Both morphometric (Nekola and Coles 2001, Pearce et al. 2007) and mtDNA sequence (Nekola etal. 2009) analyses have borne out this methodology, with the latter generally indicating at least 50 base-pair substitutions between concatenated COl and 16S mtDNA sequences of most recognized species (4.5% of all sequenced base pairs). These analyses have also demonstrated no more than 12 (and typically less than 5) base-pair differences exist between individuals within a given species at continental extents, even when they were sympatric at sub-meter scales with closely related species. 32 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Nomenclature Where possible, we have followed the nomenclature used by Elubricht (1985). However, the present account includes several taxa that have been elevated to species status since that time ( e.g ., Nekola 2001, Nekola and Coles 2001, Coles and Nekola 2007). In addition, while some pupillid “species” will probably prove to be species complexes, we are not yet in a position to provide definitive resolution regarding their taxonomy. We have provided our views, however, in the hope that other workers will be able to build on our observations. It should also be noted that we are in the process of revising the taxonomy of Vertigo gouldii group based on both conchological and DNA sequence data (Nekola et al. 2009). Although it is inappropriate to preempt these revisions, it should be noted that this work will ultimately change taxon rank in some cases. It will, however, otherwise not affect the following accounts. Taxonomic keys Using the suite of distinguishing conchological features detailed above, dichotomous taxonomic keys were written de novo to first assign a specimen to a given genus, followed by genus keys to allow assignment to a given species. An illustrated glossary for specialized pupillid conchological terms is presented (Appendix 1). The keys were written from a purely functional standpoint based solely on external shell features, and are thus artificial and should in no way be seen to construe any potential phylogenetic relationships. For accuracy and ease of use, occasionally a variable genus or species occurs multiple times in a given key. To aid use of the key, each couplet has been provided with a list of shell images which demonstrate the characters being defined. We have trialed these keys in a number of different public and academic settings and have revised them accordingly with the hope that they will provide many types of users, ranging from amateur conchologists and high school students to academic biologists, with the means to accurately identify individuals to the species level. Annotated comments for each recognized species are alphabetically arranged within each genus. Favored habitats were determined by analysis of species abundance patterns across all principle habitat types in our North American data set. Favored microhabitats were determined from our field experience. Range maps County-scale ranges for each taxon recorded east of the Rockies were constructed based on our collections in conjunction with observed lots from the Field Museum of Natural History (FMNH), Carnegie Musuem (CM), Academy of Natural Sciences, Philadelphia (ANSP), University of Michigan Museum of Zoology (UMMZ), Harvard Museum of Comparative Zoology (MCZ), Royal Ontario Museum (ROM), and the Canadian Museum of Nature (CMN). Additional occurrences were also mapped using references with known reliable pupillid identifications, in particular Hubricht (1985) but also Pilsbry (1948), Levi and Levi (1950; Wisconsin), Teskey (1954; northeastern Wisconsin), Dawley (1955; Minnesota), Frest (1981, 1982, 1987, 1990, 1991; northeastern Iowa), Frest and Dickson (1986; western Iowa), Theler (1997; western Wisconsin), and Ken Hotopp (pers. comm.; New York). As county-scale distributional data do not exist for Canada, range limits are indicated by plotting validated site occurrences in conjunction with locations provided in Brooks (1936), Brooks and Brooks (1940), Oughton (1948), and Pilsbry (1948). Figure 4. A, Columella columella alticola, Churchill, Manitoba, 58°45’6”N, 93°54’50”W, JCN 11321; B, Columella simplex (large morph), Rock Creek, Cedar Co., Iowa, 41°42’55”N, 91°9’31”W, JCN 11380; C, Columella simplex (normal morph), Haywood Landing, Jones Co., North Carolina, 34°49T0”N, 77°11’2”W, JCN 10716; D, Pupisoma dioscoricola (with high magnification inset of shell surface), Wadboo Creek, Berkeley Co., South Carolina, 33°11’50”N, 79°56’46”W, JCN 10903; E, Pupisoma macneilli (with high magnification inset of shell surface), Wadboo Creek, Berkeley Co., South Carolina, 33°H’50”N, 79°56’46”W, JCN 10904; F, Bothriopupa variolosa, Cuba, J. Bartlett, CM 62.21311 (please note that shell has faded in long-term storage); G, Sterkia eyriesi rhoadsi, Kyk-over-All, Kartabo, British Guiana, J. Bartlett, CM 62.19700 (please note that shell has faded in long-term storage). PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 33 Figure 5. A, Vertigo morsei, Woodland Fen, Aroostook Co., Maine, 46°52’45”N, 68°8’2 1 ”W, JCN 1 0324; B, Vertigo teskeyae , Huffs Island Park, Lincoln Co., Arkansas, BFC 29; C, Vertigo ovata, Epworth Fen, Dubuque Co., Iowa, 42°25’24”N, 90°54’56”W, JCN 11506; D, Vertigo binneyana, LaSalle River, Winnipeg, Manitoba, 49°38’60”N, 97°24T7”W, JCN 10987; E, Vertigo milium, Berlin Fen, Green Lake Co., Wisconsin, 43°57’47”N, 88°45’20”W, JCN 6308; F, Vertigo aff. genesi, La Grande Pointe, Duplessis District, Quebec, 50°12,6”N, 63°24’5”W, JCN 13459; G, Vertigo oughtoni. West Twin Lake Fen, Churchill, Manitoba, 58°37’46”N, 93°50’35”W, JCN 11159; H, Vertigo modesta hoppi, Churchill Northern Studies Center, Manitoba, 58°43’60”N, 93°48’25”W, JCN 11319; I, Vertigo modesta form arctica, Churchill, Manitoba, 58°44’53”N, 93°51’13”W, JCN 11092; J, Vertigo modesta, South Fork Koyukuk River, Alaska, 67°Tll”N, 150°17T9”W, JCN 15241; K, Vertigo modesta ultima. Sunny Mountain, Nunavik District, Quebec, 55°3’53”N, 67°14’5”W, JCN 13781; L, Vertigo oscariana, Wadboo Creek, Berkeley Co., South Carolina, 33°1T50”N, 79°56’46”W, JCN 10908; M, Vertigo parvula, Buffalo Mountain, Washington Co., Tennessee, 36°13’38”N, 82°24’7”W, JCN 12474; N, Vertigo tridentata. Little Maquoketa River, Dubuque Co., Iowa, 42°28T 7”N, 90°58’50”W, JCN 6375; O, Vertigo pygmaea. Kingfisher Farm, Manitowoc Co., Wisconsin, 43°57’50”N, 87°42’25”W, JCN 1770; P, Vertigo elatior, Karlstad South, Marshall Co., Minnesota, 48032’14”N, 96°29’4”W, JCN 6909; Q, Vertigo ventricosa. Portage Lake, Aroostook Co., Maine, 46°47’6”N, 68°32’27”W, JCN 15915; R, Vertigo perryi, Clinton, Kennebec Co., Maine, 44°36’40”N, 69°26’35”W, JCN 15422. Image figures A representative, fresh shell of each eastern North American pupillid taxon was chosen from the authors’ col- lections, except for Bothriopupa variolosa, Pupoides modicus, Sterkia eyriesi, and Vertigo hebardi, which were obtained from CM collections. All specimens were imaged in apertural view at 15x (for Pupilla, Pupoides, and the Gastrocopta armifera group) or 30x magnification (all remaining taxa) using a digital camera attached to a stereomicroscope. Approximately 12 separate 1388 x 1040 pixel images were made of each specimen with the image focal lengths positioned at 100 pm increments from the front to back of the shell. CombineZ5 freeware (http://www.hadleyweb. pwp.blueyonder.co.uk/CZ5/combinez5.htm) was used to assemble a final image from the focused parts of each separate image. This image was then imported into Adobe Photoshop, where brightness and contrast were optimized and the background made uniformly black. These images were then compiled into figures. Please note that because of their much older age, the four specimens imaged from the Carnegie collection have faded and do not represent the live shell color. 34 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 6. A, Vertigo rugulosa, Lock and Dam #5 Park, Jefferson Co., Arkansas, BFC 1297; B, Vertigo oralis, Rayonier Wildlife Management Area, Brantley Co., Georgia, 31°20’36”N, 81°49’34”W, JCN 12313; C, Vertigo concinnula, Neutrioso South, Apache Co., Arizona, 33°54T4”N, 109°9’43”W, JCN 14007; D, Vertigo cristata (large morph), Sunny Mountain, Nunavik District, Quebec, 55°3’51”N, 67°14’2”W, JCN 13686; E, Vertigo cristata (small morph), Sugar Camp Bog, Oneida Co., Wisconsin, 45°50’60”N, 89°17’45”W, BFC 1 1635; F, Vertigo malleata. Holly Shelter Game Lands, Pender Co., North Carolina, 34°3T57”N, 77°44’41”W, BFC NMW.Z.2005.0 11.03831 (paratype); G, Vertigo hebardi, Porgy Key, Dade Co., Florida, CM 73090 (please note that color has faded in long-term storage); H, Vertigo hannai. Happy Valley, Alaska, 69°20’8”N, 148°43’49”W, JCN 15144; I, Vertigo meramecensis, North Bear Creek, Winneshiek Co., Iowa, 43°26’52”N, 91°37’19”W, JCN 5192; J, Vertigo bollesiana, Collins Siding, Aroostook Co., Maine, 47°6’4T’N, 68°7’54”W, JCN 16137; K, Vertigo gouldii (small southern form), Tellico Gorge, Monroe Co., Tennessee, 35°19’49”N, 84°10’59”W, BFC 1332; L, Vertigo gouldii (normal form), Deer Creek, Fillmore Co., Minnesota, 43°43’56”N, 92°20’39”W, JCN 14646;M, Vertigo arizonensis, Devils Den Canyon, Eddy Co., New Mexico, 32°T59”N, 104°48T7”W, JCN 14582; N, Vertigo arthuri. Devils Lake Wayside, Manitoba, 52°24T3”N, 98°54’43”W, JCN 11289; O, Vertigo hubrichti, Blue Springs East, Winneshiek Co., Iowa, 43°24’35”N, 91°56’29”W, JCN 8883; P, Vertigo paradoxa. Caribou, Aroostook Co., Maine, 46°51’32”N, 68°0’43”W, JCN 9898; Q, Vertigo nylanderi. Sturgeon Gill Road, Manitoba, 53°28’23”N, 99°9’55”W, BFC 10708/504s; R, Vertigo alabamensis, Lanier Quarry, Pender Co., North Carolina, 34°37’49”N, 77°40’27”W, JCN 10781; S, Vertigo alabamensis Cconecuhensis' morph), Pond Creek seep, Covington Co., Alabama, 31°6T2”N, 86°32’3”W, JCN 12364; T, Vertigo clappi, Tellico Gorge, Monroe Co., Tennessee, 35°19’49”N, 84°10’59”W, BFC 110. ANNOTATED KEYS TO EASTERN NORTH AMERICAN PUPILLIDS Generic key 1. At least one lamella in the aperture (Figs. 1, 2, 6) 2 No apertural lamellae (Figs. 3A-C, 3E-H, 4A-E, 5F, 5K) 7 2. Parietal lamella fused with angular forming a complex bilobed structure (Figs. 1, 2C, 2G-P) Gastrocopta Parietal lamella a simple peg or plate (occasionally absent); angular lamellae (if present) not fused with the parietal (Figs. 2A, 2D-F, 5A-E, 5G-J, 5L-R, 6) 3 3. Fresh shells waxy white to clear (Fig. 2A, 2D-F) Gastrocopta Fresh shells pale to deep reddish brown (Figs. 3D, 4F-G, 5A-E, 5G-J, 5L-R) 4 4. Shell >3 mm tall (Fig. 3D) Pupilla Shell <3 mm tall (Figs. 4F-G, 5, 6) 5 5. Entire shell surface pitted; shell almost as tall as wide; three lamellae in aperture (Fig. 4F) Bothriopupa Shell lacking pits; shell taller than wide or if as tall as wide, then no apertural lamellae (Figs. 4G, 5A-E, 5G-J, 5L-R, 6) 6 PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 35 6. Shell apex strongly domed; body whorl % shell height (Fig. 4G) Sterkia Shell apex tapered; body whorl ~Vi shell height or less (Figs. 5,6) Vertigo 7. Shell >3 mm tall (Figs. 3A-C, 3E-H) 8 Shell <3 mm tall (Figs. 4A-E, 5F, 5K) 9 8. Shell ovoid or conical with tapered apex (Figs. 3E-H) Pupoides Shell cylindrical with domed apex (Figs. 3A-C) Pupilla 9. Adult shell height and width approximately equal; note that many immature pupillids will also key here (Figs. 4D-E) Pupisoma Adult shell distincdy taller than wide (Figs. 4A-C, 5F, 5K) 10 10. Shell ovoid or cylindrical; shell surface smooth (Figs. 5F, 5K) Vertigo Shell cylindrical or slightly conical; shell surface striate (Figs. 4A-C) Columella Annotated species keys Bothriopupa Pilsbry, 1898: Bothriopupa variolosa (Gould, 1848); Figs. 4F, 7 A Pilsbry (1948) reported this species from Little Marco Island, Key Marco, and Big Pine Key off the extreme southern Florida coast, and speculated that it might favor mossy rocks or trees. It has apparently not been seen alive in our region in over 75 years (Hubricht 1985). Columella Westerlund, 1878 (Figs. 4A-C): Shell cylindrical; apex domed; adult shell with 6-7 whorls, >2Vi mm tall (Fig. 4A) C. columella alticola Shell tapered; apex conical; adult shell with 5V4-6V& whorls, <2'A mm tall (Figs. 4B-C) C. simplex } I). Gastrocopta abbreviata ANgT E. Gastrocopta armifera A. Bothriopupa variolosa B. Columella columella alticola C. Columella Figure 7. Range maps for Bothriopupa variolosa, Columella columella alticola, Columella simplex, Gastrocopta abbreviata, and Gastrocopta armifera. 36 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Columella columella alticola (Ingersoll, 1875); Figs. 4A, 7B Individuals prefer willow and dwarf birch litter accumulations in taiga shrub carr communities and a wide variety of tundra habitats. At the southern edge of its range, it is restricted to seeps and coastal turf. Columella simplex (Gould, 1841); Figs. 4B-C, 7C This species, as currently defined, is found across a wide range of forested and open habitats, ranging from subtropical to taiga, xeric to wet, and acidic to calcareous. In the north, it is commonly found climbing on ferns and other herbaceous vegetation up to a meter above the ground. In such situations leaf litter sieving underestimates population size. In the south, however, it most commonly appears in leaf litter accumulations. Columella “ simplex ” encompasses such a large variation of shell sizes, shapes, and shell surface sculptures that Pilsbry (1948), Oughton (1948), and Hubricht (1985) all suggest this name likely refers to a problematic species complex. Our own observations confirm this view, but we have not yet resolved the problem. Large forms in this complex have been commonly confused with Columella columella in the southwestern U.S.A. (Bequaert and Miller 1973, Metcalf and Smartt 1997). Gastrocopta Wollaston, 1878 (Figs. 1-2): 1. Shell <3 Vi mm tall; ovoid-conical; color white to brown (Fig. 2) 2 Shell >314 mm tall, cylindrical or barrel-shaped; translucent white when fresh (Fig. 1; subgenus Albinula) 5 2. Angulo-parietal lamella a simple peg-like tooth (Figs. 2D-F, 2N) 9 Angulo-parietal lamellae not peg-shaped (Figs. 2A-C, 2G-M, 20-P) 3 3. Angulo-parietal lamella a single large, folded sheet (Figs. 2A) G. contracta Angular and parietal lobes of angulo-parietal lamellae distinct (Figs. 2C, 2G-M, 20-P) 4 4. Fresh shells whitish to pale horn yellow (Figs. 2C, 2G-H, 20-P) 12 Fresh shells yellow-brown to brown-red (Figs. 2I-M) 17 5. Columellar lamella triangular or round in cross section (Figs. 1A-B) 6 Columellar lamella a more or less vertical, flat plate (Figs. 1C-E) 7 6. Columellar lamella with both forward and basally pointing components, appearing more or less pyramidal in apertural view; shell usually >4 mm tall (Fig. 1A) G. armifera Columellar lamella lacking a basal lobe, making the entire structure appear as a downward-pointing peg in apertural view; shell <4 mm tall (Fig. IB) G. abbreviata 7. Columellar lamella a simple plate, with lower end slightly more deeply inserted into aperture; lower palatal lamella taller than wide, inserted at same depth into the aperture as the upper palatal lamella (Fig. 1C) G. clappi Columellar lamella creased in the middle, more deeply inserted at the top; lower palatal lamella as wide or wider than tall, inserted more deeply into aperture than upper palatal (Figs. 1D-E) 8 8. Lamellae massive, filling over % of the aperture, lower end of the parietal lamellae overlapping the upper end of the lower palatal; shell with domed apex (Fig. ID) G. ruidosensis Lamellae less massive, filling less than % of the aperture, lower end of the parietal lamellae at most approaching the upper end of the lower palatal; shell with tapered apex (Fig. IE) G. similis 9. Fresh shells brown-red (Fig. 2N) G. cristata Fresh shells white-transparent (Figs. 2D-F) 10 10. Shell approximately cylindrical (Fig. 2F) G. pilsbryana Shell ovoid-conical (Figs. 2D-E) 11 11. Shell narrowly conical, with height more than 1V4 times width; lower palatal lamella deeply entering aperture (Fig. 2E) G. pentodon Shell broadly conical, with height less than IV2 times width; lower palatal lamella not deeply entering aperture (Fig. 2D) G. tappaniana 12. Lobes of angulo-parietal lamella distinct, more or less parallel (Figs. 2B-C, 2G) 13 Angulo-parietal lobes intersecting, in form of a curved X-like structure (Figs. 2H, 20-P) 15 13. Basal and palatal lamellae absent; shell >2.4 mm tall (Fig. 2B) G. corticaria Basal and palatal lamellae present; shell <2.4 mm tall (Figs. 2C, 2G) 14 14. Angulo-parietal lamella massive, almost filling aperture; height >1% mm (Fig. 2C) G. ashmuni Angulo-parietal lamella filling only V4 of aperture; height <1% mm (Fig. 2G) G. holzingeri 15. Shell approximately cylindrical, with bottom three whorls of similar diameter (Fig. 2H) G. pelludda Shell narrowly conical, with each whorl of increasing diameter (Figs. 20-P) 16 16. Aperture margin with thickened callus (Fig. 20) G. rupicola Aperture margin unthickened (Fig. 2P) G. servilis 17. Alive or recently dead clean shells deep tan to brown, opaque (Figs. 2I-N) 18 Alive or recently dead clean shells horn-yellow to light yellow-brown, translucent (Figs. 2H, 20-P) 24 18. Angular lobe of angulo-parietal lamella reduced to a small protuberance (Fig. 2N) G. cristata Angulo-parietal lamella distinctly bi-lobed (Figs. 2I-M, 20-P) 19 19. Angulo-parietal lobes parallel throughout (Fig. 21) G. rogersensis Angulo-parietal lobes intersecting, in form of a curved X-like structure (Figs. 2J-M) 20 20. Lower palatal lamella parallel to and inserted at roughly the same depth into aperture as upper palatal lamella (Figs. 2K-L) 21 Lower palatal lamella angled away from upper palatal lamella, and inserted more deeply into shell (Figs. 2J, 2M) 22 21. Shell strongly tapered, with body whorl wider than the penultimate; callus plate of variable thickness on aperture margin in front of palatal lamellae (Fig. 2L) G. riograndensis Shell columnar to ovate, with body and penultimate whorls of approximately same width; aperture margin unthickened (Fig. 2K) G. riparia PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 37 22. Shell tapered, with body whorl wider than the penultimate (Fig. 2L) G. riograndensis Shell columnar to ovate, with body and penultimate whorls of approximately the same width (Figs. 2H, 2J, 2M, 20-P) 23 23. Angular lobe flaring upwards, triangular in shape, distinct from and crossing over the parietal lobe; lower palatal lamella inserted moderately deep, lying at a 45° angle to aperture axis; shell height usually >2.4 mm; thick callus plate present on aperture margin (Fig. 2M) G. sterkiana Angular lobe linear and fused to the middle of the parietal lobe; lower palatal lamella inserted deeply, lying parallel to aperture; shell height usually <2.4 mm; no callus (Fig. 2J) G. procera 24. Shell more or less cylindrical, with bottom three whorls of similar diameter (Fig. 2H) G. pellucida Shell narrowly conical, with each whorl of increasing diameter (Figs. 20-P) 25 25. Aperture margin with thickened callus (Fig. 20) G. rupicola Aperture margin unthickened (Fig. 2P) G. servilis Gastrocopta abbreviata (Sterki, 1909); Figs. IB, 7D This is the characteristic member of the subgenus Albinula in the central and western plains, where it is found under stones, within loess bank fissures, and in thin litter accumulations in riparian forests and grasslands. Towards the east it becomes largely limited to xeric habitats associated with bedrock outcrops or sand deposits. Our observations support Hubricht (1972) who noted that even though it fre- quently co-occurs with Gastrocopta armifera and Gastrocopta similis , intermediate individuals never occur. Gastrocopta armifera (Say, 1821); Figs. 1A, 7E This species is found in leaf litter accumulations on bedrock glades, rich rocky woodlands, and floodplain forests. As noted by Hubricht (1985), mapped reports of this species west from the eastern Plains and northwest of southern Minnesota are questionable. All material from these locations should be re- examined to determine if they represent Gastrocopta abbreviata or Gastroctopa similis, respectively. Gastrocopta ashmuni (Sterki, 1898); Figs. 2C, 8A East of the Rockies, populations are limited to leaf accumulations in somewhat mesic juniper, pinon pine, and oak forest on bedrock outcrops. Gastrocopta clappi (Sterki, 1909); Figs. 1C, 8B This member of the subgenus Albinula is a xerophile and calciphile that is found under rocks, around the base of grass tuffs, and under sparse vegetation on xeric glades and grasslands. Gastrocopta contracta (Say, 1822); Figs. 2 A, 8C Found in leaf litter and under logs in a wide range of habitats, ranging from mesic to wet, and forested to open. Although occurring in higher numbers in base-rich sites, it is also frequently present in base-poor locations. Gastrocopta corticaria (Say, 1816); Figs. 2B, 8D Large numbers of individuals can often be found on soil-covered ledges on wooded, calcareous bedrock outcrops. The species may also be frequent in deep leaf litter ac- cumulations under red cedar, and is occasionally found in wooded wetlands. Pilsbry ( 1948) noted that this species may also be found crawling on trees 0.3-0. 6 meters above the ground. Gastrocopta cristata (Pilsbry and Vanatta, 1900); Figs. 2N, 9A In central and eastern Plains, individuals are largely limited to sandy floodplains. Farther west, they are reported from thin leaf litter accumulations on more xeric sites (Pilsbry 1948, Bequaert and Miller 1973, Hubricht 1985). They can also be common in disturbed, anthropogenic habitats, such as in Albuquerque, New Mexico where they commonly occur in yards under juniper plantings and in grass turf near irrigation sprinkler heads. The disjunct populations in the southern Delmarva Peninsula appear to be adventitious (Hubricht 1985). Gastrocopta holzingeri (Sterki, 1889); Figs. 2G, 10A Individuals are found in accumulations of grass thatch on dry grassland, soil covered ledges on bedrock outcrops, leaf litter accumulations under red cedar, and under rocks on forested talus slopes. Gastrocopta pellucida (Pfeiffer, 1841); Figs. 2H, 9B In the southern Plains, populations are most often found in leaf litter accumulations under juniper and among grass tufts on xeric bedrock outcrops and riparian sand deposits. Along the Gulf Coast, individuals occur in open woodlands, parklands, roadsides, and lawns. This is one of the most arid- tolerant snails of the desert southwest, being found throughout southern Arizona and New Mexico in litter accumulations under low juniper, palo verde, or mesquite scrub (Metcalf and Smartt 1997). Gastrocopta pentodon (Say, 1821); Figs. 2E, 10B (syn. Gastrocopta carnegiei (Sterki, 1916)) This cosmopolitan species occurs in leaf litter accumula- tions from base-rich to base-poor, xeric to mesic, and open to forested, including sites as diverse as sand savanna, carbonate cliffs, bedrock glades, and tallgrass prairie to high elevation heath balds and mesic upland forests. We fully agree with Pearce et al. (2007) that Gastrocopta carnegiei simply repre- sents a somewhat wider than average G. pentodon individual for its height. Gastrocopta pilsbryana (Sterki, 1890); Figs. 2F, 10C In the southern Rockies, populations occur in leaf litter accumulations across a wide variety of dry-mesic and mesic forest sites from low to high elevation. In the far southwestern 38 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 A. Gastrocopta ashmuni B. Gastrocopta clappi D. Gastrocopta corticaria C. Gastrocopta contracta Figure 8. Range maps for Gastrocopta ashmuni, Gastrocopta clappi, Gastrocopta contracta, and Gastrocopta corticaria. Plains, it is limited to forested pockets on shaded canyon sides. Individuals from eastern New Mexico are characterized by a thick apertural callus and distinct crest. While Pilsbry (1948) and Metcalf and Smartt (1997) indicate that G. pilsbryana lacks these features, these populations clearly represent this species due to their distinctly cylindrical shape, even in sites where it co-occurs with G. pentodon. The taxonomic status of this form is unclear. Gastrocopta procera (Gould, 1840); Figs. 2J, 10D This obligate calciphile is found under stones, in thatch, and in leaf litter accumulations on scrub-covered and exposed sites such as bedrock glades, dry prairie, and roadside verges. It also occurs in sandy river floodplain scrub and forest. Gastrocopta riograndensis (Pilsbry and Vanatta, 1892); Figs. 2L, 11A We have found this species in thin soil accumulations on small ledges of xeric south-facing limestone cliffs in the Sacramento Mountains of New Mexico, where organic litter is generated from grasses and shrubs. It has also been reported from similar habitats in west Texas (Neck 1980). While much of the south Texas material at ANSP and CM represents flood wash debris, a number of these shells were also alive at time of collection, indicating the presence of extant populations in more mesic riparian habitats. This material differs from those observed in New Mexico by having a wider shell for a given height and a thinner palatal callus. In all other respects, however, these forms appear identical, suggesting that they are simply endpoints of environmentally driven clinal variation. PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 39 Figure 9. Range maps for Gastrocopta cristata and Gastrocopta pellucida. Gastrocopta riparia Hubricht, 1978; Figs. 2K, 11B Individualsarefoundindecomposedleaflitteraccumulations, often under dense shrub or vine thicket cover in mesic, disturbed sites such as railroad rights-of-way, roadside verges, vacant lots, floodplains, and other scrubland habitats. It seems more tolerant of acidic conditions than Gastrocopta procera. Gastrocopta rogersensis Nekola and Coles, 2001; Figs. 21, 11C This calciphile is found on exposed soil, under stones and in thin accumulations of leaf litter and grass thatch on dry bedrock cliffs, xeric glades, and occasionally rocky, upland forest. Gastrocopta ruidosensis (Cockerell, 1909); Figs. ID, 11D This member of the subgenus Albinula is found on bare soil, under stones, and in thin accumulations of grass thatch and juniper litter on mid-elevation carbonate cliffs and xeric limestone grasslands along the eastern slopes of the Sangre de Cristo and Sacramento mountains in eastern New Mexico. Pleistocene fossil material has been found throughout the southern Plains (Hubricht 1985). Gastrocopta rupicola (Say, 1821); Figs. 20, 12A Individuals are found in decomposed leaf litter, often under dense shrub or vine thicket cover in lowland forest, scrub, and disturbed habitats. 40 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 A. Gastrocopta holgingeri B. Gastrocopta pentodon C. Gastrocopta pilsbryana D. Gastrocopta procera Figure 10. Range maps for Gastrocopta holzingeri, Gastrocopta pentodon, Gastrocopta pilsbryana, and Gastrocopta procera. Gastrocopta servilis (Gould, 1843); Figs. 2P, 12B This species appears to favor grass thatch and decomposed leaf litter in shoreline thickets and anthropogenically disturbed habitats such as roadsides, vacant lots, yards, and railroad rights-of-way. Gastrocopta similis (Sterki, 1909); Figs. IE, 12C This obligate calciphile is the characteristic member of the genus Albinula in the upper Midwest, where it is found under stones, on bare soil, soil-covered cliff ledges, and in decomposed grass thatch and red cedar litter accumulations across a wide variety of habitats ranging from xeric grasslands to mesic forest and fens. However, it is most frequently encountered in dry, gravelly prairie and bedrock glades. Gastrocopta similis appears very similar to Gastrocopta ruidosensis, differing only by its slightly less massive apertural lamellae. The relationship be- tween these two taxa requires further investigation. Gastrocopta sterkiana Pilsbry, 1912; Figs. 2M, 12D Found under stones, on bare soil, in thin grass thatch and juniper or litter accumulations on xeric grasslands such as bare limestone outcrops in the Flint Hills of northeastern Oklahoma and pinon-juniper parkland in northeastern New Mexico. On the western limit of its range, it may also occur in accumulations of cottonwood litter in riparian forest. Gastrocopta tappaniana (C. B. Adams, 1842); Figs. 2D, 13A Found in accumulations of decomposing leaf litter in wooded and open wetland habitats such as riparian, floodplain and swamp woodlands, mesic and wet prairies, open shoreline bedrock outcrops, fens, pocosins, and Sphagnum bogs. While some have suggested that this taxon is an ecophenotypic variant of Gastrocopta pentodon (e.g., Bequaert and Miller 1973), multivariate morphometric PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 41 Figure 11. Range maps for Gastrocopta riograndensis, Gastrocopta riparia, Gastrocopta roger semis, and Gastrocopta ruidosensis. analyses of museum collections indicate that this species from across the range of both species agree fully with this is clearly distinct (Pearce etal. 2007). Our field observations conclusion. Pupilla Leach, 1828 (Figs. 3A-D): 1. Aperture with three well-developed lamellae; palatal lamella often longer than wide (Fig. 3D) P. blandi Aperture with two or fewer lamellae; palatal lamellae (if present) usually as wide as long (Figs. 3A-C) 2 2. Callus inserted into aperture; shell >3 mm tall (Figs. 3A-B) P. muscorum Massive callus at apertural margin; shell <2% mm tall (Fig. 3C) P. muscorum xerobia Pupilla blandi Morse, 1865; Figs. 3D, 13B East of the Rockies, populations of dwarfed individuals are occasionally found in xeric juniper savanna and mesic mixed conifer forest. In the mountains they occur in leaf litter accumulations in mid to high elevation oak, pine, fir, and spruce forest, becoming especially abundant in aspen groves. Pupilla muscorum (Linne, 1758); Figs. 3A-C, 13C East-central North American populations (Maine and Tennessee west to eastern Iowa) generally occur in disturbed anthropogenic habitats such as road verges, vacant lots, abandoned quarries, old fields, and concrete culverts (Hubricht 1985) although they may also occasionally inhabit less disturbed carbonate cliff, glade, and grassland sites. To 42 AMERICAN MALACOLOGICAL BULLETIN 28-1/2-2010 Figure 12. Range maps for Gastrocopta rupicola, Gastrocopta servilis, Gastrocopta similis, and Gastrocopta sterkiana. the north (Newfoundland, the north shore of the St. Lawrence to northwestern Minnesota and the southern shore of Eludson’s Bay) populations occur on bare soil, under stones, on turf, and in thin leaf litter accumulations on sandy or rocky shorelines and in tundra. Recent mitochondrial DNA sequence analyses (Nekola et al. 2009, Yon Proschwitz et al. 2009) indicate that throughout its Holarctic range this name has been applied to a species complex. Most of the populations in east-central North America (referable to P. muscorum) represent apparent European introductions, with Iowa roadside verge material being closest, for instance, to Swedish haplotypes. However, northern Plains populations represent an undescribed species distantly allied to Pupilla hebes and Pupilla pratensis. Given the morphologic variability noted Shell surface with minute spiral striae (Fig. 4D) between northern Plains, southern Plains, and arctic populations, the presence of more than one native species also appears likely. The southern Plains form, limited to arid pinon-juniper forests, has been referred to as Pupilla muscorum xerobia (Pilsbry 1948). Metcalf and Smartt (1997) suggest that this taxon may be worthy of species status given its greatly thickened apertural lip, uniformly small size and height/width ratio, and divergent habitat and range. The native arctic populations differ from P. hebes only by the weak possession of a partial callus on the uppermost margin of the palatal wall, and appear quite similar to Pupilla pratensis. Additional sequence analysis will be required to make definitive taxonomic statements regarding this group not only in North America but also in Eurasia. P. dioscoricola . P. macneilli Pupisoma Stoliczka, 1873: Shell surface pitted-granulose, lacking striae (Fig. 4E) PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 43 Pupisoma dioscoricola (C. B. Adams, 1845); Figs. 4D, 13D Populations occur in sub-tropical forest and scrub, abandoned citrus orchards and carbonate rock outcrops. Individuals are principally arboreal on the undersides of leaves, with only a scattered few occurring in leaf litter. Hubricht (1985) reports that their mucus is especially adhesive, making them less likely to be dislodged by storms as compared to other arboreal taxa. Pupisoma macneilli (Clapp, 1918); Figs. 4E, 14A Populations occur in woodlands, scrub, and carbonate rock outcrops. Hubricht (1985) reports that individuals are most often found on the trunks of smooth-barked trees and shrubs, with only scattered shells occurring in leaf litter. Pupoides Pfeiffer, 1854: 1 . Shell conical (Figs. 3G-H) 2 Shell cylindrical-ovoid (Figs. 3E-F) 3 2. Adult shell with calcified apertural margin (Fig. 3H) P. albilabris Adult shell with unthickened apertural margin (Fig. 3G) P. modicus 3. Shell surface with regular, widely-spaced ribs; aperture calcified (Fig. 3E) P. hordaceus Shell surface with irregular striations; aperture expanded but unthickened (Fig. 3F) P. inornatus Pupoides albilabris (C. B. Adams, 1821); Figs. 3H, 14B A calciphile found under stones, leaf litter under red cedar, in thin grass turf and thatch accumulations on rock outcrops, bedrock glades, xeric prairie, and old fields. It is also occasionally found in riparian forests of the western plains. 44 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 14. Range maps for Pupisoma macneilli, Pupoides albilabris, Pupoides hordaceus, Pupoides inornatus, and Pupoides modicus. Pupoides hordaceus (Gabb, 1866); Ligs. 3E, 14C Individuals occur in deep juniper litter accumulations in xeric, low elevation juniper parkland where it is often the only species present. Pupoides inornatus Vanatta, 1915; Figs. 3F, 14D Populations occur in leaf litter accumulations under small shrubs and under rocks or in thin grass thatch in xeric grassland and parkland habitats (Metcalf and Smartt 1997). It also occurs in leaf litter accumulations in riparian forest. Pupoides modicus (Gould, 1848); Figs. 3G, 14E Populations occur along roadsides and a variety of other open habitats. Sterkia Pilsbry, 1898: Sterkia eyriesi rhoadsi (Pilsbry, 1899); Figs. 4G, 15A Pilsbry (1948, pp. 1016-1018) indicates that in eastern North America this species is limited to tropical hardwood hammocks in extreme southern Florida. He reported finding only two individuals in a “great amount of woodland debris” and indicated that George Clapp only located about a dozen shells from a “bushel of rubbish.” Based on our experience with other pupillids, these low numbers suggest to us that neither researcher deduced this taxon’s preferred microsites. Hubricht (1985) reported locating a single individual crawling on a log after a shower. Vertigo Muller, 1774 (Figs. 5-6): 1. Body whorl strongly pustulose (Fig. 6F) V.malleata Body whorl lacking strong pustulose bumps (Figs. 5, 6A-E, 6G-T) 2 2. Upper palatal lamella short, low and straight, with long axis barely visible in apertural view (Figs. 5, 6A-Q) 3 Upper palatal lamella long, tall and longitudinally curved, allowing long axis to be visible in apertural view (Figs. 6R-T) 8 3. Shell surface smooth or weakly striate (Fig. 5) 4 Shell surface strongly striate (Figs. 6A, 6C-E, 6G-Q) 6 PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 45 4. Six or more apertural lamellae (Figs. 5A-E, 50, 6B) 9 Five or fewer apertural lamellae (Figs. 5F-R) 5 5. Four or fewer apertural lamellae (Figs. 5F-N, 5R) 15 Five apertural lamellae (Figs. 5D, 5P-0) 27 6. Parietal lamella pointed directly at lower palatal lamella, so that parietal, lower palatal, and columellar lamellae form a cross (Figs. 5 J, 6C-E, 61) 33 Parietal lamella pointed at upper palatal or space between the upper and lower palatals (Figs. 6A-B, 6G-H, 6J-M) 7 7. Lower palatal lamella inserted near aperture margin so that only short axis is visible when seen in apertural view (Figs. 6A-E, 6G-M) 36 Lower palatal lamella inserted more deeply into shell so that long axis is visible when seen in apertural view (Figs. 6N-Q ) 43 8. Aperture margin thickened; shell color deep yellow; imperforate (Figs. 6R-S) V. alabamensis Aperture margin not thickened; shell color light yellow to horn; narrowly umbilicate (Fig. 6T) V. clappi 9. Angular lamella absent; palatal wall with callus and light-colored crest; shell dull (Fig. 50) V. pygmaea Angular lamella present; crest not light-colored (Figs. 5A-E, 6B) 10 10. Shell weakly striate; dull; all lamellae short; shell with shallow suture and domed apex (Fig. 6B) V. oralis Shell smooth, shiny; at least some of the lamella long and blade-like (Figs. 5A-E) 11 11. Shell basally obese, with height less than 2 times width (Figs. 5B-C) 12 Shell not basally obese, with height greater than 2 times width (Figs. 5A, 5D-E) 13 12. Aperture wider than tall; columellar lip of aperture broad, more or less straight and angled away from palatal wall; infra-parietal lamella never present (Fig. 5B) Vi teskeyae Aperture as tall as wide; columellar lip of aperture rounded, not markedly broad; infra-parietal lamella often present (Fig. 5C) Vi ovata 13. Shell height >2 Vi mm; ~6 whorls; aperture less than Vs of shell height (Fig. 5A) V. morsel Shell height <2 V2 mm; ~4 whorls; aperture more than Vs of shell height (Figs. 5D-E) 14 14. Lower palatal lamella straight and not deeply entering aperture; shell height > 1 .9 mm (Fig. 5D) V. binneyana Lower palatal lamella curved and deeply entering aperture; shell height <1.9 mm (Fig. 5E) V. milium 46 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 15. No apertural lamellae; note that juvenile Vertigo species lack lamellae and may key out here; see also Columella species which are somewhat similar in form to Vertigo (Figs. 5F, 5K) 16 2-4 apertural lamellae (Figs. 5G-J, 5L-N, 5Q-R) 18 16. Crest absent; dull surface luster (Fig. 5F) V. aff. genesii Crest present; glassy surface luster (Figs. 5H, 5K) 17 17. Shell <2Vi mm tall, shell ovoid-conical (Fig. 5H) V. modesta hoppi Shell >2Vi mm tall, shell ovate (Fig. 5K) V. modesta ultima 18. One or two apertural lamellae (Figs. 5G-H) 19 Three or four apertural lamellae (Figs. 5L-0, 5Q-R) 20 19. Shell lacking palatal lamellae; ovoid-conical; crest present (Fig. 5H) V. modesta hoppi At least one strong palatal lamellae present; shell cylindrical; crest absent (Fig. 5G) V. oughtoni 20. Shell >2 mm tall, ovoid-cylindrical (Figs. 5G, 5I-J) 21 Shell 2 mm or less tall, ovoid-conical (Figs. 5L-N, 5Q-R) 23 21. Four apertural lamellae, lower and upper palatal lamellae of similar size (Fig. 5J) V. modesta Three or four lamellae, upper palatal lamella weak or absent (Figs. 5G, 51) 22 22. Shell ovoid; >2 ‘A mm tall; lower palatal lamella a short peg (Fig. 51) V. modesta form arctica Shell cylindrical; 2 Vt mm or less tall; lower palatal lamella longer than wide (Fig. 5G) V. oughtoni 23. Body whorl narrower than penultimate whorl, making shell bluntly pointed at both top and bottom; four lamellae, with an elongate vertical columellar (Fig. 5L) V. oscariana Body whorl at least as wide as the penultimate whorl; columellar lamella peg-shaped (Figs. 5M-N, 5Q-R) 24 24. Body whorl inflated, making shell height less than twice the width (Figs. 5Q-R) 25 Body whorl not greatly inflated, making shell height approximately twice the width (Figs. 5M-N) 26 25. Moderately strong sinulus; shell color red-brown; weak spiral striation on body whorl; aperture margin pale (Fig. 5Q) V. ventricosa Weak sinulus; shell color with slight greenish cast; distinct spiral striation on body whorl; aperture margin usually dark olive-brown to black (Fig. 5R) Vi perryi 26. Shell height >1% mm; a weak upper palatal lamella often present (Fig. 5N) V. tridentata Shell height <1% mm; upper palatal lamella absent (Fig. 5M) Vi parvula 27. Shell >2Vi mm tall; angular lamella present (Fig. 5J but with angular) V. modesta form parietalis Shell <2lA mm tall; angular lamella absent (Figs. 5C-D, 50-Q, 6J) 28 28. Shell height less than twice the width, with marked basal inflation (Figs. 5C, 5Q) 29 Shell height twice the width or more, ovoid (Figs. 5D, 50-Q, 6J) 30 29. Strong apertural lamellae and sinulus; shell translucent (Fig. 5C, but lacking infraparietal) Vi ovata Moderate apertural lamellae and sinulus; shell glassy and transparent (Fig. 5Q) Vi ventricosa 30. Shell weakly striate; single depression behind aperture over both palatal lamellae (Fig. 6J) Vi bollesiana Shell smooth; separate slight depressions under each palatal lamella, or none (Figs. 5D, 50-P) 31 3 1 . Shell surface dull; strong crest; light-colored callus on palatal wall (Fig. 50) Vi pygmaea Shell surface shiny; crest less prominent; callus of same color as shell (Figs. 5D, 5P) 32 32. Lower palatal lamella inserted more deeply than upper palatal; upper palatal lamella thickened towards aperture; shell shape ovoid (Fig. 5D) V. binneyana Lower palatal lamella inserted as deeply as upper; upper palatal lamella not thickened towards aperture; shell apex elongate and somewhat conical; strong callus often present in base-rich habitats (Fig. 5P) Vi elatior 33. Shell >2.3 mm tall, shiny with weak striae (Fig. 5J) Vi modesta Shell <2.3 mm tall, shell dull with distinct striae (Fig. 6) 34 34. Shell shape conical with body whorl much wider than the penultimate; color deep cinnamon-red; crest absent; shell striation irregular in strength and spacing. (Fig. 61) V. meramecensis Shell shape ovate with body whorl approximately the same width as the penultimate; color yellow-red brown; crest present; shell striation uniform (Figs. 6C-E) 35 35. Upper and lower palatal lamellae short and of same length; shell striation fine, regular; crest weak to moderate (Figs. 6D-E) V. cristata Lower palatal lamella longer than upper palatal; shell striation coarse, irregular; very strong crest (Fig. 6C) V. concinnula 36. Angular lamella present (Figs. 6A-B, 6G-H, 6M) 37 Angular lamella absent (Figs. 6G, 6J-L) 41 37. Basal lamella absent (Figs. 6G, 6M) 38 Basal lamella present (Figs. 6A-B, 6H) 39 38. Shell ovoid, <116 mm tall; angular lamella weak/vestigal (Fig. 6G) Vi hebardi Shell cylindrical, >116 mm tall; angular lamella strong (Fig. 6M) V. arizonensis 39. Shell cylindrical with bottom two whorls of same width; no callus on palatal wall (Fig. 6H) V. hannai Shell ovate, body whorl larger than penultimate whorl; weak callus present on palatal wall (Figs. 6A-B) 40 40. Shell narrowly ovate; coarsely striate; usually >1.8 mm tall; apex tapered (Fig. 6A) V. rugosula Shell broadly ovate; weakly striate; usually <1.8 mm tall; apex domed (Fig. 6B) V. oralis 41. Basal lamella absent; shell height <116 mm (Fig. 6G) ,V. hebardi Basal lamella present; shell height >116 mm (Figs. 6J-L) 42 42. Striae indistinct, with shell often appearing smooth under low ( x 10) magnification; single deep depression over both palatal lamellae; ~1% mm tall (Fig. 6J) V. bollesiana Striae distinct, with shell not appearing smooth under low (xlO) magnification; palatal depression weak or absent; most forms >1% mm tall (Fig. 6L), with small southern Appalachian forms being -1% mm tall (Fig. 6K) V. gouldii 43. Callus surrounding at least the upper palatal and often the entire palatal wall; ranging from eastern Ontario to Alaska and south to New Mexico (Fig. 6N) V. arthuri Callus absent on palatal wall (Figs. 60-Q) 44 PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 47 44. Columellar lamella more massive than the parietal; angular lamella strong; lower palatal lamella so deeply inserted that most of it is obscured by the columellar wall; striation fine and sharp (Fig. 6Q) VI nylanderi Parietal lamella more massive than the columellar; angular lamella weak or absent; lower palatal lamella less deeply inserted so that most is observable in apertural view; striae somewhat rounded (Figs. 60-P) ,, 45 45. Basal and weak angular lamellae often present; ranging from the Upper Mississippi River valley to eastern Ontario (Fig. 60) V. hubrichti Basal and angular lamellae often absent; ranging from Newfoundland and central Manitoba to northern Minnesota, northern Wisconsin, and the New England states; also in Alaska and the Yukon (Fig. 6P) V. paradoxa Vertigo alabamensis Clapp, 1915; Figs. 6R-S, 15B (syn. V. alabamensis conecuhensis in Pilsbry, 1948 and V. conecuhensis in Hubricht, 1985) An obligate acidophile occurring in well-decomposed leaf litter typically caught among low growing shrubs and vines in mesic pineland, pine-wiregrass savanna, and bay forest. This species displays a high degree of seasonality, with all individuals hatching in early spring and coming to adult age from late April to early June. Because they rapidly erode in their acidic habitats, surveys outside this period document few (if any) shells. Populations are readily eliminated by fire management, and the species is now absent from many seemingly appropriate sites which are subjected to high return frequency prescribed burning. Obese individuals with less massive lamellae, equating to V. conecuhensis as understood by Hubricht ( 1985), are found within populations throughout the range of V. alabamensis. As populations demonstrate complete intergradation between both morphotypes, we relegate this form to a synomym of Vi alabamensis. Vertigo arizonensis (Pilsbry and Vanatta, 1900); Figs. 6M, 15C (syn. V. gouldii arizonensis in Pilsbry, 1948) Individuals favor accumulations of highly decomposed leaf litter, often under maple or Douglas fir, in mid to low ele- vation forests in the southern Rockies. In the caprock canyons of far northeastern New Mexico, it is limited to mesic forest pockets. While considered a subspecies of V. gouldii by Pilsbry (1948), and of no taxonomic merit by Bequaert and Miller (1973) and Metcalf and Smartt (1997), shells of this taxon never intergrade with other sympatric V. gouldii subspecies. DNA sequence analysis confirms its status as a full species (Nekola et. al. 2009). Vertigo arthuri (von Martens, 1884); Figs. 6N, 15D (syn V. gouldii basidens Pilsbry and Vanatta, 1900 in Pilsbry 1948) Populations occur in well-decomposed leaf litter in aspen parkland, jack pine forest, and taiga, as well as mesic mixed conifer and aspen groves in the Black Hills and southern Rockies. Long known from two individuals collected in 1882 (Pilsbry 1948, Hubricht 1985), this species is now known to be the most abundant Vertigo in aspen forests at the northern limit of the Great Plains (Nekola 2002), ranging as far east as Ottawa, Ontario and as far west as Anchorage, Alaska. Observation of material from across this range demonstrates that V arthuri encompasses the entire morphological range of Vertigo gouldii basidens, including populations at the type locality in the Jemez Mountains of northern New Mexico. Vertigo binneyana Sterki, 1890; Figs. 5D, 16A Individuals occur in grass thatch and leaf litter in mesic grasslands and adjacent oak-aspen woodlands. Reports of this species from the central plains and to the west of the continental divide appear to be based on misidentified material. Vertigo bollesiana (Morse, 1865); Figs. 6J, 16B Found in leaf litter often under shrubs, on cliff-face ledges and boulder tops in mesic upland forest, and mesic microsites in northern white cedar wetlands. This species always shows a strong depression on the outside of the shell over both palatal lamellae. All purported southern Appala- chian material seen by the authors lacks this feature and rep- resents misidentified small individuals of Vertigo gouldii. As a result, all records of V. bollesiana south of Pennsylva- nia should be considered questionable and be critically reexamined. Vertigo clappi Brooks and Hunt, 1936; Figs. 6T, 16C Individuals favor well-decomposed leaf litter and fine soil on shaded boulders, talus, ledges and bases of forested bedrock outcrops. Vertigo concinnula Cockerell, 1897; Figs. 6C, 16D East of the Rockies this species is restricted to mesic lime- stone forest in the Black Hills (Hubricht 1985). To the west, it occurs in well-decomposed leaf litter in mid to high-elevation Douglas fir, aspen, and spruce-fir forests in the Rockies where it often demonstrates considerable tolerance for acidic condi- tions. While Bequaert and Miller (1973) indicate that this taxon is simply a subspecies of Vertigo modesta, both shell morphology and DNA sequence data suggest that it is worthy of species-level distinction (Nekola et al. 2009). We retain the use of concinnula as the specific epithet for this taxon on the basis of Pilsbry (1948: 979) who considered the prior name of Vertigo ingersolli to be “absurdly inadequate.” 48 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Vertigo cristata Sterki, 1919; Figs. 6D-E, 17A Found in well-decomposed leaf litter in a wide variety of northern forest habitats, ranging from wetlands to dry upland rock outcrops. It is particularly common in base- poor sites such as pine and spruce forest, heaths, and Sphagnum- dominated peatlands. Throughout its range, two size morphs are present, one with mature shells <1.9 mm tall, and a sec- ond with shell heights ranging from 2 to 2 Vi mm (Nekola 2001). The holotype at ANSP represents the latter morph. Both morphs often co-occur in sites without presence of intermediates. However, DNA sequence analyses do not support them as being distinct (Nekola et al. 2009). Shells of the large morph are similar in size to small Vertigo modesta from which they are most readily distinguished by their strong striation and sharper crest. All prior records for Ver- tigo modesta from the New England states ( e.g ., Pilsbry 1948) likely represent the large morph of Vertigo cristata, which we have seen throughout the high mountains of this region. However, we have not seen Vertigo modesta south of the Gulf of St. Lawrence. Vertigo elatior Sterki, 1894; Figs. 5P, 17B Individuals occur in well-decomposed humid leaf litter and graminoid thatch in a variety of open and wooded wetland habitats, including coastal alvar, wet prairie, fens, wet meadows, tundra, and black ash, tamarack, northern white cedar, and black spruce swamp forests. Although predominately a calciphile in the east, it tolerates acidic conditions in the upper Midwest. After rains or on dewy mornings, individuals commonly adhere to a hand run through damp leaves. PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 49 Figure 17. Range maps for Vertigo cristata, Vertigo elatior, Vertigo aff. genesii, and Vertigo gouldii. Vertigo aff. genesii (Gredler, 1856); Fig. 5F, 17C Found in accumulations of graminoid leaf litter in sedge meadows, turf, and shrub carr in tundra and taiga districts, where it extends as far west as Alaska. This taxon has previously been confused with Vertigo oughtoni (see below). However, it appears closest to the European Vertigo genesii (see Kerney and Cameron 1979) due to its ovate-conical sell, simple apertural lip without reflexion, indentation or crest, and total lack of apertural lamellae. Whether the North American populations represent V. genesii or distinct species remains to be determined. Vertigo gouldii (A. Binney, 1843); Figs. 6K-L, 17D Individuals are most abundantly encountered in well- decomposed leaf litter on shaded cliff ledges and bases and on the top of large rocks. They also occur in lower numbers throughout upland and lowland forest, and may be occasionally seen crawling on cliff faces. Small shells <1% mm in height with reduced striation and dentition from the southern Appalachians and Ozarks have often been misidentified as Vertigo bollesiana (see above). The reported populations from Jamaica represent a taxon allied with Vertigo hebardi (Gary Rosenberg, pers. comm.). Vertigo hannai Pilsbry, 1919; Figs. 6H, 18A In Churchill, Manitoba, individuals are found in well- decomposed thatch and leaf litter of upland tundra, short turf, fens, and wooded wetlands. In northern Alaska, this species also habits upland and riparian forest and parklands. Reports of this species from eastern Ontario alvars are based on a misidentified shell of Vertigo hubrichti. Vertigo hebardi V anatta, 1912; Figs. 6G, 18B This species has not been seen alive in over 75 years. Pilsbry (1948) reported it from a series of Keys off the southern Florida coast, with no associated habitat information. Presumably, populations occur in tropical hardwood forest. Hubricht (1985) suggested that it is arboreal because all museum specimens were dead when collected. Vertigo hubrichti (Pilsbry, 1934); Figs. 60, 18C (syn. V brierensis Leonard, 1972 in Frest 1991, V hubrichti variabilis Frest, 1991, V. iowaensis Frest, 1991) Found in leaf litter pockets supporting a cool summer microclimate, in particular northern white cedar groves on carbonate bedrock ledges (and occasionally uplands) near the Lake Michigan and Lake Huron shores (Nekola 2004), eastern 50 AMERICAN M ALACOLOGICAL BULLETIN 28 -1/2 -2010 Figure 18. Range maps for Vertigo hannai, Vertigo hebardi, Vertigo hubrichti, and Vertigo malleata. Ontario alvars, and algific talus slopes along the upper Missis- sippi River valley (Frest 1991). Observations of populations across its range indicate that its normal morphologic variation completely encompasses Vertigo brierensis , Vertigo hubrichti variabilis, and Vertigo iowaensis of Frest (1991). As such, we reduce these forms to synonyms. Although treated as a spe- cific taxon by Hubricht (1985), Frest (1991), and Nekola (2004), we have also noted complete intergradation of V. hubrichti with Vertigo paradoxa from northeastern Wisconsin through northern Maine. Vertigo malleata Coles and Nekola, 2007; Figs. 6F, 18D This obligatory acidophile is primarily found in humid accumulations of ericaceous and pine leaf litter in mesic to wet base-poor habitats along the eastern seaboard such as longleaf pine forest and savanna, bay and Atlantic white cedar forest, heaths, pocosins, and other acid peatlands. Vertigo meramecensis Van Devender, 1979; Figs. 61, 19A A strict calciphile found in decomposed leaf litter on fern and moss-covered ledges and open rock and lichen-covered surfaces of mesic, shaded carbonate cliffs. Vertigo milium (Gould, 1840); Figs. 5E, 19B Individuals are found in humid, well-decomposed thatch and leaf litter across a wide variety of mesic to wet sites including rocky woodland, riparian woodland, cliffs, wet prairie, sedge meadows, roadside verges, fens, and swamps. Vertigo modesta (Say, 1824) Vertigo modesta appears to be a species complex (Pilsbry 1948). Our own experience with this aggregate in boreal and arctic North America shows the presence of at least three forms that possess consistent morphology and habitat preferences over wide geographical ranges. Their ecology and distribution PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 51 Figure 19. Range maps for Vertigo meramecensis, Vertigo milium, Vertigo modesta hoppii, and Vertigo modesta modesta. are summarized below using a sub-specific nomenclature based on the names used by Pilsbry (1948). Whether they represent distinct species or merely consistent ecophenotypes is not clear, although we have never noted intermediate individuals, even in sites of co-occurrence. Note also that the large Vertigo cristata morph has often been confused with Vertigo modesta modesta by previous researchers (see above). Vertigo modesta (Say, 1824); Figs. 5I-J, 19C Individuals occur in accumulations of humid leaf litter in mesic to wet taiga, notably shrub carr dominated by willow, alder, or birch, and in willow and birch litter accumulations across the entire moisture gradient in tundra. Populations at the extreme southern margin of the range along the Lake Superior shore are limited to cool, mesic lower margins of open talus slopes. Throughout its range, we have noted the presence of individuals with an angular lamella which Pilsbry (1948) termed V. modesta form parietalis. A marked clinal reduction in palatal lamellae development is noted towards the north, with the upper palatal being absent from many tundra locations. These shells often also possess a distinctly more red color than their southern counterparts. This form, referred to as Vertigo modesta arctica in Europe (Kerney and Cameron 1979), is dominant along the southern shore of Hudson’s Bay, southern Baffin Island, and from limestone pavements along the northern shore of the St. Lawrence. It also represents the western Newfoundland material identified as Vertigo modesta castanea by Brooks and Brooks (1940). Vertigo modesta hoppii (Moller, 1842); Figs. 5H, 19D Populations occur in leaf litter and thatch accumulations in base-poor tundra, sedge meadows, and peatlands. This form has a smaller and more conical shell than is typical for 14 modesta, with the palatal lamellae being absent. We have noted some populations from Alaska which also lack parietal and columellar lamellae. These individuals can be most 52 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 readily separated from V. aff. genesi by their larger volume, apertural crest, and shiny luster. Vertigo modesta ultima Pilsbry, 1948; Figs. 5K, 20A Individuals occur in wet shrub carr and sedge meadow in northern taiga and tundra. This entity not only differs from V. modesta by lacking parietal and palatal lamellae (though a vestigal columellar may be present), but also in its larger size, more inflated whorls, more open umbilicus, and possession of a broadly reflected lip on the columellar wall of the aperture. This form may be identical to V. extima from far northern Eurasia (Pilsbry 1948, Pokryszko 2003). DNA sequence analysis will be required to determine their exact relationship. Vertigo morsei Sterki, 1894; Figs. 5A, 20B Populations occur in humid, aerated, well-decomposed leaf litter often overlying marl or carbonate bedrock in highly calcareous open wetlands including fens, alvars, and wet prairie. They may also be occasionally found crawling on Juncus stems. Vertigo nylanderi Sterki, 1909; Figs. 6Q, 20C Individuals occur in sedge and grass thatch and stick-filled depressions in a variety of wooded wetland habitats across the base-status spectrum including northern white cedar (Maine), black ash, tamarack, black spruce (upper Midwest), and shrub carr (Ontario, Manitoba), as well as fens. Vertigo oralis Sterki, 1898; Figs. 6B, 20D Populations reside in broadleaf and graminoid leaf litter accumulations, and under logs, in wet woodlands including pool margins in oak-sweetgum forest, red maple swamp, cypress swamp, and riparian and pocosin scrub. Vertigo oscariana (Sterki, 1890); Figs. 5L, 21A Individuals occur in well-decomposed accumulations of broadleaf and pine litter in mesic-wet woodlands and shaded rock outcrops. Habitats range from montane hardwood forest in the Appalachians to oak-pine-bay bottomland woodland along the Gulf Coast to acid pine forest in Arkansas, Louisiana, and Texas. Hubricht (1985) reported it from the undersides of palmetto leaves. Vertigo oughtoni (Pilsbry, 1948); Figs. 5G, 21B (syn. Vertigo alpestris oughtoni in Pilsbry, 1948) An arctic calciphile which occurs in thin grass and sedge thatch in flushes, calcareous fens, seeps, and shrub carr. Pilsbry (1948) described V. oughtoni as a subspecies of the Eurasian Vertigo alpestris Alder, 1838, but the two taxa share little in common in terms of shell morphology or habitat preferences (Kerney and Cameron 1979). It appears most closely allied to V. parcedentata (A. Braun, 1847), with which it shares a columnar shell with a blade-like lower palatal lamella, a reduced (or absent) columellar and upper palatal lamella, a simple apertural lip without reflection and marked Figure 20. Range maps for Vertigo modesta ultima, Vertigo morsei, Vertigo nylanderi, and Vertigo oralis. PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 53 Figure 21. Range maps for Vertigo oscariana, Vertigo oughtoni, Vertigo ovata , and Vertigo paradoxa. preference for moist, base-rich meadows. While V. oughtoni differs from VI parcedentata by having a glassy shell luster and lacking any trace of a depression over the lower palatal, it is unknown how much these features are under environmental control. Determination of the status of these two taxa will require additional DNA sequence analysis. Vertigo ovata Say, 1822; Figs. 5C, 21C Populations are primarily found in graminoid litter and on cattail leaves in swamps, sedge meadows, wet and mesic prairie, low calcareous meadows, river banks, lakeshores, roadside ditches, and wooded wetlands. It is also occasionally found on bedrock outcrops, upland forest, and upland grassland habitats. It can ascend vegetation to approx. 1 m off the ground. Vertigo paradoxa (Sterki, 1900); Figs. 6P, 21D Most frequently found in white cedar Utter pockets on calcareous bedrock ledges, dry microsites in white cedar wetlands, and in thatch on calcareous alvars, seaside turf, and shoreline bedrock outcrops. This taxon introgresses with both Vertigo arthuri and Vertigo hubrichti in regions of range overlap (see above). Reports from the Black HiUs (Frest and Johannes 1993) are based on V. arthuri with a poorly developed apertural callus. Vertigo parvula Sterki, 1890; Figs. 5M, 22A Individuals occur in accumulations of well-decomposed leaf litter in base-rich cove forests, rock outcrops, and talus slopes at mid-low elevations in the central Appalachians and adjacent Piedmont. Vertigo perryi Sterki, 1905; Figs. 5R, 22B Populations reside in humid accumulations of sedge leaf litter on hummock sides in base-poor wet meadows and Sphagnum peatlands as well as in deciduous leaf litter in base-poor red maple, Atlantic white cedar, and northern white cedar wetland forests. Pilsbry (1948) reported that in wet weather individuals will crawl on living vegetation over a third of a meter above the ground; we have observed this behavior to be most pronounced on dead sedge leaves. Vertigo pygmaea (Draparnaud, 1801); Figs. 50, 22C Individuals occur in graminoid thatch and leaf litter accumulations in a variety of anthropogenically disturbed grasslands including roadsides, old fields, yards, and abandoned quarries. It may also occur in more undisturbed habitats such as upland forest, bedrock cliffs, tallgrass prairie, sedge meadows, and acid bogs. We suspect that these 54 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 22. Range maps for Vertigo parvula, Vertigo perryi, Vertigo pygmaea, and Vertigo rugosula. populations represent Eurasian waifs which were brought to North America over a century ago. Vertigo rugosula Sterki, 1890; Fig. 6A, Fig. 22D (syn. Vertigo wheeleri Pilsbry, 1928) Found primarily in graminoid thatch in short turf and scrub such as prairie, mown roadsides, yards, and riparian corridors. It also occasionally occurs in rocky forest. After observing types at ANSP, we agree with Hubricht ( 1974) that V. wheeleri simply represents a population of small V. rugulosa individuals. Vertigo teskeyae Hubricht, 1961; Figs. 5B, 23A Individuals are most commonly seen crawling on open mud and water-saturated logs in floodplain forests and along river, pond, and lake shores following water level drawdown in mid to late summer. They are also occasional in leaf or grass litter adjacent to boggy pools and streams. Vertigo tridentata Wolf, 1870; Figs. 5N, 23B Populations are found in graminiod thatch on calcareous prairie and bedrock glades, in well-decomposed leaf litter accumulations on shaded cliff ledges and talus, and occasionally in upland forest. Hubricht (1985) reported it crawling on mints, while Pilsbry (1948) mentioned it foraging over a meter off the ground on “weeds”. We have seen it crawling on Sedum spp. on limestone cliff ledges in the Ozark Mountains of Arkansas. Vertigo ventricosa (Morse, 1865); Fig. 5Q, 23C Individuals occur in accumulations of humid, well- decomposed graminoid and broadleaf plant litter in moderately to highly acidic wooded and open wetlands, in particular lowland northern white cedar and red maple forest, sedge meadows, Sphagnum peatlands, and poor fens. Although reported from as far west as central Iowa (Hubricht 1985), we have observed no specimens referable to it west of central New York state and eastern Ontario in either the field or from museum collections. All of this western material represents immature Vertigo elatior with incompletely formed apertural lamellae. The two species do co-occur in some New England sites. While distinguishing them can be quite challenging, V. elatior possesses a taller PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 55 Figure 23. Range maps for Vertigo teskeyae, Vertigo tridentata , and Vertigo ventricosa. and somewhat more conical upper half of the shell, whereas V. ventricosa is ovate in outline. The apertural lamellae and sinulus are also more weakly developed in V. ventricosa. Future investigations of these two taxa should be initiated to determine whether they represent ecophenotypes of the same species. ACKNOWLEDGMENTS Tim Pearce kindly loaned us material of Bothriopupa variolosa, Pupoides modicus, Sterkia eyriesi, and Vertigo hebardi from CM for imaging in the specimen figures. Other curators and collection managers who graciously provided access to their collections include: John Slapcinsky and Jochen Gerber (FMNH), Gary Rosenberg (ANSP), Diarmaid O’Foighil (UMMZ), Claire Healy (ROM), Jean-Marc Gagnon (CMN), and Adam Baldinger (MCZ). Michal Horsak kindly provided recently collected shells of Siberian Vertigo parcedentata for comparison. Major funding for this project was provided by the National Science Foundation (EAR-0614963), the Prince Visiting Scholar Fund at the Field Museum of Natural History, Chicago, the Committee on the Status of Endangered Wildlife in Canada, the Maine Department of Inland Fisheries 56 AMERICAN MALACOLOGICAL BULLETIN 28-1/2-2010 and Wildlife, Wildlife Resource Assessment Section, the Massachusetts Natural Heritage and Endangered Species Program, and the Minnesota Nongame Wildlife Tax Checkoff and Minnesota State Park Nature Store Sales through the Minnesota Department of Natural Resources Natural Heritage and Nongame Research Program. We also wish to thank all the people who have helped us trial these keys, as their input was invaluable in their improvement. LITERATURE CITED Bequaert, J. C. and W. B. Miller. 1973. The MoUusks of the Arid Southwest. University of Arizona Press, Tucson. Brooks, S. T. 1936. The land and freshwater Mollusca of Newfound- land. Annals of the Carnegie Museum 25: 83-108. Brooks, S. T. and B. W. Brooks. 1940. Geographical distribution of the recent Mollusca of Newfoundland. Annals of the Carnegie Museum 28: 53-75. Burch, J. B. 1962. How to Know the Eastern Land Snails. Wm. C. Brown Co., Publishers, Dubuque, Iowa. Cameron, R. A. D. and B. M. Pokryszko. 2005. Estimating the spe- cies richness and composition of land mollusc communities. Journal ofConchology 38: 529-547. Coles, B. F. and J. C. Nekola. 2007. Vertigo malleata, a new extreme calcifuge land snail from the Atlantic and Gulf coastal plains of theU.S.A. (Gastropoda, Vertiginidae). The Nautilus 121: 17-28. Dawley, C. 1955. Minnesota land snails. The Nautilus 69: 56-62. Frest, T. J. 1981. Final Report, Project SE-1-2 (Iowa Pleistocene Snail). Iowa Conservation Commission, Des Moines. Frest, T. J. 1982. Final Report, Project SE-1-4 (Iowa Pleistocene Snail). Iowa Conservation Commission, Des Moines. Frest, T. J. 1987. Final Report, Project SE-1-8 (Iowa Pleistocene Snail). Iowa Department of Natural Resources, Des Moines. Frest, T. J. 1990. Final Report, Field Survey of Iowa Spring Fens. Con- tract #65-2454. Iowa Department of Natural Resources, Des Moines. Frest, T. J. 1991. Summary Status Reports on Eight Species of Can- didate Land Snails from the Driftless Area (Paleozoic Plateau), Upper Midwest. Final Report, Contract #301-01366, USFWS Region 3, Ft. Snelling, Minnesota. Frest, T. J. and J. R. Dickson. 1986. Land snails (Pleistocene-recent) of the Loess Hills: A preliminary survey. Proceedings of the Iowa Academy of Science 93: 130-157. Frest, T. J. and E. J. Johannes. 1993. Land Snail Survey of the Black Hills National Forest, South Dakota and Wyoming. Final Report, Contract #43-67TO-2-0054, USDA Forest Service and US Fish and Wildlife Service, Rapid City, South Dakota. Hubricht, L. 1972. Gastrocopta armifera (Say). The Nautilus 85: 73-78. Hubricht, L. 1974. A review of some land snails of the eastern United States. Malacological Review 7: 33-34. Hubricht, L. 1985. The distributions of the native land mollusks of the eastern United States. Fieldiana, New Series 24: 1-191. Kerney, M. P. and R. A. D. Cameron. 1979. Field Guide to the Land Snails of the British Isles and Northwestern Europe. Collins Press, London. Levi, L. R. and H. W. Levi. 1950. New records of land snails from Wisconsin. The Nautilus 63: 131-138. Metcalf, A. L. and R. A. Smartt. 1997. Land Snails of New Mexico. Bulletin 10, New Mexico Museum of Natural History and Sci- ence, Albuquerque, New Mexico. Neck, R. W. 1980. Habitat notes on Gastrocopta riograndensis Sterki. The Veliger 23: 180-182. Nekola, J. C. 2001. Distribution and ecology of Vertigo cristata (Sterki, 1919) in the western Great Lakes region. American Malacological Bulletin 16: 47-52. Nekola, J. C. 2002. Distribution and Ecology of Terrestrial Gastropods in Northwestern Minnesota. Final Report, Minnesota Depart- ment of Natural Resources, St. Paul, Minnesota. Nekola, J. C. 2004. Terrestrial gastropod fauna of northeastern Wisconsin and the southern Upper Peninsula of Michigan. American Malacological Bulletin 18: 21-44. Nekola, J. C. and B. F. Coles. 2001. Systematics and ecology of Gas- trocopta rogersensis (Gastropoda: Pupillidae), a new species from the midwest of the United States of America. The Nautilus 115: 105-114. Nekola, J. C., B. F. Coles, and U. Bergthorsson. 2009. Evolutionary pattern and process in the Vertigo gouldii (Mollusca: Pulmo- nata, Pupillidae) group of minute North American land snails. Molecular Phylogenetics and Evolution 53: 1010-1024. Oggier, P., S. Zschokke, and B. Baur. 1998. A comparison of three methods for assessing the gastropod community in dry grass- lands. Pedobiologia 42: 348-357. Oughton, J. 1948. A Zoogeographical Study of the Land Snails of On- tario. University of Toronto Studies: Biological Series #57. Pearce, T. A., M. C. Fields, and K. Kurita. 2007. Discriminating shells of Gastrocopta pentodon (Say, 1822) and G. tappaniana (C. B. Adams, 1842) (Gastropoda: Pulmonata) with an example from the Delmarva Peninsula, eastern USA. The Nautilus 121: 66-75. Pilsbry, H. A. 1948. Land Mollusca of North America (North of Mexi- co). Monographs of the Academy of Natural Sciences of Philadel- phia 3: 1-1113. Pokryszko, B. M. 1987. On aphally in the Vertiginidae (Gastropoda: Pulmonata: Orthurethra). Journal ofConchology 32: 365-375. Pokryszko, B. M. 2003. Vertigo of continental Europe - autecology, threats, and conservation status (Gastropoda, Pulmonata: Ver- tiginidae). Heldia 5: 13-25. Teskey, M. C. 1954. The mollusks of Brown County, Wisconsin. The Nautilus 68: 24-28. Theler, J. L. 1997. The modern terrestrial gastropod (land snail) fauna of western Wisconsin’s hill prairies. The Nautilus 110: 111-121. Turgeon, D. D., J. F. Quinn, Jr., A. E. Bogan, E. V. COan, F. G. Hochberg, W. G. Lyons, P. Mikkelsen, R. J. Neves, C. F. E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F. G. Thomp- son, M. Vecchione, and J. D. Williams. 1998. Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks, 2nd Edition. American Fisheries Society Special Publication 26. Bethesda, Maryland. PUPILLID LAND SNAILS OF EASTERN NORTH AMERICA 57 Von Proschwitz, T., C. Schander, U. Jueg, and S. Thorkildsen. 2009. Morphology, ecology, and DNA-barcoding distinguish Pupilla pratensis (Clessen, 1871) from Pupilla muscorum (Linnaeus, 1758) (Pulmonata: Pupillidae). Journal ofMolluscan Studies 75: 315.322. Submitted: 19 September 2008; accepted: 30 June 2009; final revisions received: 23 September 2009 APPENDIX 1. Illustrated glossary of important terms. Many of the terms used in the keys are common to the descriptions of most land snail shells. We have chosen to not define all these here as many good sources for this information exist. Interested readers are referred in particular to the excellent illustrated glossary in Kerney and Cameron (1979). However, the use of some of these terms is essentially limited to pupillids, and a review of them is essential for successful use of the taxonomic keys: Alvar: a grassland community residing on a limestone plain with thin or no soil. Angular lamella: the tooth on the parietal wall of the aperture to the right of the parietal lamella in dextral shells (Fig. 24). Apex: the uppermost 2-3 whorls of the shell (Fig. 25). Figure 24. Location of the major apertural lamellae used in pupil- lid identification, illustrated through use of a Vertigo ovata SEM image. Basal lamella: the tooth on the bottom left side (in dextral shells) of the aperture below the columellar lamella (Fig. 24). Body whorl: the final full whorl in an adult shell (Fig. 25). Callus: calcified thickening of the palatal wall of the aperture, often deposited between lamellae (Fig. 25). Columellar lamella: tooth on the columellar wall of the aperture (Fig. 24). Columellar wall: the left side of the aperture in dextral shells. Crest: a bowing out of the shell immediately in back of the aperture as seen in side view (Fig. 25). Ericaceous: plants within the Ericaceae, or heath family. Infra-parietal lamella: the tooth on the parietal wall to the left of the parietal lamella in dextral shells (Fig. 24). Lower palatal lamella: lowermost of the two major teeth often found on the palatal wall (Fig. 24). Palatal depression: indentation of the shell surface at the location of the palatal lamellae (Fig. 25). Palatal wall: the right side of the aperture in dextral shells. Parietal lamella: major tooth in the middle of the parietal wall of the aperture (Fig. 24). Parietal wall: upper side of the aperture. Penultimate whorl: the next to the last whorl in an adult shell (Fig. 25). Pocosin: a peatland of the southeastern U.S.A. with acid soils and semitropical vegetation. Shrub carr: a wetland community dominated by tall shrubs. Sinulus: indentation of the aperture margin along the palatal wall (Fig. 25). Suture: indentation of the shell surface where two whorls meet (Fig. 25). Upper palatal lamella: uppermost of the of the two major teeth often found on the palatal wall (Fig. 24). Apex Penultimate p / * ' Whorl Body I I Whorl Callus Sinulus Figure 25. Major shell features used to identify pupillid taxa, illus- trated through use of SEM images of Vertigo elatior (left), Vertigo bollesiana (upper right), and Vertigo cristata (lower right). Amer. Malac. Bull. 28: 59-80 (2010) The lesser families of Mexican terrestrial molluscs* Edna Naranjo-Garda1 and Neil E. Fahy2 1 Instituto de Biologia, Departamento de Zoologia, Universidad Nacional Autonoma de Mexico, Avenida Universidad 3000, Ciudad Universitaria, Delegacion Coyoacan Mexico D.F. C.P. 04510, Mexico 2 California Academy of Sciences, Golden Gate Park, 55 Music Concourse Drive, San Francisco, California 94118, U.S.A. Corresponding author: naranjo@servidor.unam.mx Abstract: Forty- two families of terrestrial molluscs inhabit Mexico with approx. 1 , 1 78 species and subspecies. The most diverse families are the Urocoptidae (265 species), Spiraxidae (246), Bulimulidae (140), Helicinidae (72), Polygyridae (65), Xanthonychidae (58), Humboldtianidae (49), and Pupillidae (47). The 34 medium and small families comprise about 81% of the total number of families; however, their species total only 236 or 20% of the Mexican species. When looking at their distribution, we noticed that, in general, families cover large areas, but while some species are also found in the United States and Central America, the majority seems to be endemic to Mexico. However, our knowledge of the distribution of terrestrial molluscs throughout Mexico may be incomplete because of a lack of systematic collecting in many regions, particularly in the States of Aguascalientes and Tlaxcala. Key words: Mexico, land snails, distributions This work is based on data collected from literature records with additional data from unpublished sources (Naranjo-Garda and Fahy, unpubl. data). Small families (Table 1) with fewer than 30 species were analyzed (a total of 34), and eight more families, the most diverse families with more than 30 species (indicated by *), are not included in this work. The records were mapped to view species distribution within each family. Only representative maps are included. A total of 236 species and subspecies constitute the 34 families treated in this paper: CERESIDAE The Ceresidae is a family distributed in Mexico and South America (Thompson 1980) and is found in subtropical and tropical environments. In Mexico, the Ceresidae is mainly distributed in the east, with two records in the west. It has eight species in the genera Ceres Gray, 1856 (3), Linidiella Jousseaume, 1889 (2), and Proserpinella Bland, 1865 (3). Ceres eolina (Duclos, 1834) and Ceres salleana Gray, 1856 are from the northern and the central parts of the State of Veracruz (Martens 1890-1901, Pilsbry 1891, Solem 1957, Thompson 1980). Ceres nelsoni Dali, 1898 is found in the southern half of the State of Tamaulipas (Solem 1954, Thompson 1980, Correa-Sandoval 1999, 2002, 2003) and the eastern part of the State of San Luis Potosi (Thompson 1980, Correa-Sandoval 1997, 1998, 1999). Linidiella citrina Thompson, 1987 is from the central part of the State of Veracruz (Comalapa) (Thompson 1987). Linidiella sulfureus Thompson, 1967 is known only from a single locality in the northern part of the State of Chiapas (Solosuchiapa) (Thompson 1967a). Proserpinella berendti Bland, 1865 is from the west central part of the State of Veracruz (Mirador) (Bland 1865, Martens 1890-1901). Proserpinella hannae Dali, 1926 is from Tres Marias Islands (Maria Madre Island) of the State of Nayarit (Dali 1926, Naranjo-Garcia 1994). Proserpinella edentula Naranjo-Garda, 1994 is from the Natural Reserve on the central coast of the State of Jalisco (Estacion de Biologia Chamela) (Naranjo-Garcia 1994). CYCLOPHORIDAE The family Cyclophoridae ranges in the Western Hemisphere from Mexico to the Antilles and South America (De la Torre and Bartsch 1942). In the Eastern Hemisphere, it is in Indochina, Japan, Philippines, India, Indonesia, Melanesia, Africa, Madagascar, and Australia (Boss 1982). Thompson (2008: 74) recognizes the proper name of the family to be Neocyclotidae Kobelt and Moellendorff, 1897. From the “Leslie Hubricht Memorial Symposium on Terrestrial Gastropods” presented at the meeting of the American Malacological Society, held from 29 July to 3 August 2008 in Carbondale, Illinois. 59 60 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Table 1. A list of Mexican terrestrial mollusc families (classified as in Vaught 1989) with number of species and subspecies. Family Species number Family Species number 1. *Helicinidae 72 22. Megomphicidae 1 2. Ceresidae 8 23. Systrophiidae 4 3. Cyclophoridae 23 24. Haplotrematidae 4 4. Megalomastomidae 4 25. Punctidae 7 5. Diplommatinidae 2 26. Helicodiscidae 6 6. Annulariidae 8 27. Charopidae 2 7. Truncatellidae 6 28. Discidae 2 8. Veronicellidae 2 29. Oreohelicidae 7 9. Ellobiidae 9 30. Succineidae 24 10. Carychiidae 2 31. Sagdidae 1 11. Achatinellidae 2 32. Gastrodontidae 5 12. Cochlicopidae 1 33. Euconulidae 17 13. *Pupillidae 47 34. Vitrinidae 3 14. Vallonidae 5 35. Zonitidae 18 15. Strobilopsidae 7 36. *Polygyridae 65 16. *Bulimulidae (= Orthalicidae) 140 37. Thysanophoridae 15 17. Amphibulimidae 2 38. *Humboldtianidae 49 18. *Urocoptidae 265 39. *Xanthonychidae 58 19. Ferussaciidae 4 40. Helminthoglyptidae 10 20. Subulinidae 21 41. Arionidae 1 21. *Spiraxidae 246 42. Philomycidae 3 * Family not treated in this work. The Cyclophoridae are distributed across four regions of Mexico. Amphicyclotus Crosse and Fischer, 1879 is found mainly in the southern most region, with some records in the central part of Veracruz; Aperostoma Troschel, 1847 is distributed along the Gulf of Mexico from the mid-part of the State of Tamaulipas to southern Veracruz; Dicrista Thompson, 1969 is found in western Mexico from below the central part of the State of Sinaloa to the State of Guerrero; and Neocyclotus Fischer and Crosse, 1886 is found in south- ern Mexico, in southern Veracruz, Oaxaca, southern Tabasco, Chiapas, and the entire Peninsula of Yucatan. While the dis- tribution of Dicrista does not overlap the other genera, the distribution of Amphicyclotus generally overlaps that of Neo- cyclotus in the State of Chiapas, where most of the range of Amphicyclotus occurs. Twenty-three Mexican species of Cyclophoridae have been described in the genera Amphicyclo- tus (6), Aperostoma (3), Dicrista (7), Neocyclotus (6), and Xenocyclus Thompson, 1969 (1). Amphicyclotus boucardi (Pfeiffer, 1856) is from the center of the State of Veracruz (Cordoba) (Martens 1890-1901, Diaz de Leon 1912, Thompson 1966 ). Amphicyclotus paulsonorum Thompson, 1969 is found in the south coastal area of the State of Chiapas (Thompson 1969). Amphicyclotus maleri Crosse and Fischer, 1883 is found in the south central part of the State of Tabasco (Poana Mountains) (Pilsbry 1893, Solem 1956) and in the southern part of the State of Oaxaca (Santa Efigenia, Tehuantepec) (Martens 1890-1901, Diaz de Leon 1912). Amphicyclotus megaplanus Morrison, 1955 is found in the western part of the State of Chiapas (Ocozocoautla) (Morrison 1955, Thompson 1969),(E10coteforest) (Bequaert 1957) . Amphicyclotus texturatus spiralis Thompson, 1969 is found in the southern corner of the State of Chiapas (Thompson 1969). Aperostoma mexicanum salleanum (Martens, 1865) is found throughout the State of Veracruz (Papantla) (Martens 1890-1901, Diaz de Leon 1912, Baker 1922, 1928, Thompson 1969, Solem 1956) and in the northern part of the State of Oaxaca (Tuxtepec, Playa Vicente) (Solem 1956, Chevallier 1965). Aperostoma palmeri (Bartsch and Morrison, 1942) is from the mid-region of the State of Tamaulipas (Bartsch and Morrison 1942, Solem 1954, 1956, Correa-Sandoval 1999, 2002, 2003), the southeastern part of the State of San Luis Potosi (Solem 1956, Thompson 1969, Correa-Sandoval 1997, 1998, 2003), the northern part of the State of Hidalgo, and the north central region of the State of Veracruz (Solem 1956). Aperostoma walkeri Baker, 1928 is found in the northern part of the States of Puebla and Veracruz (Baker 1928, Bartsch and Morrison 1942, Morrison 1955, Solem 1956, Correa-Sandoval 2000, 2003). Dicrista cooperi (Tryon, 1863) is from the southern region of the State of Sinaloa (Tryon 1863, Bartsch and Morrison 1942, Thompson 1969) and on the coast of the States of Jalisco, Colima, and Guerrero (Bartsch and Morrison 1942, Thompson 1969). It has also been found in the State of Oaxaca by Thompson (1969) who did not provide a precise locality. The following species are known only from their type locality: Dicrista damianensis (Solem, 1956) in the southern part of the State of Michoacan (near San Pedro Damian Naranjestilla) (Solem 1956, Thompson 1969); Dicrista flavescens Thompson, 1969 near Mazatlan in the State of Guerrero (Thompson 1969); Dicrista indentata Thompson, 1969 in the State of Michoacan (near San Vicente), Dicrista liobasis Thompson, 1969 from the State of Jalisco (near Pihuamo). Dicrista rugosa Thompson, 1969 is found in the central region of the State of Colima and Xenocyclus patulus Thompson, 1969 from the State of Colima (Tamala) (Thompson 1969). Dicrista petersi petersi (Solem, 1956) is from the State of Michoacan (La Placita = Sulatillo) (Solem 1956, Thompson 1969). Amphicyclotus palenquensis (Pilsbry, 1935) is from south central Veracruz (Solem 1956) and north and northeastern parts of the State of Chiapas (Pilsbry 1935, Bequaert 1957). Neocyclotus dysoni dysoni (Pfeiffer, 1851) is found in the central part of the State of Veracruz (Martens 1890-1901, Diaz de Leon 1912, Baker 1922, Chevallier 1965), in the MEXICAN LAND SNAILS 61 central and southern portions of the State of Oaxaca (Martens 1890-1901, Diaz de Leon 1912), in the southern part of the State of Tabasco (Martens 1890-1901), in the northern part of the State of Campeche (Thompson 1967b), very possibly all around the State of Yucatan, since various localities have been recorded from that state (Martens 1890-1901, Pilsbry 1891, Branson and McCoy 1965), and also occurs in the center of the State of Quintana Roo (Xiatil) (Thompson 1967b). Neocyclotus dysoni ambiguum (Martens, 1890) is found in the south and south center of the State of Veracruz and at the north and center of the State of Chiapas (Martens 1890-1901, Bartsch and Morrison 1942, Solem 1956, Thompson 1969). It is also found at the east central part of the State of Oaxaca (Rio Grande) and in the southern part of the State of Tabasco (Mountains of Poana, Teapa) (Solem 1956, Thompson 1969). Neocyclotus dysoni aureum (Bartsch and Morrison, 1942) is found in the southern part of the State of Oaxaca (Panistlahuaca, Gamboa) (Bartsch and Morrison 1942, Solem 1956), in the western part of the State of Campeche (Branson and McCoy 1963), and in the northeast (Laguna Ocotal) and west center (El Sumidero, Tuxtla Gutierrez) ofthe State of Chiapas (Bequaert 1957). Neocyclotus dysoni berendti (Pfeiffer, 1861) is from the center and southern part of the State of Veracruz (Baker 1922, Solem 1956, Naranjo-Garcia and Polaco 1997) and is widely distributed in the Peninsula of Yucatan (Martens 1890-1901, Baker 1928, Bequaert and Clench 1933, 1936, 1938, Richards 1937, Bartsch and Morrison 1942, Harry 1950, Branson and McCoy 1963, 1965, Rehder 1966, Thompson 1967b, Gomez-Espinosa 1999 ). Neocyclotus dysoni cookei (Bartsch and Morrison, 1942) is found in the central part of the State of Campeche (Thompson 1967b). The species Neocyclotus simplicostus Thompson, 1969 is found in the southeastern corner of the State of Chiapas (Thompson 1969). MEGALOMASTOMIDAE The family Megalomastomidae, exclusively American (Boss 1982), is distributed from southern Mexico to Guatemala and in Cuba, Hispaniola, Puerto Rico, and the Virgin Islands (De la Torre and Bartsch 1942). The Mexican distribution occurs in two separate regions: at the so-called “Los Tuxtlas” region, and south central Veracruz and eastern Chiapas in the Selva Lacandona. In Mexico it is represented by four species in the genus Tomocyclus Crosse and Fischer, 1872. Tomocyclus guatemalensis (Pfeiffer, 1851) and Tomocyclus lunae Bartsch, 1945 are from the southern part of the State of Veracruz (Martens 1890-1901, Bartsch 1945, Thompson 1963, Chevallier 1965, Naranjo-Garcia and Polaco 1997) and Tomocyclus gealei Crosse and Fischer, 1872 and Tomocyclus simulacrum (Morelet, 1849) are from the eastern part of the State of Chiapas (Lagos de Montebello) (Martens 1890-1901, Bequaert 1957, Thompson 1963, Chevallier 1965, Naranjo-Garcia et al., in press). DIPLOMMATINIDAE The family Diplommatinidae is found in Europe, from southeast Asia to Australia, Melanesia, and Micronesia, and Mexico, Central America, and the Antilles (Boss 1982). It has a somewhat subtropical and tropical distribution. Two Mexican species in the genus Adelopoma Doering, 1884 have been recognized. Adelopoma stolli Martens, 1890 is located in the southern part of the States of Tamaulipas and San Luis Potosi (Correa-Sandoval 1998, 1999, 2003). The other species is undescribed from the center of the State of Tamaulipas (Ciudad Victoria, Sierra Victoria) and reported by Correa-Sandoval (2002). ANNULARIIDAE The family Annulariidae is considered by Burch (1962) to have a tropical and subtropical distribution. It is well represented in the Antilles (Pilsbry 1948) and ranges from Florida, Mexico, Central America, northern South America (Bolivia), and the Antilles (Henderson and Bartsch 1921, Boss 1982, Watters 2006). The Annulariidae inhabit southern Mexico (Fig. 1). The genus Choanopoma Pfeiffer, 1848 occurs on the entire Peninsula of Yucatan, Chiapas and eastern Oaxaca. Chondropoma Pfeiffer, 1847 is apparently confined to two separate regions: one in the central part of Veracruz and the second in the southern part of the State of Tabasco. In Mexico the Annulariidae has eight species in two genera Choanopoma (6) and Chondropoma (2). Choanopoma andrewsae (Ancey, 1886) is found at Cozumel Island (Richards 1937), as well as the northeast, center, and southwest parts of the State of Quintana Roo (Rehder 1966), north and north-northwest portions of the State of Campeche, and the south and southeastern portions of the State of Yucatan (Thompson 1967b). Choanopoma gaigei Bequaert and Clench, 1931 is found in the entire eastern part of the State of Campeche, in the mid western (Merida; Chichen Itza), and southern parts ofthe State of Yucatan (Bequaert and Clench 1931), and in the northern, center, and southwestern portions of the State of Quintana Roo (Thompson 1967b, Gomez-Espinosa 1999). Choanopoma largillierti (Pfeiffer, 1846) is from the north to the central part of the State of Campeche (Bequaert and Clench 1936, Solem 1961, Branson and McCoy 1965, Thompson 1967b). In the State of Yucatan, it is distributed from the center (Chichen Itza) towards the west (Pilsbry 1891, Bequaert and Clench 1933, 1936, 1938, 62 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 1. Distribution of the genera Choanopoma and Chondropoma in Mexico. Richards 1937, Harry 1950, Branson and McCoy 1963, 1965, Chevallier 1965, Thompson 1967b). It is also found in the northern, center, and southwestern regions of the State of Quintana Roo (Solem 1961, Rehder 1966, Thompson 1967b). The distribution of Choanopoma martensianun (Pilsbry, 1900) is the south center of the State of Tabasco (Pilsbry 1900, Thompson 1957, Rangel-Ruiz and Gamboa 2001). Choanopoma sumichrasti Crosse and Fischer, 1874 is from about the center of the State of Chiapas (Tuxtla Gutierrez, Chiapa de Corzo) (Fischer and Crosse 1870-1902, Bequaert 1957, Solem 1961, Chevallier 1965) and from the coastal area of the State of Oaxaca (Tehuantepec) (Chevallier 1965). According to Thompson (2008) in his recent revision of the entire molluscan fauna of Mexico and Central America, the correct name of Choanopoma sumichrasti is Annularia sumichrasti (Crosse and Fischer, 1874). Choanopoma tere- costatum Thompson, 1966 is found in the western part of the State of Chiapas (Ocozocoautla) (Thompson 1966) and at the south center of the State of Tabasco (Parque Estatal La Sierra) (Rangel-Ruiz and Gamboa 2001). Chondropoma rubicundum (Morelet, 1849) is from the northern end of the State of Chiapas (Laguna Ocotal) (Bequaert 1957, Solem 1961) and in the contiguous State of Tabasco at the south central mountains of Poana (Pilsbry 1893, Solem 1961). Chondropoma cordovanum (Pfeiffer, 1856) is found in the center ofthe State ofVeracruz (Cordoba) (Martens 1890-1901, Baker 1928, Solem 1961). TRUN CATELLID AE The family Truncatellidae has a worldwide (Boss 1982) tropical and subtropical distribution (Burch 1962). Because snails of this family are amphibious intertidal inhabitants, the Truncatellidae in the western part of Mexico is distributed around the Peninsula of Baja California and all along the Caribbean coast from the State of Tamaulipas to the State of Quintana Roo. The family in Mexico has six species in the genus Truncatella Risso, 1826. Truncatella bairdiana C. B. Adams, 1852 is from the center coastal area of the State of Guerrero (Acapulco) (Martens 1890-1901). Truncatella bilabiata Pfeiffer, 1840 is on the Isla del Carmen, State of Campeche (Martens 1890- 1900) and in pools south of Progreso, State of Yucatan (Bequaert and Clench 1936, Harry 1950). Truncatella caribaeensis Sowerby in Reeve, 1842 is found along the coast of the Gulf of Mexico from the State of Tamaulipas to the State of Quintana Roo (Bequaert and Clench 1933, Garcia- Cubas 1963, 1968, 1981, Andrews 1981, Flo res- Andolais etal. 1988, Garcia-Cubas and Reguero 1990, Reguero etal. 1991). Truncatella californica Pfeiffer, 1857 is from both coasts of the Peninsula of Baja California to the upper Gulf of California and on various nearby islands (Baker 1902, Lowe 1913, Pilsbry 1927, Keen 1971, Smith etal. 1990). Truncatella guadalupensis Pilsbry, 1901 is from Guadalupe Island, State of Baja California MEXICAN LAND SNAILS 63 (Pilsbry 1901, Smith etal. 1990). Truncatella pulchella Pfeiffer, 1839 is from Dzilam de Bravo, State of Yucatan and Mujeres Island, State of Quintana Roo (Parodiz 1977). VERONICELLIDAE Veronicellidae is found in tropical and subtropical areas of America, Africa, Asia, and Oceania (Boss 1982). It has two apparently native Mexican species that belong to the genus Leidyula Baker, 1925: Leidyula floridana (Leidy and Binney in Binney, 1851) and Leidyula moreleti (Fischer, 1871). Leidyula floridana is recorded from the central region of the State of Nuevo Leon. In contrast, L. moreleti mainly inhabits the east coastal region of Mexico, except for two localities: one in the southern part of the State of Nayarit and the other in the southern part in the State of Oaxaca (Naranjo-Garcla et al. 2007). ELLOBIIDAE Ellobiidae lives in the intertidal zone and has a worldwide distribution: the Red Sea, Pacific and Atlantic Oceans, near Australia, the Philippines, Indo-Pacific, and Mediterranean Sea (Thiele 1935). It is found on both coasts of Mexico. There are nine Mexican species in four genera: Marinula King and Broderip, 1832 (1), Melampus Montfort, 1810 (4), Pedipes Bruguiere, 1792 (3), and Sarnia H. and A. Adams, 1855 (1). The species Marinula rhoadsi Pilsbry, 1910, Melampus mousleyi Berry, 1964, Melampus olivaceus Carpenter, 1857, Pedipes liratus Binney, 1860, Pedipes unisulcatus Cooper, 1866, and Sarnia mexicana (Berry, 1964) are endemic; all inhabit the northern part of the Gulf of California (Keen 1971, Gonzalez 1993). Melampus tabogensis C. B. Adams, 1852 is from the central coastal part of the State of Jalisco (Barra deNavidad) (Keen 1971). Melampus coffeus (Linnaeus, 1758) and Pedipes mirabilis (Muhlfeld, 1816) are found along the Gulf of Mexico coasts (Abbott 1974). CARYCHIIDAE The family Carychiidae inhabits northern Europe, Asia, and North America (Pilsbry 1948, Boss 1982). It is found in the eastern part of Mexico from the State of Nuevo Leon to the State of Tabasco ( Contreras- Arquieta 1995, Pilsbry 1891, 1903, Baker 1930, Correa-Sandoval 1993, 1998, 1999, 2002, 2003, Rangel-Ruiz and Gamboa 2001). Two subspecies of Carychium Muller, 1774 exist in Mexico. Carychium exile exile H. C. Lea, 1842 is from the State of Nuevo Leon, without a specific locality (Contreras-Arquieta 1995). Carychium exile mexicanum Pilsbry, 1891 (Fig. 2) is mainly in the east coastal States of Mexico: Tamaulipas Figure 2. Distribution of the species Carychium exile mexicanum in Mexico. 64 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 (Correa-Sandoval 1999, 2002), Nuevo Leon (Pilsbry 1903, Correa-Sandoval 1993, 1999), San Luis Potosi (Correa- Sandoval 1998, 2003), Puebla (Baker 1930, Correa-Sandoval 1999), Veracruz (Pilsbry 1891, 1903, Correa-Sandoval 1999), and Tabasco (Rangel-Ruiz and Gamboa 2001). ACHATINELLIDAE The general family distribution is on the Pacific Islands and contiguous areas (Boss 1982). Consequently, the family had been found on the Mexican Pacific islands. There are only two Mexican species in the genus Tornatellides Pilsbry, 1910: Tornatellides mexicana Dali, 1926 is on Socorro Island and Tornatellides clarionensis Dali, 1926 is from Clarion Island. Both islands are in the Revillagigedo Archipelago, State of Colima (Dali 1926). COCHLICOPIDAE The family Cochlicopidae is of Palearctic origin and the genus Cochlicopa Ferussac, 1821 in Risso, 1826 is considered Holarctic (Pilsbry 1948, Boss 1982) and contains a single Mexican species: Cochlicopa lubrica (Muller, 1774) (Fig. 3) that inhabits the north central states (northwestern Chihuahua, southern Nuevo Leon, and Durango) (Pilsbry 1953a, Bequaert and Miller 1973, Contreras- Arquieta 1995, Correa-Sandoval 2003). VALLONIDAE The Vallonidae is a Holarctic family (Bequaert and Miller 1973, Boss 1982) and is distributed mainly in northern Mexico, with a record in the center of Mexico. Five species, in two genera are recorded from Mexico: Erectidens Pilsbry, 1953 (1) and Vallonia Risso, 1826 (4) (Fig. 3). Erectidens trichalus Pilsbry, 1953 is found in the west central part of the State of Nuevo Leon (Rio Maurisco) (Pilsbry 1953b). Vallonia cyclophorella Sterki, 1892 is found at Sierra San Pedro Martir, State of Baja California (Smith et al. 1990). Vallonia excentrica Sterki, 1893 is from the center of Mexico (Cuernavaca) (Pilsbry 1948). Vallonia gracilicosta Reinhardt, 1883 is in Nuevo Leon, with no specific locality specified (Correa-Sandoval 2003). Vallonia perspectiva Sterki, 1892 is from along Rio Piedras Verdes, State of Chihuahua (Pilsbry 1953a). Bequaert and Miller (1973) record that it reaches northern Mexico in Sonora and Chihuahua, without giving precise localities. STROBILOPSIDAE The family Strobilopsidae is widely distributed from Eastern North America (Quebec, Ontario, and Manitoba, latitude 52°N) to South America (Venezuela and eastern Brazil), in the Galapagos Islands, Japan, Korea, China, and Philippines (Pilsbry 1948, Boss 1982). Pilsbry (1948) thought Figure 3. Distribution of the species Cochliopa lubrica and three species of Vallonia in Mexico. MEXICAN LAND SNAILS 65 that Asia might be the center of evolution of the genus Strobilops , due to the great diversity of the genus in that territory. The family is distributed mainly in eastern Mexico from the north (States of Tamaulipas, Nuevo Leon, San Luis Potosi, and Puebla) to the south in Veracruz (Los Tuxtlas). In western Mexico, it is found in the southern part of the State of Baja California and in the Tres Marias Islands (State of Nayarit). The family Strobilopsidae is represented in Mexico by seven species in the genus Strobilops Pilsbry, 1893. Strobilops aenea mexicana Pilsbry, 1927 is from the west center of the State of Nuevo Leon (Pilsbry 1953a), eastern part of the State of San Luis Potosi (Correa-Sandoval 1998, 2003), northern part of the State of Puebla (Baker 1930), and in the west central part of the State of Veracruz (Baker 1930, Correa- Sandoval 1999). Strobilops californica Miller and Christensen, 1980 is from the southern tip of the State of Baja California Sur (Miller and Christensen 1980, Smith et al. 1990). Strobilops hubbardi (Brown, 1861 ) is from the mid-southern part of the State of Tamaulipas (Correa-Sandoval 1999, 2002) and from the eastern part of the State of San Luis Potosi (Correa- Sandoval 1998, 2003). Strobilops labyrinthica (Say, 1817) is from the west central part of the State of Veracruz (Martens 1890-1901, Diaz de Leon 1912). Strobilops sinaloa Morrison, 1953 is known only from vegetables confiscated at the United States boarder coming from the State of Sinaloa (Morrison 1953); its type locality needs to be located. Strobilops strebeli (Pfeiffer, 1861) is present at the Tres Marias Islands (Socorro Island), State of Nayarit (Dali 1926). Strobilops veracruzensis Pilsbry, 1927 is from the center of the State of Veracruz (Veracruz) (Clench and Turner 1962). AMPHIBULIMIDAE The family Amphibulimidae is mainly distributed in South America and the Antilles with one genus in South Africa (Boss 1982). There are two Mexican species in the genus Simpulopsis Beck, 1837. Simpulopsis aenea Pfeiffer, 1861 is found in the center of the State of Oaxaca (La Parada) (Martens 1890-1901) and Simpulopsis Simula (Morelet, 1851) in the northeastern part of the State of Chiapas (Laguna Ocotal to El Censo) (Bequaert 1957). The genus Simpulopsis belongs to the family Orthalicidae according with Thompson (2008). FERUSSACIIDAE The family Ferussaciidae is worldwide in tropical and subtropical habitats (Boss 1982). It is mainly in eastern Mexico with some records in the western States of Sonora and Nayarit. It has four Mexican species, in three genera: Cecilioides Ferussac, 1814 (2), Coelostele Crosse, 1876, and Karolus de Folin in de Folin and Perrier, 1870. Cecilioides section Caecilianopsis Pilsbry, 1907 has a tropical American distribution; Coelostele has species in Asia, northern Africa, southern Spain, and eastern Mexico (Thiele 1935). Cecilioides ( Caecilianopsis ) jod Pilsbry, 1907 has been recorded only from Tampico, in the southern part of the State of Tamaulipas (Hinkley 1907, Pilsbry 1907). There is an undescribed species of Cecilioides from Alamos, Sonora (Naranjo-Garcia 1991). Coilostele tampicoensis (Pilsbry, 1907) is recorded from Tampico, in the southern part of the State of Tamaulipas (Clench and Turner 1962). Thompson (2008: 254) places Coelostele in the family Carychiidae and mentions that the genus is located in India, southern Arabia, Egypt, Syria, and southern Spain. Being Coilostele tampicoensis a single species in the Americas, it is most likely an introduction (Thompson 2008). Karolus consobrinus primus (De Folin, 1870) is the most widely distributed Mexican species of this family. It is found in the center and southeastern part of the State of Sonora (Pilsbry 1953a, Naranjo-Garcia 1991), in the northern part of the State of Sinaloa (Pilsbry 1953a), and in northwestern Mexico. In eastern Mexico it is recorded from the eastern part of the States of Nuevo Leon (Correa-Sandoval 1997), in the mid-southern part of the State of Tamaulipas (Harry 1950, Correa-Sandoval 2002, 2003), in the eastern part of the State of San Luis Potosi (Correa-Sandoval 1998), in northern and center parts of the State of Veracruz (Fischer and Crosse 1870-1902, Martens 1890-1900, Baker 1930, Correa-Sandoval 1999, 2000, 2003), in the northern part of the State of Puebla (Baker 1930), and in about the center of the State of Yucatan (Chichen Itza) (Richards 1937, Harry 1950). Dali (1926) recorded the species from Tres Marias Islands (Maria Madre), State of Nayarit and from the Revillagigedo Archipelago (Socorro Island), State of Colima. SUBULINIDAE The family Subulinidae has representatives in Europe as well as in tropical America (Boss 1982). The genus Subulina Beck, 1837 has various tropical species. Leptinaria Beck, 1837 and Lamellaxis Strebel and Pfeffer, 1882 inhabit the American tropics (according to Thiele [1935], Lamellaxis is a section of Leptinaria), and Opeas Albers, 1850 has a subtropical and tropical distribution (Thiele 1935). The family is mainly distributed in the Gulf of Mexico states from the north to the Peninsula of Yucatan, with some records in the Peninsula of Baja California and the Mexican Pacific States. So far, the family shows a more tropical distribution. In Mexico, the Subulinidae is represented by 66 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2 -2010 twenty-one species and subspecies in six genera: Allopeas Baker, 1935 (3), Diaopeas Haas, 1962 (1), Lamellaxis (8), Leptinaria (2), Opeas (5), and Subulina (2). Allopeas gracile (Hutton, 1834) is from the southern part of the State of Baja California Sur (Smith etal. 1990) and mid- southern part of the State of Sonora (Drake 1953, Naranjo- Garcia 1991), in eastern Mexico from the west central part of the State of Tamaulipas (Correa- Sandoval 2002, 2003), the eastern part of the State of San Luis Potosi (Correa-Sandoval 1997, 2003), the mid-northern part of the State of Campeche (Thompson 1967b), and from the northwestern and center portions of the State of Yucatan (Bequaert and Clench 1933, 1938). This species was described from India, nowadays it is distributed world wide; however, Pilsbry (1946) considered the species as native to tropical America. Allopeas micra micra (D’Orbigny, 1835) is recorded from eastern Mexico, that is from the southern mid-part of the State of Tamaulipas (Martens 1890-1901, Correa-Sandoval 2002) , the southeastern part of the State of San Luis Potosi (Correa-Sandoval 1997), from the center of the State of Veracruz (Correa-Sandoval 2000), from the southern and center of the State of Tabasco (Martens 1890-1901, Correa- Sandoval 1999), from the center of the State of Yucatan, and from the State of Quintana Roo (Cozumel Island) (Richards 1937). Allopeas micra micra (D’Orbigny, 1835) is Leptopeas micra (D’Orbigny, 1835) with the studies of Thompson (2008: 538). The following taxa have been recorded from a single locality: Allopeas micra mazatlanica (Pilsbry, 1931) is from the State of Sinaloa (Isla del Paro) (Pilsbry 1931). Prom the State of Veracruz various species were recorded: Lamellaxis argutus (Pilsbry, 1906) is from the center (Orizaba) (Pilsbry 1906) as is Leptinaria imperforata (Strebel, 1882) (Jalapa) (Martens 1890- 1901), and from the south is Lamellaxis interstriatus (Tate, 1870) (Naranjo-Garcia 2003). Lamellaxis exiguus (Martens, 1898) is from the southern part of the State of Tabasco (Teapa) (Martens 1890-1901), and Lamellaxis semitriatus (Morelet, 1851) is from the northern part of the State of Chiapas (Palenque) (Morelet 1851, Martens 1890-1901). Lamellaxis argutus (Pilsbry, 1906), Leptinaria imperforata (Strebel, 1882), and Lamellaxis semitriatus (Morelet, 1851) change to the following species names according to Thompson (2008: 536, 535, 540): Leptopeas argutus (Pilsbry, 1906), Lamellaxis imperforatus Strebel, 1882, and Leptopeas semitriatum (Morelet, 1851), respectively. Diaopeas beckianum (Pfeiffer, 1846) is from the southern part of the State of Tamaulipas (Correa-Sandoval 1999, 2002, 2003) , the southeastern part of the State of San Luis Potosi, and in the northern and southern parts of the State of Veracruz (Correa-Sandoval 1999, 2000, 2003, Naranjo-Garcia 2003), and in the State of Tabasco (no locality given) (Correa- Sandoval 1999 ). Diaopeas beckianum (Pfeiffer, 1846) changed to Beckianum beckianum (Pfeiffer, 1846) in Thompson’s work (2008: 523). Lamellaxis martensi (Pfeiffer, 1856) is from the center (Martens 1890-1901) and southern part (Naranjo-Garcia 2003) ofthe State ofVeracruz. Lamellaxis mexicanus mexicanus (Pfeiffer, 1866) is from the mid-southern part of the State of Tamaulipas (Correa-Sandoval 2002, 2003), from the eastern part of the State of San Luis Potosi (Correa-Sandoval 1997), and from the mid-northern part of the State of Veracruz (Martens 1890-1901, Correa-Sandoval 2000). Lamellaxis mexicanus abbreviatus (Martens, 1898) and Lamellaxis mexicanus turritus (Martens, 1898) are both from various sites in the center of the State ofVeracruz (Martens 1890-1901). Leptinaria tamaulipensis Pilsbry, 1903 is from the mid- southern part of the State of Tamaulipas (Pilsbry 1903, Correa-Sandoval 2002) and the southeastern part ofthe State of San Luis Potosi (Correa-Sandoval 1997). Thompson (2008: 526) considers that Leptinaria tamaulipensis Pilsbry, 1903 should be Lamellaxis tamaulipensis (Pilsbry, 1903). Opeas colimense (Crosse and Pischer, 1859) is from the State of Colima and the center of the State of Veracruz (Cordova) (Martens 1890-1901). In Thompson’s work (2008: 537), Opeas colimense (Crosse and Fischer, 1859) is Leptopeas colimense (Crosse and Fischer, 1859). Opeas odiosum Pilsbry, 1899 and O. patzcuarense Pilsbry, 1899 are from the vicinity of Patzcuaro in the State of Michoacan (Pilsbry 1899). Opeas pumilum Pfeiffer, 1840 is from the northern end of the Peninsula of Baja California (Smith et al. 1990) and the southern part of the State ofVeracruz (Naranjo-Garcia 2003). Opeas rhoadsae Pilsbry, 1899 is from the west central part of the State of Nuevo Leon (Diente) (Pilsbry 1899). Subulina octona (Bruguiere, 1789) is found in eastern Mexico from the southern part of the States of Tamaulipas (Correa-Sandoval 2002, 2003) and San Luis Potosi (Correa- Sandoval 1997, 2003), from the west center of the State of Veracruz (Martens 1890-1901, Correa-Sandoval 2000), in the southwestern part of the State of Tabasco (Martens 1890- 1 90 1 ), at the southwestern part of the State of Campeche, and at the northwestern part of the State of Yucatan (Martens 1890-1901, Correa-Sandoval 1999). It is a widely distributed species around the world (Martens 1890-1901). Subulina porrecta Martens, 1898 is from the southwestern part of the State of Tabasco (Teapa), and from the State of Yucatan (Martens 1890-1901). To Thompson (2008: 639), Subulina porrecta Martens, 1898 belongs to the family Spiraxidae under Mayaxis porrecta (Martens, 1898). MEGOMPHICIDAE The Megomphicidae is represented in Mexico with one member of the subfamily Ammonitellinae. According to MEXICAN LAND SNAILS 67 Pilsbry (1939), the subfamily Ammonitellinae is a declining ancient group present in the western United States (Washington, Montana, Oregon, and California). Glyptostoma Bland and Binney, 1873 is one of four genera that form this subfamily and the genus is found from San Gabriel Range, California to Ensenada, State of Baja California (Pilsbry 1939). Glyptostoma newberryanum depresum Bryant, 1902 is found in the northwestern corner of the State of Baja California (Smith et al. 1990). Pilsbry (1939) stated that it needs further anatomical studies and Smith et al. ( 1990) consider it a weakly characterized subspecies. SYSTROPHIIDAE The family Systrophiidae comes from South America (Boss 1982) where various genera and species thrive. The Systrophiidae inhabit the east central part of Mexico in the State of San Luis Potosi, in northern part of the State of Puebla, and at the center of the State of Veracruz. There are some records of the family in the central coast of the State of Campeche and south corner of the State of Yucatan. It has four Mexican species in the genus Miradiscops Baker, 1925. Miradiscops haplocochlion Thompson, 1967 is from the central coast of the State of Campeche (Thompson 1967b). Miradiscops maya (Pilsbry, 1920) is from the southern corner of the State of Yucatan (Thompson 1967b). Miradiscops opal (Pilsbry, 1920) and Miradiscops puncticipitis (Pilsbry, 1926) have similar distributions. They both are found in the eastern part of the State of San Luis Potosi (Correa- Sandoval 1997), the northern part of the State of Puebla, and in the west central part of the State of Veracruz (Baker 1930). Miradiscops puncticipitis is also found in the southern part of the State of San Luis Potosi (Correa-Sandoval 1997). HAPLOTREMATIDAE The Haplotrematidae is mainly in the western United States with various species. According to Burch (1962), the family comes from North America although it has repre- sentatives in the Antilles, Central America, and northern South America (Boss 1982). In Mexico, it consists of the genus Haplotrema Ancey, 1881 with four species. The species Haplotrema caelatum (Mazyck, 1886) and Haplotrema transufga (Hemphill, 1892) inhabit the north- western end of the State of Baja California (Smith etal. 1990). Haplotrema guadalupensis Pilsbry, 1927 is from Guadalupe Island, State of Baja California (Pilsbry 1927). A fourth taxon is an undescribed species reported by Correa- Sandoval (2003) from the States of Nuevo Leon and Tamaulipas on the eastern side of Mexico. PUNCTIDAE The Punctidae is mainly distributed in the northern hemisphere with some representatives in the south (South Africa) as is the case of the genus Punctum Morse, 1864 (Pilsbry 1948). The Mexican distribution of Punctidae is scattered. It is found in the northern part of the State of Baja California, in the States of Nayarit, Jalisco, Colima, and Federal District (Mexico City), northern part of the State of Puebla, southeastern part the State of San Luis Potosi, and in the States of Tamaulipas and Nuevo Leon. The Punctidae is represented in Mexico by seven species and subspecies in the genera Paralaoma Iredale, 1913 (3) and Punctum (4). Paralaoma caputspinulae caputspinulae (Reeve, 1852) is from the northern part of the State of Baja California (Smith et al. 1990). Paralaoma caputspinulae jaliscoensis (Pilsbry, 1926) is from the central part of the State of Jalisco (Pilsbry 1926, Baker 1930) and from the western part of the Federal District (Mexico City) (Baker 1927, 1930). Thompson (2008: 675) considers Paralaoma caputspinulae caputspinulae (Reeve, 1852) to be a synonym of Paralaoma servilis ; in addition, this species has been introduced in various temperate areas around the globe. Yet, Paralaoma caputspinulae jaliscoensis (Pilsbry, 1926) is Punctum conspectum jaliscoensis (Pilsbry, 1926) under Thompson’s (2008: 678) studies. Paralaoma vitreum (Baker, 1930) is from the western part of the State of Nuevo Leon (Correa-Sandoval 1997, 2003) and from the center of the States of Tamaulipas (Correa-Sandoval 2002) and Veracruz (Baker 1930). Punctum minutissimum (Lea, 1841) is from the northern part of the State of Puebla (Baker 1930, Pilsbry 1948), the eastern part of the State of San Luis Potosi (Correa-Sandoval 1997, 1998, 2003), and in the Pacific islands: Islas Tres Marias (Maria Magdalena) of the State of Nayarit and in the Revillagigedo Archipelago (Socorro Island) State of Colima [as Punctum pygmaeum (Draparnaud, 1801)] (Dali 1926). The two subspecies Punctum minutissimum rotundum (Dali, 1926), and Punctum minutissimum albeolum (Dali, 1926) , and the species Punctum planatum Dali, 1926 are from the Tres Marias Islands, State of Nayarit, both subspecies are present at Maria Magdalena Island, and P. m. albeolum and P. planatum are also present at Maria Madre Island (Dali 1926). Thompson (2008: 679) considers Punctum planatum Dali, 1926 to be a synonym of Chanomphalus pilsbryi (H. B. Baker, 1927) . HELICODISCIDAE The origin of the genus Helicodiscus Morse, 1864 is Nearctic (Bequaert and Miller 1973). Chanomphalus Strebel 68 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 and Pfeffer, 1880 is in Mexico and Central America (Thiele 1935) and according to Pilsbry (1948), Radiodiscus Pilsbry and Lerriss, 1906 comes from South America where the genus is well represented by various species. The Helicodiscidae is found on the east coast of Mexico down to the southern part of the State of Veracruz. In the west it is found in the northern states of Baja California Sur, Sonora, and Chihuahua, and at the center of Mexico in the central States of Michoacan and the Pederal District (Mexico City). The Helicodiscidae contains six Mexican species and subspecies in three genera, Chanomphalus (1), Helicodiscus (2), and Radiodiscus (3). Chanomphalus pilsbryi (H. B. Baker, 1927) is found from the mid-southern part of the State of Tamaulipas (Correa- Sandoval 1999, 2002) to the eastern part of the State of San Luis Potosi (Correa- Sandoval 1998), northern part of the State of Puebla (Baker 1927, 1930), and from the north to south in the State of Veracruz (Baker 1927, 1930, Correa- Sandoval 2000, Naranjo-Garcia 2003). Helicodiscus eigenmanni Pilsbry, 1900 is from the northern part of the States of Chihuahua and Sonora (Pilsbry 1948) and also from the northern part of the State of Puebla (Baker 1930). Helicodiscus singleyanus (Pilsbry, 1889) is from Catalina Island and the south tip of the State of Baja California Sur (Smith et al. 1990). In the State of Sonora it is found in the north central (Naranjo-Garcia 1991) and the southern parts (Branson et al. 1964). Radiodiscus millecostatus costaricanus Pilsbry, 1926 is from the western part of the Pederal District (Mexico City) (Baker 1930) and the northern part of the State of Puebla (Baker 1927, 1930). Radiodiscus millecostatus millecostatus Pilsbry and Lerriss, 1906 is from the northern part of the State of Chihuahua, the central part of the State of Michoacan (Pilsbry 1948) and from the State of Tamaulipas (without given localities) (Bequaert and Miller 1973). Radiodiscus proameri Baker, 1930 is from the northern part of the State of Puebla (Baker 1930) and the west central part of the State of Veracruz (Baker 1928, 1930). CHAROPIDAE The Charopidae is of South American origin (Miquel et al. 2004). It is found in the center of Mexico, with one Mexican species with two subspecies. The subspecies Rotadiscus hermanni hermanni (Pfeiffer, 1866) inhabits the northern part of the State of Puebla, the center of the State of Veracruz, and to the south it inhabits the center and western part of the State of Michoacan (Pilsbry 1891, 1903, Diaz de Leon 1912, Baker 1927, 1930). Rotadiscus hermanni nivatus Baker, 1930 was described from Desierto de Los Leones, west of the Pederal District (Mexico City) (Baker 1928, 1930). DISCIDAE The genus Discus Pitzinger, 1833 is of Holarctic origin, considered by Pilsbry (1948) as wide spread; Bequaert and Miller (1973) propose this genus evolved earlier somewhere in Eurasia, since that region has more fossil records and more Recent species than in America. In Mexico Discidae contains two species in two genera. Discus whitneyi (Newcomb, 1864) is found in the northern part of the State of Chihuahua (Bequaert and Miller 1973). Gonyodiscus victorianus (Pilsbry, 1904) is in the mid- southern part of the State of Tamaulipas (Pilsbry 1903, Baker 1930, Hinkley 1907, Correa-Sandoval 2002, 2003), in the eastern part of the State of San Luis Potosi (Correa-Sandoval 1998, 2003), and in the northern part of the States of Puebla and Veracruz ( Baker 1 930, Correa-Sandoval 2000) . Thompson (2008: 693) considers Gonyodiscus victorianus (Pilsbry, 1904) to be a synonym of Radiocentrum victoriana (Pilsbry, 1904) of the family Oreohelicidae. OREOHELICIDAE The Oreohelicidae is found in Canada, the Rocky Mountains, and northern Mexico (Pilsbry 1939, Boss 1982). The family is distributed in the north and northwest side of Mexico in the States of Sonora, Chihuahua, and the Peninsula of Baja California. Seven species in two genera comprise the Oreohelicidae in Mexico: Oreohelix Pilsbry, 1904 (3) and Radiocentrum Pilsbry, 1905 (4). Known from a single locality are the species Oreohelix caenosa Pilsbry, 1948 and Oreohelix labrenana Pilsbry, 1948 which are in the northwestern part of the State of Chihuahua (Pilsbry 1948, Clench and Turner 1962). Thompson (2008: 693) considers that the correct epithet of O. labrenana Pilsbry, 1948 is Radiocentrum labrenana (Pilsbry, 1948). Prom the southern part of the same State (Cueva del Diablo) is the species Radiocentrum almoloya (Drake, 1949) (Drake 1949). Oreohelix concentrata (Dali, 1895) is from the northern part ofthe State of Sonora (San Jose Mountains) (Pilsbry 1939, Bequaert and Miller 1973), and Radiocentrum orientalis Metcalf, 1980 is from the northern part of the State of Coahuila (Metcalf 1980). Radiocentrum exorbitans (Miller, 1973) is from the middle part of the State of Baja California Sur (Smith et al. 1990) and from the southern end of the same State is the species Radiocentrum discus Christensen and Miller, 1976 (Smith et al. 1990). SUCCINEIDAE The Succineidae is of worldwide distribution (Thiele 1935, Pilsbry 1948, Boss 1982). In Mexico it is distributed in MEXICAN LAND SNAILS 69 the northwestern part of the State of Baja California, the central region of Mexico over the Trans-Mexican volcanic belt morphotectonic Province (Ferrusquia-Villafranca 1993) (Fig. 4), and from the southern part of the State ofTamaulipas to the north central part of the State of Veracruz. In Mexico it has twenty-four species and subspecies in three genera Catinella Pease, 1870 (1), Oxyloma Westerlund, 1885 (1), and Succinea Draparnaud, 1815 (22). Catinella rehderi (Pilsbry, 1948) is found in the northern, center, and the southern parts of the Peninsula of Baja California (Smith et al. 1990). Oxyloma nuttallianum (Lea, 1841) is from a single locality in the central part of the State of Baja California Sur. Most of its distribution range is located in the northwestern United States (Pilsbry 1948, Bequaert and Miller 1973). Succinea ampullacea Martens, 1898 is from the center of the State of Jalisco (Ameca) (Martens 1890-1901) and from the southern part of the State of Sonora (between Guaymas and Ciudad Obregon) (Branson et al. 1964). Succinea avara Say, 1824 is from the center coast of the State of Campeche (Pantel Aguada) (Bequaert and Clench 1936). Its main distribution occurs in Canada and the northeastern United States with the closest locations to Mexico being Florida and Texas. Pilsbry (1948) mentioned that it is found in northwestern Mexico without giving specific localities. Thompson (2008: 264) considers Succinea avara Say, 1824 to be a synonym of Catinella avara (Say, 1824). Recorded from a single locality are the following species: Succinea brevis Dunker in Pfeiffer, 1850 from the northwestern part of the State of Hidalgo (Zimapan) (Martens 1890-1901); Succinea californica Crosse and Fischer, 1878 from the north end of the State of Baja California (Smith etal. 1990); Succinea carmenensis Fischer and Crosse, 1878 from the State of Campeche (Carmen Island); Succinea colorata Fischer and Crosse, 1878 and Succinea guatemalensis Morelet, 1849 at the center of the State of Tabasco (Villahermosa) (Pilsbry 1900); Succinea campestris Say, 1817 from the eastern part of the State of Mexico (Lake Texcoco) (Martens 1890-1901). However, Succinea campestris is mostly found in the United States from Florida and North and South Carolina (Pilsbry 1948). Succinea guadalupensis Dali, 1900 is from the northwestern part of the State of Baja California (Guadalupe Island) (Smith etal. 1990). Succinea pueblensis Crosse and Fischer, 1878 is from the west center of the State of Puebla (City of Puebla) (Fischer and Crosse 1870-1902, Martens 1890-1901, Diaz de Leon 1912). Succinea socorroensis Dali, 1926 is from the Revillagigedo Archipelago (Socorro Island), State of Colima (Dali 1926). Succinea tlalpamensis tlalpamensis Pilsbry, 1899 is from the southern part of the Federal District (Mexico City, Tlalpan) (Pilsbry 1899). Succinea tlalpamensis cuitseana Pilsbry, 1899 is from the north northeastern part of the State of Michoacan (Lake Cuitseo near Huingo) (Pilsbry 1899, Clench and Turner 1962). Thompson (2008: 266) considers that the correct Figure 4. Distribution in the Trans-Mexican volcanic belt of various species of Succinea. 70 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 epithets of Succinea tlalpamensis tlalpamensis Pilsbry, 1899 and Succinea tlalpamensis cuitseana Pilsbry, 1899 are Oxyloma tlalpamensis tlalpamensis (Pilsbry, 1899) and Oxyloma tlalpamensis cuitseana (Pilsbry, 1899). Succinea undulata moerchi Dunker in Paetel, 1889 is from the northwestern part of the State of Hidalgo (Zimapan) (Martens 1890-1901). Succinea concordalis Gould, 1848 is from the northern part of the State of Veracruz (Tuxpan) (Martens 1890-1901, Diaz de Leon 1912, Correa-Sandoval 1999, 2003) and southern part of the State of San Luis Potosi (Huichihuayan) (Correa- Sandoval 1999). Succinea luteola luteola Gould, 1848 is from the northern part of the State of Sonora (Branson et al. 1964), in the northern part of the State of Chihuahua (Lake Palomas, Valle de los Mimbres) (Dali 1897, Diaz de Leon 1912, Bequaert and Miller 1973), in the southern part of the State of Tamauhpas (Correa-Sandoval 1999, 2002, 2003), and in the northern part of the State of Veracruz (Correa-Sandoval 2000). It is also found in the eastern part of the State of San Luis Potosi (Ciudad Valles) (Correa-Sandoval 1999, 2003), at the center of the State of Guerrero (Venta del Zopilote) (Martens 1890-1901, Correa-Sandoval 1999), and from the northwestern part of the State of Yucatan (Progreso) (Martens 1890-1901). Pilsbry (1948) considers the species widely spread in the State of Veracruz and in the southeastern part of the State of Puebla (Tehuacan). Succinea luteola sonorensis Fischer and Crosse, 1878 is found around the northeastern part of the State of Sonora (Bequaert and Miller 1973). Succinea panucoensis Pilsbry, 1909 is from the southeastern part of the State of San Luis Potosi (Pilsbry 1909, Clench and Turner 1962, Correa- Sandoval 1999, 2003), and from the southern part of the State of Tamaulipas (Pilsbry 1909, Clench and Turner 1962). Succinea rusticana Gould, 1846 is from the southern end of the State of Baja California Sur (Smith et al. 1990). Succinea solastra Hubricht, 1961 is from the center and northeastern part of the State of Nuevo Leon, and at the northwestern part of the State of Tamaulipas (Hubricht 1961, Correa-Sandoval 2003). Succinea undulata Say, 1829 is from the southwestern part of the State of Guanajuato (Irapuato near Lerma) at the center of the State of Jalisco (Sayula), at the south central coastal part of the State of Veracruz (Santecomapan) (Martens 1890-1901, Diaz de Leon 1912), and from the southwestern part of the State of Queretaro (near the City of Queretaro) (Pilsbry 1925). Succinea virgata Martens, 1865 is from the central part of the State of Veracruz (Martens 1890-1901, Pilsbry 1903, Diaz de Leon 1912) and the southeastern part of the State of Puebla (Martens 1890-1901, Diaz de Leon 1912). SAGDIDAE According to Boss (1982), the family Sagdidae ranges from Florida south to the Antilles and Venezuela. It is a family of the tropics and subtropics of America (Pilsbry 1940). The Sagdidae in Mexico is represented only by the species Xenodiscula taintori Goodrich and van der Schalie, 1937 from the southern part of the State of Veracruz at Los Tuxtlas Biological Station (Naranjo-Garcia 2003). It was originally described from Guatemala, at El Peten (Goodrich and van der Schalie 1937) and it has recently been collected from Nicaragua (Perez et al. 2008). As far as we know, the only other species of Xenodiscula in the Americas is in Venezuela (Pilsbry 1919a). GASTRODONTIDAE The Gastrodontidae is Holarctic (Boss 1982); in addition, there are some representatives of it in Bermuda and Madeira (Pilsbry 1946). In Mexico, the Gastrodontidae is mainly in the east in various states of the Gulf of Mexico, reaching the northern part of the State of Chiapas, and in western Chiapas from the northwestern corner to the north central portion. In Mexico Gastrodontidae is represented by five species in three genera: Pseudohyalina Morse, 1864 (1), Striatura Morse, 1864 (2) and Zonitoides Lehmann, 1862 (2). Pseudohyalina cidariscus Martens, 1892 is from the northern part of the State of Chiapas (Palenque) (Martens 1890-1901). Thompson (2008: 681) considers that this taxon belongs to the family Charopidae as Choanompalus cidarisca (Martens, 1892). Striatura ( Striatura ) pugetensis (Dali, 1895) is from Guadalupe Island, State of Baja California (Pilsbry 1946, Smith et al. 1990). Striatura ( Striatura ) meridionalis (Pilsbry and Ferris, 1906) is from south central Tamaulipas (Correa-Sandoval 2002), from near the center of the State ofNuevo Leon (Pablillo) (Pilsbry 1946, Contreras-Arquieta 1995 — without giving precise localities), from the northern part of the State of Puebla (Necaxa) (Baker 1930), and from the center of the State of Veracruz (Orizaba) (Pilsbry 1946). Zonitoides arboreus (Say, 1816) is from the State of Chihuahua (Bequaert and Miller 1973), from the central western part of the State of Nuevo Leon (Pilsbry 1903, Contreras-Arquieta 1995 — no localities given), from the southeastern part of the State of San Luis Potosi (Correa- Sandoval 1997), northern part of the State of Puebla (Baker 1930), and from the center of the State of Veracruz (Martens 1890-1901, Pilsbry 1903, 1946, Baker 1930). Dali (1926) recorded Zonitoides socorroensis Dali, 1926 from the Revillagigedo Archipelago (Socorro Island), State of Colima. Thompson (2008: 721) places Zonitoides socorroensis Dali, 1926 under Nesovitrea subhyalina socorroensis (Dali, 1926) of the family Zonitidae. MEXICAN LAND SNAILS 71 EUCONULIDAE The genus Euconulus Reinhardt, 1883 is Holarctic with the genera Guppya Morch, 1867 and Habroconus Fischer and Crosse, 1872 occurring in tropical and subtropical Americas. They are found from latitude 25°38’N south to eastern Mexico (Pilsbry 1939). The Euconulidae is distributed mainly on Mexico’s east coast, with some western records in the States of Baja California, Sonora, Jalisco, and Michoacan. The Euconulidae has seventeen Mexican species in three genera: Euconulus (1), Guppya (12), and Habroconus (4). Euconulus fulvus (Muller, 1774) is on the northwestern side of Mexico in the northern part of the State of Baja California (Smith etal. 1990), in the southern part of the State of Sonora (Naranjo-Garda 1991), and in the northwestern part of the State of Chihuahua (Bequaert and Miller 1973). There is one record of the species, as a fossil from the Pleistocene, in the northern part of the State of San Luis Potosi (Oliver a- Carrasco 2007). Guppya biolleyi Martens, 1892 is from the northern part of the State of Puebla (Baker 1930) and the south coastal part of the State of Veracruz (Naranjo-Garda 2003). The following species are known only from their type locality: Guppya capsula Dali, 1926 and Guppya montanicola Dali, 1926 are from Socorro Island of the Revillagigedo Archipelago, State of Colima (Dali 1926); Guppya jalisco Pilsbry, 1919 from the center of the State of Jalisco (Guadalajara) (Pilsbry 1919b), and Guppya sterkii punctum Baker, 1930 from the northern part of the State of Puebla (Necaxa) (Baker 1930). Guppya elegans (Strebel, 1880) is from the west central part of the State of Veracruz and at the northwestern part of the State of Morelos (Martens 1890-1901, Pilsbry 1919b). Guppya gundlachi gundlachi (Pfeiffer, 1840) is from the west central part of the State of Tamaulipas (Pilsbry 1903), the southeastern part of the State of San Luis Potosi (Correa- Sandoval 1997), at Carmen Island (Martens 1890-1901, Diaz de Leon 1912), along the central coast of the State of Campeche (Thompson 1967b), at Cozumel Island (Rehder 1966, Thompson 1967b), and the southwestern part of the State of Quintana Roo (Thompson 1967b). Guppya gundlachi orosciana Martens, 1892 is found in the west central part of the State ofVeracruz (Baker 1930). Guppya miamensis Pilsbry, 1903 is found in the south coastal part of the State ofVeracruz (Naranjo-Garda 2003). Guppya micra Pilsbry, 1903 is from the west central part of the State of Tamaulipas (Pilsbry 1903), the eastern part of the State of San Luis Potosi (Correa- Sandoval 1997), and from the central part of the State of Michoacan (Pilsbry 1903). Guppya perforata Dali, 1926 and Guppya socorroana Dali, 1926 are from the Tres Marias Islands, State of Nayarit; Guppya perforata from Maria Madre, both from Maria Magdalena (Dali 1926), and in addition Guppya socorroana is recorded from the Revillagigedo Archipelago (Socorro Island), State of Colima (Dali 1926). Habroconus pittieri (Martens, 1892) is from the southern tip of the State of Yucatan, is scattered throughout the State of Campeche, and occurs in the central and southwestern portions of the State of Quintana Roo (Thompson 1967b). Habroconus trochulinus (Morelet, 1851) is from the northern part of the State of Puebla (Baker 1930), and the west central part of the State of Veracruz (Martens 1890-1901, Pilsbry 1903, Baker 1930). Habroconus elegantulus (Pilsbry, 1919) is on both the Gulf of Mexico and Pacific coasts of Mexico. It is known from the west central part of the State of Nuevo Leon (Pilsbry 1919), from the southwestern part of the State of Tamaulipas (Pilsbry 1919b), the eastern part of the State of San Luis Potosi (Correa-Sandoval 1997), the northern part of the State of Puebla (Baker 1930), the west central part of the State ofVeracruz (Baker 1930), and from the center of the States of Jalisco and Michoacan (Pilsbry 1919b). Habroconus selenkai (Pfeiffer, 1866) is from the northern part of the State of Puebla and from the west central part of the State of Veracruz (Baker 1930). VITRINIDAE The Vitrinidae is Holoarctic (Boss 1982) and is distributed mainly in eastern Mexico with various records in the State of Sonora. In Mexico the Vitrinidae contain three species in two genera: Pycnogyra Strebel and Pfeffer, 1880 (1) and Hawaiia Gude, 1911 (2). Pycnogyra berendti (Pfeiffer, 1861) is found in the northern part of the State of Puebla (Baker 1930) and at the central part of the State ofVeracruz (Martens 1890-1901, Pilsbry 1903, Baker 1928, 1930). Pycnogyra is only known from Mexico (Thiele 1935). Hawaiia minuscula (Binney, 1840) is from the eastern (north and south) part of the State of Sonora (Branson et al. 1964, Naranjo-Garda 1991), from the central part of the State of Tamaulipas (Correa-Sandoval 1999, 2002, 2003), the eastern part of the State of San Luis Potosi (Correa-Sandoval 1997), the northern (Correa-Sandoval 2000), and southern parts (Naranjo -Garcia and Polaco 1997, Naranjo-Garda 2003) of the State ofVeracruz, from the western (Martens 1890-1901), the central coastal portions (Thompson 1967b) of the State of Campeche, and the northern part of the State of Chiapas (Palenque) (Bequaert 1957). In addition, Bequaert and Miller (1973) mentioned that Hawaiia minuscula has been recorded from the States of Baja California, Nayarit, Puebla, and Yucatan without giving exact localities. Reddell (1981) found it in caves from the States of Campeche, Veracruz, and Yucatan. Smith et al. (1990) recorded an undescribed Hawaiia from the State of Baja California and commented on the necessity of further 72 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 studies to determine its specific epithet. Hawaiia is a native Nearctic snail but had been widely dispersed by humans (Bequaert and Miller 1973). Thompson (2008: 735) places both genera Pygnogyra and Hawaiia in the family Zonitidae. ZONITIDAE The Zonitidae is of worldwide distribution (Thiele 1935, Burch 1962, Boss 1982). In Mexico, the distribution is mostly in the eastern part of the country from the center of the States of Nuevo Leon and Tamaulipas down to central Veracruz with a few localities in the States of Guerrero, Oaxaca, and Chiapas. In the northwest, there are various records from the State of Sonora. The family Zonitidae, in Mexico, has eighteen species and subspecies in four genera Glyphyalinia von Martens, 1892 (2), Mesomphix Rafinesque, 1819 (14), Retinella Fischer in Suttleworth, 1877 (1), and Zom'fes Montfort, 1810 (1). Glyphyalinia indentata indentata (Say, 1822) is from the east side of the State of Sonora (Naranjo-Garcia 1991); there are some records of the species in the eastern central part of the State of Nuevo Leon (Correa-Sandoval 1993, Correa- Sandoval and Salazar 2005), at the center of the State of Tamaulipas (Correa-Sandoval 2002), and at the eastern part of the State of San Luis Potosi (Correa-Sandoval 1998). In these last three states, the species is less frequent (Correa- Sandoval 2002). The subspecies Glyphyalinia indentata paucilirata (Morelet, 1851) is from the northern part of the State of Puebla and from the eastern part of the State of Mexico (Teotihuacan) (Baker 1930). Bequaert and Miller (1973) recorded it from the States of Baja California, Sonora, and Chihuahua without giving specific localities. Mesomphix zonites (Pfeiffer, 1845) is from the southern part ofthe State ofVeracruz (Tuxtla) (Martens 1890-1901) and from the west central part of the State of Chiapas (Martens 1890-1901, Bequaert 1957). Mesomphix lucubratus caducus (Pfeiffer, 1846) and Mesomphix lucubratus lucubratus (Say, 1 829) are from the central part of the State ofVeracruz (Martens 1890-1901, Baker 1930). Mesomphix lucubratus strebelianus (Martens, 1892) is also found around the same region plus at the northern part of the State of Puebla (Necaxa) (Martens 1890-1901, Baker 1930 ). Mesomphix montereyensismontereyensis Pilsbry, 1899 is from the west central part of the State of Nuevo Leon (Diente) (Pilsbry 1899, 1903) and in the west central part ofthe State of Tamaulipas (Correa-Sandoval 2002). Mesomphix montereyensis victorianus (Pilsbry, 1903) is from the west central part of the State of Tamaulipas (Correa- Sandoval 1999, 2002) and from the eastern part of the State of San Luis Potosi (Correa-Sandoval 1997). Thompson (2008: 733) considers that Mesomphix montereyensis montereyensis Pilsbry, 1899 and Mesomphix montereyensis victorianus (Pilsbry, 1903) are Patulopsis montereyensis montereyensis (Pilsbry, 1899) and Patulopsis montereyensis victorianus (Pilsbry, 1903), respectively. Mesomphix paradensis (Pfeiffer, 1860) is from the central part of the States ofVeracruz and Oaxaca (Martens 1890-1901). Various species are recorded from a single locality: at the center of the State of Veracruz are Mesomphix salleanus Martens, 1892 (Cordoba) (Martens 1890-1901) and Mesomphix tuxtlensis Crosse and Fischer, 1870 (Tuxtla [sic]) (Baker 1930). Mesomphix sculptus (Martens, 1892) is from the center of the State of Guerrero (Omilteme) (Martens 1890-1901), and Zonites tehuantepecensis Crosse and Fischer, 1870 is from the coastal part of the State of Oaxaca (Tehuantepec) (Martens 1890-1901). Mesomphix veracruzensis (Pfeiffer, 1856) and Mesomphix modestus Martens, 1892 are found at various localities at the center of the State ofVeracruz (Martens 1890-1901, Baker 1930). Mesomphix carinatus (Strebel and Pfeffer, 1880) and Retinella subhyalina subhyalina (Pfeiffer, 1867) are from the central part of the State of Veracruz and from the northern part of the State of Puebla (around Necaxa) (Martens 1890-1901, Baker 1930). Mesomphix bilineatus (Pfeiffer, 1845) is from the west central part of the State ofVeracruz (Cordoba) (Martens 1890-1901, Baker 1930) and in the northeastern part of the State of Chiapas (Laguna Ocotal) (Bequaert 1957). From the preceding paragraph, the species that change considering Thompson’s (2008: 734, 720, 732, 717) ideas are: Mesomphix salleanus Martens, 1892 to Patulopsis salleanus (Martens, 1892), Mesomphix veracruzensis (Pfeiffer, 1856) to Patulopsis veracruzensis veracruzensis (Pfeiffer, 1856), Mesomphix carinatus (Strebel and Pfeffer, 1880) to Patulopsis carinatus Strebel and Pfeffer, 1880, Retinella subhyalina subhyalina (Pfeiffer, 1867) to Nesovitrea subhyalina subhyalina (Pfeiffer, 1867), and Zonites tehuantepecensis Crosse and Fischer, 1870 to Zonitoides tehuantepecensis (Crosse and Fischer, 1870) of the family Gastrodontidae. THYSANOPHORIDAE The family Thysanophoridae is distributed from the southern United States to northern South America. It is well represented in Mexico and the Antilles (Pilsbry 1940, Boss 1982). In Mexico, Thysanophoridae is patchily spread (Figs. 5, 6). There are various records from the eastern part of the State of Baja California Sur, over the entire State of Sonora, and very few records in other western states. On the eastern side, the family is in the mid-southern part of the State of Tamaulipas, over the entire State ofVeracruz, in the eastern part of the State of San Luis Potosi, the northern part of the State of Puebla, at the northeastern part of the State of Chiapas, and the northwestern part of the state of Yucatan. Over the Trans-Mexican volcanic MEXICAN LAND SNAILS 73 Figure 5. Distribution of various species of Microconus and Thysanophora in Mexico. Figure 6. Distribution of various species of Thysanophora in Mexico. 74 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 belt few records occur. The family in Mexico has fifteen species in two genera: Microconus Strebel and Pfeffer, 1880 (2) and Thysanophora Strebel and Pfeffer, 1880 (13). Microconus wilhelmi (Pfeiffer, 1866) is from the northern part of the State of Puebla (Necaxa) and in the center of the State of Veracruz (Mirador) (Baker 1927, Thompson 1958). There is an undescribed species of Microconus recorded by Correa-Sandoval (1999) from the southern part of the State of San Luis Potosi (Xilitla). Thysanophora caecoides (Tate, 1870) is from the center of the State of Veracruz (Baker 1927) and at the northwestern part of the State of Yucatan (Progreso) (Pilsbry 1891, 1920). Thysanophora clarionensis Dali, 1926 is from Isla Clarion and various other localities of the same island, Revillagigedo Archipelago, State of Colima (Dali 1926). Thysanophora conspurcatella (Morelet, 1951) is from the center of the State of Veracruz (Antigua) (Pilsbry 1903) and west central part of the State of Yucatan (Martens 1890-1900). Thysanophora fuscula (C. B. Adams, 1849) is from the mid-southern part of the State of Tamaulipas (Pilsbry 1903, Correa-Sandoval 1999, 2002), northern part of the State of Veracruz (Correa-Sandoval 2000), the eastern part of the State of San Luis Potosi (Correa- Sandoval 1997), and the northeastern part of the State of Chiapas (Laguna Ocotal and Monte Libano) (Bequaert 1957). Thysanophora hornii (Gabb, 1866) is from the eastern part of the State of Baja California Sur (Smith et al. 1990), most of the State of Sonora (Naranjo-Garda 1991), from the southern half of the State of Tamaulipas (Pilsbry 1903, Correa-Sandoval 2002), from the mid-eastern part of the State of San Luis Potosi (Correa-Sandoval 1997), from the central coast of the State of Jalisco (Bequaert and Miller 1973, Naranjo-Garda, unpubl. data), from the mid-northern part of the State of Veracruz (Correa-Sandoval 2000), and from the States of Sinaloa, Chihuahua and Nuevo Leon (Bequaert and Miller 1973 — without localities given). Bequaert and Miller (1973) consider the genus native of America and Thysanophora hornii as an invader from the south. Metcalf and Smartt (1997) also believe that this species has a neotropical origin. Thysanophora impura (Pfeiffer, 1866) is from the eastern part of the State of Morelos (Yautepec) (Pilsbry 1891), the center of the State of Veracruz (Pilsbry 1891, 1926, Baker 1927), from the west central region of Yucatan (Pilsbry 1926), and from the northwestern part of the State of Chiapas (Ocosingo) (Bequaert 1957). Species from a single locality are: Thysanophora intonsa (Pilsbry, 1891) from the central part of the State of Veracruz (Orizaba) (Pilsbry 1891), Thysanophora mazatlanica (Pfeiffer, 1856) from the south coastal part of the State of Sinaloa (Mazatlan) (Martens 1890-1901), and Thysanophora minuta Baker, 1927 from northern part of the State of Puebla (near Necaxa) (Baker 1927). Thompson (2008: 678) places Thysanophora mazatlanica (Pfeiffer, 1856) under Punctum mazatlanica (Pfeiffer, 1856) of the family Punctidae. Thysanophora materna Dali, 1926 is from the Tres Marias Islands (Maria Madre and Maria Madgalena) of the State of Nayarit (Dali 1926). Thysanophora paleosa (Strebel and Pfeiffer, 1880) is from the northern part of the State of Puebla (Necaxa) and from the center of the State of Veracruz (Baker 1927). Thysanophora proximo Pilsbry, 1899 was found by Branson et al. (1964) in the northern part of the State of Sonora and it is also found in the northern and central portions of the State of Michoacan (Pilsbry 1903). Thysanophora plagioptycha (Shuttleworth, 1854) is from the northern part of the State of Puebla (Necaxa) (Baker 1927) and the central and southern portions of the State of Veracruz (Pilsbry 1940, Naranjo-Garda 2003). HELMINTHOGLYPTIDAE The Helminthoglyptidae is an American family found in the western United States (Boss 1982) and northwestern Mexico. The family Helminthoglyptidae had about 60 Mexican species before Schileyko (1991) revised the superfamily Helicoidea. He suggested the family Helminthoglyptidae be confined to the genera Helminthoglypta Ancey, 1887 and Eremarionta Pilsbry, 1913. The other genera, such as Sonorella Pilsbry, 1900, Sonorelix Berry, 1943, and Greggelix Miller, 1972 (formerly part of the Helminthoglyptidae) he placed in the Xanthonychidae. The Mexican Helminthoglyptidae now contains ten species and subspecies in two genera Eremarionta (3) and Helminthoglypta (7). The Helminthoglyptidae distributed in the State of Baja California are: Eremarionta indioensis (Yates, 1890), Eremarionta rowelli bechteli (Emerson and Jacobson, 1964), and Helminthoglypta tudiculata (Binney, 1843), also found at the northern part of the State of Baja California relatively close to the border with the United States are the species Helminthoglypta coelata (Bartsch, 1916), Helminthoglypta misiona Chace, 1937, and Helminthoglypta reederi Miller, 1981 (Sierra San Pedro Martir) (Smith et al. 1990). From the islands of the same state, two subspecies were recognized. Helminthoglypta hannai hannai Pilsbry, 1927 and Helminthoglypta hannai diodon Pilsbry, 1927 inhabit Guadalupe Island (Pilsbry 1927, Clench and Turner 1962), and Helminthogypta traskii coronadoensis (Bartsch, 1916) is found in the Coronado Island (Bartsch 1916). The species Eremarionta rowelli rowelli (Newcomb, 1865) is the only other member of the family that is found in the northwestern coast of the State of Sonora (Drake 1957, Bequaert and Miller 1973). MEXICAN LAND SNAILS 75 ARIONIDAE The family Arionidae has a Holarctic origin (Pilsbry 1948); however, they may have spread to the south since members of the family are present in Africa (Burch 1962, Boss 1982). In the United States several genera are mainly distributed in the west (Pilsbry 1948). There is one Mexican species in the Arionidae, Binneya guadalupensis Pilsbry, 1927 from Guadalupe Island, State of Baja California (Pilsbry 1927, Smith et al. 1990). Thompson (2008: 739) recognizes the family Binneyidae Cockerell, 1891 in the superfamily Arionoidea instead of the family Arionidae. PHILOMYCIDAE The family Philomycidae is distributed in parts of Canada, eastern United States, tropical Central America and South America and parts of Asia (Pilsbry 1948, Burch 1962, Boss 1982). The family is patchily distributed in Mexico: in the northwestern part of the State of Sonora, in the center (Mexico City), in the northeastern region of the State of Puebla, and at the center of the State of Veracruz. In Mexico, it is composed of three subspecies in the genus Pallifera Morse, 1864. Pallifera arizonensis arizonensis (Pilsbry, 1917) is found in the northwestern part of the State of Sonora (Sierra Purica) (Bequaert and Miller 1973). Pallifera costaricensis alticola Baker, 1930 is found in the western part of the Lederal District (Mexico City) (Desierto de los Leones) (Baker 1930). Pallifera costaricensis crosseana (Strebel and Pfeffer, 1880) is from the northern portion of the State of Puebla to the west central region of the State of Veracruz (Necaxa to Cordoba) (Baker 1930). SUMMARY OF DISTRIBUTION RECORDS The terrestrial molluscs reported from Mexico consist of 1,178 species and subspecies in 42 families. The 236 species in 34 medium and small families represent 81% of a total of 42 families found there. The diversity of the lesser families ranges from one species in Cochlicopidae, Megomphicidae, Arionidae, and Sagdidae to 23 species in Cyclophoridae and 24 in Succineidae. General patterns of the distribution of the lesser families of terrestrial molluscs in Mexico are shown (Table 2). To the north-western side of the country (Fig. 3) are the families Megomphicidae, Arionidae, Haplotrematidae (in part, 1 record intheeast),Vallonidae,Helminthoglyptidae,andOreohelicidae. The Cochlicopidae is localized in the north-center (Fig. 3) of Mexico; the Discidae is concentrated in the north-east center. The Carychiidae, Ceresidae (in part, 2 species in the west), Veronicellidae (in part, 1 record in the west, and 1 record in the south), Diplommatinidae, Systrophiidae, Sagdidae are observed mainly in the eastern side (Fig. 2). The Charopidae are established in central Mexico. The Philomycidae are patchily distributed but fairly widespread. The families Annulariidae, Amphibulimidae, and Megalomastomidae are localized towards the southern side (Fig. 1). The Achatinellidae are Table 2. General geographic distribution of some taxa occurring South Palearctic Holarctic American in Mexico. Nearctic (North America) Worldwide distribution Other Strobilops possibly its origin is in Asia Discus Charopidae Haplotrematidae Strobilopsidae Annulariidae Antilles, Central America, South America Carychiidae Punctum Systrophiidae Helminthoglyptidae Zonitidae Cyclophoridae same data as above plus other places Cochlicopidae Hawaiia Veronicellidae Oreohelicidae Philomycidae Achatinellidae Pacific Islands Arionidae Euconulidae Vallonidae Ceresidae Megomphicidae Middle America Thysanophoridae Truncatellidae Diplommatinidae Ellobiidae Ferussaciidae Succineidae Megalomastomidae American 76 AMERICAN M ALACOLOGICAL BULLETIN 28-1/2-2010 representative of the Pacific islands. The families Ellobidae and Truncatellidae are dwellers of the intertidal zone of both coasts. In addition, the families Cyclophoridae, Helicodiscidae, Punctidae (more toward the north), Thysanophoridae, Succi- neidae, Euconulidae, Ferussaciidae, Subulinidae, Vitrinidae ( Hawaiia ), Gastrodontidae, Strobilopsidae, and Zonitidae are of wide distribution in the country (Figs. 4-6). The distributions of the diverse genera appear to be related to the most humid, sub-humid, and semiarid areas of Mexico. The Diplommatinidae, Ceresidae, Vitrinidae, Eu- conulidae, Helicodiscidae, Succineidae, and Thysanophoridae occur in tropical and subtropical environments, while the Annulariidae, Megalomastomidae, and Amphibulimidae are mostly tropical. The Veronicellidae (Naranjo-Garcia et al. 2007), Charopidae (Miquel et al. 2004), Amphibulimidae (Thiele 1 93 5 ), along with the Systrophiidae are much better represented in South America. Typical American continent families are Megalomastomidae, Sagdidae, and the genus Thysanophora. In Table 3, interestingly, the number of species Mexico shares with the United States (51) and with Central America (54) is about the same, and the taxa shared among the three areas include 14 species. Thus, it would appear that 146 spe- cies are endemic to Mexico and 90 are also found elsewhere. It must be kept in mind, however, that many areas of Mexico and adjacent regions are under-collected. CONCLUSIONS Distribution patterns are suggestive but are somewhat preliminary because extensive areas of Mexico have not been studied for land molluscs. Under-collected areas are the States of Nayarit, Sinaloa, Tlaxcala, Guanajuato, Durango, Coahuila, Aguascalientes, Guerrero, Zacatecas, and parts of the State of Chihuahua. Regarding micromolluscs, Solem’s (1976: 2) statement continues to be valid around the world: “The species are very small (96.4 % are <7 mm in maximum size), secretive inhabitants of litter or may be found on moss-covered tree trunks in dense and undisturbed forests. They are found only by the most skilled collectors”. Some species have limited data information, others are known from very few records, and some from only a single locality. The apparently eastern distribution of most families is a consequence of the extensive collecting by explorers of the late nineteenth and early twentieth centuries whose travels were on the east side of the country (Naranjo-Garcia and Polaco 1997). Studies of the distribution of native species, including terrestrial molluscs, are of utmost importance because of the Table 3. Mexican (MX) families with number of species shared with the United States (USA) and Central America (CA). United Central Shared Family States America USA/MX/CA Amphibulimidae 1 Annulariidae 1 Carychiidae 2 1 1 Charopidae 1 Cochlicopidae 1 Cyclophoridae 4 Diplommatinidae 1 Discidae 1 Ellobiidae 3 4 Euconulidae 3 6 1 Ferussaciidae 2 Gastrodontidae 3 2 2 Haplotrematidae 2 Helicodiscidae 3 3 1 Helminthoglyptidae 4 Megomphicidae 1 Megalomastomidae 3 Oreohelicidae 1 Philomycidae 1 Punctidae 4 Sagdidae 1 Strobilopsidae 2 Subulinidae 3 8 4 Succineidae 8 1 Systrophiidae 3 Thysanophoridae 2 7 2 Truncatellidae 1 3 Vallonidae 2 Veronicellidae 1 2 1 Vitrinidae 1 1 1 Zonitidae 2 1 1 Total species and 51 54 14 subspecies many threats to their existence. Habitat is destroyed by urban development, land and stream pollution, and cutting of native forests. Many type localities have disappeared due to habitat destruction. Habitat changes in both developed and underexplored areas may lead to the extinction of entire species. The introduction of non-native species presents the potential for competition with native species, which can and does lead to their extinction. Without adequate records, it is impossible to evaluate the magnitude and effect of such changes. The future of biological life will be controlled by governmental decisions. The study of terrestrial molluscs and other living things is essential if we are to influence those governmental decisions. MEXICAN LAND SNAILS 77 ACKNOWLEDGMENTS This long term project received financial support from UNIBIO [Unidad de Bioinformatica para la Biodiversidad of Instituto de Biologia, UNIBIO is a unit of the larger project SIBA (Sistema de Informatica sobre la Biodiversidad y el Ambiente) of the Universidad Nacional Autonoma de Mexico] , from 2005 to 2007 (E. N.-G.). 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Some Mexican and Central American land snails of the family Cyclophoridae. Zoologica, New York Zoo- logical Society 54: 35-77. Thompson, F. G. 1980. Proserpinoid land snails and their rela- tionships within the Archaeogastropoda. Malacologia 20: 1-33. Thompson, F. G. 1987. A new prosobranch land snail from Eastern Mexico (Archaeogastropoda: Ceresidae). Archiv fur Mollusken- kunde 117: 159-162. Thompson, F. G. 2008. An Annotated Checklist and Bibliography of the Land and Freshwater Snails of Mexico and Central America. On-line publication, http://www.flmnh.ufI.edu/malacology/ mexico-central_america_snail_checklist/; accessed 9 December 2009. Tryon, G. W. 1863. Description of two new species of Mexican land shells. Proceedings of the Academy of Natural Sciences of Phila- delphia 6: 281. Vaught, K. C. 1989. A Classification of the Living Mollusca. American Malacologists Inc., Melbourne, Florida. Watters, G. T. 2006. The Caribbean Land Snail Family Annulariidae: A Revision of the Higher Taxa and a Catalog of the Species. Back- huys Publishers, Leiden, Netherlands. Submitted: 10 January 2009; accepted: 29 June 2009; final revisions received: 10 December 2009 Amer. Malac. Bull. 28: 81-90 (2010) Diversity and conservation of the land snail fauna of the western Pacific islands of Belau (Republic of Palau, Oceania)* Rebecca J. Rundell1,2,t 1 Committee on Evolutionary Biology, University of Chicago, 1025 East 57th Street, Chicago, Illinois 60637, U.S.A. 2 Division of Invertebrates, Department of Zoology, Field Museum of Natural History, 1400 South Lake Shore Drive, Chicago, Illinois 60605, U.S.A. Corresponding author: rrundell@interchange.ubc.ca Abstract: Pacific land snails are among the most threatened animals on Earth, and basic information on the number of extant species is lacking for many island groups. The isolated western Pacific archipelago of Belau comprises 586 islands, most of which have suitable land snail habitat; yet little has been published on the land snail fauna. I undertook surveys throughout Belau, searching trees and emergent vegetation, leaf litter, and limestone rock. Survey results from selected, geographically representative islands are presented here. The total number of species found in these areas (117) indicates that there may be ca. 200 extant Belau land snail species. This number far exceeds previous estimates. Most species are endemic to Belau (95% in this survey), and species endemic to one or a few islands are not uncommon. Leaf litter and rock dwelling diplommatinid land snails are a large component of the snail biota: only 26 Belau diplommatinids have been described, and 8 1 species were collected in this survey. Although caenogastropod land snails comprise the most obvious portion of the fauna, notable pulmonates include the partulids and endodontoids, two land snail groups that have suffered extinction throughout the Pacific region. Belau has one of the most spectacularly diverse extant land snail faunas in the Pacific region, and the restricted ranges of many species highlight the need for conservation attention, particularly on the island of Babeldaob, which is undergoing increased deforestation. Key words: biodiversity, distribution, management, biogeography, Micronesia Pacific oceanic islands have drawn European explorers’ vessels for hundreds of years, including the Beagle and the ships of Captain Cook although many islands remained vir- tually unknown to the rest of the world until a little over a century ago. Major inroads have been made in understanding diversification patterns in some of the larger island groups, such as the Hawaiian islands (Wagner and Funk 1995), Galapagos (Grant 1986), and French Polynesia (Clarke and Murray 1969), yet the fact that we are still uncovering many undescribed species (particularly of lesser-studied taxa) in these areas suggests that not only does much work remain in understanding the evolution of species on these islands, but there is a seemingly endless need for natural history informa- tion on species, including species distributions. The need for species surveys and natural history data for all Pacific island species is particularly urgent because of the high extinction rate for Pacific island biotas (Quammen 1996). The high levels of species richness found among oceanic island land snails was noted by Darwin (1859), and since then, Pacific island land snail extinction has accelerated to the point where almost 90% of the 763 described Hawaiian land snail species are now extinct (Cowie 2001a), and the snail fau- nas of other archipelagos have suffered a similar fate {e.g., Samoa [Cowie 2001b, Cowie and Cook 2001]; French Polynesia [Cowie 1992]). Habitat destruction, introduction of rats and mice, introduction of snail predators, invasive species, and in a few cases, over-collecting, have all contrib- uted to the disaster that is land snail extinction in the Pacific (Hadfield 1986, Hadfield et al. 1993, Lydeard etal. 2004). Early land snail specialists such as C. Montague Cooke and Yoshio Kondo of the Bishop Museum (Honolulu, Hawai‘i) were likely aware of these threats and fortunately col- lected some of the few remaining examples of many land snail species whose populations were later decimated. These collec- tions, in addition to those of Alan Solem (Field Museum of Natural History, Chicago, Illinois) are an invaluable resource for future research; however, we still frequently lack basic information such as the number of recorded or extant endemic * From the “Leslie Hubricht Memorial Symposium on Terrestrial Gastropods” presented at the meeting of the American Malacological Society, held from 29 July to 3 August 2008 in Carbondale, Illinois. t Present address: Department of Botany, University of British Columbia, #3529-6270 University Blvd., Vancouver, British Columbia V6T 1Z4, Canada. 81 82 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 species currently on an island. Such information is necessary for developing and implementing conservation management strategies (e.g., alien species prevention and eradication efforts) that are sorely needed on many islands (Lydeard et al. 2004). Given the rate at which extinction is occurring on Pacific islands (Solem 1990, Cowie 1992, Lydeard etal. 2004), we do not have the luxury of time in documenting faunas at such great risk of complete decimation. The land snail fauna of the western Pacific islands of Belau (Fig. 1, and described in Materials and Methods) is not A. Belau T Caroline Islands V jjU^' V BABELDAOB well documented, and there is urgent need for baseline data on species distributions. Smith (1993) lists 69 described spe- cies (including both indigenous/endemic and introduced species). Cowie et al. (1996) suggest that there may be 40-50 indigenous/endemic species in Belau, based on a survey of the literature and a scan of the Bishop Museum (Honolulu, Hawai‘i) collections. I present the first results of an intensive survey effort on the islands of Belau, which began in 2003. Documentation of the rock and leaf litter dwelling diplommatinid land snails was the priority of this work (Rundell 2008), and this paper represents the first of sev- eral detailing surveys of Belau’s diverse land snail fauna. Many species are unde- , scribed and some, particularly those <5 ..' i mm, are cryptic. Results indicate that endemism is high (at least 95% for the present survey of 26 areas), and many species, particularly the rock and leaf lit- ter dwelling diplommatinids, are single or few-island endemics. Based on my surveys, Belau land snail species richness has been seriously underestimated, par- ticularly for the tiny diplommatinids. Few invasive species were collected, a rarity for Pacific islands, and representa- tives of two of the most endangered land snail groups in the Pacific, namely the endodontoids and the partulids, were found. Belau’s land snail fauna ranks among the most diverse in the Pacific relative to island area and is worthy of conservation protection. MATERIALS AND METHODS Figure 1. A, (Inset) Western Pacific region, showing the location of Belau. The Belau archipel- ago is centered at 7°20'N and 134°E. B, The islands of Belau (Republic of Palau, Oceania). The archipelago is 160 km in length. Island names follow Motteler (2006). Numbers represent the following islands and localities: (1) island ofNgcheangel, (2) island ofNgerechur, (3) Ngerch- elong State, island of Babeldaob, (4) near waterfall, Ngatpang State, island of Babeldaob (5) near Ngardok Lake, island of Babeldaob, (6) Oikull and Ngerduais, Airai State, island of Ba- beldaob, (7) island of Ngerekebesang, (8) Ngermid, island of Oreor, (9) island of Malakal, ( 10) island of Ulebsechel, (11) an unnamed northern Rock island, Koror State, (12) an unnamed Rock island, south of 1 1, Koror State, (13) island of Ngeruktabel, ( 14) Medukriikuul, island of Ngeruktabel, (15) island of Ngchus, (16) island of Ulong, (17) island of Eudelchol, (18) near Jellyfish Lake, island of Mecherchar, (19) western Mecherchar, (20) island of Omekang, (21) near island of Kmekumer, (22) one island of Ngemelis, (23) large island of Ngemelis, (24) Techakl, island of Beliliou, (25) island of Beliliou, and (26) island of Ngeaur. Survey area The islands of Belau (comprising the independent Republic of Palau; Fig. 1), spanning 160 km, are centered at 7°20'N and 134°E in the western Pacific region known as Micronesia. Belau is a crest of an arc ridge (Kobayashi 2004) and has never been in contact with a continental landmass. The closest large islands to Belau are: Mindanao in the Philippines, the Moluccas and New Guinea (800 km), and Borneo (1500 km) (Crombie and Pregill 1999). The oldest emergent rocks in Belau are ca. 30 Ma, and its 586 islands (415 km2 BELAU LAND SNAIL DIVERSITY 83 total land area) are composed of volcanic rock and limestone. There are several different island types: volcanic (highest ele- vation is 242 m), high limestone, low limestone, reef or atoll, and a combination of volcanic rock and limestone (Crombie and Pregill 1999). Although most of the land area (333 km2) is the volcanic island of Babeldaob, the Miocene (23 to 5 Ma) to Pleistocene limestone islands to the south, many of which are <1 km2, also harbor great diversity. The karst mushroom- shaped Rock Islands in particular (Kelletat 1991) were sus- pected to harbor many calciphilic species; land snail species richness has been shown to increase with increasing pH and calcium (Emberton etal. 1997), and similar tropical limestone rock outcrops are known to harbor diverse land snail faunas (Vermeulen 1993, 1994, Schilthuizen etal. 2005, 2006). The Belau islands are humid tropical (350-400 cm annual precipitation; rainfall can be heavy year-round), and forested with mostly native species, even at low elevations (Crombie and Pregill 1999). The island of Babeldoab has the largest tract of pristine rainforest in Micronesia. Field methods and specimens Land snail collections occurred in 2003, 2005, and 2007. I selected 26 localities throughout Belau to represent a diver- sity of island and habitat types as well as geographical spread, and collected land snail specimens using timed searches. Island localities are numbered and shown in Fig. 1 . 1 under- took separate searches in the trees/emergent vegetation, leaf litter, and rocks (both live and dead shells were collected in the different habitats). Where there were no limestone out- crops ( e.g ., on volcanic substrates), searches included only the first two habitats. Snail sampling involved thorough search of each habitat, locating snails by eye, and using forceps or hands to place snails in separate vials, each labeled according to habitat. Sorting several collections of bulk leaf litter in 2003 and 2005 revealed that this procedure could be abandoned in favor of the field-based hand capture method since bulk leaf litter col- lections did not result in additional species diversity. I killed snails in 95% ethanol on the day of capture and replaced ethanol in each vial two times post-killing to ensure proper preservation. Following shipment to the Field Museum of Natural History (FMNH), I sorted adult specimens to species using a dissecting microscope, shell characters {e.g., shape, pres- ence/absence of keel, spines, number of ribs), species descrip- tions (and additional species accounts, i.e., Kobelt 1902, Wagner 1905, Thiele 1927, Cooke and Kondo 1960, Thompson and Huck 1985), and comparative and type material from the Bishop Museum (BPBM) and FMNH. For Diplommatinidae, species identifications based on shell morphological differences were corroborated by molecular data (Rundell 2008). Undescribed diplommatinid species were listed by letter code, reflecting the order in which they were collected. To impose order on the enormous diversity of diplommatinids, each undescribed spe- cies was assigned a temporary genus name, reflecting its habitat and morphology. Palaina Crosse, 1866 species are generally leaf litter dwelling, ovate conical, brown, and with different rib pat- terns. Diplommatina Benson, 1849 species are rock dwelling, pointed, heavily calcified, and whitish or yellowish. Hungerfordia Beddome, 1889 species are rock dwelling with dramatic shell spines or lamellae and are whitish and heavily calcified. I depos- ited land snail specimens at the FMNH, which holds one of the world’s largest collections of Pacific island land snails. RESULTS Faunal composition Described species in the current survey are listed in Table 1. Species localities are listed by island and locality (as indicated by number: see Fig. 1 ) in Table 2. Table 2 is most representative of species presence on an island, rather than absence (particu- larly for some widespread species); in other words, some com- mon species such as the assimineid Omphalotropis cheynei Dohrn and Semper, 1862 and the helicinid Palaeohelicina het- erochroa A. J. Wagner, 1906 occur on most islands and I expect, as additional data come to light, species matrices will include additional taxa. A large number of species have been recorded in the current study: i.e., the island of Beliliou has 36 species and the Medukriikuul site on the island of Ngeruktabel has 20 species, 18 of which are diplommatinids (Tables 1-2). The total number of species documented in this survey was 117. These species represent 14 families (Tables 1-2). Approximately 95% of the species found were Belau endemics, and many species occur on only one or a few islands (Table 2). The amount of undescribed diversity in Belau is great. Out of the 117 total species found in this survey, only 36 spe- cies are described. The families with the most undescribed species were: Diplommatinidae (only 10 of the 81 species recorded here were described), Hydrocenidae (none of the four species recorded were described), and Helicarionidae (two of the 10 species recorded were undescribed). Some species are relatively widespread, while others are endemic to single islands. The Diplommatinidae in particular are composed of some species that occur on only a single island (e.g., the leaf litter dweller Palaina AU), and others that occur on multiple islands {e.g., the rock dweller Hungerfordia A; Table 2). DISCUSSION The total number of species recorded in this survey (117) exceeds previous estimates by Smith (1993), who listed 69 described species (including both native and introduced 84 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Table 1. Described species included in survey, authors, and habitat (Ground and Gr. = leaf litter, Veg. = trees and emergent vegetation, Rock = limestone rock). Some endodontoids occur under rocks as well as in leaf litter. Under the category of “status,” endemic refers to species that are native to only the Belau islands, indigenous refers to species that are native to Belau, but also found elsewhere, and introduced refers to all species that are non-indigenous to Belau, having been introduced either accidentally or deliberately. Species marked with an asterisk are listed in the 1994 Red List of Threatened Animals (Groombridge 1993, IUCN 2004). Family Species Habitat Status Caenogastropoda ASSIMINEIDAE Kubaryia pilikia Clench, 1948 Ground Endemic Omphalotropis cheynei (Dohrn and Semper, 1862) Ground/ Veg. Endemic DIPLOMMATINIDAE Diplommatina lutea Beddome, 1889* Rock Endemic Palaina albata (Beddome, 1889)* Ground Endemic Palaina dimorpha (Crosse, 1866)* Ground Endemic Diplommatina inflatula (Crosse, 1866)* Rock Endemic Palaina moussoni Crosse, 1866* Ground Endemic Palaina patula (Crosse, 1866)* Ground Endemic Diplommatina polymorpha (Crosse, 1866)* Rock Endemic Diplommatina ringens (Crosse, 1866)* Rock Endemic Palaina striolata Crosse, 1866* Ground Endemic Hungerfordia pelewensis Beddome, 1889* Rock Endemic HELICINIDAE Palaeohelicina heterochroa (Ancey?) Ground/Veg. Endemic Pleuropoma pelewensis (Sykes, 1901) Gr./Rock/Veg. Endemic PUPINIDAE Pupina difficilis Semper, 1864* Ground Indigenous TRUNCATELLIDAE Truncatella guerinii A. and J. B. Villa, 1841 Ground Indigenous Pulmonata ACHATINELLIDAE Elasmias ovulatum (Mollendorff, 1900) Vegetation Indigenous ACHATINIDAE Achatina fulica Bowdich, 1822 Ground Introduced CHAROPIDAE Semperdon kororensis (Beddome, 1889)* Ground Endemic Semperdon uncatus Solem, 1982* Ground Endemic Semperdon xyleborus Solem, 1982* Ground/Rock Endemic ELLOBIIDAE Pythia scarabaeus (Linnaeus, 1758) Ground Indigenous ENDODONTIDAE Aadonta constricta (Semper, 1874)* Ground Endemic Aadonta fuscozonata depressa Solem, 1976* Ground Endemic Aadonta irregularis (Semper, 1874)* Ground Endemic Aadonta kinlochi Solem, 1976* Ground Endemic HELICARIONIDAE Coneuplecta turrita Belauensis Baker, 1941 Ground Endemic Kororia palaensis (Semper, 1870) Ground/Rock Endemic Liravidena lacerata (Semper, 1974) Ground/Rock Endemic Palaua minor (Semper, 1873)* Ground/Rock Endemic Videna electra (Semper, 1873)* Gr./Rock/Veg. Endemic Videna oleacina (Semper, 1873)* Ground/Rock Endemic Videna pagodula (Semper, 1873)* Ground/Rock Endemic Videna pumila (Semper, 1873)* Ground Endemic PARTULIDAE Partula calypso Semper, 1865* Veg. Endemic Partula thetis Semper, 1865* Veg. Endemic SUBULINIDAE Subulina octona (Bruguiere, 1789) Ground Introduced species), and Cowie et al. ( 1996), who suggested that there may be 40-50 indigenous/endemic species in Belau. Given the fact that only 26 areas were included in the present study, and that many of the species are very small and morphologically cryptic (e.g., diplommatinids, hydrocenids, and helicarionids), an over- all estimate of ca. 200 species for the Belau fauna is reasonable. Perhaps the most significant result of this survey was the discovery of the extraordinary species richness among the diplommatinid land snails of Belau. Previously, only 26 spe- cies were described, and yet 8 1 species were recorded in this study. Most of these species are undescribed. Several of the described diplommatinid species were not collected in this BELAU LAND SNAIL DIVERSITY 85 survey, and it is unknown whether these species are extinct or merely await collection as more islands are added to the results. Given that most habitat destruction has occurred on the islands of Babeldaob and Oreor, species from these islands may be most likely to have suffered extinction. A surprising result was the high species diversity on the island of Beliliou, despite the island’s notoriety as the site of one of the bloodiest battles of World War II (Crombie and Pregill 1999). Although much of the island was burned dur- ing this conflict, apparently enough habitat survived to main- tain high endemic species richness there, since Beliliou is home to 36 Belau endemics (Table 2). Much of this diversity is concentrated along the limestone spine of the island, popu- larly known as “Bloody Nose Ridge.” This study also uncovered at least seven endodontoid species, which is noteworthy since endodontid and charopid species are generally rare or extinct throughout most Pacific islands where they were once known to occur (Cowie 1996). Another Pacific endemic family, the Partulidae, was repre- sented by at least one species, which is likely Partula calypso. However, there could be an additional extant endemic Partula species present among some of my records (D. 6 Foighil, pers. comm. ) . Partulids were recorded from two of the islands presented in this paper. Given the extinction of Partulidae throughout the Pacific (Cowie 1992), all Belau partulid local- ities warrant conservation attention. Threats and conservation recommendations Although the impacts of rats and introduced predatory land snails ( e.g ., the rosy wolf snail Euglandina rosea Ferussac, 1821) have, rightly, been emphasized for their role in the dec- imation of land snail faunas throughout the Pacific (e.g., Cowie 1992, Lydeard et al. 2004), habitat destruction is the most significant looming threat to Belau’s endemic land snails. For example, deforestation caused by the construction of subsidiary roads to Babeldaob’s Compact Road (also men- tioned by Cowie et al. 1996, in the early stages of its construc- tion), development near the new capital building in the state of Melekeok on Babeldoab as well as planned tourist-centered areas on Babeldoab and low limestone islands could have rapid and far-reaching consequences for Belau’s snail species. Once forests are cut, a cascade of effects ensues, including erosion, soil and microclimate changes, and the subsequent invasion of non-indigenous species suited to disturbed areas. This inevitably leads (and likely has led, prior to species records) to the extinction of endemic snails, most of which have very small geographic ranges and can survive only under a narrow set of ecological conditions under which their spe- cific fungal and detrital food accumulates. Mining of limestone outcrops is another important threat to Belau’s land snails since many species, particularly diplommatinids, but also some of the Videna Semper, 1873 species and endodontoids, are endemic to limestone karst and limestone karst forests. Although to date most of Belau’s Rock Islands are protected by a management plan (e.g., Koror State) and are considered one of Belau’s most important nat- ural resources — not just for tourism, but for future genera- tions of Belauans — the limestone excavation pressure that exists in similar tropical areas, such as Malaysian Borneo (Schilthuizen et al. 2005) indicates that diligence should be exercised in maintaining Belau Rock Island protection, both in Koror and Airai States. The environs of Oikull and Ngerduais in southeastern Babeldoab are especially impor- tant since they are home to one of the few remaining popula- tions of endemic partulid tree snails. Predation by rats may be an issue for some Belau land snails, such as the slow- reproducing partulids (Cowie 1992). Broken shells of some indigenous land snail species (though not partulids), indicative of rat predation, were found on Ulong (Rundell, unpubl. data, 2003, 2005, and 2007). Given the decimation by rats of achatinelline and partulid land snails elsewhere in the Pacific (Hadfield 1986, Cowie 1992, Hadfield et al. 1993), this threat should not be ignored though further study of Belau partulid populations is needed to understand whether current populations are healthy or in decline. While partulids may have co-occurred with rats on Ulong for many years (e.g., since the shipwreck of the English vessel the Antelope centuries ago), it is unknown whether the current pockets of partulids are fragments of more widespread populations that are now in decline. Comparisons with past collections and generation of more complete current baseline data are vital in this regard. It is also unknown whether rats on other partulid islands, such as Ngeruktabel, could have a negative effect on partulids, particularly if combined with other pressures, such as other invasive plant and animal species. Evidence from other Pacific islands suggests that the answer is “yes.” Summary and future directions Future work on Belau land snail faunal diversity should include species descriptions and enumeration of geographic distributions with the eventual aim of producing a species catalogue and thorough conservation management plan for Belau land snails. Given the decimation of endodontoids and partulids throughout the Pacific region, these two groups should be paid particular attention. In addition, undescribed diversity in groups such as the helicarionids and hydrocenids indicates that these families warrant taxonomic effort. Belau’s land snails are incredibly species rich (ca. 200 extant species) relative to the small island area they inhabit, and the people of the Republic of Palau are in a unique position to conserve the many species that still remain. The presence of endemic and indigenous species throughout the Belau archi- pelago and the rarity of invasive land snail species, relative to many other Pacific island groups, indicate that if native forests 86 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Table 2. Species occurrences on each surveyed island. Taxa are indicated on vertical axis. Undescribed or unassignable species were given letter codes, which simply reflect the order of identification. Localities (individual islands in most cases) are represented by numbers. Key to island numbers is shown in Figure 1. Species presence at a locality is indicated with a “+”. Island Taxon 1 2 3 4 5 6 7 8 9 10 11 12 13 Assimineidae Omphalotropis cheynei Diplommatinidae + + + + Palaina patula Palaina dimorpha Palaina striolata Palaina K + + + + + Palaina L Palaina albata Palaina O Palaina T Palaina V Palaina moussoni Palaina AI Palaina AJ Palaina AM Palaina AN Palaina AS + + + + + + + + + + + + + Palaina AU Palaina AW Palaina BJ Palaina BK Palaina BM Palaina BN Palaina BO Palaina BP + + + + + + + + Palaina BQ Palaina BR Palaina BS + + + + + Palaina CE Palaina CL + + Palaina sp. Diplommatina lutea Diplommatina AF Diplommatina AG + + + + + + + Diplommatina AH Diplommatina AK Diplommatina AL + + + + + Diplommatina AV Diplommatina BA + + + Hungerfordia A Hungerfordia C + + + + + + Hungerfordia D Hungerfordia K Hungerfordia pelewensis Hungerfordia T Hungerfordia P + + + + + + (continued) BELAU LAND SNAIL DIVERSITY 87 Table 2. (Continued) Island Taxon 1 2 3 4 5 6 7 8 9 10 11 12 13 Helicinidae Palaeohelicina heterochroa + + + Pleuropoma pelewensis Hydrocenidae Georissa sp. 1 Georissa sp. 2 Georissa sp. 5 Pupinidae Pupina difficilis Truncatellidae Truncatella guerinii Achatinellidae Elasmias ovulatum Achatinidae + + + + + + + + + Achatina fulica Charopidae Semperdon uncatus + + + Semperdon sp. + Endodontoid Ellobiidae + Pythia scarabaeus Endodontidae + + + Aadonta fuscozonata depressa Aadonta irregularis Helicarionidae + + Kororia palaensis Liravidena lacerata Palaua minor + + + + + + + Palaua sp. 1 + Videna electra Videna oleacina Partulidae + + Partula sp. Subulinidae + + Subulina octona + + + + Island Taxon 14 15 16 17 18 19 20 21 22 23 24 25 26 Assimineidae Kubaryia pilikia + Omphalotropis cheynei + + + + Unidentified sp. 1 Diplommatinidae Palaina dimorpha + + Palaina K + Palaina albata + + + + + + ++ + + Palaina N Palaina O Palaina Q + + + + + + + + + + + + + + + + + + (continued) 88 AMERICAN M ALACOLOGICAL BULLETIN 28 • 1 12 • 20 1 0 Table 2. (Continued) Island Taxon 14 15 16 17 18 19 20 21 22 23 24 25 26 Palaina AJ Palaina AP Palaina AR Palaina AS + Palaina AY + Palaina AZ + Palaina BB + Palaina BC + Palaina BD + Palaina BE Palaina BF + Palaina BI + Palaina CE Palaina CG Palaina CH Palaina sp. Diplommatina inflatula Diplommatina polymorpha Diplommatina ringens Diplommatina AG + Diplommatina AL Diplommatina AO + Diplommatina AQ Diplommatina AT Diplommatina AX + Diplommatina BA + Diplommatina BG Diplommatina BH Diplommatina BL Diplommatina CD Diplommatina CM Diplommatina CR Diplommatina CS Hungerfordia A + Hungerfordia C Hungerfordia D Hungerfordia E Hungerfordia J + Hungerfordia K + Hungerfordia L + Hungerfordia M + Hungerfordia N Hungerfordia S Hungerfordia U Helicinidae Palaeohelicina heterochroa Pleuropoma pelewensis + + + + + + + + + + + + + + + + -+ + + + + + + + + + (continued) BELAU LAND SNAIL DIVERSITY 89 Table 2. (Continued) Taxon Hydrocenidae Georissa sp. 1 Georissa sp. 2 hydrocenids Pupinidae Pupina difficilis Truncatellidae Truncatella guerinii Achatinellidae Elasmias ovulatum Elasmias sp. 1 Charopidae Semperdon xyleborus Semperdon kororensis Endodontoids Ellobiidae Pythia scarabaeus Endodontidae Aadonta constricta Aadonta irregularis Aadonta kinlochi Helicarionidae Coneuplecta turrita Kororia palaensis Palaua minor Palaua sp. 3 Videna electra Videna oleacina Videna pagodula Videna pumila Subulinidae Subulina octona Island 14 15 16 17 18 19 20 21 22 23 24 25 26 + + + + + + + + + + + + + + + + + + can be preserved in Belau and non-indigenous species can be kept out of the country, Belau has every hope of conserving this fauna for future generations. Land snails will be not only of interest to evolutionary biologists and ecologists (though this interest should be substantial), but land snails’ presence coin- cides with healthy indigenous forests, which support water- sheds, plants, and animals critical for human survival, and that make the islands of Belau one of the most beautiful, unique, and instantly recognizable places on the planet. ACKNOWLEDGMENTS I thank the people and governments of the Republic of Palau, particularly the governors of the 16 States, who kindly granted permission to work on the lands under their admin- istration. I especially thank the Palau Bureau of Agriculture, Republic of Palau’s Office of Environmental Response and Coordination, Palau Conservation Society, Belau Cares, Inc., Belau National Museum, and Coral Reef Research Poundation for supporting my field research and/or providing facilities. I thank B. Sakuma for his continued enthusiastic and inspira- tional support of this project. Special thanks are owed to J. Miles, A. Eledui, T. Holm, L. Colin, P. Colin, R. Crombie, A. Olsen, M. Etpison, A. Kitalong, the Koror State Rangers (Belau), Ibedul Y. Gibbons, and to my field assistants J. Czekanski-Moir, A. Gawel, D. Mulroney, R. Orben, S. Wilkinson, R. Brewer, and C. Carroll. Thanks to D. Ngirkelau and additional field personnel too numerous to list. R. Kawamoto and A. Suzumoto (Bishop Museum), V. Heros and P. Bouchet (Museum National d’Histoire Naturelle), J. Slapcinsky and P. Thompson (Plorida Museum of Natural History) provided access to specimens. R. Bieler, J. Gerber, M. Pryzdia, and J. Jones provided FMNH collections support. 90 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2 -2010 E. Thompson, C. Christensen, R. Cowie, and R. Crombie provided helpful advice early in the project. I thank K. Perez for her interest in this study. Funding was provided by the National Geographic Society Committee for Research and Exploration (NGS CRE Grant 7972-06), American Malacological Society, Unitas Malacologica, Conchologists of America, FMNH Zoology Department Marshall Field Fund, and the University of Chicago Hinds Fund. LITERATURE CITED Clarke, B. C. and J. J. Murray. 1969. Ecological genetics and specia- tion in land snails of the genus Partula. Biological Journal of the Linnean Society 1: 31-42. Cooke, C. M., Jr. and Y. Kondo. 1960. Revision of Tornatellinidae and Achatinellidae (Gastropoda, Pulmonata). Bernice P. Bishop Museum Bulletin 221: 1-303. Cowie, R. H. 1992. Evolution and extinction of Partulidae, endemic Pacific island land snails. Philosophical Transactions of the Royal Society of London (B) 335: 167-191. Cowie, R. H. 1996. Pacific island land snails: Relationships, origins and determinants of diversity. In: A. Keast and S. E. Miller, eds., The Origin and Evolution of Pacific Island Biotas, New Guinea to Eastern Polynesia: Patterns and Processes. SPB Academic Pub- lishing, Amsterdam. Pp. 347-372. Cowie, R. H. 2001a. Can snails ever be effective biocontrol agents? International Journal of Pest Management 47: 23-40. Cowie, R. H. 2001b. Decline and homogenization of Pacific faunas: The land snails of American Samoa. Biological Conservation 99: 207-222. Cowie, R. H. and R. P. Cook. 2001. Extinction or survival: Partulid tree snails in American Samoa. Biodiversity and Conservation 10: 143-159. Cowie, R. H„ A. Allison, F. G. Howarth, G. A. Samuelson, and N. L. Evenhuis. 1996. Impacts of Construction of the Palau Com- pact Road: Survey of the Non-Marine Fauna of the Island ofBa- beldaob. Unpublished Report. Will Chee Planning, Honolulu, Hawaii. Crombie, R. I. and G. K. Pregill. 1999. A checklist of the herpeto- fauna of the Palau Islands (Republic of Belau), Oceania. Herpe- tological Monographs 13: 29-80. Darwin, C. 1859. The Origin of Species. Penguin Classics 1985 Edi- tion. Penguin Books, London. Emberton, K. C., T. A. Pearce, P. F. Kasigwa, P. Tattersfield, and Z. Habibu. 1997. High diversity and regional endemism in land snails of eastern Tanzania. Biodiversity and Conservation 6: 1 123- 1 136. Grant, P. R. 1986. Ecology and Evolution of Darwin’s Finches. Princ- eton University Press, Princeton, New Jersey. Groombridge, B., ed. 1993. 1994 Red List of Threatened Animals. IUCN, Gland, Switzerland and Cambridge, U.K. Hadfield, M. G. 1986. Extinction in Hawaiian achatinelline snails. Malacologia 27: 67-81. Hadfield, M. G., S. E. Miller, and A. H. Carwile. 1993. The decima- tion of endemic Hawaiian tree snails by alien predators. Ameri- can Zoologist 33: 610-622. IUCN. 2004. 2004 IUCN Red List of Threatened Species. Available at: http://www.iucnredlist.org; accessed 23 February 2009. Kelletat, D. 1991. Main trends of Palau Islands’ coastal evolution, identified by air and ground truthing. Geojournal 24: 77-85. Kobayashi, K. 2004. Origin of the Palau and Yap trench-arc systems. Geophysical Journal International 157: 1303-1315. Kobelt, W. 1902. Tierreich: Cyclophoridae. Verlag von R. Friedlander und Sohn, Berlin [In German]. Lydeard, C., R. H. Cowie, W. F. Ponder, A. E. Bogan, P. Bouchet, S. A. Clark, K. S. Cummings, T. J. Frest, O. Gargominy, D. G. Herbert, R. Hershler, K. Perez, B. Roth, M. Seddon, E. E. Strong, and F. G. Thompson. 2004. The global decline of non- marine molluscs. BioScience 54: 321-330. Motteler, L. S. 2006. Pacific Island Names. 2nd Edition. Bishop Mu- seum Press, Honolulu. Quammen, D. 1996. The Song of the Dodo: Island Biogeography in the Age of Extinctions. Touchstone, New York. Rundell, R. J. 2008. Cryptic diversity, molecular phylogeny and biogeography of the rock and leaf litter dwelling land snails of Belau (Republic of Palau, Oceania) Philosophical Transactions of the Royal Society of London (B) 363: 3401-3412. Schilthuizen, M., T. S. Liew, B. Elahan, and I. Lackman-Ancrenaz. 2005. Effects of karst forest degradation on pulmonate and prosobranch land snail communities in Sabah, Malaysian Bor- neo. Conservation Biology 19: 949-954. Schilthuizen, M., A. van Til, M. Salverda, T-.S. Liew, S. S. James, B. Bin Elahan, and J. J. Vermeulen. 2006. Microgeographic evolu- tion of snail shell shape and predator behavior. Evolution 60: 1851-1858. Smith, B. D. 1993. Working List of the Terrestrial Gastropods of Palau, Caroline Islands. Working List No. 4, Dickinson Memorial Mollusc Collection. Marine Laboratory, University of Guam, Guam. Solem, A. 1990. How many Hawaiian land snail species are left? and what we can do for them. Bishop Museum Occasional Papers 30: 27-40. Thiele, J. 1927. 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Submitted: 16 December 2008; accepted: 25 February 2009; final revisions received: 12 November 2009 Amer. Malac. Bull. 28: 91-95 (2010) Analysis of museum records highlights unprotected land snail diversity in Alabama* Russell L. Minton1 and Kathryn E. Perez2 1 Department of Biology, University of Louisiana at Monroe, 700 University Avenue, Monroe, Louisiana 71209-0520, U.S.A. : Department of Biology, University of Wisconsin at La Crosse, 1725 State Street, La Crosse, Wisconsin 54601, U.S.A. Corresponding author: perezke@gmail.com Abstract: In order to address the conservation status and needs of Alabama’s land snail species, we examined their diversity and distribution using 1 1,816 museum records representing 226 land snail species. The Chao-1 statistic identified seven areas of high species richness. The areas with the highest richness contain an estimated 200 species of land snail. These seven areas are not currently well protected by state or federal lands. While taxonomic misidentification and geo-referencing quality may be inflating our results, we suggest that studies like ours provide valuable baseline diversity estimates and launching points for continued studies. Key words: biodiversity, Alabama, land snail, conservation In comparison to their better-studied island relatives (Cowie 2001, Chiba 2003), the conservation status of main- land North American land snails remains relatively unknown. Of the over 2,000 recognized species in North America, 75 are thought to be extinct, and all but seven of these were endemic to Hawai'i (NatureServe 2008). Nine of the 75 snails listed as threatened and endangered by the U.S. Fish and Wildlife Service are terrestrial species in the contiguous 48 states (USFWS 2008). Land snail conservation has recently gained interest as population declines and extirpations continue to be documented (Lydeard et al. 2004, Steinitz et al. 2005). As with other mollusc groups, anthropogenic effects including habitat modification, urbanization, and land use practices can have strong negative effects on land snails (Graveland et al. 1994, Orstan et al. 2005, Lange 2006) given their low dispersal abilities and limited species ranges (e.g., Burch 1955, Riggle 1976, Flubricht 1985, Hotopp 2002). Land snail con- servation is important for many reasons. Terrestrial gastro- pods (snails and slugs) can serve as critical indicator species for a number of ecosystems (Prezio et al. 1999, Ovaska and Sopuck 2005). They may play significant roles in food webs and nutrient cycling through decomposition (Mason 1970, Richter 1979), and some species are known dispersers of plant seeds and fungal spores (Richter 1980, Gervais et al. 1998). Finally, they are contributors to the overall biodiversity and health of communities (Richter 1980). Nearly 200 species of land snails are estimated to occur in Alabama (Shelton 1998), and this fauna has been intensively collected for the better part of a century (Clapp 1920, Archer 1939, Hubricht 1985). The state’s land snails were last treated in detail eighty years ago (Walker 1928), and have been over- shadowed in recent times by the decline of Alabama’s fresh- water mollusc species. The most recent study dealing with the state’s terrestrial molluscs comprised a survey of the 25,000- acre Sipsey Wilderness Area in north-central Alabama (Waggoner et al. 2006). The study yielded 58 species from a small portion of the Bankhead National Forest and increased the known richness of the area four fold. The study also stressed the need for assessment of the conservation status of the state’s land snails, given their restriction to specific envi- ronments and extensive human activity in those same areas. In order to address the conservation status and needs of Alabama’s land snail species, we examined their diversity and distribution using museum records from four institutions. Museum records are useful in determining historic patterns of species composition and can provide baseline data when such information is lacking (Mikkelsen and Bieler 2001, Ponder et al. 2001). Using estimated richness values and information on the state’s protected areas, we hoped to deter- mine how diverse Alabama’s land snails are, if discrete areas of high species richness could be identified, and to what extent federally and state protected lands offered the snails protection. MATERIALS AND METHODS Museum records for Alabama land snail species were obtained from the following institutions: Auburn University Museum and Natural History Learning Center, Auburn; * From the “Leslie Hubricht Memorial Symposium on Terrestrial Gastropods” presented at the meeting of the American Malacological Society, held from 29 July to 3 August 2008 in Carbondale, Illinois. 91 92 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Delaware Museum of Natural History, Wilmington; Field Museum of Natural History, Chicago; and Florida Museum of Natural History, Gainesville. Individual records possessing detailed collection information were geo-referenced using GeoLocate (Tulane University Museum of Natural History) to generate latitude and longitude coordinates. Localities that could not be determined automatically were identified manu- ally on topographic maps. Taxonomy generally followed Turgeon et al. ( 1998), collapsing subspecies into their parent species. While most North American land snails are diag- nosed on the basis of shell characteristics and geographic dis- tribution, and few recent studies have been conducted that attempt to revise species and sub-specific classifications, we opted for a more conservative approach toward recognizing taxa. Potential effects of this decision are treated in the discus- sion. Amphibious species in the genera Melampus Montfort, 1810 and Pomatiopsis Tryon, 1862 were excluded from the analysis, while alien and invasive species were included. While most alien land snails species exist only in isolated pockets that do not spread (Dundee 1974), species like Bradybaena similaris (Ferrusac, 1821 ) are widespread throughout the U.S. and have become part of the fauna. Museum record localities were projected on to a state map of Alabama using DIVA-GIS 5.4 (Hijmans et al. 2008). Using the analysis functions in DIVA-GIS, we calculated cor- rected Chao 1 richness (Chao 1984) estimations (SI) for the entire state with a grid size of 0.2 degrees. The corrected esti- mator Si is calculated as Sobs + (FI2 / 2[F2+1]) - (F1F2/ 2[F2+1]2), where Sobs is the number of species observed in a sample, and Fi is the number of species represented by exactly i individuals (i=l for the frequency of singletons [FI ] , i=2 for the frequency of doubletons [F2] ). This allowed us to identify areas of high estimated richness with less inherent bias. To determine if the estimated high richness areas are potentially protected, we overlaid our richness estimates with maps of federally and state protected lands including national parks, national forests, reservoirs, and Alabama state parks. Only sites of 640 acres or more are identified in the federal coverage. We then used the reserve selection function in DIVA-GIS to identify sets of grid cells (theoretical “reserves”) that would capture a maximum of species diversity in as few cells as possible. The procedure is based on the algorithm by Rebelo (1994), where the cell with highest diversity is chosen first; for cells with equally high diversity, the starting cell is chosen randomly. Additional nearby cells are then chosen iteratively based on the first cell. The result is that cells with high diversity may not contribute much to the overall pro- tected diversity based on their proximity to the first cell. This is a non-linear optimization problem, and the solution of Rebelo and Sigfried (1992) is utilized in DIVA-GIS. We used a smaller grid size (0.1 degrees) to more closely reflect the min- imum size of tracts of federally and state protected land, and compared it to the location of our high richness and pro- tected areas. RESULTS A total of 11,816 museum records from the four muse- ums were geo-referenced, representing 226 land snail species. Localities were broadly distributed across the state, with some concentrated collections near major metropolitan areas (Fig. 1). Using corrected Chao 1 values, we identified seven areas where estimated land snail diversity was highest (Fig. 2). Many of these areas were near major metropolitan areas. We then overlaid federally protected lands on the estimated rich- ness with the result that none of our highest estimated rich- ness areas corresponded with protected areas. When Alabama state parks were added, parts of one of the high richness areas would be protected by Monte Sano Park near Huntsville. Other state parks, including Cheaha Mountain in the Talladega National Forest and Blanton Springs, were found in Figure 1. Collection sites based on geo-referenced museum collec- tions. Open circles represent major metropolitan areas (identified in Fig. 2). ALABAMA LAND SNAIL DIVERSITY 93 Figure 2. Estimated corrected Chao-1 species richness based on mu- seum records. Hatched areas represent federally protected lands; cir- cles represent major metropolitan areas. Two potentially protected areas of high diversity, Redstone Arsenal and Monte Sano Park, are identified by arrows. areas of medium estimated richness. The reserve function in DIVA-GIS identified one theoretical reserve in the same high richness area as Monte Sano Park. DISCUSSION Our analyses included museum records for 226 species of land snail in Alabama, counting invasive and alien taxa. This value is a bit higher than those generated previously by Hubricht (1985; 145 species) and Shelton (1998; 194 species). We found isolated areas of high land snail diversity throughout Alabama, with few patterns of richness being readily apparent (Table 1). The locations of the highest estimated richness areas show no relationship between the number of collection locali- ties and species diversity. One of the two highest estimated diversity areas is in northwestern Alabama near the Tennessee River, where only a few collections have been made. Repeated sampling of the same species likely explains areas with lower Table 1. Locations and corrected Chao-1 richness estimations (SI) for the seven most diverse areas predicted by museum records. Ecoregions refer to level III areas designated by the U.S. Environ- mental Protection Agency. County Ecoregion SI Colbert/Franklin Southeastern Plains 200 Mobile Southern Coastal Plain 163 Madison Southwestern Appalachians 151 Butler/Wilcox Southeastern Plains 145 Montgomery 1 Southeastern Plains 140 Montgomery 2 Southeastern Plains 128 Wilcox Southeastern Plains 128 estimated richness despite more numerous collection sites. Areas around the Tennessee River in the northern part of the state showed high overall diversity compared with other areas. This was expected, as the area around the river tends to be higher altitude with exposed limestone and is more densely wooded than other parts of the state. These high-calcium for- ested areas have been shown to have high snail diversity (Gardenfors 1992, Hotopp 2002, Jurickova et al. 2008). Only one area of high diversity occurred near either federal or state lands, the region east of Huntsville represented primarily by Monte Sano Park, just outside of the Redstone Arsenal. The high diversity observed in some of our areas may be a result of including invasive and alien species in our analyses. We feel this is not a serious issue as fewer than ten non-native species occurred together in any one area. Most introduced snails with Alabama records were found in and around Mobile, supporting the notion that invasive and alien species enter through commercial ports and may become established near them (Dundee 1974). A few single widespread records of alien species likely reflect greenhouse species found on imported plant material. While some introduced species have become ubiquitous components of the ecosystem, such as the previously mentioned Bradybaena, our inclusion of the occa- sional non-native should not be interpreted as support for the notion that introduced species are beneficial by increasing species diversity. While invasives may increase diversity on a small temporal and spatial scale, their importance has been well documented in the overall decline of native diversity and overall richness (Davis 2003, Keeley et al. 2003). More likely is that our species diversity and distribution figures in Alabama suffer from two of the limitations identified by Guralnick et al. (2007) in using museum specimens. First, taxonomic misidentifications may have inflated our richness estimates. Specimen misidentification rates may be as high as 60% in some groups (Meier and Dikow 2004), producing mis- leading results. Second, since best practices for geo-referencing are still relatively new (Chapman and Wieczorek 2006), issues of 94 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2 -2010 accuracy arise. Although we used GeoLocate as a means for stan- dardization, we did not treat low and high accuracy references differently. Thus, the accuracy of our points varies among data and may further alter our estimates (Guralnick et al. 2006). In biodiversity estimates, misidentifications also complicate accu- rate delineation of areas of endemism and other hotspots (Ng and Tan 2000). Thus, analyses like ours should be seen as starting points for continued studies, and not the final word on richness or distribution. Using museum records for diversity estimation can be fruitful, but there are also significant biases that may exist in our data. Finding high diversity areas near cities is a common bias encountered when using collection data for richness calculations (Hijmans et al. 2000). This potential non-representative sampling bias is the most difficult source of error to correct for in natural history data (Williams etal. 2000). Museum data also provide presence-only data and may not reflect the true distri- bution of a species (Graham et al. 2004), either in historical or modern times. The scope of the museum data is over -150 years of collections, and land use changes have surely affected diversity. In a poorly studied group with morphologically delin- eated species like land snails, identification errors can skew rich- ness in both directions. Combined with inexactness in collection locality information, misidentifications introduce the most error (Chapman 1999). Even with these potential shortcom- ings, the increase in availability of museum records has led to a corresponding increase in their incorporation into conserva- tion studies, with positive results (Ponder et al. 2001, Hugall etal. 2002, Raxworthy et al. 2003). Studies like ours can play an important role in discover- ing biodiversity hotspots, which are areas with high numbers of endemic species along with specific biotic characteristics (Myers 2003). These hotspots are usually based on floral and vertebrate-oriented estimates, with the assumption that pro- tecting diversity in those two groups will protect a similar number of invertebrates. This is unfortunate, since land snails, as part of the “other 99%” of global diversity (Ponder and Lunney 1999), have been shown to predict vertebrate conservation priorities but not vice versa (Moritz et al. 2001 ). 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Submitted: 22 October 2008; accepted: 4 August 2009; final revisions received: 25 September 2009 Amer. Maine. Bull. 28: 97-103 (2010) Strategies for collecting land snails and their impact on conservation planning* Marla L. Coppolino* Department of Zoology, Southern Illinois University, Carbondale, Illinois 62901, U.S.A. Corresponding author: marlacoppolino@gmail.com Abstract: Land snail collecting methods are each designed with a particular purpose. This paper provides a brief overview of land snail collecting strategies, suggests the use of multiple methods and collection of ecological data to broaden the knowledge of land snail life habits and relationships with their environment, and indicates approaches that are most relevant to land snail conservation. Key words: search methods, historical data, terrestrial snails, Pulmonata, Gastropoda Land snails exhibit a wide range of habitat preferences, varying with species and region, but these generally reflect their main survival requirements: moisture, food, shelter, and a source of calcium for shell building and physiological pro- cesses (Burch and Pearce 1990). As a consequence, some of the greatest species abundance and richness occurs in moist, deciduous forest (Pearce and Orstan 2006). Nevertheless, snails have evolved to survive also in arid environments (Bequaert and Miller 1973). Fens, algific talus slopes, wetlands, and rock, particularly limestone bluffs are also habitat to various snail species (Nekola 1999). Most species require a humid environment and seek shelter in microhabitats such as under logs, rocks, leaf litter, in and around bryophytes, coarse woody debris, and moist vegetation. Other places that sup- port snails include the interface regions of the forest floor, such as the crevices between a log and the ground litter and between exposed tree roots. It is often in these places that vegetation is found in a decomposing state, so these micro- habitats are also important as areas for feeding because most land snails are generalist herbivores (Burch and Pearce 1990). Many approaches for sampling terrestrial molluscs have been suggested in the literature. Most of these studies discuss methods in the context of a research project (Grime and Blythe 1969, Riggle 1976, Coney etal. 1982, Alvarez and Michael 1993, Emberton etal. 1996, Hawkins etal. 1998, Nekola 1999, Tattersfield etal. 2001, Hotopp 2002, Millar and Waite 2002), while others specifically address collection methods and com- parisons between them (Bishop 1977, Boag 1982, Emberton et al. 1996, Cameron and Pokyryzko 2005). Historically speaking, most of the land snail collecting in North America has been qualitative, and many distributional records are simply the recorded presence of a species (Hubricht 1985). These records offer collectors useful baseline information on species, habitats, and ranges, and what habitats might still be best for populations of endangered species, where collection of live specimens is forbidden. Since the mid-20th century, collecting efforts in North America and elsewhere have turned more toward quantitative studies, often perceived as having greater scientific value. Such studies require that land snails are collected using some standard measurement, either by time, volume sampled ( e.g ., of leaf litter, soil, etc.), or area, and often by some combination of these factors. Most often, snails collected in this type of study are also accompanied by habitat or microhabitat data, which is also measured in a quantifiable, repeatable way. When a collecting method is quantitative, the snail abundance and diversity counts can be legitimately compared, and more effectively used in statistical analyses. Many contemporary studies use quantitative methods of collection, from which population dynamics can be studied (Bishop 1977). The choice of collecting method(s) is dependent upon ( 1 ) the extent and scope of the investigation, (2) the type(s) of terrain to be sampled, and (3) the intent of the investigation. Among the basic questions that require answers before making this choice are: Is the study exploratory in nature or does it have specific goals? Can the objectives be met with qualitative observations or does the study require quantitative data to answer specific questions? Does the investigation require (or would it benefit from) ecological and/or environmental data for comparison with species data? Regardless of the responses to these questions and the choices made, the most important consideration is that the methods are repeatable and that they have scientific rigor. * From the “Leslie Hubricht Memorial Symposium on Terrestrial Gastropods” presented at the meeting of the American Malacological Society, held from 29 July to 3 August 2008 in Carbondale, Illinois. + Present address: 384 Pleasant Valley Road, Groton, NY 13073, U.S.A. 97 98 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 LAND SNAIL COLLECTING STRATEGIES Qualitative visual searching Qualitative observations are historically relevant, and in modern times often provide the first indication that a land snail survey is necessary, especially when unexpected species are casually encountered. Direct visual search is the simplest and least expensive method, but is of course restricted to species and specimens that are large enough to be seen in field situations. Qualitative data also have limited application in statistical analyses. Even when quantitative methods are also used, supple- mental qualitative searching can recover elusive or rare species not collected within the measured and/or timed searches. Most importantly, snails and slugs present on tree trunks and limbs or vertical rock ledges are usually not effectively sampled in quadrat samples. Hence, the inclusion of such qualitative collections, i.e., collections made by direct visual searching, can maximize estimates of diversity. Quantitative quadrats and transects Many examples of quantitative sampling methods exist in the literature (Riggle 1976, Bishop 1977, Coney et al. 1982, Bengtsson et al. 1994, Emberton et al. 1996, Nekola 1999). For these types of surveys, it is most common to section an area into plots or linear transects, which usually comprise a gradient of the landscape. Quadrats, most often areas of one square meter or less, are demarcated on the ground using a frame (Bishop 1977, Emberton et al. 1996). Each transect, plot, or quadrat is visually searched for a predetermined length of time, e.g., 30 minutes (Emberton et al. 1996). In areas of low- slope soft substratum, the entire contents of the quadrats (leaf litter, coarse woody debris, etc.) can be collected as a bulk sample and returned for sorting in the laboratory (Coney et al. 1982, Bengtsson etal. 1994). Replicate quadrats can be arranged along a linear transect (Alvarez and Michael 1993), randomly placed, or located using a stratified random sampling method (Bishop 1977, Emberton etal. 1996), in which replicate locations are randomly selected from areas most likely to include snails. The latter school of thought follows the understanding that snails tend to be patchy in distribution. Pearce and Orstan (2006) noted that they are most likely to be found under the bark of trees, near bases of rocks, under logs, and in other microhabitats that might be missed in a completely random selection of samples — but this approach has been criticized as biased preferential sampling (Cameron and Pokyryzko 2005). The obvious major advantage of quantitative samples is that inter- and intra-site comparisons and statistical analyses can be performed. The most serious disadvantage is the time required to process bulk samples (and replicate samples) in the laboratory. In a recent survey of the land snails of southwestern Illinois (Coppolino 2009), a total of 240 one-liter leaf litter samples were collected from 60 sites, as one portion of the sampling method. These required an average of 30 minutes to sort and identify per sample, or a total of 120 hours for the entire study (not including the time required to identify and tabulate the specimens). Nevertheless, this amount of effort was necessary to achieve the goals of that investigation. Cardboard sheets Cardboard sheets or masonite boards have been advocated by some researchers for attracting land snails (Boag 1982, Hawkins et al. 1998). The sheets are placed at intervals within a site and left for predetermined periods of time, after which snails are recovered from the undersides of the sheets. Boag (1982) and Hawkins etal. { 1998) each claimed that this method recovered species, especially slugs, which might not otherwise be collected. However, cardboard sheets have a number of limitations that can, in certain circumstances, outweigh the advantages. ( 1 ) A cardboard sheet is arguably not quantitative because attracted snails can migrate from outside the two- dimensional area, thus sampling more than its dimensions. (2) A sheet can be moved, by wind or the actions of animals, if not anchored in place. (3) A sheet is prone to warping and/or disintegration from excessive rainfall, reducing the effective surface area in contact with the substratum (although its effectiveness might in fact depend on a certain degree of moisture [Boag 1982, Hawkins et al. 1998], for better or for worse). (4) Some investigators have claimed that cardboard sheets have been ineffective at collecting any land snails (T. A. Pearce, pers. comm.). Nevertheless, they are valuable in qualitative surveys, especially if complementing other types of samples, for determining species richness or diversity. Bulk sampling It is well known that much of the terrestrial snail fauna constitutes microsnails, defined as individuals measuring 5 mm or less in greatest shell dimension as adults or juveniles. In many areas, tiny snails comprise most of the resident species (Pearce and Orstan 2006). As such, microsnails are often missed by visual searching. A frequently used method is bulk sampling leaf litter (and usually the top 2 cm of soil), where microsnails are most likely to be found (Pearce and Orstan 2006). This material is later sorted in the laboratory under magnification, sometimes assisted by passing through a gradu- ated series of sieves to provide size fraction data (Emberton et al. 1996, Hotopp 2002, Pearce and Orstan 2006). This method can be conducted qualitatively or quantitatively, in the latter case by collecting and fully sorting a predetermined amount of leaf litter for each sample. The time required for processing bulk samples can be, as in the case of bulk material from quantitative quadrats, the greatest disadvantage. LAND SNAIL COLLECTING 99 LIVE VERSUS DEAD COLLECTING In both qualitative and quantitative studies, the question arises whether to collect only empty shells, only live snails, or both. The choice again depends on the goals of the research, and each has its advocates. In studies with the goal of providing an accurate current estimate of snails living in a given area, only living specimens are counted (Boag 1982, Hawkins et al. 1998, Sulikowska-Drozd 2005). Preserved tissues or intact specimens from such studies can provide material for morpho- logical and molecular investigations, potentially identifying species new to science, including cryptic species that often remain overlooked (Bickford etal. 2006). Two exceptions restrict the use of live-collecting, even if goals suggest otherwise: (1) the presence of rare or endangered species and (2) endemic species, even if in high abundance. The best examples of this are freshwater mussels, arguably among the most endangered invertebrates in North America (Cummings and Bogan 2006). In such cases, population densities should be estimated in some manner, but collecting severely limited to one or a few repre- sentatives (with proper permits, of course), or to empty shells only. Other studies have included both live snails and empty shells to maximize diversity assessment. In the marine realm, Bieler and Mikkelsen (2004) judged dead shells to be essential to describing bivalve diversity in the Florida Keys, recovering 50.8% of a predicted species list through original samples including dead shells, compared to only 30.4% recovered by live-collected specimens alone. For land snails (Coppolino 2009), only 13% of the total snails recovered were live- collected, and most of the species considered rare in southern Illinois were collected from empty shells. In a land snail diversity study in Turkey, Orstan et al. (2005) found fourteen species (58% of total collected) as empty shells only and two species (8% of total) as live snails only. Results such as these indicate that total diversity is best estimated by collecting both live snails and empty shells. In some cases, species identification can be achieved from shell fragments. However, among the disadvantages of dead-collected shells is that the amount of time that the snail has been dead is largely unknown. The numbers of dead shells collected for a study can bias its results as abundances do not truly reflect those of living individuals (Pearce 2008). Pearce (2008) found several factors that influence the rate of shell decomposition, namely dissolution, breaking, and bioerosion; additionally, shells of different species were shown to decompose at different rates. Another uncertainty lies in translocation of empty shells, which, for instance, can roll downhill from a higher elevation, distorting accurate microhabitat records for those specimens. At some sites in Illinois (Coppolino 2009), great numbers of empty shells were found at the base of rock bluffs, as opposed to far fewer found at the tops. ASSOCIATED DATA Environmental data In addition to basic locality information, the collection of associated data often includes ecological variables, both biotic and abiotic, in the land snail habitat. Shimek (1930) first recorded land snail associations with ecological variables. He described, for example, how snail assemblages changed over a geographic region in relation to the changing vegetation types. He suggested that forest types could serve as an index for snail populations, and vice versa. Since that time, numerous other examples have been published, commonly including soil factors, vegetation, topography, and/or climate data (Burch 1955, 1956, Wareborn 1970, Coney etal. 1982, Emberton etal. 1996, Nekola 1999, Nekola and Smith 1999, Tattersfield et al. 2001). Some general correlations with ecological factors are recognized, such as high-calcium, high-pH soils supporting greater snail abundance and diversity (Burch 1955, Riggle 1976, Hotopp 2002), and such assumptions often steer project design. In practice, however, time and budgetary constraints are often the strongest determinants of which data to collect or exclude. Habitat complexity in the broad sense has been suggested as influencing land snail abundance and diversity. A habitat complexity index was devised for use in a survey of southwestern Illinois land snails (Coppolino 2009), combin- ing vegetation diversity, topographical changes, and exposed rock, which showed positive associations with species diversity and abundance in multiple regression analyses. Soil variables such as moisture, calcium, and pH have been shown to positively influence land snail abundance and diver- sity (Burch 1955, Riggle 1976, Nekola 1999, Nekola and Smith 1999, Tattersfield et al. 2001, Hotopp 2002, Millar and Waite 2002, Coppolino 2009). Not surprisingly, calcium was strongly correlated with species diversity (although to a lesser degree with species abundances) in southwestern Illinois (Coppolino 2009). Soil factors that could negatively impact land snail populations, such as iron, were shown in a multiple regression as negatively correlated with species diversity (Coppolino 2009) , and could also be worthy of measurement. Radioactive strontium, a product of nuclear fallout, behaves chemically like calcium and could therefore be used as a marker, suggesting further investigation of this topic (T. H. Nation, pers. comm.) . Rock type can be closely related to soil calcium content, and has also been shown to be a significant factor in both land snail abundance and diversity (Nekola 1999, Nation 2005). As an alternative to collecting original soil and rock data for each station in a study, maps are often available for soil types from the U.S. Forestry Service (http:// www.fs.fed.us/) and for underlying and exposed rock through the United States Geological Survey (http://www. usgs.gov/). The limitation of these data, however, is that 100 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 they seldom cover the study area exactly, thus requiring extrapolation or use of over-generalized data. Vegetation type, at both broad and microhabitat levels, has been shown to influence snail abundance and diversity (Burch 1956, Karlin 1961, Grime and Blythe 1969, Beyer and Saari 1977, Nation 2007), and much remains to be learned about the relationships between land snails and particular plant species. In light of this concern, Karlin (1961) urged that vegetation records be reported in conjunction with land snail records. This could be an important element to a study despite the fact that soil type, climate, and other environmental factors in turn influence vegetation type, making exact associations between vegetation and snails difficult to discern. The complexity and interrelatedness of these and other biotic and abiotic variables, as well as their relative importance to land snail diversity and abundance, have been discussed by other authors (Wareborn 1970, Coney et al. 1982, Nekola 2003, Nation 2007). Ecological relationships Most professional and public attention in wildlife conser- vation (and thus most of the available funding) lies above land snails in the food web, i.e., land snail consumers. Nevertheless, recognition of imperiled “charismatic megafauna” that are also snail consumers could warrant a more serious approach to land snail conservation. One example of such a relationship was found by Graveland et al. (1994), in which a decline in forest passerines (song birds) was correlated with a decline in one of their main sources of dietary calcium — land snails — presumably due to acidic rainfall. Another good example is wild turkeys, which are popular game birds that easily draw the attention of conservation officials, wildlife managers, and the general public; it has been shown that female turkeys consume 40% more land snails during the egg-laying phase of the year (Beasom and Pattee 1978). Lunding for land snail studies can also be facilitated by recognition of snail-predator relationships in which the predator is specialized or dependent on the snail for survival. One such example was a review of the state of knowledge about freshwater Llorida Applesnails ( Pomacea paludosa (Say, 1829)), which are the sole food item for the endangered Everglades or Snail Kite ( Rostrhamus sociabilis), funded in 1989-1990 by Florida Game and Fresh Water Fish Commission (Turner and Mikkelsen 2004). Thus, recording data on birds and other malacophagous predators in a land snail study area could an important consideration for conservation. Historical data Because land snails have a “survive where you are” strategy, they cannot escape rapid alterations to their environ- ment. Thus, when surveying an area of land for snails, it is worthwhile to investigate the history of the land, its use, and the major impact events upon it that potentially affect snail populations. Natural disturbances such as floods can alter the composition of snail populations in an otherwise amenable habitat (Coppolino 2009). The irreversible effects of human- induced interference can also greatly reduce or eliminate snail populations. For example, housing and commercial construc- tion, agricultural use of land, roads and golf courses, and the chemicals used on them, potentially affect snail numbers (Kay 1995), and even recreational activities such as rock- climbing have been shown to adversely affect snail populations (McMillan 2003). Fire, whether from natural causes or human-induced, can negatively impact snail abundance and diversity (Nekola 2002, Kiss and Magnin 2003, Severns 2005). Artificial modifications to rivers cause permanent changes in a landscape, which can have detrimental consequences to the native snail species (Cejka et al. 2008). As studies such as these help us to understand changes in snail communities over history, it is prudent that we use this information to preserve areas that naturally support land snail populations. For example, in considering future land use plans, Orstan etal. (2005) emphasized that the land surrounding a unique snail habitat in Turkey was being threatened by the rapid human development. Museum holdings and the literature can provide data to generate a baseline to compare with current investigations and are often the only reference baselines available. Hubricht’s (1985) distribution guide remains the most comprehensive work of its kind, supplying locality data of land snails by county in each state in the eastern continental United States. However, caution must be exercised when interpreting presence/ absence data in comparison with any kind of past collection records, because the absence of a species in a collection does not necessarily translate to absence in the area; it might merely have not been collected or recorded (Waggoner etal. 2006). RECOMMENDATIONS FOR CONSERVATION PLANNING Land snails are among the world’s fastest declining faunas (Lydeard et al. 2004). As such, any land snail surveyor should feel obligated to identify species and populations that are most at risk in his or her survey area. Orstan et al. (2005) identified land snail populations in Turkey that were being reduced at the edges of a large, sprawling urban area. Further- more, populations of some species, such as Euchemotrema hubrichti (Pilsbry, 1940) thrive in great abundance, but only over an extremely localized area (Anderson and Smith 2005). If the land supporting this endemic population is damaged, there could be little chance of recovery. As in any biological study with complex variables and goals, multiple sampling methods are recommended in any survey that aims to improve land snail conservation. Combined LAND SNAIL COLLECTING 101 Table 1. Land snail collection methods, with advantages and disadvantages. Method Advantages Disadvantages Qualitative visual searching - Simplest, least expensive - Recover rare or elusive species - Sample hard-to-sample habitats (e.g., rock ledges) - Restricted to large-bodied species visible in the field - Limited statistical applications Quantitative quadrats and/or transects - Most effective inter- and intra-site comparisons and statistical analyses - Can have time limit for the area to standardize search effort - Location choice can be a potential bias Bulk (leaf litter) samples - Recover microsnails - Qualitative or quantitative applications - High time investment required to sort samples Cardboard sheets - Attract elusive species, especially slugs - Quantitative nature questionable - Moveable over time - Prone to warping and/or disintegration - Ineffective in some studies quantitative/qualitative survey methods are most effective in maximizing diversity assessment, usually important in such cases. The technique itself need not be elaborate; for example, the combination of visual searching, quantitative quadrats, and bulk leaf litter/soil samples for microsnails might provide excellent coverage. The added inclusion of ecological data increases the breadth and potential usefulness of the study. Partnerships between malacologists and botanists, ornithol- ogists, geologists, soil scientists, and others are almost always fruitful; some examples include works by Grime and Blythe (1969), Beyer and Saari (1977), Beasom and Pattee (1978), Coney et al. (1982), Graveland etal. (1994), Tattersfield etal. (2001), Kiss and Magnin (2003), McMillan et al. (2003), and Cejka etal. (2008). Such collaborative studies are disseminated to a larger body of scientists, and importantly, more readily reach the interest of land and wildlife conservation officials. Although the definitions of “rare” species vary with context, a rare land snail is generally considered to occupy a limited geographical range or niche, regardless of local abundance (Bengtsson et al. 1994). Land snail “hotspots,” or localities that support rare or endemic species, have been identified to assist conservation efforts; Solymos and Feher (2005) indicated areas of priority by analyzing relationships between the protec- tion status and rarity of each land snail species in the study area. However, it should be remembered that “rarity” can change over time, e.g., actually increasing with changing environ- mental conditions or effectively decreasing with sampling that reveals additional populations. Cameron and Pokryszko (2005) noted that most areas are visited only once for formal land snail surveys. Although such one-time surveys provide valuable baseline inventory data, the effects of season, weather, and other natural events can bias results. Multiple surveys of one area over time can correct for this bias. Ideally, a survey should be performed at regular intervals, so that not only a baseline inventory can be achieved, but also so that natural fluctuations in population composition can be understood. As a final note, regular communication with natural resource and wildlife officials is essential to alert them of land snail species in need of conservation. The fact that snails represent a vital part of the terrestrial ecosystem can be taught through educational programs and publications. A malacologist’s pres- ence among state and federal wildlife agencies can foster only a better understanding of the importance of land snails. Efforts to continually assess the conservation status of land snails should be implemented whenever possible. CONCLUSIONS In summary, land snail surveys can be effectively accom- plished using a variety of methods (Table 1), each with its own advantages and disadvantages. The choice of which method(s) to use depends on a myriad of questions, related to physical factors of the survey area, the questions that the survey is designed to address, and the practical limitations {e.g., budget and time constraints) of the project. Given the vulnerability of land snail populations across the globe, surveyors should keep conservation issues in mind during the design phase, to include appropriate environmental and ecological parameters that will increase the usefulness of the results for future applications. ACKNOWLEDGMENTS I wish to thank Kathryn Perez for the gracious invitation to participate in the Land Snail Symposium at the American Malacological Society meeting in 2008. 1 also 102 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2-2010 extend my gratitude toward Paula M. Mikkelsen and Travis H. (Rocky) Nation for helpful comments and suggestions that greatly improved this paper. Linally, I would like to express appreciation for my opportunity to work under U.S. Pish and Wildlife Service/Illinois Department of Natural Resources State Wildlife Grant T-32-P, A quanti- tative study of land snail diversity across multiple habitat types in southern Illinois, received by Frank (Andy) Anderson, Department of Zoology, Southern Illinois University, Carbondale. LITERATURE CITED Alvarez, J. and W. Michael. 1993. Effects of treefall gaps on the den- sity of land snails in the Luquillo Experimental Forest of Puerto Rico. Biotropica 25: 100-110. Anderson, F. E. and D. A. Smith. 2005. A redescription of the cari- nate pillsnail, Euchemotrema hubrichti (Pilsbry, 1940) (Pulmo- nata: Polygyridae), with notes on habitat and genetics. Zootaxa 807: 1-11. Beasom, S. L. and O. H. Pattee. 1978. Utilization of snails by Rio Grande turkey hens. Journal of Wildlife Management 42: 916-919. Bengtsson, J., S. G. Nilsson, and S. As. 1994. 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Oikos 21: 285-291. Submitted: 9 February 2009; accepted: 2 July 2009; final revisions received: 14 October 2009 Amer. Make. Bull. 28: 105-112 (2010) Surfing snails: Population genetics of the land snail Ventridens ligera (Stylommatophora: Zonitidae) in the Potomac Gorge* Colleen S. Sinclair Department of Biological Sciences, Towson University, 8000 York Road, Towson, Maryland 21252, U.S.A. Corresponding author: csinclair@towson.edu Abstract: The population structure of the land snail Ventridens ligera (Say, 1821) was investigated in the Potomac River Basin (Virginia, Maryland, U.S.A.). Animals were collected from two islands and the adjacent riverbanks along an 8.8-km stretch of the river. Four landlocked populations in Illinois and Maryland were also sampled to provide a comparison to the river populations. A total of 246 individuals were genotyped with five newly designed species-specific microsatellite primers. Low pairwise FgT values (<0.0477) among the Potomac River sites suggest high levels of gene flow between the populations. In contrast, the landlocked populations had high FST values (0.0738 to 0.6004) which suggest genetic structuring, most likely due to physical isolation, because FST values >0.2 indicate population structuring. Low-level isolation by distance was found among the Potomac River populations, and the low FST suggests that the river is facilitating gene flow rather than acting as a barrier. Key words: genetic diversity, microsatellite, isolation by distance Distribution of genetic variation in natural populations often varies across a species range. Separation due to distance or physical barriers {i.e., bodies of water, fragmented habitat) can further subdivide a species into local subpopulations (Selander and Kaufman 1975, Selander and Ochman 1983, McCauley 1995, Pfenninger et al. 1996, Arnaud et al. 1999a, 1999b). Isolation can often lead to lower genetic diversity and a higher risk of extinction (Crozier 1997, Saccheri et al. 1998, Allendorf and Luikart 2007). Due to a limited ability to disperse, land snails tend to live in discrete populations with neighboring populations genetically more similar than distant populations (Pfenninger et al. 1996, Arnaud et al. 2001, Schweiger et al. 2004). Arnaud et al. (2001) found a positive correlation between geographic distance and genetic distance in the land snail Helix aspersa (Muller, 1774) sampled along a 900-m ditch in the polders of the Bay of Mont-Saint-Michel in France (FST = 0.055 to 0.02). Land snails isolated by water {i.e., on islands) face a greater barrier to gene flow than distance. Gene flow may occur only during flooding, when debris carrying snails washes ashore, or if the snails are inadvertantly transported by other organisms. Migration of a river channel, connecting or separating pieces of land and the populations on them, may also facilitate or disrupt gene flow. Therefore land snail populations separated by barriers, such as rivers, would be expected to show the same or greater genetic isolation than populations in continuous environments. Few studies have been done on land snails to assess the effect of physical barriers on gene flow. Arnaud et al. (2003) reported that the population sub-structuring found in Helix aspersa in the polders of the Bay of Mont-Saint-Michel was due to isolation by distance and not by habitat fragmentation as might be expected. My study focused on Ventridens ligera (Say, 1821) populations living along the riverbanks and on two islands in the Potomac River Basin. Ventridens ligera is a small (11-15.6 mm) snail in the subfamily Gastrodontinae (Burch 1962). Individuals are usually found in the leaf litter of wooded areas. The species range extends from New York to Florida and west to Michigan and Oklahoma. Species- specific microsatellite markers were developed to assess the spatial partitioning in VI ligera populations located along the Potomac River and to investigate whether the river acts as an effective barrier to gene flow. MATERIALS AND METHODS Sample collection Ventridens ligera were collected at 14 locations along a 8.8-km stretch of the Potomac River north of Washington D.C., including Bear Island and Plummers Island, and both riverbanks adjacent to the islands located in the Chesapeake and Ohio Canal National Historical Park (Maryland) and George Washington Memorial Parkway/ Great Falls Park * From the “Leslie Hubricht Memorial Symposium on Terrestrial Gastropods” presented at the meeting of the American Malacological Society, held from 29 July to 3 August 2008 in Carbondale, Illinois. 105 106 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 (Virginia) (Eig. 1A-B). Samples were collected during the summer months in 2005 and 2007. Additional samples were collected on the campus of Southern Illinois University in Carbondale, Illinois and a site outside of Carbondale, Illinois (Fig. 1C), the campus of Towson University in Towson, Maryland and a site in southwestern Baltimore County, Maryland (Fig. 1A) in July 2008. Ten to twelve individuals were collected at each site. Samples were frozen at -80 °C upon return to the laboratory at Towson University. Alcohol preserved specimens of congeneric Ventridens were provided by the Field Museum of Natural History (Chicago, Illinois), the Carnegie Museum of Natural History (Pittsburgh, Pennsylvania), the Florida Museum of Natural History (Gainesville, Florida), and the Delaware Museum of Natural History (Greenville, Delaware). Microsatellite marker development Ten previously published land snail microsatellite markers (Guiller et al. 2000, Wirth 2000, Depraz et al. 2008) were tested on Ventridens ligera, but all failed to amplify or give a clean product suitable for genetic analysis. Therefore, Ventridens specific primers were developed. Microsatellite markers were developed following the protocol described by Hamilton et al. (1999). Briefly, genomic DNA from five different V. ligera individuals was pooled and fragmented with restriction enzymes Haelll, Rsal, and Nhel. Fragments between 200-1000 bp in length were isolated by gel electrophoresis and ligated to special linker fragments that facilitate isolation and cloning. Probes containing microsatellite repeats were then hybridized to the linker-DNA assembly in order to isolate fragments that contained repeats. The resulting fragments were ligated into pBluescriptll KS plasmids (Stratagene, La Jolla, California) and the recom- binants transformed into competent DH5a Escherichia coli cells. Plasmid DNA was extracted from clones positive for inserted DNA using a Qiagen MiniPrep Spin Plasmid Kit (Qiagen, Valencia, California). Insert DNA was sequenced on an ABI 3130 XL Genetic Analyzer with plasmid primers T7 and M13 Reverse at the Center of Marine Biotechnology (Baltimore, Figure 1. Map of collection sites. A, Map of the state of Maryland showing the location of collection sites TU, MD, and the Potomac River sites; B, map of the collection sites located in the Potomac River Basin below Great Falls; C, map of the state of Illinois showing the location of Carbondale and collection sites SIU and MC. POPULATION GENETICS OF VENTRIDENS LIGERA 107 Maryland). Primers were designed for each microsatellite locus using Primer3 (Rozen and Skaletsky 2000). Plasmid DNA was purified from 305 bacterial colonies and sequences obtained for 125 plasmids containing appropriately sized inserts. Twenty of the sequences were used for primer development and five were ultimately found informative. All five microsatellite loci were highly polymorphic (Table 1) with the number of alleles ranging from 17 at locus VL301 to 27 at locus VL10. Observed and expected heterozygosities for each locus ranged from 0.420 to 0.699 and 0.697 to 0.784, respectively. Primers were also tested on fourteen additional species of Ventridens and found to amplify comparably sized products in nine of the fourteen (Table 2). DNA preparation Genomic DNA was purified from a small piece of foot tissue following the protocol described in Miller et al. (2000). A 2-4 mm length of foot was removed with a sterile razor blade and placed in 460 pi of extraction buffer (10 mM Tris pH 8.0, 50 mM EDTA, 1% SDS) along with 30 pi Proteinase K (10 mg/ml - final concentration 0.6 pg/pl) and 10 pi 0.1M DTT. Samples were incubated at 55 °C, 225 rpm for 2-3 h. Once the tissue had been completely digested, 5M NaCl was added to a final concentration of 1.5M and the sample incubated at 55 °C, 225 rpm for 30 min. Samples were centrifuged for 10 min at 0.87 RCF to pellet any remaining undigested material or debris and the supernatant transferred to a sterile microcentrifuge tube. An equal volume of chloroform was added and the mixture incubated at room temperature for 30 min at 100 rpm on an orbital shaker. The mixture was then centrifuged at 0.87 RCF for 10 min. The aqueous phase of the suspension was transferred to a sterile microcentrifuge tube and an equal volume of isopropanol was added. The tube was inverted several times to mix and centrifuged at 0.94 RCF for 15 min to pellet the DNA. Following centrifugation, the DNA pellet was washed with 70% ethanol, air-dried, and re-suspended in 75 pi Tris-EDTA buffer pH 8.0. Microsatellite genotyping Individual genotypes were obtained using five polymorphic microsatellite loci (Table 1). Polymerase chain reaction (PCR) was conducted in 15 pi volumes under the following condi- tions: IX PCR buffer (Gene Choice, Continental Lab Products, San Diego, California), 200 pM dNTPs (Roche, Indianapolis, Indiana), 0.2 pM of each primer, 1.25 units of Taq polymer- ase (Gene Choice), and 30 ng of template DNA. The PCR thermocycling program consisted of 94 °C for 3 min, 35 cycles of 94 °C for 30 s, TA (dependent on primer pair, Table 1) for 30 s, and 72 °C for 45 s followed by a final extension of 72 °C for 5 min. Primers for locus VL542 required a touchdown PCR with the TA stepping down 2 °C from 70 °C to 62 °C with 5 It £ § o , •S § £ G-S r£> \p o h G 6 § LG & af R: 5’-CACCTACGGTAATCGGCACT-3’ VL301 (GT)4(GTT)22TGATGGTT(GT)5(GTT)5 212-262 58 F: 5’-GCTTCAGTTTTCAGGGCATC-3’ 246 17 0.640 0.724 R: 5’-GGGTCCTGGACTCATAGCAA-3’ VL306 (CAA)35TAACAA(TAA)2(CAA)27 224-338 60 F: 5’-GATCGCGCCTTCAAATAACT-3’ 246 25 0.597 0.784 R: 5’-CGACCCGTCACCTAGGATCT-3’ 108 AMERICAN MALACOLOGICAL BULLETIN 28-1/2-2010 Table 2. Amplification of microsatellite loci in other species of Ventridens. The sym- bol + indicates that a product was amplified and verified by gel electrophoresis and fragment analysis. Locus 542 failed to amplify in all species examined. Species VL10 VL1A VL301 VL306 VL542 V. acerra (J. Lewis, 1870) + + + V. arcellus (Hubricht, 1976) + + + + V brittsi (Pilsbry, 1892) + + + + V. cerinoideus (Anthony, 1865) + V. collisella (Pilsbry, 1896) + + V. demissus (A. Binney, 1843) + + + + V. intertextus (A. Binney, 1841) + V. lawae (A. Binney, 1892) + + V volusiae (Pilsbry, 1900) -1- + + -1- cycles at each step and 20 cycles at 60 °C; the rest of the program was the same. PCR products were verified by gel electrophoresis on a 1.5% agarose gel with ethidium bromide. The forward primer for each locus was labeled with a fluorescent dye (Sigma Proligo, St. Louis, Missouri) to facilitate fragment analysis on a CEQ8000 Genetic Analysis System (Beckman Coulter, Fullerton, California). Statistical analysis The number of alleles (NJ, expected heterozygosity (He), observed heterozygosity (Ho) and allelic richness (R$) were calculated for each population and locus using GeneAlex6 (Peakall and Smouse 2006) following the methods of Hard and Clark (1997). An analysis of molecular variance (AMOVA) (Excoffier et al. 1992) was used to determine partitioning of genetic variation within the region, among populations, and among individuals within populations from microsatellite allele frequencies according to Weir and Cockerham (1984) using Arlequin ver. 3.01 (Excoffier et al. 2005). Population structure was also tested using STRUCTURE v2.1 (Pritchard etal. 2000) with a burn in period of 5,000 and 50,000 reiterations after burn in. STRUCTURE uses a Bayesian approach to assign individuals to clusters based on their genotypes while estimating population (cluster) allele frequencies (Pritchard etal. 2000). Global and pairwise Fsx (Weir and Cockerham 1984) values were calculated using FSTAT v2.9.3 (Goudet 2001) to measure population differentiation. Significance was determined using 1,000 permutations with Bonferroni multiple corrections at the 1/1,000 nominal level in FSTAT. To test for population-level deviations from Hardy-Weinberg equilibrium, Wright’s F,s was calculated for each population using the estimator 0 (Weir and Cockerham 1984) in GENETIX v4.05 (Belkhir et al. 2004). Significance of F[S was tested with 1,000 permutations. Isolation by distance was tested using a Mantel test of the association between genetic distance and the logarithm of geographic distance in the IBD program (Bohonak 2002). Genetic distance was calculated as Fsx / (1 - Fsx) and geographic distances (km) were based on GPS coordinates. The Mantel test was based on 10,000 permutations of spatial locations among the sample populations. RESULTS Population analysis Levels of polymorphism in each population varied with the average number of alleles ranging from 2.2 in the TU population to 11.6 in the PLN population (Table 3). Allelic richness (Rs), which accounts for sample size biases, showed little difference among the Potomac River populations with values ranging from 5.4210 to 6.2102. In contrast, Rs for the four landlocked populations ranged from 1.8704 to 3.9520. Average observed (H ) and expected (H ) heterozygosities for the Potomac River populations ranged from 0.4666 to 0.7810 and 0.7692 to 0.8496, respectively. Values for the landlocked populations ranged from 0.2466 to 0.3820 (Ho) and 0.2600 to 0.6774 (He). Nine of the 18 populations had significantly positive multilocus FIS values indicating an overall heterozygote deficiency. AMOVA results estimated the global Fsx for all 18 populations to be 0.091 (P < 0.0001) (Table 4). Analysis of all populations found that 70% of the variation was among the individuals of the populations whereas only 9.11% was between the populations. A separate analysis of the 14 Potomac River populations found that 79.33% of the variation was among individuals in contrast to 1.64% among the populations. Analysis of the four landlocked groups alone showed that 43.88% of the variation was among the individuals with 34.86% among the populations. Pairwise analysis of Fsx estimates between the Potomac River populations (Table 5), resulted in values ranging from -0.0003 to 0.0477. Pairwise analysis of the four landlocked populations with the Potomac River populations and each other resulted in Fsx estimates ranging from 0.0738 to 0.6004. Fsx values for several Potomac River populations were significant; however, no patterning was evident (i.e., island sites vs. mainland sites) whereas nearly every value for the four landlocked sites was significant. Analysis by STRUCTURE clearly delineated the samples into three clusters: the Potomac River sites, the Illinois sites (SIU, MC), and the Maryland sites (TU, MD) (data not shown). Based on the Mantel test, there was a significant association between the pairwise genetic and geographic POPULATION GENETICS OF VENTRIDENS LIGERA 109 Table 3. Intrapopulation genetic diversity at five microsatellite loci for 18 Ventridens ligera sampling sites. For each population I provide the sample size, average number of alleles (NJ, the observed heterozygosity (Ho), the expected heterozygosity (He), F|S, which indicates deviation from random mating, and allelic richness (Rs). * Indicates significant values. Population Sample size Na Ho He F,s R PIS 18 10.4 0.7056 0.8018 0.1490* 5.7526 PIM 19 9.6 0.5632 0.8004 0.3218 5.6158 PIN 19 9.8 0.6032 0.7842 0.2572 5.4210 PLS 19 10.0 0.6892 0.8232 0.1906* 5.7702 PLM 21 10.0 0.6704 0.8052 0.1914 5.6662 PLN 21 11.6 0.7810 0.8218 0.0740* 5.8826 PUP 11 9.0 0.6096 0.8232 0.2067 6.0056 PD 9 9.0 0.7684 0.8288 0.1367* 6.4610 VAP 9 6.8 0.6402 0.8092 0.2787* 5.6228 BMN 10 7.6 0.7200 0.8036 0.1568* 5.6386 BMS 8 7.6 0.6250 0.7672 0.2489* 5.8708 BIS 15 8.8 0.4666 0.8022 0.4463 5.5286 BIN 7 6.4 0.7428 0.7818 0.1261* 5.4942 VAB 18 10.6 0.5706 0.8496 0.3548 6.2102 SIU 10 2.8 0.3288 0.4678 0.3455 2.6040 MC 11 5.2 0.3510 0.6774 0.5194 3.9520 TU 10 2.2 0.2466 0.2600 0.1066* 1.8704 MD 11 3.8 0.3820 0.6768 0.4737 3.4042 distances between the sampled populations {R2 = 0.0599, P — 0.0281). The plot of Fsx / (1 - FST) against log (geographic distance) (Fig. 2) shows a statistically significant positive relationship between the geographic distance and genetic similarity of the Ventridens ligera populations sampled (P < 0.001, 10,000 permutations). DISCUSSION We analyzed genetic variation in land snail, Ventridens ligera, populations along the Potomac River to determine if the river acts as an effective barrier against gene flow. Analysis of five V. ligera specific microsatellite loci showed no partitioning of genetic variation between individuals on Bear Island, Plummers Island, or the adjacent riverbanks but did show strong partitioning of individuals among the four landlocked sites (SIU, MC, TU, and MD). These results suggest that V. ligera in the Potomac River populations experience higher levels of gene flow than the landlocked populations. The global Fsx estimate among the 18 populations was low (Fsx = 0.091; P < 0.0001) which suggests a lack of genetic partitioning and that gene flow is occurring between the sampled populations. However, analysis of the Potomac River populations and the landlocked sites separately revealed Fsx values of 0.016 (P not significant) and 0.350 (P < 0.0001), respectively. FSJ values >0.2 are considered to reflect population structuring (Beebee and Rowe 2008). The FST for the Potomac River populations only is much lower than the value when all populations or the landlocked populations only are evaluated, further supporting the suggestion that gene flow is occurring among these river populations. In contrast, the Fsx for the landlocked sites suggests that there is structuring between the four populations and there is no gene flow occurring. Analysis in STRUCTURE found no partitioning among the Potomac River populations; instead the results suggest that all 14 sites are part of one large homogenous population. Pairwise Fsx values were very low (-0.0003 to 0.0483) among the Potomac River populations while values for the landlocked sites ranged from 0.0738 to 0.6004 with nearly every value significant. These results also indicated that the four landlocked populations were significantly different not only from the Potomac River populations but also from each other, while the Potomac Rivers populations are highly similar to each other. A Mantel test showed a significant positive correlation between genetic distance and geographic distance. Over short distances (< 1 km) there is a linear pattern of isolation that is expected as gene flow homogenizes the populations. The pairwise comparisons that fall within this range are those on the same islands ( i.e ., Plummers Island to Plummers Island, Bear Island to Bear Island). The more distant comparisons (>2 km; i.e., Bear Island to Plummers Island) display a significant level of isolation by distance most likely due to a limited gene flow and genetic drift. 110 AMERICAN MALACOLOGICAL BULLETIN 28-1/2*2010 Table 4. Analysis of molecular variance (AMOVA) describing the partitioning of genetic variation for 18 Ventridens ligera sampling sites. All samples were analyzed then reanalyzed in two groups: Po- tomac River populations and the landlocked populations. Source - All 18 populations d.f Estimated variance Percentage tD H Fsx q U Among populations 17 0.154 9.11 V 3 Among individuals Oh Within populations 228 0.352 20.77 U SIU Within individuals 246 1.189 70.12 Total 491 1.696 100 xS P5 Global Fst = 0.091, P < 0.0001 y > Source - Potomac River only 7C 2 Fst 3 Among populations 13 0.027 1.64 'J 'B Among individuals .B c/3 3 Within populations 190 0.317 19.03 Within individuals 204 1.324 79.33 > A Total 407 1.668 100 P3 Global Fst = 0.016, P = 0.934 ± 0.006 JO 2 Source - 4 landlocked sites 22 3 CQ Fst f Among populations 3 0.586 34.86 2 CC < Among individuals S > Within populations 38 0.358 21.26 > Within individuals 42 0.738 43.88 C o '-a Q a, Total 83 1.682 100 & Global F = 0.3486, P < 0.0001 The Potomac River Basin floods frequently and may act as a catalyst for unidirectional gene flow. It is highly possible that Ventridens ligera from upstream are “rafting” on debris downstream during both normal and flooding periods. Chiappero et al. (1997) reported this phenomenon in the sigmodontine rodent Oligoryzomys flavescens. The authors reported a downstream flow of genes over 250 km of the Parana River in Argentina with the highest heterozygosity found at the site furthest upstream. They suggest that the animals may be passively transported downstream on raffs of floating plants. This method of gene flow for V. ligera is certainly possible, and a larger study to further elucidate gene flow is planned with sampling from populations near the source of the Potomac River, at the mouth of the river where it enters the Chesapeake Bay, and at sites along the length of the river. While I did identify upstream sites with higher '■O in to on fo oooooooo O H VO V© »-H O Os rTi Os © nr>min hONONNONNcoNincooNO\om mmnninini(N(NnjfSfnrjn|M dddddddddddddd O OO NO NO m On O rd O -H OOOOOOOOOOOOO ddddddddddddd — (N -- o o o o m »— » d> d> 5 q q q q q o d d d d d q q q q q d d d d d o o o o d d d d q q q odd co 2 £ P-* cu Qh OhPhPhDh^>WCQCQCQ> < 2 POPULATION GENETICS OF VENTRIDENS LIGERA Figure 2. Genetic distance vs. log( geographic distance) in Ventridens ligera populations along the Potomac River. Genetic distance is shown as FST / (1 - FST). Circles refer to all possible pairwise com- parisons among the Potomac River populations. heterozygosity than downstream sites, my findings were not consistent. I suggest this is potentially due to the small distance analyzed, and that the study spanning the entire Potomac River will show higher heterozygosities upstream near the source. These are rather small land snails and the cost of locomotion across even several meters would be extremely high. We collected samples along an 8.8 km stretch of the Potomac River and it is highly doubtful that V. ligera are traveling this distance on land. Land snails typically migrate only a few meters per year (Pfenninger et al. 1996, Arnaud et al. 1999b). However, passive displacement (i.e., other animals, water) could potentially move individuals a much larger distance (Dorge et al. 1999). Previous land snail studies have shown that land snails live in neighborhoods where populations are genetically distinct from one another ( Selander and Kaufman 1975, Pfenninger etal. 1996,Arter 1990, Johnson and Black 1995, Arnaud et al. 2001). Pfenninger et al. (1996) used RAPD analysis to determine that the land snail Trochoidea geyeri (Soos, 1926) lived in genetically distinct neighborhoods of 13-21 m2. Arnaud et al. (2001) reported significant genetic structuring between neighborhoods of Helix aspersa only 40 m apart using microsatellites. Both of these studies focused on populations located in habitats without moving water. Here I report the first evidence of gene flow among land snails in the Potomac River Basin. My findings suggest that the river is not an effective barrier to gene flow in the land snail Ventridens ligera. Evidence of isolation by distance between the populations on two separate islands is present; however, because the overall FSJ values are close to zero, I suggest that the river is actually facilitating gene flow. ACKNOWLEDGMENTS Funding for this study was provided by the Washington Biologists Field Club, the Maryland/District of Columbia Chapter of the Nature Conservancy, and the Faculty Devel- opment and Research Committee of Towson University, Towson, Maryland. The author would like to thank the Chesapeake & Ohio Canal National Historic Park and the George Washington Memorial Parkway for collection permits. The author would like to thank Butch Norden for his help getting the project off the ground, Michael Lloyd for his assistance with the statistical analysis, and Roland Roberts and his laboratory for their assistance with microsatellite development. The author would also like to thank Jochen Gerber of the Field Museum of Natural History, Liz Shea of the Delaware Museum of Natural History, Tim Pearce of the Carnegie Museum of Natural History, and John Slapcinsky of the Florida Museum of Natural History for providing additional Ventridens species for the study. The author would like to acknowledge the Towson University undergraduate students whose help was crucial to the success of this project, Rebecca McConnell, Oluwarotimi Folorunso, and Tony Basel. LITERATURE CITED Allendorf, F. W. and G. Luikart. 2007. 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Genetica 101: 105-113. Crozier, R. H. 1997. Preserving the information content of species: Genetic diversity, phylogeny and conservation worth. Annual Review of Ecology and Systematics 28: 243-268. Depraz, A., E. Rathey, and J. Hausser. 2008. Characterization of 13 polymorphic microsatellite loci for two land snail species, Trochulus villosus and T. sericeus (Gastropoda: Pulmonata: Hy- gromiidae). Molecular Ecology Resources 8: 704-706. Dorge, N., C. Walther, B. Beinlich, and H. Plachter. 1999. The significance of passive transport for dispersal in terrestrial snails (Gastropoda, Pulmonata). Zeitschrift fur Okologie und Naturschutz 8: 1-10. Excoffier, L., P. Smouse, and J. Quattro. 1992. Analysis of molecu- lar variance inferred from metric distances among DNA hap- lotypes: Application to human mitochondrial DNA restriction data. Genetics 131: 479-491. Excoffier, L„ G. Laval, and S. Schneider. 2005. 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Effects of population dynamics on genetics in mosaic landscapes. In: L. Hansson, L. Fahrig, and G. Merriam, eds., Mosaic Landscapes and Ecological Processes. Chapman 8c Hall, London. Pp. 178-198. Miller, M. P„ L. E. Stevens, J. D. Busch, J. A. Sorensen, and P. Keim. 2000. Amplified fragment length polymorphism and mito- chondrial sequence data detect genetic differentiation and relationships in endangered southwestern U.S.A. ambersnails [Oxyloma spp.). Canadian Journal of Zoology 78: 1845-1854. Peakall, R. and P. E. Smouse. 2006. GENALEX6: Genetic analysis in Excel. Population genetic software for teaching and research. Molecular Ecology Notes 6: 288-295. Pfenninger, M., A. Bahl, and B. Streit. 1996. Isolation by distance in a population of a small land snail Trochoidea geyeri: Evidence from direct and indirect methods. Proceedings of the Royal Soci- ety of London (B) 263: 1211-1217. Pritchard, J. K., M. Stephens, and P. Donnelly. 2000. Inference of population structure using multilocus genotype data. Genetics 155: 945-959. Rozen, S. and H. J. Skaletsky. 2000. Primer3 on the WWW for gen- eral users and for biologist programmers. In: S. Krawetz and S. Misener, eds., Bioinformatics Methods and Protocols: Meth- ods in Molecular Biology. Humana Press, Totowa, New Jersey. Pp. 365-386. Saccheri, I., M. Kuussaari, M. Kankare, P. Vikman, W. Fortelius, and I. Hanski. 1998. Inbreeding and extinction in a butterfly metapopulation. Nature 392: 491-494. Schweiger, O., M. Frenzel, and W. Durka. 2004. Spatial genetic structure in a metapopulation of the land snail Cepaea nemora- lis (Gastropoda: Helicidae). Molecular Ecology 13: 3645-3655. Selander, R. K. and D. W. Kaufman. 1975. Genetic population struc- ture and breeding systems. Isozymes 4: 145-167. Selander, R. K. and H. Ochman. 1983. The genetic structure of pop- ulations as illustrated by mollusks. Isozymes 10: 93-123. Weir, B. S. and C. C. Cockerham. 1984. Estimating E-statistics for the analysis of population structure. Evolution 38: 1358-1370. Wirth, T. 2000. Isolation and characterization of microsatellite loci in the land snail Helicella itala, and cross-species amplification within the family Helicidae. Molecular Ecology 9: 501-502. Submitted: 8 January 2009; accepted: 29 June 2009; final revisions received: 6 October 2009 Amer. Malac. Bull. 28: 1 13-120 (2010) Reproductive biology and annual population cycle of Oxyloma retusum (Pulmonata: Succineidae)* Aydin Orstan Section of Mollusks, Carnegie Museum of Natural History, 4400 Forbes Ave., Pittsburgh, Pennsylvania 15213-4080, U.S.A. Corresponding author: pulmonate@earthlink.net Abstract: I studied the reproduction and the population turnover of a succineid land snail living by a small lake in central Maryland. The identity of the snail, deduced from its external characteristics and the genitalia, comes closest to Oxyloma retusum (Lea, 1834). The species has a semelparous life cycle. The snails that survive the winter grow and reproduce from late March until the end of June when they reach their maximum size and die off. Their offspring (the spring generation) grow throughout the spring and the summer and reproduce briefly near the end of August. In the fall, the survivors from the spring generation and their offspring hibernate from November until the end of March. Snails mate by shell-mounting. In 89% of pairs, mating was anatomically reciprocal. During courtship, one snail climbs on the shell of a prospective mate and circles the shell to initiate mating. In mating pairs with a shell length difference of more than 1 mm, the smaller snail was always on top. This suggests that one function of shell-circling during courtship may be to help the top snail judge its potential partner’s size. Key words: land snail, genitalia, semelparity, mating, courtship Oxyloma retusum (Lea, 1834) is one of the most common and widely distributed succineid species of the northeastern U.S. (Hubricht 1985). It is found in the vicinity of lentic and lotic habitats, usually very close to the water (Shimek 1935, Baker 1939, Miles 1958, Hubricht 1985, Lannoo and Bovbjerg 1985). The reproductive organs (Pilsbry 1948, Miles 1958, Franzen 1963, Wu 1993), mating anatomy (Webb 1977b), dispersion (Lannoo and Bovbjerg 1985), and diet (Shrader 1972, Orstan 2006) of O. retusum have been studied. However, no published information is available on the life cycle of Oxyloma retusum. Mostly anecdotal information in the literature suggests that the continental North American succineids have semelparous life cycles. For example, Pilsbry (1948) stated that Novisuccinea ovalis (Say, 1817) often reached “ordinary” size in one season and Grimm (1971) noted that in Maryland, O. subeffusa Pilsbry, 1948 matured in early spring and died off before the summer. Strandine ( 1941 ) showed that N. ovalis indeed had an annual life cycle in Illinois. The only relevant published information regarding the life cycle of O. retusum is Pilsbry’s (1948) comment that probably most of the large individuals of Oxyloma decampi gouldi (Pilsbry, 1948; synonymized with O. retusum by Hubricht 1985) died by the end of the summer and that he could find only half-grown shells in the fall. Besides Webb’s (1977b) work on the mating anatomy of Oxyloma retusum , no other research on the reproductive biology of the species has been published. Furthermore, Webb’s (1977b) brief accounts of courtship and whether or not penis intromission was reciprocal seem incomplete and need to be supplemented with more thorough observations. Pulmonate gastropods have evolved a wide spectrum of lifestyles. A better understanding of the biology of any one species will contribute to the solution of the puzzle of pulmonate evolution. Furthermore, information obtained with common species such as Oxyloma retusum could be useful for the development of more effective conservation strategies for less common and endangered succineids, for example, Oxyloma haydeni kanabensis Pilsbry, 1948 (Spamer and Bogan 1997-1998). In this paper, I present the results of my studies on the mating biology and the annual population cycle of O. retusum in Maryland. MATERIALS AND METHODS The study took place along the shore of Lake Churchill, a small (-640 m by -200 m) lake (39.188°N, 77.280°W) in Germantown, Maryland. Besides a small forest remnant along its eastern shore, the lake is surrounded by a residential neighborhood. The lake is connected by pipes to the nearby, larger Little Seneca Lake in Black Hill Regional Park. Both lakes occupy former agricultural fields and were filled in the 1980s by the damming of Little Seneca Creek that runs through the park. * From the “Leslie Hubricht Memorial Symposium on Terrestrial Gastropods” presented at the meeting of the American Malacological Society, held from 29 July to 3 August 2008 in Carbondale, Illinois. 113 114 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 During preliminary field trips, I found Oxyloma retusum within stands of cattails ( Typha sp.) at several points along the lake. I selected as the study locations two cattail stands, one at the western shore of the lake and the other about 580 m to the east near the northeast corner of the lake. From April through October both stations were covered with dense growths of cattails, various herbaceous plants, and grasses. The shore of the lake was always muddy and had a layer of decomposing leaves. During the winters the lake often froze. During the monitoring of the sizes of the snails at approx. 2-month intervals from early April to late October, I collected all the live Oxyloma retusum I could find at either one or both of the stations until I had a total of at least 40 snails. The collections were usually done over two days. The snails were brought to the laboratory and the shell length (height) of each snail was measured with calipers. Especially during the spring, the edges of the aperture lips of juvenile snails were thin and fragile and easily crushed by the caliper jaws. Con- sequently, accurate measurements of shell lengths were diffi- cult and the histograms were constructed using classes 1 mm wide. The permanent removal of large numbers of live snails, especially from a small population, may modify population dynamics (Heller 2001). Since I did not know the total popu- lation sizes of Oxyloma retusum at the study locations, to avoid interfering with the natural population cycles, I returned the live snails to their respective stations within a few days, occasionally keeping only a few of the largest snails for dissection. Those were killed in ~5% ethanol and saved in 70% ethanol. To study their mating, I brought snails to the laboratory in individual containers and kept them isolated from each other except during periods of observation. I kept the snails in small, numbered plastic containers lined with damp toilet paper and fed them dead cattails, tree leaves, and carrots. Although some young snails were kept up to 14 days before they were returned to the field, they did not grow much in captivity. To induce snails to mate, I placed them in pairs or in groups of three or four in a small container or on a large glass plate held horizontally by a clamp. The advantage of the latter method was that the plate could be rotated, permitting exam- ination and photography of a mating pair from different angles. If the plate was rotated slowly, snails remained attached to it even when upside down. I sprayed water on the plate occasionally to prevent the snails from drying. To determine if mating was anatomically reciprocal, that is, if each snail in a pair was inserting its penis into its partner’s vagina simultaneously, I developed a forced withdrawal method: I gently picked up a pair that had been conjoined for at least about 15 minutes, pulled them apart slightly to dis- rupt their mating, and then waited for the snails to complete withdrawing their penises while watching them under a dissecting microscope. I have deposited two alcohol-preserved specimens of Oxyloma retusum in the Field Museum of Natural History, Chicago, Illinois (FMNH 312423, FMNH 312424). Addition- ally, I have deposited four alcohol-preserved specimens in the Carnegie Museum of Natural History, Pittsburgh, Pennsyl- vania (CM93115, CM93116, CM96781, CM96782), including one specimen with an everted penis (CM93115) and one specimen in 95% ethanol (CM96781). RESULTS Identification External characteristics The mantle, especially in the front and along the sides, was densely stippled with small, black spots. But the spots did not form any distinct patterns (Figs. 1-2). The head was light brown. The head and the sides of the foot were also stippled with dark spots. The spots coalesced into longitudinal bands in front of the head between the tentacles (Fig. 3) and formed splotches along the sides of the foot (Fig 2). Genitalia I dissected five snails collected in June 2001, June 2002, June 2003, and May 2008. The general characteristics of the genitalia (Fig. 4) agreed in all specimens. The penis and the epiphallus were encased within a translucent sheath. The vas Figure 1. A mating pair of Oxyloma retusum. The larger snail’s shell has a varix (arrow) across its body whorl. REPRODUCTION AND POPULATION CYCLE OL OXYLOMA RETUSUM 115 the epiphallus (Pilsbry 1948), was present in all specimens; its approximate length varied from 0.3 to 0.5 mm (N = 5). The ratio of the combined lengths of the penis and the epiphallus to the length of the vagina varied from 1.2 to 1.5 (AT = 5); the smallest ratio was obtained with a specimen in which both the penis and the epiphallus were coiled, while the largest ratio was obtained with a specimen in which neither the penis nor the epiphallus was coiled. The bursa copulatrix had a bulbous bursa (Pig. 4A). The ratio of the combined lengths of the peduncle and the bursa of the bursa copulatrix to that of the penis and the epiphallus varied from 1.1 to 1.6 ( N = 3). Along retractor muscle originating from the back of the body cavity forked near the vagina (Pig. 4A); one arm of it passed between the penis and the vagina and fused into the right upper tentacle, while the other arm, which was loosely connected to the base of the penis as it passed by it, inserted into the body wall near and to the left of the right upper tentacle. Live of the snails placed in ~5% ethanol died with their penises everted. The distal ends of four of the penises narrowed down to a finger-like tip that was probably the everted appendix (Pig. 4B). The fifth penis had a blunt end; what appeared to be the uneverted appendix was visible inside the penis. The epiphallus was visible through the translucent wall of each penis and its opening was subterminal when the appendix was everted (Fig. 4B). The surfaces of the everted penises were covered with very small, crystalline, scattered granules. The mean length of four penises, including their everted appendices was 3.88 mm ( SD = 0.27 mm). The mean length of the four appendices was 0.55 mm (SD = 0.17 mm). One specimen (CM93115) that had mated shortly before it was killed had an additional 0.78-mm long, yellow appendage loosely attached to the surface of its penis (ps in Fig. 4B). The everted penis of its mate that was also killed lacked that appendage. The external characteristics and the genitalia of the snails agreed with the literature descriptions of Oxyloma retusum (Pilsbry 1948, Miles 1958, Franzen 1963, Wu 1993). Mating The only time I saw mating Oxyloma retusum in the field was in May 2008. However, mating pairs were observed many times among the snails brought to the laboratory between late March and early September. The earliest matings took place on 24 March 2006 among the snails collected on the same day, while the latest matings were obs- erved on 1 September 2002 among the snails collected on that day. None of the snails collected on 13 September 2003, deferens entered the sheath near the retractor muscle (Fig. 4A). The sheath around the epiphallus and the surface of vas deferens, before it entered the sheath, were stippled with black dots. Within the sheath, the epiphalluses of all but one specimen and the penises of two specimens were coiled. An appendix, traditionally taken to be demarcating the penis and Figure 3. Courtship behavior preceding the mating of Oxyloma retusum. A, the smaller active snail is climbing onto the shell of the larger passive snail; B, the active snail is turning around the apex of the passive snail and moving towards the latter’s head; C, having failed to initiate mating, the active snail is crawling across the head of the passive snail to repeat its cycle. Figure 2. A pair of Oxyloma retusum that flipped over the edge of a glass plate after they started mating. The snail that was on top is now hanging below. Arrow is pointing at the junction of the genital openings. 116 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 4. Lower genitalia of Oxyloma retusum. A, genitalia removed from a snail with the penis and the epiphallus still within the sheath; B, everted penis of a snail killed soon after mating. Abbreviations: a, appendix; be, bursa copulatrix; e, epiphallus; oe, opening of epiphallus; p, penis; rp, retractor of penis; rt, retractor of tentacle; ps, possible paraspermatophore; v, vagina; vd, vas deferens. Therefore, I assumed that in those two pairs mating was anatomically unilateral. On one occasion, I photographed two pairs mating simultaneously on top of each other. The examinations of the snails during mating and of their photographs later indicated that each snail was mating with only one other snail. I timed the uninterrupted matings of 11 pairs at 19-21 °C. The median mating duration was 79 min (mean = 81 min) with a range of 63 to 97 min. At the completion of mating, the snails separated, the top snail climbed down and the snails crawled away from each other. 25-26 October 2003, and 23 October 2004 mated in the laboratory. Courtship (the sequential behavioral steps prior to mating) started when an active snail climbed on the shell of a passive snail (Fig. 3A). If the snail on top started out on the left side of the bottom snail, it crawled around the apex and then along the right side of the shell of the bottom snail towards its head (Fig. 3B). After the top snail reached the head of the bottom snail, either the snails started copulating or the top snail continued to crawl, first across the head of the bottom snail (Fig. 3C) and then along the left side of its shell towards the back. The top snail then crawled around the apex of the shell of the bottom snail one more time and back to its head to attempt to initiate mating again. The snails that could not initiate copulation were observed to circle their partners’ shells two or three times before climbing down and leaving. In one case, an active snail took approx. 10 minutes to circle its partner’s shell almost three times before they started mating. A pair started copulating when the top snail twisted its head and brought its genital opening against that of the bottom snail (Fig. 1). After a pair started mating, the bottom snail stopped crawling and the snails remained motionless for the rest of their mating. During mating, the snails’ tentacles were partially or completely withdrawn (Figs. 1-2). Once the two snails’ genital openings were closely pressed against each other, no external parts of their genitalia were visible and I could not ascertain if mating was anatomically reciprocal by looking at the close junction of the genital openings, even under magnification (Fig. 2). Nineteen pairs of snails were separated during mating and forced to withdraw their penises. In 17 pairs, mating was anatomically reciprocal (Fig. 5). In the remaining two pairs, I could see only the penis of the top snail being withdrawn. Significance of shell size Casual observations indicated that the smaller snail in a mating pair was usually on top (Figs. 1 and 3). To support those observations, I measured the shell length and noted the position of each snail in 40 mating pairs (Table 1). When the shell length difference between the snails in a pair was 1 mm or less, the respective positions of the snails in each pair were not different than a random distribution (Chi-square test, %2 = 1.8, P = 0.180). But when the shell length difference between the snails in a pair was more than 1 mm, the positioning of the smaller snail on top in all cases was significantly different than a random distribution (/2 = 20, P < 0.0001). Oviposition I watched two snails while they were laying eggs in the laboratory. Prior to oviposition, a snail’s head-foot outside its shell became swollen; the snail stopped moving and appeared moribund. One snail took approx. 41 minutes to lay 22 eggs. Five clutches from captive snails had 4 to 22 eggs. The eggs did not have hard shells but were enclosed inside a jelly matrix. Annual population cycle From April 2002 through March 2006, live Oxyloma retusum were abundant at both stations from early spring until the end of October. Snails were either on cattail stalks or on muddy areas next to the water. However, during the final season of field work in April and May 2008, casual observations indicated that the populations at both stations had decreased significantly although no physical alterations of the landscape were evident. In contrast to their visibility and abundance from late March through October, the snails were rare and difficult to locate during the rest of the year. During the nine searches carried out between early November and early REPRODUCTION AND POPULATION CYCLE OF OXYLOMA RETUSUM 117 disappeared by August and were presumed dead (Fig. 6). The smallest mating snail observed had a shell length of 6.2 mm. Therefore, I took 6 mm to be the arbitrary minimum shell length for sexually mature snails. Snails with shells 6 mm or longer made up more than 80% of the population in April and August (Fig. 6). DISCUSSION Figure 5. Forced withdrawal of the penises of a mating pair of Oxyloma retusum. A, the penises of both snails are still inserted into each other’s vaginas; B, the snail on the right (bottom) has withdrawn its penis (right arrow), while the penis of the snail on the left (top) is still inserted (left arrow); C, the snail on the left has withdrawn its penis. Table 1. Positioning of snails in mating pairs according to their shell sizes. There were 20 pairs in each size group. Shell size difference Smaller snail on top Larger snail on (mm) (# of pairs) top (# of pairs) <1 13 7 >1 20 0 March from 2002 through 2008 lasting a total of about 3 hours, I found only five live Oxyloma retusum. Active snails between spring and early fall seemed incapable of withdrawing fully into their shells even when they were handled repeatedly. In contrast, all of the snails found in the winter were fully withdrawn into their shells and dormant. The edges of their apertures were loosely attached to dead plant material, such as tree or cattail leaves. The shell lengths of these five snails were as follows; 6.6 and 7.6 mm ( 10 November 2006), 4.2 mm (27 December 2005), 5.4 and 5.7 mm (12 January 2002). The shells of the snails collected early in the spring usually had a flange-like ridge across their body whorls (Fig. 1), indicating that the aperture, normally thin and fragile in growing snails, had thickened prior to winter dormancy and formed what is referred to as a varix (Moore 1952). Out of a total of 57 snails collected on 4-5 April 2003, 38 had a prominent varix across their body whorls, while the rest had a less distinct varix. The frequency distributions of the shell lengths from April through October for 2003 and 2004 are given in Fig. 6. The snails attained their largest shell sizes near the end of June. The largest snail encountered, on 13 June 2004, had a shell length of 14.1 mm. All of the snails larger than ~11 mm Some uncertainty remains about the identity of the snails despite the anatomical similarities between them and the published descriptions of Oxyloma retusum (Pilsbry 1948, Miles 1958, Franzen 1963, Wu 1993). The primary reason for this uncertainty is that prior to Pilsbry (1948), species descriptions were based exclusively on shells, which not only lack distinct and species-specific features but also are variable (Franzen 1963). The subsequent anatomical studies have shown variability also of the reproductive organs of snails identified by different authors as O. retusum (Miles 1958, Franzen 1963, Wu 1993). One characteristic of O. retusum is its coiled epiphallus (Pilsbry 1948, Franzen 1963); however, both Miles (1958) and Wu (1993) noted that in some specimens the epiphallus was straight. One of the five snails I dissected also had a straight epiphallus. Likewise, the penis may also be straight or coiled (Miles 1958, Franzen 1963). Coiled penises were present in two of the five snails I dissected. Franzen’s (1963) recommendation that anatomies of all described Oxyloma species be studied and compared with each other before we can determine which species are valid remains in effect today more than 45 years later. The shell-mounted mating position of Oxyloma retusum is the same as those of other succineid species (Hecker 1965, Webb 1977b, Villalobos etal. 1995, Rundell and Cowie 2003). Webb (1977b) did not mention the circling of an active snail around the shell of its passive prospective mate that I observed during the courtship of O. retusum. Hecker (1965) reported a similar courtship behavior prior to mating of Succinea putris (Linnaeus, 1758) and suggested that the active snail’s circling may stimulate the passive snail. However, as Hecker (1965) also noted, courtship does not always lead to copulation; I observed pairs of O. retusum separate after the active snail had circled its partner’s shell two or three times. Reverse-size-assortive mating observed in Oxyloma retusum (Table 1) has been noted in other succineids (Jackiewicz 1980). Jordaens etal. (2005) also showed that the majority of the mating pairs of Succinea putris consisted of a 118 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 SHELL LENGTH (MM) Figure 6. Frequency distributions of the shell lengths of live Oxyloma retusum from April through October for 2003 and 2004. Shell lengths are grouped in 1-mm classes. small active snail and a large passive snail. Their explanation for this behavior, that it is easier for a larger snail to carry a smaller one than vice versa, especially on vertical surfaces, makes sense for several pairs of O. retusum that I observed mating vertically (Fig. 1) or upside down (Fig. 2). Mating between snails of unequal sizes requires that the active snail be able to judge, relative to its own size, the size of its prospective partner. Therefore, shell-circling during courtship, in addition to its possible stimulatory action, may also help an active snail compare its size with that of its partner. Webb (1977b) gave a mean mating duration of 67 min with a range of 37 to 108 min for Oxyloma retusum at an unspecified temperature. In comparison, the mean mating duration measured in this study was 81 min (median, 79 min) with a range of 63 to 97 min at 19-21 °C. Hecker (1965) compared the copulation times of Succinea putris , Succinella oblonga (Draparnaud, 1801), O. elegans (Risso, 1826), and O. sarsii (Esmark, 1886). The copulations of S. putris lasted up to 8 hours, while the mean copulation times of the other three species were approx. 90 min (Hecker 1965, fig. 15). Jordaens et al. (2005) reported the median mating duration of S. putris at 20 °C as 487 min with a range of 230 to 725 min. The biological significance of the much longer copulation time of S. putris than those of other succineid species is unclear. Eighty-nine percent of the interrupted matings of Oxyloma retusum were anatomically reciprocal. This agrees with Webb’s (1977b) statement that mating between O. retusum was “mostly reciprocal,” although, it is not clear from Webb’s account how many pairs of snails he had examined. As Jordaens et al. (2005) and Davison and Mordan (2007) noted, anatomical reciprocity during mating does not necessarily imply reciprocal sperm transfer. Nevertheless, Jordaens et al. (2005) showed that in Succinea putris, whose matings were also anatomically reciprocal, 86% of matings involved reciprocal sperm exchange. Therefore, they stressed that the active snail on top cannot necessarily be considered to be the “male” and the passive snail on the bottom the “female.” Their findings, if extrapolated to my results, strongly suggest that in O. retusum reciprocal penis insertion also results in reciprocal sperm exchange and that each snail in an anatomically reciprocal pair performs both as a male and a female. In contrast, Brown et al. (2003, 2006) noted that during the matings of the Hawaiian Succinea thaanumi Ancey, 1899 and S. newcombiana Garrett, 1857 the snails on top were “acting as the male,” which, presumably, meant that the sperm exchange was unilateral. Brown etal. (2003) also reported one case of three individuals of S. thaanumi mating simultaneously with the middle snail “acting as both a male and a female.” The only case of simultaneous mating of more than two individuals of Oxyloma retusum that I observed involved four snails on top of each other, but each snail mated with only one other snail. The anatomy of the everted penis of Oxyloma retusum ( Fig. 4B ) , including the subterminal opening of the epiphallus, agrees with Webb’s ( 1977b) descriptions. The identity and the function of the granular, crystal-like formations on the surfaces of everted penises of O. retusum, also noted by Webb REPRODUCTION AND POPULATION CYCLE OF OXYLOMA RETUSUM 119 (1977b), are unknown. Webb (1977a) proposed the term paraspermatophore for the “softer, non-preformed” masses of semen of succineids. He described the paraspermatophore of Catinella avara (Say, 1824) as a “short cylindrical spindle” that adhered to the everted penis and noted that the semen of O. retusum was likewise discharged as a paraspermatophore. Jackiewicz (1980) also noted that the spermatozoa of the European succineids formed a “compact mass” in the genitalia of preserved specimens and Jordaens et al. (2005) described the sperm package of S. putris as “cohesive but plastic.” These descriptions indicate that in succineids the discharged semen forms a recognizable mass that may indeed be conveniently referred to as the paraspermatophore. It is likely that the appendage attached to the everted penis of one of the O. retusum killed after copulation is a paraspermatophore (ps in Fig. 4B). The resemblance of the overall shape of the putative paraspermatophore to that of the appendix (a in Fig. 4B) suggests that the paraspermatophore was formed inside the tip of the everted penis as Webb (1977a) speculated. The comparison of the frequency distributions of the shell sizes of Oxyloma retusum measured at the end of October 2003 and early in April 2004 (Fig. 6) indicates that very little, if any, shell growth takes place during the winter. The increase in the proportion of larger shells in April probably results from the rapid growth that starts in late March. This conclusion is supported by the observations that the five snails found during the November-February period were dormant and that their shell lengths (4.2-7.6 mm) were within the ranges of the October measurements. Furthermore, the presence of a varix across the body whorls of the shells of most of the snails in early spring indicates that growth stops during the winter. The snails that survive the winter become sexually mature (shell length >6 mm) by the end of March and early April when they make up more than 80% of the population (Fig. 6). The snails continue to grow throughout the spring, reaching maximum shell sizes near the end of June when they start to die off. None of the winter-survivors seem to reach August. Their offspring (the spring generation) grows during the spring and the summer and apparently goes through a brief reproductive period near the end of August, giving rise to a summer generation. This is implied by the increase in the proportion of sexually mature snails at the end of summer and indicated by the appearance of very small (shell length <4 mm) snails in August and early September (Fig. 6). At the same time, the shifts towards smaller sizes observed in the histograms for September and October indicate that some of the snails making up the spring generation, presumably those that reproduce at the end of the summer, die during that period. In other words, the population goes through a partial turnover near the end of the summer and the survivors from the spring generation are joined by the summer generation to create the group that goes through the winter dormancy to become the following spring’s reproducing adults. Pilsbry’s (1948) comments regarding Oxyloma decampi gouldi (synonymized with O. retusum by Hubricht 1985), that he could find only half-grown shells in autumn and that most of the large individuals probably died by the end of summer, accord with the life cycle of O. retusum presented here. Strandine (1941) studied the life cycle of Novisuccinea ovalis in Illinois during one year and interpreted his data as showing that N. ovalis hibernated during the winter, reproduced the following spring, and died off by the early summer. Therefore, the annual life cycle of N. ovalis is also similar to that of O. retusum. The data of Brown et al. (2003, 2006) show that growth, and to a lesser extent, reproduction are continuous throughout the year in the two species of semelparous Hawaiian succineids they studied. Likewise, Villalobos and Monge-Najera (2004) noted that Succinea costaricana von Martens, 1898, also a semelparous species, had continuous growth and reproduction throughout the year in Costa Rica. In contrast, Oxyloma retusum, undoubtedly because of the much colder winters in Maryland than in southerly climates, goes through a dormant period between November and early March. Nevertheless, the northern and southern American succineid species all seem to have semelparous life cycles, despite the differences in their habitats and the climates they experience. LITERATURE CITED Baker, F. C. 1939. Fieldbook of Illinois Land Snails. Illinois Natural History Survey, Manual 2, Urbana, Illinois. Brown, S. G., B. K. Spain, and K. Crowell. 2003. A field study of the life history of an endemic Hawaiian succineid land snail. Mala- cologia 45: 175-178. Brown, S. G., J. M. Spain, and M. Arizumi. 2006. A field study of the life history of the endemic Hawaiian snail Succinea newcombi- nana. Malacologia 48: 295-298. Davison, A. and P. Mordan. 2007. A literature database on the mating behavior of stylommatophoran land snails and slugs. American Malacological Bulletin 23: 173-181. Franzen, D. S. 1963. Variations in the anatomy of the succineid gas- tropod, Oxyloma retusa. The Nautilus 76: 82-95. Grimm, F. W. 1971. Annotated checklist of the land snails of Maryland and the District of Columbia. Sterkiana 41: 51-57. Heller, J. 2001. Life history strategies. In: G. M. Barker, ed„ The Biology of Terrestrial Molluscs. CABI Publishing, New York. Pp. 413-445. Hecker, U. 1965. Zur Kenntnis der mitteleuropaischen Bernstein- schnecken (Succineidae). I. Archiv fur Molluskenkunde 94: 1-45 [In German]. Hubricht, L. 1985. The distributions of the native land mollusks of the eastern United States. Fieldiana 24: 1-191. 120 AMERICAN MALACOLOGICAL BULLETIN 28*1/2-2010 Jackiewicz, M. 1980. Some observations on biology of reproduction of Succinea Draparnaud (Gastropoda, Pulmonata). Annales Zoologici 35: 65-73. Jordaens, K., J. Pinceel, and T. Backeljau. 2005. Mate choice in the hermaphroditic land snail Succinea putris (Stylommatophora: Succineidae). Animal Behaviour 70: 329-337. Lannoo, M. J. and R. V. Bovbjerg. 1985. Distribution, dispersion, and behavioral ecology of the land snail Oxyloma retusa (Suc- cineidae). Proceedings of Iowa Academy of Science 92: 67-69. Miles, C. D. 1958. The family Succineidae (Gastropoda: Pulmo- nata) in Kansas. The University of Kansas Science Bulletin 38: 1499-1543. Moore, R. C. 1952. Gastropods. In: R. C. Moore, C. G. Lalicker, and A. G. Fischer, eds., Invertebrate Fossils. McGraw-Hill, New York. Pp. 276-334. Orstan, A. 2006. Natural diet of Oxyloma retusa (Pulmonata: Suc- cineidae). Triton 13: 36-37. Pilsbry, H. 1948. Land Mollusca of North America (North of Mexico), Vol. 2, part 2. The Academy of Natural Sciences of Philadel- phia, Philadelphia, Pennsylvania. Rundell, R. J. and R. H. Cowie. 2003. Growth and reproduction in Hawaiian succineid land snails. Journal ofMolluscan Studies 69: 288-289. Shrader, A. L. 1972. Feeding behavior of three species of succineid snails. Malacological Review 5: 12-13. Shimek, B. 1935. The habitats of Iowa succineas. The Nautilus 49: 6-10. Spamer, E. E. and A. E. Bogan. 1997-1998. Contrasting objectives in en- vironmental mediation, reconnaissance biology, and endangered species protection - a case study in the Kanab ambersnail, Oxylo- ma haydeni kanabensis Pilsbry, 1948 (Gastropoda: Stylommato- phora: Succineidae). Walkerana 9: 177-215. Strandine, E. J. 1941. Quantitative study of a snail population. Ecol- ogy 22: 86-91. Villalobos, C. M., J. Monge-Najera, Z. Barrientos, and J. Franco. 1995. Life cycle and field abundance of the snail Succinea costaricana (Stylommatophora: Succineidae), a tropical agri- cultural pest. Revista de Biologia Tropical 43: 181-188. Villalobos, C. M. and J. Monge-Najera. 2004. Yearly body size dis- tribution in the terrestrial snail Succinea costaricana (Stylom- matophora: Succineidae). Brenesia 62: 47-50. Webb, G. R. 1977a. On the sexology of Catinella ( Mediappendix ) avara (Say) or C. (M.) vermeta (Say). Gastropodia 1: 100-102. Webb, G. R. 1977b. Some sexologic observations on Oxyloma retusa (Lea). Gastropodia 1: 102-104. Wu, S. 1993. Notes on the succineid land snails of New Mexico. Malacological Review 26: 91-94. Submitted: 8 December 2008; accepted: 24 March 2009; final revisions received: 12 October 2009 Amer. Malac. Bull 28: 121-126 (2010) Distribution, density, and population dynamics of the Anthony Riversnail (. Athearnia anthonyi) in Limestone Creek, Limestone County, Alabama Jeffrey T. Garner1 and Thomas M. Haggerty2 1 Division of Wildlife and Freshwater Fisheries, Alabama Department of Conservation and Natural Resources, 350 County Road 275, Florence, Alabama 35633, U.S.A. department of Biology, University of North Alabama, UNA Box 5182, Florence, Alabama 35632, U.S.A. Corresponding author: bleufer@aol.com Abstract: Athearnia anthonyi (Redfield, 1854) is a federally endangered gastropod endemic to the Tennessee River drainage in Alabama and Tennessee. It occurs in only three small populations, the most robust in Limestone Creek, Limestone County, Alabama. In 1996, this population was restricted to the lower 14.5 km of unimpounded stream, confined to riffle and run habitats. A follow-up survey in 2006 suggested no change in range within Limestone Creek. In 1996-97, quantitative data were collected from 4 selected sites in the reach and mean A. anthonyi density was 83.9 ± 9.9 SE per m2 (N = 90). Although density did not vary among months, the proportion of individuals within four size classes differed. New recruits appeared in the population between May and July, and a significant die-off of older individuals occurred during the same period. Many individuals were suspected of having at least two breeding seasons. Increasing urbanization within the Limestone Creek watershed necessitates monitoring of A. anthonyi. Key words: endangered species, recruitment, freshwater gastropod Freshwater snails of the family Pleuroceridae comprise a prominent element of the benthic fauna of many southeast- ern U.S. streams (Richardson et al. 1988, Brown et al. 2008). Species within the family are generally large compared to other benthic organisms and often occur in high densities. However, many species have undergone dramatic declines during the past century, following impoundment of major rivers and other negative impacts on their habitats (Lydeard et al. 2004, Lysne et al. 2008). Nowhere have these declines been more evident than Alabama. Of approx. 112 pleurocerid species known from the state, 29 are considered extinct and five are federally protected (Mirarchi 2004). Athearnia anthonyi (Redfield, 1854) (= Leptoxis crassa anthonyi , Anthony Riversnail) is a pleurocerid that has under- gone drastic declines during the past century and was listed as endangered under the federal Endangered Species Act in 1994 (Federal Register 1994). This species is endemic to the Tennessee River drainage and was historically widespread in upper and middle reaches of the Tennessee River proper, ranging from Knoxville in eastern Tennessee, downstream to Muscle Shoals in northwestern Alabama, as well as in the lower reaches of major tributaries (Goodrich 1931, Burch 1989, Garner 2004). Three extant populations are currently known (Minton and Savarese 2005). Two of the remaining populations occur in close proximity, in lower reaches of the Sequatchie River and adjacent Tennessee River in the Nickajack Dam tailwaters, Marion County, Tennessee, and Jackson County, Alabama. The third and most robust popu- lation is found in Limestone Creek, Limestone County, Alabama. In 2001, a Nonessential Experimental Population (NEP) was designated for the Tennessee River in Wilson Dam tailwaters by the U. S. Fish and Wildlife Service (Federal Register 2001). From 2003 to 2008 a total of 4,000 A. anthonyi from the Limestone Creek population were released into the NEP area at Tennessee River mile (TRM) 249. The first obser- vation of reproduction among the reintroduced snails was recorded during late summer 2008 (Garner, pers. obs.). Although the locations of extant Athearnia anthonyi populations are known, little has been published about the species, and life history information is needed for its recovery (U. S. Fish and Wildlife Service 1997). Therefore, the objec- tives of this research were to: (1) determine distributional limits of A. anthonyi in Limestone Creek, (2) estimate density within a reach of Limestone Creek to better understand pop- ulation vigor and provide baseline data for future studies, and (3) obtain information concerning population recruit- ment and dynamics. MATERIALS AND METHODS Habitat description Limestone Creek is approx. 72 km long and has a drain- age area of 290 km2. Most of the drainage lies within Lime- stone County, Alabama, but headwaters originate in Madison County, Alabama, and Lincoln County, Tennessee. It is a third order stream within the Tennessee Valley District of the Interior Low Plateau Physiographic Province (Sapp and 121 122 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Emplaincourt 1975). Underlying geology is composed of Eort Payne Chert and Tuscumbia Limestone, with some upstream reaches containing exposed sediments of the Ordovician System (Osborne et al. 1988, Szabo et al. 1988). Substrata of runs and riffles are mostly gravel with interstitial silt whereas pools and marginal areas often have deposits of mud and beds of Waterwillow ( Justicia americana). Accumulations of leafy detritus are often encountered in pools. Exposed bed- rock occurs at some sites, but outcrops are generally not extensive. Land use surrounding Limestone Creek is primar- ily agricultural, with scattered forested areas, but residential areas have increased considerably in the last decade. Riparian vegetation is generally intact and banks are stable. Canopy cover in most reaches is extensive, spanning the stream in many areas. The stream empties into Limestone Creek embay- ment which enters Wheeler Reservoir of the Tennessee River at TRM 311 (Pig. 1). Qualitative survey methods A qualitative survey to determine the distribution Athearnia anthonyi in Limestone Creek was performed pri- marily at road access points 7 May-24 June 1996. A 9-km float trip, employing mask and snorkel searches in runs and riffles at approx. 25 sites, was carried out between Limestone Creek mile (LCM) 14.6 (Limestone County Rd. 24) and LCM 9.0 (Limestone County Road 12) (Pig. 1). This survey provided access to sites between bridge crossings and allowed determi- nation of the upstream limit of A. anthonyi. A more extensive qualitative survey was conducted in August and September 2006 at 13 road access points between LCM 4.5 and LCM 38 (Table 1, Fig. 1 ). The site identified as the upstream limit of Athearnia anthonyi in 1996 was not accessible during the 2006 survey. However, a visit to LCM 12.5 was made in December 2007 to confirm the continued presence of the species. Qualitative sampling in reaches up Figure 1. Study area showing survey sites on Limestone Creek, Limestone and Madison counties, Alabama. Circles denote road crossing survey sites and squares denote sites used to obtain density and demographic estimates. Inset shows more specific location of density and demographic estimate sites. Survey site numbers correspond to those used in Table 1. ATHEARNIA ANTHONYI LIFE HISTORY 123 Table 1. Sample localities and survey results from 13 sites on Limestone Creek, Limestone and Madison counties, Alabama, obs, number of observers. Sample ID Date Latitude (°N) Longitude (°W) Individuals/hr/ obs obs time (min) LC1 14- Aug-2006 34°37’09.26” 86°51’40.78” 0 3 72 LC2 7-Aug-2006 34°37’53.72” 86°52’01.06” 500+ 2 60 LC3 14-Aug-2006 34°40’31.36” 86°52’42.56” 500+ 3 83 LC4 14-Aug-2006 34o40’17.72” 86°51 ’52.99” 2 3 49 LC5 14-Aug-2006 34°43’46.38” 86°50’37.43” 0 3 63 LC6 18-Aug-2006 34°46’22.33” 86°47’58.16” 0 3 40 LC7 18-Aug-2006 34°48T0.40” 86°48’57.67” 0 3 50 LC8 18-Aug-2006 34°50’06.36” 86°48’31.28” 0 3 35 LC9 25-Aug-2006 34°51 ’06.66” 86°48’54.68” 0 2 65 LC10 25-Aug-2006 34°53’03.30” 86°47’02.18” 0 2 55 LC11 1 -Sep-2006 34°54’57.96” 86°44’54.17” 0 3 70 LCD 1 -Sep-2006 34°54’51.80” 86°43’50.19” 0 3 70 LC13 1 -Sep-2006 34°55’58.87” 86°43’10.99” 0 2 55 to 100 m long was carried out for an average of 2.65 person hours per site (range 1.75-4.15, N - 13). Most A. anthonyi were located by visually searching the stream bottom, by col- lecting substratum with a 1-mm mesh dipnet, or by hand and sorting them in a white pan. Quantitative methods Quantitative data were collected at four sites, selected based on presence of Athearnia anthonyi and stream accessi- bility. Sites were located at LCM 6.7 (34°38’36.60”N, 86°52’ 9.84”W) near Belle Mina, LCM 11.1 (34°4r55.32”N, 86°52’9.83”W) on the Tennessee Valley Research and Extension Station (TVRES), LCM 12.4 (34°42’46.80”N, 86°52’4.08”W) near Anderson Cemetery and in an artificial side channel also located on the TVRES [approx, adjacent to LCM 9.5] (34o40’52.31”N, 86°52’43.31”W), all within Limestone County, Alabama (Fig. 1). Since studies of other pleurocerids (Houp 1970, Miller-Way and Way 1989, Huryn etal. 1994) found little growth and reproductive activity dur- ing winter months, the sites were sampled in May and July 1996, and in May, July, and September 1997. Because some sites were not sampled for all three months [TVRES site (September only), Anderson Cemetery (July and September only), Belle Mina (May and September only)], data from all four sites were pooled to characterize the population within this relatively small reach (approximately 9 km). At each site and sampling interval, three samples were collected one to three meters from both banks (depending on stream width and marginal depths) and at midstream for a total of nine samples per visit. Each sample was collected using a Surber sampler (0.09 m2), from which all substrate was removed to a depth of approx. 10 cm. The material was washed in a brass 2-mm sieve with creek water and contents placed into a shallow white pan. Due to the federal protection of Athearnia anthonyi, all samples were processed and snails enumerated and measured streamside. Each sample was re- examined by a different worker as a quality control measure. Since the tip of the spire was often eroded in larger individu- als, width measurements were used to determine size class. Width was defined as the greatest distance between outer sur- faces of the body whorl as measured on the same plane as the aperture. Measurements were made to the nearest millimeter using dial calipers. All A. anthonyi were then returned to the area from which they were collected. A one-way ANOVA was performed to compare tempo- ral differences in snail densities. Size-frequency plots were used to analyze population dynamics. In addition, size fre- quency distributions per sample were compared among months by dividing individuals into four size classes based on maximum width measurements and size frequency plots. The percentage of individuals within each size cohort per sample was compared among months using a Kruskal -Wallis test. RESULTS During the 1996 qualitative survey, Athearnia anthonyi was found in Limestone Creek from near the mouth of Martin Branch, approx. LCM 13.2, downstream to the upstream limit of impoundment at LCM 4.2, a distance of 14.5 km (Fig. 1). In this reach, A. anthonyi was common to abundant (hundreds usually observed) in almost all riffles and runs. During the more extensive 2006 survey, A. antho- nyi was found in roughly the same reach, from LCM 5.6 to LCM 9.0, and was confirmed to continue upstream to at least LCM 12.5 in 2007. 124 AMERICAN MALACOLOGICAL BULLETIN 28-1/2-2010 During quantitative sampling in 1996 and 1997 the mean density for all 4 sample locations and collection dates was 83.9 ± 9.9 SE per m2 (N = 90). Although density varied some- what among months (May: 88.6 ± 10.9, N = 27; July: 1 13.2 ± 28.8, N= 27; September: 58.4 ± 7.8, N = 36), differences were not significant (ANOVA, F = 2.7, P = 0.07). Size-frequency distributions indicated that in May, sizes ranged from 5 to 16 mm with two clear peaks in frequency, one composed of 8—9 mm sized individuals and the other of 1 1-12 mm snails (Fig. 2). Also in May, on average, 93% of the samples contained individuals that ranged in size from 6-15 mm (Fig. 3). This changed dramatically in July when sizes ranged from 2-17 mm, and a third mode composed of 3 mm individuals occurred, with the two peaks noted from May less pronounced (Fig. 2). Examination of the 4 size classes in the July samples showed that the percentage of individuals in the 1-5 mm size class increased significantly (Chi-square test, X2 = 16.9, P = 0.0002), whereas individuals in the 11-15 mm 100 < LU 5 ■ 1 - 5 mm 27(284) S 6 -10 mm July Sept Figure 3. Mean percentage of individuals per sample for four shell width size classes for three months in 1996 and 1997. Numbers out- side and inside parentheses above bars are sample sizes and number of individuals measured for each month, respectively. Error bars represent standard error. size class decreased (x2 - 16.0, P = 0.0003) since May (Fig. 3). Little change in the percentages of the other two ages classes (6-10 mm and 16-20 mm) were noted between May and July (Fig. 3). Although the range of sizes in September was also large (3-18 mm), only one clear mode in frequency was noted (10 mm, Fig. 2). Also, the percentage of individuals in the 1-5 size class that was predominant in July decreased dramati- cally by September, and there was an increase in the percent- age of snails in the 11-15 (x2 = 16.0, P = 0.0003) and 16-20 mm size class (y2 = 13. 2, P = 0.001) (Fig. 3). The percentage of individuals in the 6-10 mm size class from September was similar to those of the other two months (x2 = 1.6, P = 0.4) (Fig- 3). DISCUSSION WIDTH (mm) Figure 2. Size-frequency histograms indicating shell width range of Athearnia anthonyi for three months in 1996 and 1997 in Limestone Creek, Limestone County, Alabama. The distribution of Athearnia anthonyi within Limestone Creek was similar between 1996 and 2006. Its absence from the lowermost site in 2006 is not surprising because it lies within the transition zone between free-flowing stream and reservoir and habitat is marginal (few individuals were found there in 1996). During the more extensive 2006 survey, A. anthonyi was not found above the site where it was noted in 1996, even though appropriate habitat (i.e., riffles and runs) was examined. The upstream-most site for A. anthonyi in 1996 was again inaccessible during 2006, but since A. antho- nyi was again observed just downstream in December 2007, little range reduction appears to have occurred between the, ATHEARNIA ANTHONYI LIFE HISTORY 125 two surveys. Factors responsible for limiting A. anthonyi to the lower 14 km of free-flowing stream are unclear. Overall average density observed during 1996 quantita- tive sampling falls within the range of densities observed in other pleurocerid species. Examples of low pleurocerid densi- ties reported in the literature were 24 and 38 per m2 for Pleurocera acuta Rafinesque, 1831 and 38 per m2 for Elimia livescens (Menke, 1830) (Dazo 1965, Houp 1970). In contrast, densities of Elimia semicarinata (Say, 1829) up to 1,706 per nr in specific habitats have been reported (Johnson and Brown 1997). Densities were not measured in 2006-07, but observations in suitable habitat suggested healthy popula- tions {i.e., all size classes represented, hundreds of individuals present). The two peaks in the May size-frequency histogram suggest that at least two age cohorts were present: one com- posed of 1 year olds (i.e., hatched the previous year) and another of at least two year olds. The presence of a third mode in the July size-frequency histogram and a significant increase in the proportion of 1-5 mm snails suggest that sig- nificant recruitment occurred between May and July. The timing of new-recruit appearance varies among pleurocerid species, but ranges from June and July to October (Houp 1970, Aldridge 1982, Miller-Way and Way 1989, Huryn et al. 1994, Richardson and Scheiring 1994). Van Cleave (1933) reported newly hatched Pleurocera acuta to measure 0.4 mm in diameter and surmised that they were hatched in June or July, but not collected until later, at a diameter of 6 mm. Some species have extended periods in which recruits appear, lasting three to four months, with numbers peaking toward the end of the period (Aldridge 1982, Richardson and Scheiring 1994). However, in this study it was difficult to tell whether small numbers of individuals in the 1-5 mm size class observed in May and September were new hatch- lings or slow-growing individuals. Growth rates can be highly variable within some pleurocerid species (Huryn et al. 1994) and earlier-hatching captive pleurocerids often grow considerably faster than those that hatch later in the season (Paul Johnson, pers. comm.). Although predation may have contributed to the drop in the proportion of 1-5 mm individuals between July and September (Haag and Warren 2006), many of the hatch- lings likely grew into the 6-10 mm size class by September. This would explain why the proportion of 6-10 mm indi- viduals was similar between July and September, even though the percentage of individuals in the 11-15 mm and 16-20 mm size classes also increased due to growth of the 6-10 and 11-15 mm cohorts of July, respectively. Rapid growth during early months, relative to the remainder of the life span, has been reported for other pleurocerids (Dazo 1965, Stiven and Walton 1967, Richardson etal. 1988, Huryn etal. 1994). The relatively high percentage of individuals in the 6-10 mm size class in May suggests that many individuals do not exceed 10 mm during their hatching year and this size class is composed primarily of year-old snails. Many of these 1 year olds apparently grew into the 11-15 mm size class by September. The significant decrease in frequency of individu- als in the 11-15 mm size class between May and July suggests mortality of many individuals within the class. Magruder (1934) reported a die-off of Pleurocera canaliculatum undula- tum (Say, 1829) in the spring. Some pleurocerid species are biennial, including Leptoxis carinata (Bruguiere, 1792), Leptoxis dilatata (Conrad, 1835), and Pleurocera acuta (Houp 1970, Aldridge 1982, Miller-Way and Way 1989). However, some live even longer, such as Elimia clara (Anthony, 1854) and Elimia cahawbensis (Lea, 1861) that may live as long as 10 and 11 years, respectively (Richardson et al. 1988, Huryn etal. 1994), and Leptoxis foremani (Lea, 1843) which has lived 6+ years in captivity (Paul Johnson, pers. comm.). Data from this study are insufficient to specifically determine life span, but assuming that individuals mature by the end of their first year and reproduce until death, A. anthonyi has at least two breeding seasons. Variation in growth can make it difficult to identify older cohorts (Huryn et al. 1994). Mark- recapture studies are needed to help identify age cohorts and obtain longevity estimates for A. anthonyi. Johnson and Brown (1997) found that the densities and population size structure of E. semicarinata varied with abiotic factors ( e.g ., current velocity). However, limited sampling in this study prevented such comparisons for A. anthonyi and variation in habitat features among sites could have contributed to the temporal variation observed. More specific ecological research is needed to characterize A. anthonyi habitat requirements, and this study provides preliminary estimates for density, recruitment time, and demographics. In summary, the distribution Athearnia anthonyi has remained stable over the past decade. Densities observed at four sites and from three months in 1996 and 1997 averaged 83.9 per m2. New recruits appeared in the population between May and July, a significant die-off of older individuals occurred during the same period, and adults may have at least two breeding seasons. Regular monitoring will be required to protect the Limestone Creek population of A. anthonyi from rapid urbanization of its watershed. Preliminary observations indicate that A. anthonyi is most abundant in riffle and run habitats and therefore anthropogenic activities (e.g., excessive irrigation, sedimentation, pollution) that affect these high- density habitats could be detrimental to the species. A better understanding of the ecological factors limiting A. anthonyi populations is thus needed to help in the species recovery and to identify sites for reintroduction efforts (U. S. Fish and Wildlife 1997). 126 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 ACKNOWLEDGMENTS This study was funded by the U. S. Fish and Wildlife Service and Alabama Division of Wildlife and Freshwater Fisheries and supported by the University of North Alabama. Thanks to P. Ekema, K. Floyd, G. Gaston, L. Gilbert, C. Graydon, R. Jackson, P. Kittle, and T. Richardson for their help during various stages of this study. Thanks also to Ken Brown and Paul Johnson for reviewing and improving the manuscript. LITERATURE CITED Aldridge, D. W. 1982. Reproductive tactics in relation to life-cycle bioenergetics in three natural populations of the freshwater snail, Leptoxis carinata. Ecology 63: 96-208. Brown, K. M., B. Lang, and K. E. Perez. 2008. The conservation ecology of North American pleurocerid and hydrobiid gastro- pods. Journal of the North American Benthological Society 27: 484-495. Burch, J. B. 1989. North American Freshwater Snails. Malacological Publications, Hamburg, Michigan. Dazo, B. C. 1965. The morphology and natural history of Pleurocera acuta and Goniobasis livescens (Gastropoda: Cerithiacea: Pleu- roceridae). Malacologia 3: 1-80. Federal Register. 1994. Endangered and threatened wildlife and plants; determination of endangered status for the Royal Snail and Anthony’s Riversnail. Federal Register 59: 17994- 17998. Federal Register. 2001. Establishment of nonessential experimen- tal population status for 16 freshwater mussels and 1 fresh- water snail (Anthony’s Riversnail) in the free-flowing reach of the Tennessee River below the Wilson Dam, Colbert and Lauderdale counties, AL. Federal Register 66: 32250-32264. Garner, J. T. 2004. Anthony’s Riversnail Athearnia anthonyi. In: R. E. Mirarchi, J. T. Garner, M. F. Mettee, P. E. O’Neil, eds., Alabama Wildlife. Vol. 2. Imperiled Aquatic Mollusks and Fishes. The University of Alabama Press, Tuscaloosa. P. 124. Goodrich, C. 1931. The pleurocerid genus Eurycaelon. Occasional Papers of the Museum of Zoology, University of Michigan, 223: 1-9. Haag, W. R. and M. L. Warren, Jr. 2006. Seasonal feeding special- ization on snails by river darters (Percina shumardi) with a review of snail feeding by other darter species. Copeia 2006: 604-612. Houp, K. H. 1970. Population dynamics of Pleurocera acuta in a cen- tral Kentucky limestone stream. The American Midland Natu- ralist 83: 81-88. Huryn, A. D., J. W. Koebel, and A. C. Benke. 1994. Life history and longevity of the pleurocerid snail Elimia: A comparative study of eight populations. Journal of the North American Benthologi- cal Society 13: 540-556. Johnson, P. D. and K. M. Brown. 1997. The role of current and light in explaining the habitat distribution of the lotic snail Elimia semicarinata (Say). Journal of the North American Benthological Society 16: 545-561. Lydeard, C., R. H. Cowie, W. F. Ponder, A. E. Bogan, P. Bouchet, S. A. Clark, K. S. Cummings, T. J. Frest, O. Gargominy, D. G. Herbert, R. Hershler, K. E. Perez, B. Roth, M. Seddon, E. E. Strong and F. G. Thompson. 2004. The global decline of nonmarine mollusks. Bioscience 54: 321-330. Lysne S. J., K. E. Perez, K. M. Brown, R. L. Minton, and J. D. Sides. 2008. A review of freshwater gastropod conservation: challeng- es and opportunities. Journal of the North American Benthologi- cal Society 27: 463-470. Magruder, S. R. 1934. Notes on the life history of Pleurocera canali- culatum undulatum Say. The Nautilus 48: 26-28. Miller-Way, C. A. and C. M. Way. 1989. The life history of Leptoxis dilatata (Conrad) (Prosobranchia: Pleuroceridae) from the Laurel Fork River, West Virginia. American Midland Naturalist 122: 193-198. Minton, R. L. and S. P. Savarese, Jr. 2005. Consideration of genetic relationships in management decisions for the endangered Anthony’s riversnail, Leptoxis crassa anthonyi (Redfield, 1854) (Gastropoda: Pleuroceridae). The Nautilus 119: 11-14. Mirarchi, R. E. 2004. Alabama Wildlife. Vol. 1. A Checklist of Ver- tebrates and Selected Invertebrates: Aquatic Mollusks, Fishes, Amphibians, Reptiles, Birds, and Mammals. The University of Alabama Press, Tuscaloosa, Alabama. Osborne, W. E., M. W. Szabo, T. L. Neathery, and C. W. Copeland, Jr. 1988. Geologic map of Alabama, northeast sheet. Special Map number 220, Geological Survey of Alabama, Tuscaloosa, Alabama. Richardson, T. D. and J. F. Scheiring. 1994. Ecological observations of two pleurocerid gastropods: Elimia clara (Lea) and£. cahawbensis (Say). The Veliger 37: 284-289. Richardson, T. D„ J. F. Scheiring, and K. M. Brown. 1988. Secondary production of two lotic snails (Pleuroceridae: Elimia). Journal of the North American Benthological Society 7: 234-245. Sapp, C. D. and J. Emplaincourt. 1975. Physiographic regions of Alabama. Map number 168, Geological Survey of Alabama, Tuscaloosa, Alabama. Stiven, A. E. and C. R. Walton. 1967. Age and shell growth in the freshwater snail Goniobasis proximo (Say). American Midland Naturalist 78: 207-214. Szabo, M. W., W. E. Osborne, and C. W. Copeland, Jr. 1 988. Geologic map of Alabama, northwest sheet. Special Map number 220, Geological Survey of Alabama, Tuscaloosa. U. S. Fish and Wildlife Service. 1997. Recovery plan for Anthony’s riversnail. Atlanta, GA. P. 21. Available from: http://ecos.fws .gov/docs/recovery_plan/9708 13.pdf. Van Cleave, H. J. 1933. Studies on snails of the genus Pleurocera. II. The young of P. acuta. The Nautilus 47: 48-49. Submitted: 5 February 2009; accepted: 25 June 2009; final revisions received: 13 October 2009 Amer. Malac. Bull. 28: 127-133 (2010) Epiphyton or macrophyte: Which primary producer attracts the snail Hebetancylus moricandi ? Roger Paulo Morrnul1, Sidinei Magela Thomaz2, Marcio Jose da Silveira3, and Liliana Rodrigues2 1 Pos-gradua^ao em Ecologia de Ambientes Aquaticos Continentals, Nucleo de Pesquisas em Limnologia, Ictiologia e Aquicultura - Nupelia, Universidade Estadual de Maringa - UEM, Bloco H90, Av. Colombo, 5790, CEP 87020-900, Maringa, Parana, Brazil 2 Nucleo de Pesquisas em Limnologia, Ictiologia e Aquicultura - Nupelia, Universidade Estadual de Maringa - UEM, Bloco H90, Av. Colombo, 5790, CEP 87020-900, Maringa, Parana, Brazil 3 Pos-graduaqao em Biologia Comparada, Departamento de Biologia, Universidade Estadual de Maringa - UEM, Bloco H90, Av. Colombo, 5790, CEP 87020-900, Maringa, Parana, Brazil Corresponding author: smthomaz@nupelia.uem.br Abstract: Relationships between snails, epiphyton, and macrophytes are widely studied because epiphytes decrease light for macrophytes, and snails may benefit the latter when they consume epiphytes. Thus, organic compounds released by macrophytes that attract snails could be an evolutionarily advantageous mechanism. This hypothesis was tested with three species of submerged macrophytes (natives: Egeria najas and Cabomba furcata; exotic: Hydrilla verticillata), which were maintained in microcosms in the presence of ancylid snails. However, the hypothesis of limpet attraction by macrophytes was rejected. Instead, epiphyton attached to E. najas attracted more snails than that attached to the other species. This attraction could be explained by chemical signals (organic compounds), released by certain species of algae that are detected by snails. Key words: food web, grazers, periphyton, herbivory, chemoreception Aquatic macrophytes are an important component of aquatic ecosystems. For example, they increase habitat com- plexity (Bronmark and Vermaat 1998, Thomaz et al. 2008) and contribute organic matter to food webs (Esteves 1998, Benedito-Cecilio et al. 2004, Lopes et al. 2007). Interactions among macrophytes, epiphyton, and invertebrates may affect the composition and distribution of epiphytic algae and invertebrate species as well as submerged macrophyte pro- ductivity and longevity (Bronmark 1985, 1989, Lodge 1986). Wetzel (2001) suggested a subtle symbiosis between these three groups and reported that herbivory is mediated by organic compounds, and that a compound that acts as an inhibitor for some snail species can stimulate others, making this type of interaction species-specific. Several authors criti- cized experiments using artificial macrophytes to test these interactions because unlike natural plants, artificial plants provide only substrate without chemical signals (Cattaneo and Kalff 1978, 1979, Cattaneo 1983). The presence of these interactions has been indicated by many studies that showed that epiphyton reduced growth of macrophytes due to shading plant surfaces (Sand-Jensen 1977, Bulthius and Woelkerling 1983, Bronmark 1985, 1989). Some freshwater invertebrate grazers, especially gastropods, are attracted by chemical signals that they detect with chemoreceptors, and food preferences between macrophytes and/or microalgae species differ among snail species. Snails also differ in assimilating different types of food (Townsend 1973, Calow and Calow 1975, Croll 1983). Many gastropods have strongly developed chemoreception and responses to signals from submerged plants. There is evidence that some species of macrophytes release compounds that attract snails, which, in turn, graze the epiphyton that colonizes the plant (Bronmark 1985, 1989). This interaction, based on chemore- ception, benefits both species: the snail uses the plant as habi- tat for spawning, feeding, and hiding while the plant benefits from reduced competition with epiphyton. The epiphyton is also benefited, because algal removal by herbivores, a physical disturbance, promotes the regeneration of the community (Rodrigues and Bicudo 2001). Recent studies have shown that the size of plant fragments, plant age, and density of herbivores are factors that affect the submerged plant-snail interactions (Eiger et al. 2007). In Neotropical aquatic habitats, like the Upper Parana River in Brazil, native snails of the family Ancylidae are reported in association with native macrophytes, such as Egeria najas and Cabomba furcata (Thomaz et al. 2008). The submerged macrophyte Hydrilla verticillata, native from Africa and Asia, has also been recorded in habitats of the Upper Parana River. However, specific interactions between these native snails and native and exotic plant species are unknown. In this study, we tested the hypothesis that the snail Hebetancylus moricandi (d’Orbigny, 1837) is attracted by particular submerged macrophytes in response to the release of chemical compounds. We expected that if this 127 128 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2-2010 hypothesis were true, more snails would be found in treat- ments with macrophytes than in epiphyton alone or in absence of macrophytes and epiphyton. The assumption behind this hypothesis is that plants and herbivores are ben- efited, based on previous experiments with north-temperate species (Thomas 1982, Bronmark 1985). An alternative hypothesis (also tested in our experiments) is that snails are attracted by epiphyton itself. In this case, we expected that more snails would be found in treatments with epiphyton alone than in macrophytes or in absence of macrophytes and epiphyton. MATERIALS AND METHODS Snails ( Hebetancylus moricandi) and macrophytes ( Egeria najas, Cabomba furcata, and Hydrilla verticillata) were sampled in a backwater in the Upper Parana River floodplain, Brazil (22°45’S, 53°16’W). Invertebrates and macrophytes were collected randomly inside macrophyte stands. All sampling and experiments were completed from September to December of 2007, and each individual experi- ment was initiated less than three hours after sampling. Plant fragments were maintained inside transparent plastic bags at ca. 10 °C, to reduce their metabolism, until beginning the experiment. Our experiments were based on Bronmark’s (1985) work. All experiments were carried out in five “labyrinth type” acrylic aquaria measuring 30 x 15 x 15 cm. Each aquar- ium was created with five connected compartments (Fig. 1). Aquaria were filled with two liters of water, added from cen- ter to periphery. We then added plant fragments and algae to the compartments. Since chemicals specific to plants or algae were more concentrated in each specific compartment, we Treatments Figure 1. Aquarium used in experiments. predicted snails would choose the compartment with the pre- ferred food. In the first experiment, two species of submerged macro- phyte were used (the native Egeria najas and the exotic Hydrilla verticillata). Three compartments were considered treatments, and the snails were introduced into the fourth, at the center of the aquarium, equidistant from the three treatments. The remaining compartment was isolated. We placed three plant fragments with epiphyton in one compart- ment and three without epiphyton in the other, leaving one empty compartment (control). Epiphytic material was removed from the macrophyte surface carefully with a brush and filtered water. Four snails were placed in the central com- partment. The procedure was repeated nine times for each macrophyte species and each set of experiments had five aquaria, each aquarium was considered as a replicate, yield- ing 45 independent replicates (nine aquaria repeated five times each). The replication of each treatment was random- ized in the compartments. In a pilot experiment, we found 30 minutes to be suffi- cient for snail movement, and this interval was used in all experiments. Every 30 minutes, the snails’ positions were recorded, plants were removed, and the water replaced. The aquaria were carefully washed with distilled water to remove attached material before every new replicate. In a second experiment, another native submerged mac- rophyte ( Cabomba furcata ) was added to assess the consis- tency of results. The procedure described previously was repeated, but epiphyton was removed from the macrophytes and added in the compartment that had been previously iso- lated. At this stage, there were a total of 30 repetitions per macrophyte species. In addition to the control (“C”; com- partment without material), the following treatments were: Egeria najas (En), E. najas + epiphyton (En + E), epiphyton removed from E. najas (E - En), Hydrilla verticillata (Hv), H. verticillata + epiphyton (Hv + E), epiphyton removed from H. verticillata (E - Hv), Cabomba furcata (Cf), C. furcata + epiphyton (Cf + E), and epiphyton removed from C. furcata (E-Cf). The experiments were repeated a third time, but epiphy- ton was sampled and preserved in lugol acetic 0.5% (Rodrigues and Bicudo 2001), for further analysis of the main species of algae and their relative abundances. The organisms were counted according to Utermohl (1958). This experiment had 15 independent replicates per macrophyte species. Two sam- ples of epiphyton for each macrophyte species were preserved for algal identification. The number of snails found in each compartment after 30 minutes was considered the response variable. The effects of the attraction by macrophytes and epiphyton were evalu- ated by analysis of variance (one-way ANOVA), considering significant values of P < 0.05. All assumptions were met and SNAIL ATTRACTION BY PRIMARY PRODUCER 129 thus it was not necessary to transform data before statistical analysis. When differences were significant, an a poste- riori Tukey test was applied. The composition of epiphytic algae on the three macrophyte species identi- fied in the third experiment was evalu- ated by a Detrended Correspondence Analysis (DCA). This analysis was applied to assess whether algae composition diff- ered between macrophyte species. RESULTS In the first experiment, in which we tested the effects of Egeria najas on the attraction of Hebetancylus mori- candi, the snails were more attracted by the compartment En + E (ANOVA, F2 42 = 17.62, P < 0001; Fig. 2). In con- trast, for the experiment with Hydrilla verticillata, there was no preference for any specific compartment (F2 42 = 0.13, P = 0.87). The results of the second experi- ment showed that the treatment En + E resulted in greater colonization by snails than the treatments C and En. However, according to the Tukey test, the treatment E - En presented the highest mean snail numbers {F} 26 = 12.86, P < 0001; Fig. 3). For Hydrilla verticillata, a slightly higher number of snails was attracted to the E - Hv com- partment, but this difference was only marginally significant (F} 2g = 2.89, P - 0.04). In fact, these differences were not detected by Tukey tests (P > 0.05). In the experiment with Cabomba furcata, there was no significant difference between the numbers of snails attracted by each compartment (P3 26 = 1.57, P = 0.20) (Fig. 3). Thus, the hypothesis that the plants attract snails was rejected and the alternative hypothesis of attrac- tion of snail by epiphyton was not rejected. Because the second experiment showed that snails were attracted by epiphyton attached to a specific macro- phyte ( Egeria najas), we carried out a 1.4 1.2 w 1.0 53 ■g 0.8 « 0.6 lo, 0.2 0.0 Figure 2. Mean and standard error of snail numbers attracted in the first experiment for the aquarium compartments. A, Egeria najas ; B, Hydrilla verticillata. En + E, E. najas + epiphyton; En, E. najas; Hv + E, H. verticillata + epiphyton; Hv, H. verticillata; C, control compartment. Figure 3. Mean and standard error of snail numbers attracted in the second experiment for the aquarium compartments. A, Egeria najas; B, Cabomba furcata; C, Hydrilla verticillata. En, E. najas; En + E, E. najas + epiphyton; E - En, epiphyton removed from E. najas; Hv, H. verticil- lata; Hv + E, H. verticillata + epiphyton; E - Hv, epiphyton removed from H. verticillata; Cf, C. furcata; Cf + E, C. furcata + epiphyton; E - Cf, epiphyton removed from C. furcata; C, control compartment. 130 AMERICAN MALACOLOGICAL BULLETIN 28-1/2-2010 third experiment. Similar results were obtained in this third experiment, i.e. snails were also more attracted by compartment E - En (F3 M = 6.64, P < 0.01; Fig. 4). Again, for Hydrilla verti- cillata and Cabomba furcata there were no significant differences between treatments {F} n = 0.79, P = 0.52 and F3 u = 0.15, P = 0.92, respectively) (Fig. 4). The epiphyton from the three spe- cies of macrophytes was dominated by different species of algae (Table 1). The results of the DCA showed that the algal composition differed consistently among macrophytes (Fig. 5). In the upper right quadrant, only the algae associated with Egeria najas are present; in the upper left quadrant, algae associated with Hydrilla verticillata predominated, while in the lower quadrants algae species associated with C. furcata predominated. Among the main taxa associated with E. najas were Cosmarium abbreviatum , Cyano- phyceae 2, and Cymbella sp. In contrast the epiphyton associated with Cabomba furcata was represented instead mainly by Chlorophyceae 3, Chlorophyceae 4, and Cocconeis placentula, while the algae asso- ciated with H. verticillata was dominated by Chlorophyceae 1, Cocconeis pediculus, and Cyanophyceae 3. DISCUSSION The hypothesis that macrophytes attract grazing snails to remove epiphyton, benefitting the plant by reducing compe- tition for light and nutrients was rejected, and the results of three independent experiments indicated food preference by IK I Hv E-Hv Figure 4. Mean and standard error of snail numbers attracted in the third experiment for the aquarium compartments. A, Egeria najas; B, Cabomba furcata; C, Hydrilla verticillata. The acronyms are given in previous figure legends. Table 1. Main algal taxa encountered on macrophytes. Egeria najas Cabomba furcata Hydrilla verticillata Cyanophyceae 2 Chlorophyceae 3 Chlorophyceae 1 Cosmarium abbreviatum Chlorophyceae 4 Cyanophyceae 3 Eunotia sp.2 Cocconeis placentula Gomphonema sp.l Gomphonema gracile Encyonema minutum Synedra goulerdi Gomphonema angustum Cyclotela stelligera Cocconeis pediculus Cymbella sp.2 Scenededesmus brevispida Cyanophyceae 1 Cymbella sp. 1 Gomphonema parvalum Eunotia sp.3 Synedra sp.3 Synedra sp. 1 Synedra sp.2 snails for Egeria najas epiphyton. These results were evi- denced by significant differences shown by ANOVA, revealing that a greater number of snails were attracted when epiphy- ton alone remained in the compartment. In addition, both the analyses of dominant groups and the DCA showed that the composition of epiphytic algae differed among the three species of macrophytes and thus, some specific group of algae attached to E. najas exerted more attraction (probably dia- toms, which occurred in greater densities). The importance of abundance, composition, and dis- tribution of epiphyton for snail dis- tribution and to the interactions between snails, epiphyton, and mac- rophytes in freshwater ecosystems has been shown in several papers (Lamberti and Resh 1983, Kohler 1984, McAuliffe 1984). Snails are able to select epiphyton, preferring certain algae species (Calow 1970, 1973a, 1973b, 1974). Similar results were found by Lodge (1986), who suggested significant selective her- bivory by snails, since they may SNAIL ATTRACTION BY PRIMARY PRODUCER 131 Figure 5. DCA with algal taxa and macrophyte species, a, Achenan- thidium sp.; b, Achenanthidium minutissimum; c, Aulacoseira sp.; d, Bulbochaete sp.; e, Characium sp.; f, Cyanophyceae 6; g, Chlorophy- ceae 1; h, Chlorophyceae 2; i, Chlorophyceae 3; j, Chlorophyceae 4; к, Cocconeis placentula; 1, Cocconeis pediculus; m, Cosmarium sp.; n, Cosmarium abbreviation ; o, Chrysophyceae; p, Cyanophyceae 1; q, Cyanophyceae 2; r, Cyanophyceae 3; s, Cyanophyceae 4; t, Cyano- phyceae 5; u, Cyclotela stelligera; v, Cymbella sp2; w, Cymbella spl; x, Dynophyceaea; y, Encyonema minutum ; z, Encyonema silesianunv, аа, Eunotia spl; bb, Eunotia sp2; cc, Eunotia luveris ; dd, Eunotia mi- nor, ee, Eunotia sp3; ff, Euglenophyceae; gg, Fragilaria rumpens ; hh, Gomphonema gracile-, ii, Gomphonema spl; jj, Gomphonema sp2; kk, Gomphonema sp3; 11, Gomphonema angustum-, mm, Gomphonema sp4; nn, Gomphonema parvalum; oo, Navicula sp.; pp, Nitzschia sp.; qq, Nitzschia sigmoide ; rr, Oedogonium spl; ss, Oedogonium sp2; tt, Oedogonium sp3; uu, Scenededesmus sp.; w, Scenededesmus brevis- pida; ww, Synedra spl; xx, Synedra goulerdi; yy, Synedra sp2; zz, Syn- edra sp3; aaa, Ulnaria ulna-, ▲, Cabomba furcata-, U, Egeria najas-, •, Hydrilla verticillata. choose the local area or microhabitat where they feed. Although Lodge ( 1986) did not analyze epiphytic composi- tion, he emphasized that in general diatoms were more readily consumed. Organic compounds are released by macrophytes and algae (Gross etal. 1996, Mulderij etal. 2007, Yun etal. 2007), but most research has dealt with compounds that play a major role in inhibiting colonization by algae on macro- phytes, or herbivory of algae by grazers. Hilt (2006) showed that native species of algae and macrophytes may have inter- actions related to specific compounds, and that algae can tol- erate allelopathic compounds released by macrophytes. Juttner (2005) showed that some diatoms are able to reduce the activity of planktonic grazers. However, as suggested by Wetzel (2001), compounds that inhibit the activity of certain species stimulate others, and the relationship emphasized by Hilt (2006) between species of algae and macrophytes may also occur between algae and herbivores. Thus, herbivorous species would evolve towards tolerance of the inhibitory compounds released by algae, and use them as stimuli. Macrophytes may also determine the composition of epi- phyton through release of organic compounds that inhibit certain species of algae ( Bronmark and Vermaat 1 998 ) . T ogether with plant age and limnological features, the release of these compounds can explain why different macrophyte species are colonized by distinct epiphytic communities. The architecture and rugosity of macrophyte species are other features that also explain the dissimilarity of epiphyton found in our samples. In the area chosen for sampling macrophytes and snails, native species of macrophytes preferentially colonized areas where light transmission is low (S. M. Thomaz, unpubl. data). At least for Egeria najas, the reduction of epiphyton by snails may increase light intensity over the plant surface, and facili- tate the success of this species. Although this is only an infer- ence, it has been shown that the abundance of snails can indirectly affect the pattern of macrophyte distribution, mainly in regions where the availability of light is a limiting factor (Bronmark 1989). The composition of epiphyton attached to macrophyte species may also have been affected by their distribution in the backwater habitats. The exotic species ( Hydrilla verticil- lata) is found only in the Parana main channel, where water velocity and light transmission are higher and nutrient con- centrations are lower. Native species ( Egeria najas and Cabomba furcata) colonize mainly backwater habitats (Thomaz et al. 2009). In general, habitats with more light availability and nutrient concentrations sufficient for repro- duction support elevated epiphyton growth rates (Liboriussen etal. 2005, Liboriussen and Jeppesen 2006). In summary, experiments with two native species and one exotic macrophyte indicated the absence of chemorecep- tion associated with macrophytes, and the epiphyton associ- ated with only one species ( Egeria najas) being responsible for the attraction of Hebetancylus moricandi. Based on these results, we suggest identification of organic compounds that attract snails might be searched for in the algal taxa that occurred on E. najas epiphyton. These results differ from those obtained in temperate ecosystems, where chemorecep- tion seems to be associated with chemical compounds released by macrophytes {e.g., Bronmark 1985). However, further studies involving other tropical macrophyte and snail species are necessary to assess whether these results are repre- sentative for tropical aquatic communities. ACKNOWLEDGMENTS We acknowledge with appreciation funds provided by the Brazilian Council of Research (CNPq), through the Long 132 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Term Ecological Research Program (LTER). S. M. Thomaz is a CNPq Productivity Researcher and acknowledges this agency for constantly funding his research. R. P. Mormul is grateful to CAPES (Coordenadoria de Aperfeiqoamento do Pessoal de Nivel Superior) for a scholarship. M. J. Silveira is grateful to PTI (Parque Tecnologico ITAIPU) and PDTA (Programa de Desenvolvimento Tecnologico Avanqado) for a scholarship. We acknowledge with appreciation the comments and cor- rections of Dr. Kenneth M. Brown and the suggestions of one anonymous reviewer. LITERATURE CITED Benedito-Cerilio, E., C. A. Lopes, G. I. Manetta, M. F. Gimenes, A. C. E. A. Faria, E. C. S. Dourado, R. P. Pinheiro, and L. A. Martinelli. 2004. Trophic structure of the fish assemblage in the floodplain of the upper Parana River: Stable isotopes. In: A. 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Academic Press, Burlington, Massachusetts. Yun, H. Y., J. Cruz, M. Treitschke, M. Wahl, and M. Molis. 2007. Testing for the induction of anti-herbivory defences in four Portuguese macroalgae by direct and water-borne cues of graz- ing amphipods. Helgoland Marine Research 61: 203-209. Submitted: 14 April 2009; accepted: 5 August 2009; final revisions received: 8 October 2009 Amer. Malac. Bull. 28: 135-150 (2010) Distribution and environmental influences on freshwater gastropods from lotic systems and springs in Pennsylvania, USA, with conservation recommendations Ryan R. Evans1’* and Sally J. Ray2 1 Pennsylvania Natural Heritage Program, Pittsburgh office, 209 Fourth Ave., Pittsburgh, Pennsylvania 15205, U.S.A. 2 Pennsylvania Natural Heritage Program, Middletown office, 208 Airport Drive, Middletown, Pennsylvania 17057, U.S.A. Corresponding author: ryan.evans@ky.gov Abstract: We examined current distributions of and influential variables on aquatic gastropods in streams and springs across the state. Our research located 37 species representing 7 families. This inventory included rare species such as Somatogyrus pennsylvanicus Walker, 1904 and several populations of the Ohio Pebblesnail Somatogyrus Integra (Say, 1829). Despite targeted surveys, no collections were made of the Buffalo Pebblesnail Gillia altilis (I. Lea, 1841). We also examined the influence of rapid bio-assessment habitat measurements, reach and basin hydrological variables, and selected water chemistry variables on the freshwater snail communities of Pennsylvania. Several measures of habitat quality, drainage area, and water chemistry were among the more important variables explaining patterns in species richness. Several species appear rare and 7 species are recommended for conservation consideration. Further work is needed to better understand the diversity of freshwater gastropods in Pennsylvania. Key words: zoogeography, rare species, snails, ecology Freshwater gastropods are of increasing conservation concern in the United States. Of the nearly 800 species in North America, 60 species of freshwater snails are believed extinct (Paul Johnson, pers. comm.) and the status of many more species is poorly known. Recent estimates by NatureServe (2008) show that over 70% of the North American gastropod fauna is imperiled (species extirpated or at-risk). Despite recent efforts by resource managers to identify species of concern in the state (PFBC and PGC 2005), Pennsylvania lacks information on the present distributional status of its aquatic gastropod fauna. The mussel fauna of the state has been better documented, with early inventory efforts by Ortmann (1919) and more recently by others (Art Bogan, pers. comm.). Pennsylvania has a rich legacy in malacology. Most of the workers in the state have historically focused on distribution and systematics of freshwater mussels, sphaeriid clams, and land snails. A. E. Ortmann, Timothy Conrad, Henry Pilsbry, Victor Sterki, and other prominent malacologists collected freshwater snails in Pennsylvania (often incidental to mussel collecting) largely in the early 20th century. There has been little focus on the group in subsequent years and contemporary survey efforts are needed. Recent research by Evans and Ray (2008) provided support for 63 known or potential species in the state. Pennsylvania is one of the most diverse states in the country in terms of drainage basins and contains 83,184 miles of streams (PFBC and PGC 2005) across its 67 counties. The northeastern edge of the Ohio Basin is contained in the state (Fig. 1). Nearly half of the Delaware and Susquehanna River basins flow through the state. The headwaters of the Elk and Northeast Creeks as well as the Genesee and Potomac River Basins originate in Pennsylvania. Previous workers have determined temperature and pH to be useful determinants of freshwater snail communities (Pip 1987, Brown et al. 1998), while other have shown the importance of drainage area (and concomitant increase in habitat diversity) to be useful (Prezant and Chapman 2004). To help inform the conservation effort for freshwater snails, we initiated an inventory of the Pennsylvania fauna of streams and springs. This study represents the first dedicated, modern effort aimed specifically at the entire aquatic gastropod fauna across the state of Pennsylvania, focused on lotic and spring habitats. We also wanted to examine the influence that habitat data, geology, land use, physical characteristics, and water quality data have on the lotic species in the state. MATERIALS AND METHODS Field sampling was primarily in streams and, to a lesser extent, accessible surface springs. Stream sampling sites were Current address: Kentucky State Nature Preserves Commission, 801 Schenkel Lane, Frankfort, Kentucky 40601, U.S.A. 135 136 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 1. Major river basins of Pennsylvania. selected based on accessibility and areas that would provide the maximum number of microhabitats for maximizing species richness. Sampling sites were initially broken into USGS HUC 8 level watersheds (which are watershed level groupings) and then chosen by using USGS 7.5 topographic maps and a Pennsylvania Gazateer (DeLorme Corporation, Maine). Our goal was to get 8-10 sampling sites per HUC 8 unit. Springs were located from USGS maps and from examining the USGS Geographic Names Information System using ArcView 9.1 and ArcView3.2 software (ESRI, Redlands, California). Hand collecting was done on large woody debris substrates, and handheld sieves were used to sweep vegetation and sample small accumulations of detritus. D-frame nets were used in deeper areas. Stones and trash were examined at each site for species. Although searches were not made in a standardized timed fashion, sampling was typically conducted until no new species were collected. Museum records were obtained from Academy of Natural Sciences in Philadelphia (ANSP), Carnegie Museum of Natural History (CMNH), Delaware Museum of Natural History (DMNH), Florida Museum of Natural History (FMNH), Illinois Natural History Survey (INHS) and The U.S. National Museum (Smithsonian Institution - USNM). To examine local and landscape variables associated with gastropod species richness, sites were assigned physical variables from a modified EPA Level 3 Reach File (RF3) geographic information system (GIS) stream layer produced by The Nature Conservancy (A. Olivero, unpubl. data). This layer contains values assigned for every stream segment in the state of Pennsylvania and contains data representing various measures of land cover types, upstream dams, upstream road crossings, and watershed area for both the reach scale and HUC 12 scale of Pennsylvania streams. Dissolved oxygen, pH, tem- perature, calcium hardness, total hardness, and total alkalinity were measured at many sites by the inves- tigators. At many sites, conductivity and total dissolved solids (TDS) data were also collected. Because not all parameters were collected for all sites, data were supplemented with infor- mation from the Pennsylvania Natural Heritage Program Aquatic Database, one of the largest sources of aquatic habitat and water quality data for the state of Pennsylvania (Nightingale etal. 2004) and averaged for the respective stream segment. To evaluate associa- tions with physical habitat structure, a modified Rapid Bioassessment Protocol (RBP) developed by Pennsylvania Department of Environmental Protection was used (modified from Barbour et al. 1999). A Kruskal-Wallis test was used to examine if there were any statistical differences between species richness at the large basin (HUC 6) level. Due to limited sampling data, Lake Erie basin stream sites were excluded from all analyses. Pearson correlations were run to examine correlations among variables. If two variables showed a correlation coefficient >0.6, the more useful of the two was retained. This reduced the dataset from 43 to 10 variables for RF3 and RBP, respectively. All 7 water chemistry variables were retained from the initial dataset to examine possible influences on community structure. Backward stepwise multiple regressions were used to examine which variables appeared strongest in explaining gastropod species richness. Analyses were conducted separately for landscape variables, land use, water chemistry variables, and habitat variables. In examining land use, species richness values were averaged at the HUC 12 level and assigned a land use value from the National Land Cover Dataset (2000 version). To examine relationships for selected, highly significant variables on presence-absence data, we used PC-ORD software (ver. 4.32) to run Nonmetric Multidimensional Scaling (NMDS) tests. A Sorenson (Bray-Curtis) distance measure was used with 200 maximum iterations and 30 runs against a Monte-Carlo simulation. Prior to analysis, Beals Smoothing was used to minimize the influence of a large number of zero values, as is the case with presence-absence data (McCune et al. 2002). Predicted versus observed species richness was examined with a species-area curve of statewide taxa (for 35 species) using PENNSYLVANIA SNAILS 137 a Sorensen distance measure against all stream and spring sites. Two species, Pleurocera acuta Rafinesque, 1831 and Pleurocera canaliculata (Say, 1821), were excluded from the master taxa dataset as records for these species were from contributed specimens and not a result of a dedicated survey and, thus, could artificially weight species richness data as compared to community data. We also examined species-area curves at the HUC 8 level where adequate sampling data existed. Species richness estimates were conducted using two estimators, Jackknife 1 (Palmer 1990) and Jackknife 2 (Palmer 1991): j ,, . S + rl(n -1) Jackknife 1 = n where S = the observed number of species, rl = the number of species occurring in one sample unit, and n = the number of sample units and Jackknife 2 = S + rl(2n-3) n r2(n - 2)2 n(n-l) where r2 = the number of species occurring in two sample units. Taxonomy generally followed Turgeon et al. (1998), Smith (2000), and Wethington and Lydeard (2007). We chose to follow the name Galba over Fossaria based on the ruling by ICZN (1998). Identifications of collections and distribution ranges were confirmed using Walker (1904), Goodrich (1940), Basch (1963), La Rocque (1968), Burch (1989), Jokinen (1992), and with comparison to museum specimens. Voucher specimens collected in the course of this study were deposited with Carnegie Museum of Natural History in Pittsburgh, Pennsylvania. Species richness across all Pennsylvania streams was highest overall in the Ohio Basin (N = 25) and lowest in the Lake Erie Basin ( N = 4). Pulmonates were more common than caenogastropods, totaling 23 species, or 62%, of the species. The most common pulmonate taxa were Physa acuta Draparnaud, 1805 and Ferrissia rivularis (Say, 1817), while Leptoxis carinata (Bruquiere, 1792) was the most common caenogastropod. A significant difference was observed between major river basins in terms of species richness (Kruskal-Wallis test statistic = 17.01, P = 0.004). Examining richness patterns by HUC 8 region (Pig. 4), the greatest species richness was found in the Northwestern, Glaciated Plateau Physiographic Region in the Lrench Creek watershed of northwest Pennsylvania (Allegheny River system) with 17 species present, followed by Middle Allegheny River-Tionesta Creek, Lower Susquehanna River-Swatara Cr., and Lower Juniata River each with 14 species. Despite a large number of sampling sites, only 21 species were located from the Susquehanna River system. Proportionately, the Potomac River watershed was very species rich, with 10 species located from only 23 sites. Zero species were reported from two USGS 8-digit HUCs in north- central Pennsylvania. No specimens of Pomatiopsidae were located in this study, despite investigating marginal habitats where they have been reported. Regarding the freshwater limpets (Ancylidae; Pig. 5), Ferrissia rivularis (Say, 1817) was one of the most common species in the state, ranging from headwaters to larger rivers. Ferrissia parallelus (Haldeman, 1841), a species typical of slower-flowing stream sections or lentic environments, was reported in 3 collections: Marsh Creek (Adams County), Tobyhanna Creek, and Tunkhannock Creek (both in Monroe RESULTS Survey results We sampled a total of 398 stream and spring sites across Pennsylvania (Pig. 2) in the Delaware, Lake Erie, Ohio, Potomac, and Susquehanna River systems. Due to time constraints, no sampling occurred in the Genesee River basin. A total of 121 sites sampled (30% of total) had no snails (Pig. 2). In streams and springs that were sampled, a total of 37 species were collected (Table 1). Overall, sampling appears reasonably effective in charactering species richness across the state, with first-order jackknife estimates of 38 species and second-order jackknife estimates of 33 species (Pig. 3). 138 AMERICAN MALACOLOGICAL BULLETIN 28-1/2-2010 Table 1. Distribution of species by major basin. List is for nominal taxa; due to taxonomic uncertainties in certain groups, not all species reported may be valid taxa. Total no. of Species Delaware Lake Erie Ohio Potomac Susquehanna sites present Amnicola limosus (Say, 1817) X X X 19 Bellamya chinensis (Reeve, 1863) X X 6 Campeloma decisum (Say, 1817) X X X X 28 Cincinnatia Integra (Say, 1829) X 3 Elimia livescens (Menke, 1830) X 6 Elimia virginica (Say, 1817) X X 45 Ferrissia fragilis (Tryon, 1863) X X 5 Ferrissia parallelus (Haldeman, 1841) X X 5 Ferrissia rivularis (Say, 1817) X X X X X 125 Fontigens nickliniana (I. Lea, 1838) X 3 Galba exigua (I. Lea, 1841) X 1 Galba modicella (Say, 1825) X X X 15 Galba obrussa (Say, 1825) X 8 Galba parva (I. Lea, 1841) X 3 Galba rustica (I. Lea, 1841) X 2 Gyraulus deflectus (Say, 1824) X X 5 Gyraulus parvus (Say, 1817) X X 8 Helisoma anceps (Menke, 1830) X X X X 31 Laevapex fuscus (C. B. Adams, 1841) X X X 16 Leptoxis carinata (Bruguiere, 1792) X X 60 Lithasia obovata (Say, 1829) X 17 Lyogyrus granum (Say, 1822) X X 7 Micromenetus dilatatus (Gould, 1841) X X X 8 Physa acuta (Draparnaud, 1805) X X X X X 133 Physa ancillaria (Say, 1825) X X X X 6 Physa gyrina (Say, 1821) X X X X 40 Planorbella campanulata (Say, 1821) X X 6 Planorbella trivolvis (Say, 1817) X X X 12 Pleurocera acuta Rafinesque, 1831 X 1 Pleurocera canaliculata (Say, 1821) X 1 Promenetus exacuous (Say, 1821) X 1 Pseudosuccinea columella (Say, 1817) X X X 5 Somatogyrus integra (Say, 1829) 16 Somatogyrus pennsylvanicus Walker, 1904 X 1 Stagnicola catescopium (Say, 1867) X X 3 Stagnicola elodes (Say, 1821) X 1 Stagnicola emarginata (Say, 1821) X 1 Total number of species 18 3 26 10 21 Number of null sites 43 0 26 1 51 County). Lurther sampling of low-gradient, slow-flowing stream habitats would likely yield additional specimens. Various taxa of Lymnaeidae were located in the study (Lig. 6). In particular, species of Galba were common inhabitants of mud flats, cobble bars, and floodplains. Galba exigua (I. Lea, 1841) was reported from Enlow Eork, an upper Ohio River tributary, while G. rustica (I. Lea, 1841) was observed only from the Clarion River, Lorest/Jefferson County and Cussewago Creek, Erie County. Galba parva (I. Lea, 1841) was collected from Aughwick Creek in Huntington County and Beaverdam Creek, Somerset County. Specimens of Stagnicola, in general, were somewhat less common in PENNSYLVANIA SNAILS 139 Figure 3. Species-area curve showing species versus distance. Lines around the curves represent ± 1 standard deviation. floodplain and channel margin areas of flowing waters than Galba although collections were made of Stagnicola elodes (Say, 1821) in spring runs in Cumberland County. This species was observed to be more numerous in ditches, seeps, and seasonal pools (Evans and Ray, pers. obs.). Stagnicola catescopium (Say, 1867) was found in Little Sugar Creek in northwestern Pennsylvania, Aughwick Creek in Huntingdon County (central Pennsylvania) and in the lower mainstem Susquehanna River, Lancaster County. Stagnicola emarginata (Say, 1821) were observed in a slow-flowing, muddy section of Brokenstraw Creek in Warren County; museum records were obtained (Northampton County; CMNH #62.12594) that extended the range across the state to the Delaware River system. Planorbidae were among the more commonly encountered groups in this study (Fig. 7); Helisoma anceps (Menke, 1830) and Planorbella trivolvis (Say, 1817) were generally common and distributed nearly statewide. Promenetus exacuous (Say, 1821) was found at one site along a muddy edge in an upper Potomac River tributary in Bedford County. We located a few museum records for this species in the habitats of focus in this study (ANSP #21562, 27284, 122810; CMNH #62.18003; DMNH #081596). Although representatives of Gyraulus were less common than Helisoma, specimens were obtained from drainages across the state. Interestingly, most records of Micromenetus dilatatus (Gould, 1841) were clustered in the funiata and lower Susquehanna drainages although other collections were made outside these areas. Physidae were distributed statewide (Fig. 8). The Pumpkin Physa Physa ancillaria (Say, 1825) had the fewest records among the physids although the species was broadly distributed and located in all but the Lake Erie system. We observed P. acuta and P. gyrina (Say, 1821) in a variety of habitats, ranging from forested headwaters to lowland agri- cultural streams and harsh environments including extremely ephemeral waterbodies (such as small ponds). Within the Hydrobiidae (Fig. 9), Cincinnatia integra (Say, 1829) was collected only from the mainstem Allegheny River and French Creek (Appendix 1 ) in backwater or edge habitats. In spring systems in Centre County, three collections were made of Fontigens nickliniana (I. Lea, 1838). One of these records (from a spring in Bellefonte, Centre County) had been previously reported (Hershler et al. 1990). Other records exist for the species (USNM#121387; 783938; 791479) which extend the range north to Clinton County and south to Huntingdon County, all within the Central Appalachian province. The Blue Ridge Springsnail Fontigens orolibas Hubricht, 1957 was not located in our sampling of springs. Database records from the Smithsonian Museum of Natural History (USNM #522087; 522846; 853173) of Fontigens antroecetes Hubricht, 1972 are actually F. orolibas (fide Robert Hershler, Smithsonian Museum of Natural History). The Shale Pebblesnail Somatogyrus pennsylvanicus (Walker, 1904) was collected from 1 location in the Susquehanna River from the type locality at Columbia, Lancaster County (Walker, 1 904 ) which represents the most recent record for the species in the state. We located Somatogyrus integra (Say, 1829) in 16 collections from the French Creek watershed, SPPRJCH IB unsampled □ 0 55511-4 ITTTTTTlg- 8 I==|9- 14 S3 15- is Figure 4. Species richness by HUC 8 watershed. 140 AMERICAN MALACOLOGICAL BULLETIN 28-1/2-2010 ★ Laevapex fuscus Figure 5. Distribution of Ancylidae in streams and springs of Pennsylvania. Conewango Creek, Brokenstraw Creek, Oil Creek (all Allegheny River system), the Allegheny River mainstem, and Neshannock Creek (Beaver River system). It has been previously reported from the town of Warren (Warren County) by A. E. Ortmann (CMNH #62.7485) which could be a record from either the Allegheny River or Conewango Creek. Despite targeted surveys, we were unable to locate Gillia altilis (I. Lea, 1841) during this study. In Pennsylvania, Gillia has been reported from the middle Delaware River (Northampton County; FMNH #35020) to the lower Susquehanna River (Lancaster County; FMNH #88524). The distributions of Pleuroceridae species fell out by basin (Fig. 10). In the Ohio Basin, Pleurocera acuta Rafinesque, 1841 was found in Muddy Creek, a tributary of the French Creek system, and along the shoreline of Lake Erie. Pleurocera canaliculata (Say, 1821) was reported from the mainstem Ohio River around Phyllis Island in Beaver County (fide Patty Morrison, US Fish and Wildlife Service). Museum records exist from the work of Brooks (1931) from the middle Allegheny River in Warren County (CMNH #62.7902) to the lower Allegheny River (Allegheny County; CMNH #62.661 1 ), the lower Youghiogheny River (CMNH #62.6796), and o 25 so loo a Galba exigua mc=^^1 1011 • Galba modicella ★ Galba obrussa © Galba parva □ Galba rustica • Stagnicola catescopium ★ Stagnicola elodes © Stagnicola emarginata Figure 6. Distribution of Lymnaeidae in streams and springs of Pennsylvania. several records for the mainstem Ohio River. Our records of Elimia livescens (Menke, 1830) were restricted to Beaver, Lawrence, and Crawford Counties although additional work may lead to additional contemporary records. In the Atlantic Slope, Elimia virginica (Say, 1817) and Leptoxis carinata (Bruguiere, 1792) were found across the Susquehanna River basin, with L. carinata also occurring broadly across the upper Potomac. Within the Viviparidae (Fig. 11), Campeloma decisum (Say, 1817) was distributed across the state, but principally in northwestern and south central Pennsylvania. Due to time constraints, we were not able to devote adequate time to search for Lioplax sub carinata (Say, 1816), known principally from the Delaware River basin. Although a record exists for the West Branch Susquehanna River (CMNH #62.7527), the species was not collected in this study. The non-native Chinese Mystery snail Bellamya (= Cipangopaludina) chinensis (Reeve, 1863) was located at 7 sites in the Atlantic Slope, PENNSYLVANIA SNAILS 141 ▲ Gyraulus deflectus • Gyraulus parvus ★ Micromenetus dilatatus 0 Promenetus exacuous Figure 7. Distribution of Planorbidae in streams and springs of Pennsylvania. Delaware, and Ohio Basin. Most notably, B. chinensis was particularly abundant at the mouth of Conodoguinet Creek as well as a site 16 km upstream in the mainstem Susquehanna River in Harrisburg. Results of statistical analyses Stepwise multiple regression models identified 23 variables that appeared to have the strongest influence on gastropod species richness (Table 2). The most important variables were total upstream area, riparian area - barren ground, total number of upstream pollution point sources, percentage of mixed forest cover at the HUC 12 level, and pH. Across all models, the water chemistry model was the strongest at explaining species richness patterns (adjusted r2 = 0.26) with RF3 being the weakest model (adjusted r2 = 0.175). Collectively, multiple regression models were able to explain 55.6% of the variation with regard to snail species richness. NMDS was useful in describing the RF3 variables. As with stepwise multiple regression, the strongest trend from Figure 8. Distribution of Physidae in streams and springs of Penn- sylvania. NMDS ordination appeared to be drainage area, with pulmonate assemblages grouping to lower-order streams and caenogastropod assemblages in higher-order streams (Fig. 12). The NMDS also showed a particular correspondence to larger drainage areas for Elimia virginica and Leptoxis carinata ; these species were very common in a large number of our samples in higher order stream sites in the Atlantic Slope. In addition, the number of upstream road crossings also followed these trends and although this variable was not statistically correlated with drainage area, values increased with drainage area and its importance should be carefully interpreted. Species of Physa were found to be associated with increasing amounts of upstream point source pollution sources, with Physa gyrina showing the clearest trend along this gradient. DISCUSSION We found 37 species from streams, rivers, and spring habitats across the state. However, several species were found in a restricted number of locations. Promenetus exacuous, a species of temporary waterbodies, lakes, and sluggish sections of streams, was undersampled in this study. Additional 142 AMERICAN MALACOLOGICAL BULLETIN 28-1/2-2010 o 25 so ioo ^ Amnicola limosus I km A Cincinnatia integra • Fontigens nickliniana ★ Lyogyrus granum © Somatogyrus integra □ Somatogyrus pennsylvanicus Figure 9. Distribution of Hydrobiidae in streams and springs of Pennsylvania. sampling targeting those habitats would likely produce more records. The first modern record for Somatogyrus pennsylvanicus is in this study. It was located during a search for an older record of Gillia altilis from the Susquehanna River (Florida Museum of Natural History 88524). Specimens were found in only 1 location in the river; given the length of the mainstem Susquehanna River within the state, more field sampling is needed as well as demographic studies to derive population estimates for S. pennsylvanicus. Hershler et al. (1990) noted that Fontigens orolibas, which was not located in this study, was known from springs at elevations above 182 meters in Virginia, and also in caves. Few specimens of Pleurocera, a genus restricted to the Mississippi and Ohio River basin (Burch 1989), were found. Additional sampling efforts in the lower Allegheny River, upper Ohio River, and Monongahela Rivers, using scuba, would allow better sampling of this genus in the state. Somatogyrus integra, a species of potential conservation concern in the Ohio Basin (unpublished list, Ohio River ▲ Elimia livescens • Elimia virginica ★ Pleurocera canaliculata Figure 10. Distribution of Pleuroceridae in streams and springs of Pennsylvania. Valley Ecosystem Recovery Team) and listed as globally vulnerable (G3) by NatureServe (2008), was located in several streams of the Allegheny River basin, frequently in large numbers. Additional sampling in other Allegheny River system streams could yield new specimen records. Given that this species was often encountered in large numbers, and that hydrobiids are easy to overlook in surveys, this species might not be as rare as reported in the Ohio Basin. We recommend that this species be targeted for additional survey work across the upper Ohio River basin to refine its global conservation status. Low species diversity was observed in Lake Erie tributary streams, as most sampling was from bedrock-bottomed shallow streams which typically provide limited freshwater snail habitat. Additional stream sampling in Conneaut Creek, a larger watershed that exhibits sand/gravel glacial deposits, stable substrates, and pool development, would likely reveal new species not reported from the watershed. Because of the recent work of Walther et al. (2006) resolving the taxonomy of the genus Laevapex, specimens obtained from the Allegheny River in this study initially identified as Laevapex diaphanus are now considered Laevapex PENNSYLVANIA SNAILS 143 0 25 so loo • Campeloma decisum ★ Bellamya chinensis Figure II. Distribution of Viviparidae in streams and springs of Pennsylvania. fuscus (C. B. Adams, 1841). The degree of shell variation in North American ancylids has recently been shown to be considerable, varying not only temporally but also spatially (McMahon 2004). Albrecht et al. (2007) contended that the Ancylidae is a polyphyletic family that falls basally within the Planorbidae. Taxonomy of the various species of Galba is muddled. In a recent treatment of the freshwater gastropods of Virginia, Stewart and Dillon (2004) grouped all putative species under Fossaria (= Galba). While we followed currently accepted species names for members of the genus, the use of modern molecular taxonomic techniques as well as anatomical comparisons would allow better estimates of the true diversity of the freshwater gastropods of Pennsylvania. Bellamya chinensis has been documented from the Delaware Basin (ANSP #392419, 401246; this study) and was sampled in this study from the Susquehanna River around Harrisburg, Dauphin County. It was also observed in several reservoirs across the state. The first records of this species in the United States were from the San Francisco Bay area in the early 20th century (Hannibal 1911). Johnson (1915) reported the first eastern United States records, and the species was first reported from Pennsylvania by Richards and Adams (1929). In stream ecosystems, little is known regarding the effects that this species can have on native fauna, underscoring the need for basic monitoring and inventories to locate other populations. Other exotic freshwater snails previously doc- umented in Pennsylvania lotic and spring systems, including Physa skinneri Taylor, 1954 (Smithsonian #375998) and Radix auricularia (Linnaeus, 1758) (Smithsonian #47868), were not located in this study although these species were not speci- fically targeted. Future studies should focus on refining the knowledge of invasive species within the state. Recent work has documented the presence of the highly invasive New Zealand Mud Snail Potamopyrgus antipodarum (J. E. Gray, 1853) at Presque Isle in Lake Erie (Levri etal. 2007) Table 2. Variables used for developing stepwise multiple regression models, with species richness used as the dependent variable. Only the variables retained in the final models are shown. All tests were evaluated at the a = 0.05 level of significance. NS, not significant. Variables E-statistic E-value Substrate riffle/run 5.38 0.02 Sediment deposition - high 8.21 0.005 gradient streams Channel sinuosity 5.69 0.01 Total upstream drainage area 26.337 <0.001 Degree of channel alteration 2.93 NS Riparian zone vegetation 7.88 0.006 Riparian zone width 3.59 NS Riparian area - barren 12.35 0.001 Riparian area - forest 1.770 NS Riparian area - wetland 2.217 NS Upstream dam density 0.797 NS Number of upstream pollution 27.899 <0.001 point sources Number of upstream road 5.276 0.02 stream crossings % calcareous geology - HUC 12 4.062 0.04 % unconsolidated geology - HUC 12 0.938 NS % evergreen forest - HUC 12 1.298 NS % mixed forest - HUC 12 17.452 <0.001 % rowcrop - HUC 12 2.312 NS % emergent wetland in 1.139 NS watershed - HUC 12 % non-row crop agriculture - HUC 12 2.598 NS pH 18.79 <0.001 Dissolved oxygen 3.5 NS Specific conductance 2.71 NS in Erie County. Additional records now exist from Lake Ontario (Zaranko et al. 1997) and Lake Superior (Grigorovich et al. 2003). This finding has significant biodiversity implications, given the negative effects that this species has had on stream macroinvertebrates (Kerans et al. 2005) in the western United States. Efforts to educate the public about the New Zealand Mud Snail and to monitor streams in this region for its presence will be needed to guard against further spread of the species. The Northwestern Glaciated Plateau region of the northwestern section of the state, heavily influenced by the Wisconsin Glaciation approx. 17,000 years ago (Sevon and Fleeger 2002), exhibited the greatest species richness. The wide array of habitat types and calcareous geology in the Northwestern Glaciated Plateau is important to the rich diversity of this region. Given the unique gastropods, freshwater mussels, and fish assemblages, we recommend the Middle Allegheny River system and larger tributaries such as Conewango Creek and French Creek be priorities for conservation. 144 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Higher species richness of snails was associated with fewer upstream point sources of pollution, fewer upstream road crossings, a higher percent of mixed forest in the watershed, and lower sediment deposition. Although our overall model for habitat measures was weak (r2 = 0.121), we did not examine several potentially important microhabitat or local scale measures in detail. Lor instance, we observed that the amount of woody debris and microhabitats increased snail diversity although this relationship could be explained in terms of upstream drainage area, which was also found to be statistically significant. Harman (1972) found significant correlations between gastropod species richness and substrate diversity, which also occur in many other invertebrate groups (Hynes 1970). Regarding water chemistry, pH was a strong predictor of species richness, as previously reported for freshwater gastropods (Russell-Hunter 1978, Jokinen 1987, Pip 1987, Miller et al. 2000). Pyron et al. (2009) determined that most species were found at sites with pH values ranging from 7.8 to 8. Temperature, a variable we expected to be important in influencing species richness, was not significant. Greater sampling intensity in warmer, lower-gradient stream systems could possibly reveal the importance of this variable, considered influential for freshwater snails (Aldrich 1983, McMahon 1983). Another highly significant measure was total upstream drainage area. Total drainage area is also an important predictor of species richness in other studies of snails in streams and lakes (Dillon and Benfield 1982, Jokinen 1987, Prezant and Chapman 2004) and in freshwater mussels (Ortmann 1920, Strayer 1983). We found headwaters domi- nated by pulmonate species, and larger streams dominated by CM w x < 2-dimensional solution Final minimum stress = 18.9 Phygyr Gyrpar A M Phyanc Mlcdila L Ferrriv |gV Phyacut cpiantn. — ■" v? Lyogran Laefus Lithobo Amnlimo Somatoint Galba _G Gyrdefl M IVI \ ^ Helancep Lepcar Elimvir Elimlive Bellchi Campdec Psecoll Ferrpara Increasing Drainage Area Axis 1 Increasing riparian forest in HUC 12 Figure 12. Plot of taxa against retained RF3 variables area using NMDS. PENNSYLVANIA SNAILS 145 pleurocerids and viviparids, as reported in the literature (Brown et al. 1998, Minton et al. 2008, Pyron et al. 2009). Barnese et al. (1990) documented resource partitioning between pulmonate and prosobranch snails, while Johnson and Brown (1997) found that current regime and light penetration were important in Elimia semicarinata (Say, 1829). Other potentially important stream measures for snail community composition include sheer stress (Gore 1983) and spates (Stewart and Garcia 2002) for pleurocerid snails. We also discovered that calcareous geology was a significant predictor of species richness. Huryn et al. (1994) found Elimia to be absent from streams with alkalinity values below 20 mg/L CaC03, while Pip (1987) found alkalinity to be significant in explaining species richness of gastropods. However, 0kland (1983) and Miller et al. (2000) found snails in very low calcium environments in lakes. Snails must actively transport ions in low calcium environments, which can limit production (Allan 1995). Increased diversity with increasing amounts of riparian forest may be attributed to greater forest cover found in smaller watersheds across the state. We were often able to collect Galba within small watersheds in streams that maintained an active floodplain connection, in many cases connected to intact floodplain forest. It was also associated with Ohio River species of Hydrobiidae, which could underscore the importance of forest cover to this group. Increased forest cover could maintain stream bank integrity, reduce sedimentation, shade streams, mediate spates, and increase woody debris. Several snail taxa may actually respond to stable local environments. The increased amounts of point source pollution in association with Physidae also corresponds to reported pollution tolerances for the group (Barbour et al. 1999). Ecoregions have also been found to be a useful variable for predicting vertebrate (McCormick et al. 2000) and invertebrate (Johnson 2000) diversity. We recommend future studies of ecoregion associations for aquatic gastropods, coupled with variables significant in this and other studies, as suggested by Sandin and Johnson (2000). In the Northern Appalachian Plateau (northcentral region of the state), we suspect acid precipitation may depress freshwater snail populations; areas within the northwestern portion of the Susquehanna Basin had no species of freshwater snails. While this could have been the result of inadequate sampling, most streams in this region are of sandstone geology with poor buffering capacity. We located sampling sites with high amounts of forest cover, no acid mine drainage impacts, little human development, but few snails despite available habitat. Observed pH values can drop into the low 4 range after rain events, and chronic impacts of acid precipitation include depleted cations and elevated levels of S04and N03 (DeWalle and Swistock 1994, Driscoll et al. 2001) The same reduced diversity has also been determined for fishes (Van Sickle et al. 1996, McClurg et al. 2007). Based on our work and examination of museum records, we recommend listing 7 species of freshwater snails from lotic and spring environments as species of conservation concern (Table 3). Due to limited sampling efforts for the mainstem Delaware River, we did not feel that adequate survey data were available to give a conservation status recommendation for Lioplax subcarinata. A comprehensive survey of lentic taxa is sorely needed in Pennsylvania as well as a survey of the state’s stygobitic gastropod fauna and additional sampling of springs. Our inclusion here of Fontigens orolibas (Table 3) is based on only 1 museum record for the species from Pennsylvania (Hershler et al. 1990); we did not detect this species in our sampling, but given its peripheral range in the state it should be considered for conservation. Leptoxis dilatata (Conrad, 1835) also has a peripheral range in Pennsylvania, and given the damage caused by coal mining and impoundment of the Lower Cheat River, it should be considered endangered, if not extirpated in Pennsylvania. Schwartz and Meredith (1962) also did not collect this species in the state. We hope this publication will be useful for conservation and planning efforts. Nightingale et al. (2004) for example Table 3. Pennsylvania Species of Conservation Concern from lotic systems and springs based on results from this study as compared to historical records. Species Family Current global rank’*' Recommended state rank* Cincinnatia integra Hydrobiidae G5 S3S4 - between Special Concern and Apparently Stable Fontigens orolibas Hydrobiidae G3 S1S2 - S1S2 - Between Endangered and Threatened Gillia altilis Hydrobiidae G5 S1S2 - Between Endangered and Threatened Somatogyrus integra Hydrobiidae G3 S2 - Threatened Somatogyrus pennsylvanicus Hydrobiidae G3 SI - Endangered Leptoxis dilatata Pleuroceridae G3 SI - Endangered Pleurocera acuta Pleuroceridae G5 S2 - Threatened * using conservation ranking methodology of NatureServe (2008). 146 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2 -2010 focused on using survey information to delineate natural community groupings and associated abiotic and biotic factors in Pennsylvania. According to NatureServe (2008), more than 275 snail species in North America are considered G1 (Globally Critically Imperiled) or G2 (Globally Threatened). ACKNOWLEDGMENTS We would like to thank the Pennsylvania Department of Conservation and Natural Resources Wild Resources Conser- vation Program, which provided funding. We would like to thank Charles Bier, Amy Bush, Katrina Morris, Patricia Morrison, Betsy Nightingale, and Tamara Smith for con- tributing specimens and locality information. Thanks to Delaware Museum of Natural History, Carnegie Museum of Natural History, and the Academy of Natural Sciences in Philadelphia for sharing museum records; special thanks to Tim Pearce at Carnegie Museum for allowing access to the specimen collection at the Carnegie Museum. Brad Georgic of Western Pennsylvania Conservancy assisted in the prepa- ration of maps. The comments from two anonymous reviewers greatly improved this manuscript. LITERATURE CITED Albrecht, C., K. Kuhn, and B. Streit. 2007. A molecular phylogeny of Planorboidea (Gastropoda: Pulmonata): Insights from en- hanced taxon sampling. Zoologica Scripta 36: 27-39. Aldrich, D. 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Lydeard. 2007. A molecular phylogeny of Physidae (Gastropoda: Basommatophora) using mitochondri- al DNA sequences. Journal ofMolluscan Studies 73: 1-17. Zaranko, D.T., D. G. Farara, and F. G. Thompson. 1997. Another exotic mollusk in the Laurentian Great Lakes: The New Zealand native Potamopyrgus antipodarum (Gray, 1843) (Gastropoda, Hydrobiidae). Canadian Journal of Fisheries and Aquatic Sci- ences 54: 809-814. Submitted: 14 March 2008; accepted: 14 September 2009; final revisions received: 24 October 2009 PENNSYLVANIA SNAILS 149 Appendix 1. Species occurrences by USGS 8-digit HUC. Species HUC Amnicola limosus Upper Delaware R., Lehigh R., Upper Juniata R., Lower Juniata R., Middle Allegheny R.-Tionesta Cr., French Cr., Middle Allegheny R.-Redbank Cr., Lower Monongahela R., Shenango R. Bellamya chinensis Campeloma decisum Middle Delaware R.-Mongaup Cr.-Brodhead Cr., Lower Susquehanna R.-Swatara Cr. Lackawaxen R., Middle Delaware R.-Mongaup Cr.-Brodhead Cr., Upper Susquehanna R.-Lackawanna R., Lower luniata R., Lower Susquehanna R.-Swatara Cr., Lower Susquehanna R., Conococheague Cr.-Opequon Cr., Middle Allegheny R.-Tionesta Cr., French Cr., Middle Allegheny R.-Redbank Cr. Cincinnatia integra Elimia livescens Elimia virginica Middle Allegheny R.-Tionesta Cr., French Cr. French Cr., Shenango R., Connoquenessing Cr. Schuylkill R., Upper Susquehanna R.-Tunkhannock Cr., Upper Susquehanna R.-Lackawanna R., Lower Susquehanna R.-Penns Cr., Upper Juniata R., Lower Juniata R., Lower Susquehanna R.-Swatara Cr., Lower Susquehanna R. Ferrissia fragilis Ferrissia parallelus Ferrissia rivularis French Cr., Schuylkill R. Lehigh R., Monocacy R. Upper Delaware R., Lackawaxen R., Lehigh R., Schuylkill R., Upper Susquehanna R.-Tunkhannock Cr., Upper Susquehanna R.-Lackawanna R„ Lower West Branch Susquehanna R., Lower Susquehanna R.-Penns Cr., Upper Juniata R., Raystown Br„ Lower Juniata R., Lower Susquehanna R.-Swatara Cr., Lower Susquehanna R., North Branch Potomac R., Cacapon R.-Town Cr., Conococheague Cr.-Opequon Cr., Chautauqua Lake- Conneaut Cr., Upper Allegheny R., Conewango Cr., Middle Allegheny R.-Tionesta Cr., French Cr., Clarion R„ Middle Allegheny R.-Redbank Cr., Conemaugh R., Lower Allegheny R., Lower Monongahela R., Youghiogheny R., Shenango R., Connoquenessing Cr., Upper Ohio R. -Wheeling Cr. Fontigens nickliniana Galba exigua Galba modicella Bald Eagle Cr., Lower Susquehanna R.-Penns Cr. Upper Ohio R. -Wheeling Cr. Upper Juniata R., Lower Juniata R., Lower Susquehanna R.-Swatara Cr., Upper Allegheny R., Middle Allegheny R.-Tionesta Cr., French Cr., Clarion R., Lower Monongahela R., Youghiogheny R., Connoquenessing Cr. Galba obrussa Upper Juniata R., Lower Juniata R., Middle Allegheny R.-Tionesta Cr., Middle Allegheny R.-Redbank Cr., Upper Ohio R.-Wheeling Cr. Galba parva Galba rustica Gyraulus deflectus Lower Juniata R., Conemaugh R. Clarion R. Middle Delaware R.-Mongaup Cr.-Brodhead Cr., Middle Allegheny R.-Tionesta Cr., Shenango R. Gyraulus parvus Helisoma anceps Lehigh R., Schuylkill R., Upper Allegheny R., French Cr. Middle Delaware R.-Mongaup Cr.-Brodhead Cr., Crosswicks Cr.-Neshaminy Cr., Upper Susquehanna R., Pine Cr., Lower West Branch Susquehanna R., Lower Susquehanna R.-Penns Cr., Raystown Br., Lower Juniata R., North Branch Potomac R., Cacapon R.-Town Cr., Conococheague Cr.-Opequon Cr., Upper Allegheny R., Conewango Cr., Middle Allegheny R.-Tionesta Cr., French Cr., Clarion R., Middle Allegheny R.-Redbank Cr., Conemaugh R., Youghiogheny R. Laevapex fuscus Lower Susquehanna R.-Swatara Cr., Cacapon R.-Town Cr., Upper Allegheny R., Conewango Cr., Middle Allegheny R.-Redbank Cr., Middle Allegheny R.-Tionesta Cr., French Cr., Clarion R. Leptoxis carinata Upper Susquehanna R., Upper Susquehanna R.-Tunkhannock Cr., Upper Susquehanna R.-Lackawanna R., Lower Susquehanna R.-Penns Cr., Upper Juniata R., Raystown Br., Lower Juniata R., Lower Susquehanna R.-Swatara Cr., Lower Susquehanna R„ North Lithasia obovata Branch Potomac R., Conococheague Cr.-Opequon Cr., Monocacy R. Upper Allegheny R., Conewango Cr., Middle Allegheny R.-Tionesta Cr., French Cr., Middle Allegheny R.-Redbank Cr., Shenango R., Connoquenessing Cr. Lyogyrus granum Upper Delaware R., Lackawaxen R., Middle Delaware R.-Mongaup Cr.-Brodhead Cr., Crosswicks Cr.-Neshaminy Cr., Lower Susquehanna R.-Swatara Cr. 150 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2 -2010 Appendix 1. (continued) Species HUC Micromenetus dilatatus Upper West Branch Susquehanna R., Upper Juniata R., Raystown Br., Lower Juniata R., Cacapon R.-Town Cr., Youghiogheny R., Shenango R. Physa acuta Lackawaxen R., Middle Delaware R.-Mongaup Cr.-Brodhead Cr., Crosswicks Cr.-Neshaminy Cr., Brandywine Cr.-Christina R., Tioga R., Upper Susquehanna R.-Tunkhannock Cr., Upper Susquehanna R.-Lackawanna R., Pine Cr., Lower West Branch Susquehanna R., Lower Susquehanna R.-Penns Cr., Upper Juniata R., Raystown Br., Lower Juniata R., Lower Susquehanna R.-Swatara Cr., Lower Susquehanna R., Conococheague Cr.-Opequon Cr., Monocacy R., Chautauqua Lake-Conneaut Cr., Upper Allegheny R., Conewango Cr., Middle Allegheny R.-Tionesta Cr., French Cr., Clarion R., Middle Allegheny R.-Redbank Cr., Lower Allegheny R., Lower Monongahela R., Youghiogheny R., Shenango R., Connoquenessing, Upper Ohio R.-Wheeling Cr. Physa ancillaria Upper Delaware R., Middle Delaware R.-Mongaup Cr.-Brodhead Cr., Raystown Br., Lower Susquehanna R.-Swatara Cr., Conococheague Cr.-Opequon Cr., Upper Ohio R. Physa gyrina Upper Delaware R., Lackawaxen R., Lower Delaware R., Schuylkill R., Upper Susquehanna R.-Lackawanna R., Upper Juniata R., Raystown Br. Br., Lower Juniata R., Lower Susquehanna R.-Swatara Cr., North Branch Potomac R., Cacapon R.-Town Cr., Middle Allegheny R.-Tionesta Cr., French Cr., Middle Allegheny R.-Redbank Cr., Conemaugh R., Lower Monongahela R., Youghiogheny R., Shenango R., Upper Ohio R.-Wheeling Cr. Planorbella campanulata Planorbella trivolvis Middle Delaware R.-Mongaup Cr.-Brodhead Cr., Crosswicks Cr.-Neshaminy Cr. Lower West Branch Susquehanna R., Lower Juniata R., Lower Susquehanna R.-Swatara Cr., Lower Susquehanna R., Upper Allegheny R., Middle Allegheny R -Tionesta Cr., French Cr., Lower Monongahela R., Shenango R. Pleurocera acuta Pleurocera canaliculata Promenetus exacuous Pseudosuccinea columella French Cr. Upper Ohio R., French Cr. Cacapon R.-Town Cr. Lehigh R., Upper Susquehanna R.-Tunkhannock Cr., Lower West Branch Susquehanna R., Lower Susquehanna R., French Cr. Somatogyrus Integra Upper Allegheny R., Conewango Cr., Middle Allegheny R.-Tionesta Cr., French Cr., Middle Allegheny R.-Redbank Cr. Somatogyrus pennsylvanicus Stagnicola catescopium Stagnicola elodes Stagnicola emarginata Lower Susquehanna R. Lower Juniata R., Lower Susquehanna R., French Cr. Lower Susquehanna R.-Swatara Cr. Upper Allegheny R. Amer. Malac. Bull. 28: 151-158 (2010) Fish hosts of the Carolina heelsplitter ( Lasmigona decorata ), a federally endangered freshwater mussel (Bivalvia: Unionidae) Chris B. Eads1, Robert B. Bringolf2,Renae D. Greiner1, Arthur E. Bogan3, and Jay F. Levine1 1 Aquatic Epidemiology and Conservation Laboratory, Department of Population Health and Pathobiology, College of Veterinary Medicine, North Carolina State University, Raleigh, North Carolina 27606, U.S.A 2 Warnell School of Forestry and Natural Resources, University of Georgia, Athens, Georgia 30602, U.S.A. 3 North Carolina State Museum of Natural Sciences, Raleigh, North Carolina 27607, U.S.A. Corresponding author: Jay_Levine@ncsu.edu Abstract: Two laboratory trials were conducted to determine the required host fish for the Carolina heelsplitter ( Lasmigona decorata (Lea, 1852)), an endangered freshwater mussel (Unionidae). The first trial used glochidia from a female collected from the Yadkin-Pee Dee River basin, and the second trial used the glochidia of an adult collected from the Catawba River basin. Two different techniques were utilized for glochidia extraction: flushing and serotonin-induced release. The first female tested (Yadkin-Pee Dee) packaged most of its glochidia attached to unfertilized eggs, and extraction of glochidia by flushing the marsupia with a syringe yielded few glochidia and caused extensive tearing of the gill tissue. In the second trial (Catawba) the female was immersed in 500 mg/L serotonin creatinine sulfate, and the glochidia were readily released without injury to the adult. Several species of minnows (Cyprinidae) from both basins served as hosts. Some sunfish species (Centrarchidae) supported transformation of a few juveniles, but differences in transformation success were observed between the two basins on these species. Key words: Cyprinidae, glochidia, serotonin, conglutinate, Unionidae Artificial propagation and culture has been identified as an important tool for conservation of native freshwater mussels (Unionidae) in the United States (National Native Mussel Conservation Committee 1998). The larval stage (glochidia) of most species of freshwater mussels must attach to the gills or fins of a fish to complete their metamorphosis into juveniles (Williams et al. 2008). Both male and female adult mussels and the appropriate fish hosts must be avail- able for reproduction and the recruitment of new juveniles into the population. This obligate, parasitic relationship between mussel larvae and fish host can be a bottleneck to population expansion. As mussel populations decline, the probability of successful mating and subsequent population recruitment also declines (McLain and Ross 2005). Union- ids reared in captivity can be used to augment declining populations, and moving this process to a controlled envi- ronment for artificial propagation can help bypass the bottle- neck. Captive reared individuals may also serve as a resource for sentinel field or controlled laboratory studies and provide animals for educational exhibits. One of the obstacles to the propagation of mussels native to the Atlantic slope is that the required host fishes are unknown for many species (Bogan 2002). Fish hosts can be determined in a variety of ways. Fish can be collected from the wild during the time of year they are most likely to be infected with glochidia from a specific species of mussel (Zale and Neves 1982a, 1982b). Fish gills and fins are then exam- ined closely for glochidial attachment, and those glochidia are then identified by taking physical measurements of the shell (Surber 1912) or through molecular techniques (White et al. 1994, Gerke and Tiedemann 2001, Kneeland and Rhymer 2007). While this method provides documentation of attach- ment to a specific species in the wild, attachment to a fish does not necessarily reflect successful transformation. Mus- sels commonly have some host species that yield only a few transformed individuals, in contrast to other more effective fish-hosts that successfully support the transformation of large numbers of glochidia to juvenile mussels (Khym and Layzer 2000, Rogers et al. 2001, Van Snik Gray et al. 2002). Wild-caught fish infected with glochidia can also be held in aquaria until transformation is completed (Surber 1913, Coker et al. 1921). When fish are held individually in aquaria or as a species cohort, the ability of a specific species to sup- port transformation is confirmed. However, both field-based methods may require extensive fish sampling to find the glochidia of a rare mussel species because the percentage of fish infected with glochidia approaches 0% at sites with low mussel densities (McLain and Ross 2005). Laboratory-based host determinations are conducted by acquiring non-infested fish that co-occur with a mussel spe- cies, exposing them to glochidia obtained from gravid females, and harvesting juveniles after transformation is complete (Zale and Neves 1982a). After transformation, glochidia that attach to individually housed cohorts or 151 152 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 individuals can be collected, and the relative ability of different species of fish to support transformation can be compared. Laboratory-based infestations may not necessarily reflect field conditions, but they confirm transformation — a necessary step before proceeding with propagation. The Carolina heelsplitter ( Lasmigona decorata (Lea, 1852)) was historically found from small streams to large riv- ers and even millponds in the Pee Dee, Catawba, Savannah, and the Saluda River systems (Bogan 2008). Now federally endangered, it has been restricted to a few fragmented popu- lations, primarily in headwater streams, in North and South Carolina (U.S. Pish and Wildlife Service 1996). The remain- ing populations in North Carolina and some in South Caro- lina lie in head-water streams near the rapidly expanding Charlotte, North Carolina metropolitan area. As outlying areas have been developed, changing land-use practices have altered stream habitat and degraded water quality. These watershed practices and a decade long drought have imper- iled L. decorata populations, and this rare mussel now risks being extirpated from North Carolina (John Pridell, U.S. Pish and Wildlife Service, pers. comm.). The recovery plan for Lasmigona decorata called for life history research as well as augmentation and reintroduction efforts (U.S. Fish and Wildlife Service 1996). Efforts to pre- serve and potentially augment populations through captive culture and propagation are dependent on our ability to select the most appropriate fish species to serve as fish hosts. Accordingly, we examined the ability of 27 species of fish, representing six families, from two North Carolina river basins to serve as potential hosts for captive L. decorata cul- ture and propagation. MATERIALS AND METHODS Collection and holding of adult Lasmigona decorata In August 2006, five Lasmigona decorata were collected from Duck Creek (Yadkin-Pee Dee River Basin) in Union County, North Carolina and immediately transported in a cooler wrapped in a wet cloth to the Table Rock Fish Hatch- ery near Morganton, North Carolina, where we maintained four troughs for culturing mussels. The hatchery was located on Irish Creek (Catawba River Basin), and the main water supply was a small (1.4 hectare) reservoir on the creek. Water pumped into the hatchery was mixed with water from a warmer, more nutrient-rich culture pond (0.4 hectare) and was piped continuously through the mussel culture trough. After exiting the trough, water flowed into a concrete raceway (4.5 m long x 1 m wide x 0.9 m deep), to allow settling of any larvae released by the mussels. The water was then pumped through a sand filter containing commercial filter media fil- tering down to 20-40 pm (Aquatic Ecosystems, Inc., Apopka, Florida). This prevented potential escape of mussels and their larvae not indigenous to the Catawba River Basin that were reared in the hatchery. In January 2007, two of the five adults were found to be gravid. One of those adults was transported in an aerated cooler of water to our propagation laboratory at North Carolina State University to be used as the source of glochidia for the initial host trial. In February 2007, three adult Lasmigona decorata were collected in Sixmile Creek (Catawba River Basin) on the Union/Mecklenburg County line in North Carolina. One of those individuals was gravid and was maintained in the prop- agation laboratory for one week at 6 ± 1 °C before being used as the source of glochidia for the second trial. Voucher repre- sentatives of both adults and glochidia of both the Duck Creek and Sixmile Creek stock were deposited in the North Carolina Museum of Natural Sciences, Raleigh, North Carolina. Host fish trials Yadkin-Pee Dee River Basin On 18 January 2007, potential host fish representing 20 species (Table 1) were collected by seine and backpack electro- fishing from Big Bear and Island Creeks in Stanly County, North Carolina and Irish Buffalo Creek in Cabarrus County, North Carolina (all in the Yadkin-Pee Dee River Basin). These streams were selected because they held few to no mussels, with no Lasmigona decorata , and naive fish of a variety of species could be easily collected. Fish were transported in aerated cool- ers of stream water back to the propagation laboratory and maintained in various aquaria at 13 ± 1 °C. In January 2007, we extracted the glochidia from the single gravid adult from the Duck Creek stock by flushing the marsupium with a syringe filled with water from the mussel’s tank. Mature glochidia were adhered to unfertilized eggs, forming conglutinates (Fig. 1). The size of these conglutinates made extraction somewhat dif- ficult, and the repeated flushing necessary to complete the task caused some trauma to the gill tissue. This animal died within a week after glochidial extraction, and we believe it was a result of the gill trauma. We freed glochidia from conglutinates by repeatedly drawing the packets in and out of a plastic, 1-ml pipette. A small subsample (<100 individuals) of larvae were then tested for viability by exposing them to a salt solution (Zale and Neves 1982a). The brood was deemed viable when 100% of glochidia snapped shut in response to the salt solu- tion. All fish were then placed together with the glochidia in approx. 12 liters of water for 30 minutes, and strong aeration was used to keep glochidia in suspension (approx. 2000 glochidia/liter). After infestation, fish were divided into differ- ent aquaria by species and maintained at 13 ± 1 °C. We used up to four aquarium replicates for some fish species when enough individuals of that species were available. We siphoned FISH HOSTS OF LASMIGONA DECORATA 153 Table 1. Yadkin-Pee Dee River Basin host trial for Lasmigona decorata from Duck Creek, Union County, North Carolina. No. of aquaria No. of juveniles Mean percentage of Common name (species) No. of fish tested replicates produced glochidia transformed Aphredoderidae Pirate perch ( Aphredoderus sayanus ) 5 3 1 0.9 Catostomidae Creek chubsucker ( Erimyzon oblongus ) 5 2 0 0.0 Centrarchidae Largemouth bass ( Micropterus salmoides) 3 3 1 4.8 Redbreast sunfish ( Lepomis auritus) 3 3 0 0.0 Green sunfish ( Lepomis cyanellus ) 3 3 0 0.0 Bluegill ( Lepomis macrochirus) 15 3 185 23.3 Redear sunfish ( Lepomis microlophus) 1 1 0 0.0 Cyprinidae Satinfin shiner ( Cyprinella analostana) 1 1 6 54.5 Bluehead chub ( Nocomis leptocephalus) 4 2 67 73.7 Golden shiner ( Notemigonus crysoleucas) 1 1 64 64.0 Whitemouth shiner ( Notropis alborus) 9 3 2 16.7 Highfin shiner ( Notropis altipinnis) 15 3 23 44.0 Redlip shiner ( Notropis chiliticus) 7 2 1 3.1 Spottail shiner ( Notropis hudsonius) 6 3 66 32.2 Ictaluridae Yellow bullhead ( Ameiurus natalis ) 1 1 2 10.5 Margined madtom ( Noturus insignis) 6 2 0 0.0 Percidae Carolina darter ( Etheostoma collis ) 1 i 0 0.0 Fantail darter (Etheostoma flabellare) 12 4 1 0.1 Tessellated darter ( Etheostoma olmstedi ) 12 2 1 0.4 Piedmont darter ( Percina crassa) 5 2 0 0.0 the bottom of each aquarium routinely (every 1-2 days) through a 150-mm mesh sieve to capture dead glochidia and transformed juvenile mussels for enumeration. Transforma- tion success for each replicate was calculated by dividing the number of transformed juveniles by the sum of transformed juveniles and sloughed glochidia. Individuals were observed under a stereomicroscope and considered to be transformed if two adductor muscles were visible or if there was foot movement. Catawba River Basin On 1 March 2007, we collected potential host fish repre- senting 17 species by seine and backpack electrofishing in the Twelvemile Creek system (Catawba River Basin) in Union County, North Carolina (Table 2). As with the Yadkin-Pee Dee trial, these streams were selected because they contained few mussels. Lasmigona decorata did not occur in the stream, and naive fish of a variety of species could be easily collected. Fish were transported to the laboratory in aerated coolers of stream water and maintained in aquaria at 13 ± 1 °C. Because of the mortality caused by flushing of the marsupia in the initial trial, we elected to use chemical methods to induce release of glochidia in this trial. We immersed the mussel in 500 mg/L serotonin creatinine sulfate (Acros Organics, Geel, Belgium, 99% purity). The adult began releasing a few glochidia after 1 hour, but not nearly enough to conduct a host fish trial. After 3 hours, with some swelling of the mus- sel’s foot tissue and few glochidia released, the mussel was removed from the serotonin and placed in chilled (13 °C) fresh water overnight. By the following morning (6 March 154 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 1. Viable glochidia of Lasmigona decorata adhered to unde- veloped eggs. 2007), the entire brood had been released into the aquarium. Viability of the glochidia was again confirmed (100%) by exposing a small subsample of the brood to a saline solution as described above. Host fish and glochidia were then placed in 12 liters of water for 30 minutes and aerated vigorously to keep the glochidia in suspension (approx. 2500 glochidia/ liter). Fish were then divided into separate aquaria by species and maintained at 13 ± 1 °C. We siphoned the bottom of each aquarium routinely (every 1-2 days) through a 150-mm mesh sieve to check for transformed juvenile mussels as described above for the Yadkin-Pee Dee trial. Because sloughed glochid- ia were not counted in this trial, we could not calculate the percentage of glochidia that transformed. RESULTS Description of glochidia Glochidia were hooked, relatively large (mean = 305 ± 5 pm long x 256 ± 5 pm high), amber in color, and had the sub-triangular shape typical of the subfamily Anodontinae. Table 2. Catawba River Basin host trial for Lasmigona decorata from Sixmile Creek, Union/Mecklenburg County, North Carolina. No. of aquaria No. of juveniles Mean no. of Common name (species) No. of fish tested replicates produced juveniles per fish Catostomidae White sucker ( Catostomus commersonii ) 2 2 0 0 Creek chubsucker ( Erimyzon oblongus ) 1 1 0 0 Striped jumprock ( Scartomyzon rupiscartes ) 1 1 0 0 Centrarchidae Warmouth (Chaenobryttus gulosus) 1 1 19 19 Redbreast sunfish ( Lepomis auritus) 1 1 1 0.1 Green sunfish ( Lepomis cyanellus ) 1 1 7 7 Bluegill ( Lepomis macrochirus) 2 2 0 0 Redear sunfish ( Lepomis microlophus) 1 1 0 0 Cyprinidae Rosyside dace {Clinostomus funduloides) 3 1 37 12.3 Whitefin shiner ( Cyprinella nivea) 2 1 29 14.5 Bluehead chub ( Nocomis leptocephalus ) 9 5 160 30.6 Golden shiner ( Notemigonus crysoleucas) 32 5 541 20.7 Sandbar shiner ( Notropis scepticus) 13 5 280 23.5 Creek chub ( Semotilus atromaculatus) 1 1 41 41 Ictaluridae Flat bullhead ( Ameiurus platycephalus) 2 3 1 0.3 Margined madtom ( Noturus insignis) 2 1 0 0 Percidae Tessellated darter ( Etheostoma olmstedi) 3 1 1 0.3 FISH HOSTS OF LASMIGONA DECORATA 155 Table 3. Time to complete transformation for each species of fish tested in trial 1 (Yadkin- Pee Dee River Basin). Common name (species) N Median transformation time with quartiles (days) Range (days) Bluegill ( Lepomis macrochirus) 185 27 (24, 30) 22-44 Satinfin shiner ( Cyprinella analostana) 6 35 (30,37) 29-41 Bluehead chub ( Nocomis leptocephalus) 67 34 (34, 36) 24-44 Golden shiner ( Notemigonus crysoleucas) 64 36 (34, 38) 27-44 Highfin shiner ( Notropis altipinnis) 23 30 (29, 35) 24-41 Spotfin shiner ( Notropis hudsonius) 66 29 (27, 30) 22-41 Most of the glochidia extracted by needle from the Yadkin-Pee Dee individual came out in semi-rectangular and relatively flat masses of eggs and glochidia (approx. 5-20 mm in size). These conglutinates were neutrally buoyant. The large glo- chidia were readily visible to the naked eye when viewed against the whitish, non-viable eggs to which they were attached (Fig. 1). The gravid adult from the Catawba River Basin that released its glochidia more naturally also produced conglutinates, but they were much smaller (<5 mm) and lacked a defined shape. When the released brood was observed the following morning, most glochidia were free of the unfertilized eggs, but were still somewhat stranded togeth- er in a heavy amount of mucous released by the adult. Host fish trials Yadkin Pee-Dee River Basin A total of 420 transformed juveniles were collected from 13 of the 20 fish species tested; however, several of those spe- cies were poor hosts, yielding only one or two juveniles (Table 1). Five species of Cyprinidae, including the satinfin shiner ( Cyprinella analostana), bluehead chub ( Nocomis lep- tocephalus), golden shiner ( Notemigonus crysoleucas), highfin shiner ( Notropis altipinnis ), and the spottail shiner ( Notropis hudsonius), acted as the most efficient hosts, with the per- centage of glochidia transforming ranging from 32.2 to 73.7%. As a species, bluegill ( Lepomis macrochirus) produced the most juveniles, but this production was attributed to the use of 15 individuals of this species. Two replicates contain- ing individual bluegill produced only four (13.3% transfor- mation) and six (33.3% transformation) metamorphosed individuals, respectively. Another aquarium of bluegill containing 13 individuals produced 175 juveniles. When individual fish from this tank were examined during the encystment period, several glochidia were readily visible on the gills and fins of some fish while other fish appeared to have no larvae attached. The length of time attached to all fish ranged from 22 to 44 days (Table 3). Catawba River Basin Some pedal swelling and gaping was observed when the adult mussel was exposed to serotonin, but this completely sub- sided within 24 hours once the animal was removed from the exposure. No other ill effects were noted, and the animal was maintained at the hatchery for six months after exposure. The animal was observed to have good adductor muscle strength and burrowing ability during this time and was presumed healthy. A total of 1,117 juveniles were collected from 1 1 of the 17 fish species tested; however, as with the Yadkin-Pee Dee trial, several species of Cyprinidae supported the greatest percentag- es of transformation (Table 2). Rosyside dace ( Clinostomus funduloides), whitefin shiners ( Cyprinella nivea ), sandbar shiners ( Notropis scepticus ), and the creek chub ( Semotilus atro- maculatus) represented new hosts not tested in the Yadkin-Pee Dee trial. Bluehead chubs and golden shiners served as hosts again as in the initial trial. A smaller number of juveniles trans- formed on the green sunfish ( Lepomis cyanellus) and warmouth ( Chaenobryttus gulosus). Unlike in the Yadkin-Pee Dee Trial, no juveniles transformed on the two bluegills tested. DISCUSSION Prior to this study, very little was known about the life history of this species (Bogan 2002). One gravid individual had been observed in October (Bogan 2002). Since we found the species gravid in January and late February, it is appar- ently bradytichtic, spawning in late summer or fall, and releasing glochidia in late winter or spring of the following year. Determination of the required host fish of Lasmigona decorata will allow propagation and captive culture of this rare species to move forward for conservation purposes. We can only speculate as to the reason why the glochidia appeared to be packaged differently between the two trials. Both indi- viduals yielded glochidia attached to non-viable eggs, but the individual that was chemically induced to release produced much smaller conglutinates. We also found more glochidia unattached to eggs released by the mussel several hours after the serotonin exposure. Because much of that release occurred overnight, we do not know what percentage of the released glochidia were originally attached to eggs, but came detached before they were observed. It may be that both indi- viduals naturally packaged their glochidia with non-viable eggs similarly within the marsupium, but the mussel exposed 156 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2-2010 to serotonin simply broke the mass into smaller pieces than those that were manually removed by syringe. None of the conglutinates had a well-defined shape, and they may have broken up differently based on how they left the mussel. Aside from possible variation between individuals or between basins, another possible explanation could be differences in fertilization success between the two females. Because of our need to use as many of these rare glochidia as possible as quickly as possible on the host fish, we did not attempt to quantify the total number of glochidia produced or the ratio of viable glochidia to non-viable eggs. Lasmigona decorata juveniles were successfully trans- formed from glochidia attached to multiple fish species. Between the two trials, a total of nine species from the Cyprini- dae family served as efficient hosts. The redlip shiner ( Notropis chiliticus ) and whitemouth shiner ( Notropis alborus) were the only cyprinid species considered poor hosts due to the small proportion of larvae that completed a metamorphosis on those species. A small number of juveniles were recovered from the highfin shiner ( Notropis altipinnis), but only a small num- ber actually attached — likely due to the small size of the indi- viduals (<40 mm) used in the trial. However, 44% of those that attached successfully transformed. In the Centrarchidae, the bluegill served as a host in the Yadkin-Pee Dee trial but yielded no metamorphosed juveniles when tested with the Catawba adult, and the green sunfish successfully supported the meta- morphosis of seven individuals in the Catawba trial but failed as a potential host in the Yadkin-Pee Dee trial. Because only a small number of bluegill were used in the Catawba trial, and only one green sunfish was used in each trial, the reason for the variation in their ability to serve as hosts is not known. Our results, however, may indicate differences in host suitability between two river basins, which has been previously demon- strated (Rogers etal. 2001, Bigham 2002, Eckert 2003). Because these two river basins drain separately to the Atlantic Ocean, the fish tested in this study were genetically isolated and may have different susceptibilities to infection by a given mussel species. We were limited to a single female mussel from each basin because L. decorata is such a rare species, and the lack of replication prevented us from making a definitive conclusion. In addition, the glochidia extraction method may have accounted for a portion of the variability between the two tri- als. To accurately determine host fish requirements, future studies should strive to use fish from the same river basin as the mussel being tested and use multiple brooding females if possible. Several other mussel species in the subfamily Anodonti- nae are relative host generalists, using fish from multiple families (Watters et al. 1998, McGill et al. 2002, Van Snik Gray et al. 2002) as hosts. While Lasmigona decorata did transform on three species of Centrarchidae, cyprinids were far more effective at facilitating transformation. The heel- splitter is typically found in deep runs as well as pools, and occurs in a variety of substrates from soft mud to clean san- dy-gravel and even occurs between large boulders within pools (Keferl and Shelley 1988, and the authors’ pers. obs.). These habitat preferences should overlap considerably with the preferences of both the centrarchids and the variety of cyprinids we identified as viable hosts. Though some identi- fied hosts, such as the bluehead chub, are more benthic in nature and may come in closer contact with the mussel, the neutrally buoyant conglutinates produced by this species should also provide dispersal of glochidia to fish up in the water column. We believe these small, white packages of glochidia would likely be attractive as a potential food source to many cyprinid species, but attempts by the fish to eat them would yield an infestation of glochidia rather than a true meal. The bluehead chubs and golden shiners appeared to be the best choices as fish hosts for captive propagation. The bluehead chub yielded the largest proportion of glochidia undergoing successful metamorphosis (73.7%), and this spe- cies is widespread, abundant and relatively easy to capture by electrofishing or seining. Golden shiners, the second most effi- cient host (64.0% transformation), produced large numbers of juveniles and can be easily purchased from a local bait shop in large numbers at a relatively low price. However, baitfish used for captive propagation should preferably be obtained from baitfish suppliers who adhere to recommended health certification guidelines. Extraction of glochidia from conglutinate-producing mussel species can be difficult, and flushing of the gills requires additional effort compared to species whose glochid- ia are not connected in packets. Relatively large openings must be made in gill tissue to extract large conglutinates from the marsupia, and this can be damaging to the gill tissue and even fatal, as in our study. Allowing a gravid mussel to release glochidia on its own often results in release of a small per- centage of the brood over several days or weeks (Eads, pers. obs.), making propagation more difficult and well-controlled host determination experiments almost impossible. There- fore, we elected to expose the gravid Lasmigona decorata in our second trial (Catawba Basin) to serotonin, a neuro- transmitter known to affect many aspects of bivalve repro- duction including induction of parturition (abortion of immature glochidia; Fong and Warner 1995) and glochidial release (Dimock and Strube 1994). Additionally, selective serotonin reuptake inhibitor (SSRI) drugs, which increase serotonin neurotransmission, have been shown to cause par- turition or glochidia release in the fingernail clam Sphaerium striatinum (Lamarck, 1818) (Fong et al. 1998), the zebra mussel Dreissena polymorpha (Pallas, 1769) (Fong etal. 2003), and the freshwater mussel Anodonta cygnea (Linnaeus, 1758) (Cunha and Machado 2001). FISH HOSTS OF LASMIGONA DECORATA 157 This study does not provide a thorough direct comparison between physical and chemical glochidia extraction methods, but it does identify an area of potential research for advance- ment in propagation methodologies. In our work, the use of serotonin was effective in allowing the collection of all glochidia in the brood apparently without lasting harm to the adult mussel. Glochidia also retained their viability, which was documented by a 100% response to a salt solution and their transformation on host fish. There were short- term physical effects observed in the mussel. We observed a strong increase in volume of the foot in Lasmigona decorata during early stages of exposure to serotonin, consistent with the response in Anodonta cygnea reported by Cunha and Machado (2001), who attributed the increase in foot size to relaxation of foot muscles and the resulting favorable condi- tions for uptake and storage of water. The L. decorata used in our trial recovered (z'.e., foot returned to pre-serotonin expo- sure size) within 24 hours of placement in clean water. Though our work with serotonin was successful, we suggest additional experimentation with varied concentrations and immersion times to optimize induction of glochidia release with minimal serotonin exposure. Additional studies are needed to determine the effects of serotonin on glochidial viability, gill respiration, the potential for future reproduc- tion, and the long-term overall health of gravid females. As the use of serotonin or other similar compounds is refined, it may prove a useful tool in the simple and safe extraction of glochidia for propagation of freshwater mussels. ACKNOWLEDGMENTS Funding for this research was provided by the North Carolina Wildlife Resources Commission (NCWRC). From the staff of the NCWRC, we thank Scott Van Horn, Brena Jones, Ryan Heise, Mark Fowlkes, Marla Chambers, Steve Fraley, and T. R. Russ for their support and field assistance. John Fridell of the U.S. Fish and Wildlife Service also offered important guidance and field assistance. Tim Savidge and the Catena Group collected the mussels from Sixmile Creek and kindly transported them to our laboratory. We thank Gene Wilson and the NCWRC staff at the Table Rock Fish Hatchery for maintaining the mussel culture trough at their facility. LITERATURE CITED Bigham, B. E. 2002. Host specificity of freshwater mussels: A critical factor in conservation. M.S. Dissertation, Southwest Missouri State University, Springfield, Missouri. Bogan, A. E. 2002. Workbook and Key to the Freshwater Bivalves of North Carolina. North Carolina Museum of Natural Sciences, Raleigh, North Carolina. Bogan, A. E. 2008. Workbook and Key to the Freshwater Bivalves of South Carolina, 2nd Edition. North Carolina Museum of Natu- ral Sciences, Raleigh, North Carolina. Coker, R. E., A. F. Shira, H. W. Clark, and A. D. Howard. 1921. Natural history and propagation of fresh-water mussels. Bul- letin of the Bureau of Fisheries [Issued separately as U.S. Bureau of Fisheries Document 893] 37: 77-181. Cunha, E. M. and J. Machado. 2001. Parturition in Anodonta cygnea induced by selective serotonin reuptake inhibitors (SSRIs). Canadian Journal of Zoology 79: 95-100. Dimock R. V. and R. W. Strube. 1994. Exogenous neurotransmitter induces unionid glochidial release. American Zoologist 34: 96A. Eckert, N. L. 2003. Reproductive biology and host requirement differences among isolated populations of Cyprogenia alberti. M.S. Disserta- tion, Southwest Missouri State University, Springfield, Missouri. Fong, P. P. and M. Warner. 1995. Serotonin-induced parturition in the fingernail clam Sphaerium ( Musculium transversum) (Say). Journal of Experimental Zoology 272: 163-166. Fong, P. P., P. T. Huminski, and L. M. D’Urso. 1998. Induction and potentiation of parturition in fingernail clams ( Sphaerium stri- atinum) by selective sertotonin re-uptake inhibitors (SSRIs). Journal of Experimental Zoology 280: 260-264. Fong, P. P., C. M. Philbert, and B. J. Roberts. 2003. Putative sero- tonin reuptake inhibitor-induced spawning and parturition in freshwater bivalves is inhibited by mammalian 5-HT2 receptor antagonists. Journal of Experimental Zoology 298A: 67-72. Gerke, N. and R. Tiedemann. 2001. A PCR-based molecular iden- tification key to the glochidia of European freshwater mussels (Unionidae). Conservation Genetics 2: 285-287. Keferl, E. P. and R. M. Shelley. 1988. The final report on a status survey of the Carolina Heelsplitter, Lasmigona decorata and the Carolina Elktoe, Alasmidonta robusta. U.S. Department of the Interior, Fish and Wildlife Service, 27 pp. Khym, J. R. and J. B. Layzer. 2000. Host fish suitability for glochidia of Ligumia recta. American Midland Naturalist 143: 178-184. Kneeland, S. C. and J. M. Rhymer. 2007. A molecular identification key for freshwater mussel glochidia encysted on naturally para- sitized fish hosts in Maine, USA. Journal ofMolluscan Studies 73: 279-282. McGill, M., M. Hove., T. Diedrich, C. Nelson, W. Taylor, and A. Kapuscinski. 2002. Several fishes are suitable hosts for creek heelspiltter glochidia. Ellipsaria 4: 18-19. McLain, D. C. and M. R. Ross. 2005. Reproduction based on local patch size of Alasmidonta heterodon and dispersal by its darter host in the Mill River, Massachusetts, USA. Journal of the North American Benthological Society 24: 139-147. National Native Mussel Conservation Committee. 1998. National strategy for the conservation of native freshwater mussels. Jour- nal of Shellfish Research 17: 1419-1428. Rogers, S. O., B. T. Watson, and R. J. Neves. 2001. Life history of the endangered tan riffleshell ( Epioblasma florentina walked) (Bivalvia: Unionidae). Journal of the North American Bentho- logical Society 20: 582-594. 158 AMERICAN MALACOLOGICAL BULLETIN 28*1/2*2010 Surber, T. 1912. Identification of the glochidia of freshwater mus- sels. Report and Special Papers of the U.S. Bureau of Fisheries [Issued separately as U.S. Bureau of Fisheries Document 771 ] . Surber, T. 1913. Notes on the natural hosts of fresh-water mussels. Bulletin of the Bureau of Fisheries [Issued separately as U.S. Bureau of Fisheries Document 778] 32: 103-116. U.S. Fish and Wildlife Service. 1996. Carolina Heelsplitter Recovery Plan. U.S. Fish and Wildlife Service, Atlanta, Georgia. Van Snik Gray, E., W. A. Lellis, J. C. Cole, and C. S. Johnson. 2002. Host identification for Strophitus undulatus (Bivalvia: Unioni- dae), the Creeper, in the Upper Susquehanna River Basin, Pennsylvania. American Midland Naturalist 147: 153-161. Watters, G. T., S. H. O’Dee, and S. Chordas. 1998. New potential hosts. Triannual Unionid Report 15: 27-29. White, L. R., B. A. McPheron, and J. R. Stauffer, Jr. 1994. Identi- fication of freshwater mussel glochidia on host fishes using restriction fragment length polymorphisms. Molecular Ecology 3: 183-185. Williams, J. D„ A. E. Bogan, and J. T. Garner. 2008. Freshwater Mus- sels of Alabama and the Mobile Basin of Georgia, Mississippi, and Tennessee. The University of Alabama Press, Tuscaloosa, Alabama. Zale, A. V. and R. J. Neves. 1982a. Fish hosts of four species of Lamp- siline mussels (Mollusca: Unionidae) in Big Moccasin Creek, Virginia. Canadian Journal of Zoology 60: 2535-2542. Zale, A. V. and R. J. Neves. 1982b. Identification of a host fish for Alasmidonta minor (Mollusca: Unionidae). American Midland Naturalist 107: 386-388. Submitted: 1 December 2008; accepted: 15 September 2009; final revisions received: 4 October 2009 Amer. Malac. Bull. 28: 159-165 (2010) Population studies of an endemic gastropod from waterfall environments Diego E. Gutierrez Gregoric, Veronica Nunez, and Alejandra Rumi Division Zoologia Invertebrados, Museo de La Plata, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Paseo del Bosque s/n, 1900, La Plata, Argentina Corresponding author: dieguty@fcnym.unlp.edu.ar Abstract: Chilinidae is a family endemic to South America, ranging from the Tropic of Capricorn to Cape Horn and the Falkland Islands, and includes 32 species. However, there are few population studies on the Chilinidae. We study aspects of the ecology of an endemic species, Chilina megastoma Hylton Scott, 1958, from the Arrechea Falls in the Iguazu National Park, Argentina, such as density and individual annual growth trends. Nine samplings were carried out between December 2003 and December 2005, using two transects that crossed the waterfall. Individual annual growth rate was analyzed according to length, following von Bertalanffy's model. Six cohorts were identified, some in the same climatic season but successive years (two in winter and two in summer). The winter and autumn cohorts reached 85% of their last whorl length in the first year. Compared to other families of gastropods from subtropical climates, these populations have several recruitment events per year, but never in winter. Key words: Argentina, Chilina, growth rate, Iguazu Falls, von Bertalanffy model In Argentina, there are forty (40) endemic species for freshwater Gastropoda: Thiaridae (3 species), Ampullariidae (1), Cochliopidae (10), Lithoglyphidae (11), Chilinidae (11), Lymnaeidae (2), and Physidae (2). Of the species of Litho- glyphidae, Cochliopidae, and Chilinidae, 50, 62.5, and 68.7 %, respectively, are endemic (Rumi et al. 2006). Although most of these endemic species are considered vulnerable to extinction, they are in serious need of bio-ecologic studies to allow more precision in the identification of their conserva- tion status. Chilinidae (Pulmonata, Basommatophora) is a family exclusive to South America, ranging from the Tropic of Capricorn to Cape Horn and the Falkland islands. The fam- ily includes a single genus, Chilina Gray, 1828 with about 32 species, 17 of which have been recorded in Argentina (Castellanos and Miquel 1980, Castellanos and Gaillard 1981, Gutierrez Gregoric and Rumi 2008) while the rest are distributed in Chile. Many Argentinean species of Chilina are endemic, and their biology and ecological strategies are practically unknown. In general, only a few studies have been recently reported in this family: Miquel (1986) studied the life cycle of Chilina flu- minea (Maton, 1809) and its gonad evolution; Bosnia et al. (1990), analyzed the growth of Chilina gibbosa G. B. Sowerby I, 1841, and its density; Estebenet et al. (2002) analyzed the natural diet for Chilina par chappii (d’Orbigny, 1835). Quijon and Jaramillo (1999) and Quijon et al. (2001), worked on Chilina ovalis (Sowerby, 1841) from southern Chile, empha- sizing spatial distribution and growth. The dominant landscape in Iguazu National Park (INP) includes waterfalls and rapids of the Iguazu River, all surrounded by subtropical forest. The park is home to four endemic species of molluscs: Eupera iguazuensis Ituarte, 1989 and Eupera elliptica Ituarte and Dreher-Mansur, 1993 (Bivalvia: Sphaeriidae), and Chilina megastoma Hylton Scott, 1958 and Chilina iguazuensis Gutierrez Gregoric and Rumi, 2008 (Gastropoda: Chilinidae). Chilina megastoma and Acrorbis petricola Odhner, 1937 (Planor- bidae) are exclusively found on waterfalls. Acrorbis petricola has been recorded at only three localities, one in Brazil and the other two in Argentina — one in grounds of the INP. Upstream dam and reservoir construction during recent years have modified the hydrologic regime of the river. These changes could induce irretrievable losses in biodiversity, especially among these types of environments. In this sense, a paradigmatic example is that of Aylacostoma Spix, 1827 (Gastropoda: Thiaridae), which included four described species known only in the rapids along the upper Parana River, in the sector now occupied by the Yacyreta Reser- voir (Argentina - Paraguay). According to Quintana and Mer- cado Laczko (1997) these species can be considered extinct in their natural habitat. The same may have occured with Chilina guaraniana Castellanos and Miquel, 1980 and Acrorbis sp., also collected before the flooding of the reservoir (Rumi 1986, Rumi etal. 2006). The scarce information available on gastropods from white-water rivers with waterfalls such as the Iguazu deals with those that lived upstream and downstream from waterfalls, or with the description of new species from this kind of environ- ment (Ponder 1982, Gloer et al. 2007). However, population dynamics of these species remain largely unknown. This work focuses on estimating population patterns such as density, individual growth rate, and recruitment times of a population of Chilina megastoma. This species is endemic to 159 160 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 the waterfalls in the Iguazu National Park, Misiones province, Argentina. MATERIALS AND METHODS Study area Iguazu National Park is located in Misiones province, at the northeastern corner of the country. This park was char- tered in 1934, and in 1984 was declared a World Natural Heritage Park because of the high biological diversity of the subtropical Paranensean Forest and the numerous waterfalls that average 75 m high. The climatic characteristics of INP (1600 mm annual precipitation, 21.1 °C mean annual tem- perature) allow wide habitat diversity and support a varied flora and fauna. The waterfalls harbor plants and animals that are especially adapted to the constant humid conditions and the force of water. Abundant islands covered by a distinct type of forest exist along the upper course of the Iguazu River. The population study was done in the Arrechea Falls of the INP (25°39’S, 54°27’W) (Fig. 1). Sampling methodology The study comprised nine seasonal samplings between December 2003 and December 2005 (December 2003, 2004, 2005; February 2004, 2005; June 2004, 2005; September 2004, 2005). Water temperature (°C), conductivity (pS), hardness (°f, calculated as conductivity / 20), total dissolved solids (mg/1), pH, dissolved oxygen (mg/1), and saturated oxygen (%) were measured during most samples. Two transects, each approx. 10 meters long, were fol- lowed from the bottom of Arrechea Falls behind the waterfall, and into the driest vegetated zone at the top (Fig. 1). The snails were hand collected from rocks exposed along the two transects. Squares of 0.15 m (0.0225 m2), placed at 0.40 m along the transect, were employed as a sample unit (SU). For each sampling date, we calculated the average density of snails and its standard deviation. Analysis of growth For individual growth rate, snails were measured on site using 0.01 mm precision calipers. As the apex of these gastro- pods is usually damaged by water current, only the length of last whorl (LWL) was recorded. Once measured, snails were returned to their natural environment. In all samples, size frequency intervals of 1 mm were used. The size frequency distribution of each sample was separated by its modes, which were assumed to correspond to coexisting cohorts. Frequency distributions corresponding to each cohort were fitted to a normal curve, for which mean and standard deviation were calculated. These values were used in the growth analysis. Individual growth rate was analyzed according to length, Figure 1. Transects in the Arrechea Falls, Iguazu National Park, Argentina. Table 1. Mean water quality parameters at Arrechea Falls. Cond, conductivity (pS); TDS, total dissolved solids (mg/1); T, tempera- ture (°C); DO, dissolved oxygen (mg/1); Sat, % oxygen saturation; n/d, no data. Date Cond TDS T pH DO Sat Dec-03 490 25.8 23.3 6.5 4.9 69.3 Feb-04 95.2 42.6 22.4 6.6 6.5 73 Jun-04 235.4 122.6 14.5 7.3 8.7 81 Sep-04 57.2 29 20.4 7 9.2 101 Dec-04 39.7 19.8 21.3 6.8 8.4 94 Feb-05 137.9 65.7 23.7 8.3 5.3 62.5 Jun-05 66 32.2 18.6 6.6 n/d n/d Sep-05 n/d n/d 14.5 7.4 2.4 24 Dec-05 n/d n/d n/d n/d n/d n/d WATERFALL GASTROPOD POPULATIONS 161 Table 2. Mollusc density (8: ind/m2), including Chilina megastoma oviposition density (ovipositions/nr). N, total individual; x, mean calcu- lated per SU (sample unit = 0.0225 m2); SD, standard deviation. Chilina C. megastoma Acrorbis Potamolithus Uncancylus Date N of SU megastoma ovipositions petricola spp. Succinea sp. concentricus N 233 14 787 75 9 0 CO 6 45 X 5.18 0.31 17.49 1.67 0.20 0 Q SD 5.39 0.84 13.75 4.7 0.68 0 8 230 14 111 74 9 0 N 184 21 829 121 17 0 o X 51 X 3.61 0.41 16.25 2.37 0.33 0 Ph SD 4.17 1.11 20.55 5.78 1.07 0 8 160 18 111 105 15 0 N 210 24 296 378 4 2 c 50 X 4.2 0.48 5.92 7.56 0.08 0.04 SD 6.86 1.74 7.31 23.01 0.27 0.4 8 187 21 263 336 4 2 N 176 11 629 244 45 3 Cu 53 X 3.32 0.21 11.87 4.6 0.84 0.06 CO SD 5.25 0.79 17.62 13.5 2.04 0.42 8 148 9 527 205 38 2 N 251 9 632 256 5 0 50 X 5.02 0.18 12.64 5.12 0.1 0 Q SD 7.5 0.6 24.06 11.53 0.46 0 8 223 8 562 228 4 0 N 53 6 142 556 1 0 o Xi 26 X 2.04 0.23 5.46 21.38 0.04 0 Ph SD 2.44 0.65 7.66 21.94 0.2 0 8 91 10 243 950 2 0 N 35 1 254 467 2 0 LO i 51 X 0.69 0.02 4.98 9.16 0.04 0 SD 1.47 0.14 12 23.82 0.2 0 8 30 1 221 407 2 0 N 74 15 338 124 55 0 o X 1.64 0.33 7.51 2.75 1.22 0 Oh 45 co SD 6.84 1.65 22.11 8.38 2.8 0 8 73 15 333 111 54 0 N 79 10 113 120 16 2 o )), ( 1 ) where LWLoo = maximum length of last whorl, k = growth rate constant, t = time, and tQ = hypothetical time in which length = 0. From linear transformation of the logarithmic equation (1) we obtained: In (1 - LWLt/LWLoo)= -kt + kt0 (2) A regression equation between In (1- LWLt/LWLoo) and t was calculated. In (2), ktQ = origin of ordinate (a), t — a/k, and the slope (b) = (-)k. Time measured for each sample was divided into parts of one year, following Basso and Kher ( 1991 ) and Rumi et al. (2007), in the equation: T — [ (month - 1 )*30 + sampling day] / 360 + A, where A = sampling year. The year in which the study started was considered as A = 0, the following year is A = 1 , etc. Thus, t = 0 corresponded to January 1st, t = 0.5 to July 1st, and t- 1 corresponds approx, to December 31st. Maximum length of the whorl was calculated on mean values of cohorts obtained from decomposition, using Walford's method (1946). Only means from cohorts beginning at the time of sam- pling were considered; the cohorts with the two highest val- ues (December 2003 and February 2004) and lowest values (June, September, and December 2005) were not consid- ered. In order to compare these data with those from other species or from the same species in different environments, tQ was considered as 0 and growth rate was expressed as per- centage of maximum length of last whorl, as reported by Rumi et al. (2007). RESULTS The averages of water quality parameters were: tem- perature (N = 8) = 19.8 °C ( SD = 3.7); pH (N = 8) = 7.1 ( SD = 0.6); conductivity (N = 7) = 160.2 pS/cm (SD = 159.8); total dissolved solids (TDS) (N = 7) = 48.2 mg/1 (SD = 36.7); dissolved oxygen (N = 7) = 6.5 mg/1 (SD = 2.5); oxygen sat- uration = 72.1% (Table 1). Water hardness (°f) was 8.01 (soft water). Gastropod assemblages in addition to Chilina megastoma in the Arrechea Falls included: Potamolithus sp. 1 and Pota- molithus sp. 2 (Lithoglyphidae); Uncancylus concentricus (d’Orbigny, 1835) (Ancylidae) ; Acrorbis petricola (Planorbi- dae); and Succinea sp. (Succineidae). Acrorbis petricola had the greatest densities, whereas there were only isolated specimens of U. concentricus in three samples (Table 2). Individuals of C. megastoma from classes 2 to 4 (1 to 3 mm) were observed throughout the entire year, except for February and June of 2005 (Fig. 2). Specimens from classes 10 to 14 (9 to 13 mm) were also observed year round at similar frequencies (Fig. 2). 0 2 4 6 8 10 12 14 16 18 size classes size classes Figure 2. Size-frequency distributions of Chilina megastoma, ex- pressed as a percentage of the sample total N, in intervals of 1 mm, among samples at Arrechea Falls. WATERFALL GASTROPOD POPULATIONS 163 Table 3. Means ( x ), standard deviation (SD), and number of individuals (N) fitted to each monthly shell-size frequency distribution for cohorts of Chilina megastoma in Arrechea Falls. Date Cohort I Cohort II Cohort III Cohort IV Cohort V Cohort VI X SD N x SD N X SD N X SD N X SD N X SD N Dec-03 7.73 0.62 34 4.79 0.93 76 Feb-04 12.21 1.07 20 7.31 0.74 12 3.76 1.06 48 Jun-04 14.54 0.76 6 11.32 1.04 50 7.9 1.08 39 3.49 1.15 49 Sep-04 16.02 1.35 11 11.21 1.4 40 6.95 0.81 25 3.37 1.08 33 Dec-04 15.2 1.13 9 11.23 1.36 23 6.39 0.66 21 Feb-05 16.21 0.90 7 12.04 1.36 31 5.6 0.5 20 fun-05 16.27 1.08 6 8.65 0.8 16 Sep-05 15.95 0.72 6 11.79 1.6 34 Dec-05 14.77 0.7 9 Polymodal size frequency distributions indicated three or four well-defined cohorts (Table 3 and Fig. 3). Six cohorts were identified (Table 4). The LWL calculated was 18.47mm. Cohorts I and V correspond to winter recruitment, and cohorts III and VI correspond to summer recruitment. Con- sidering the information of both seasonal cohorts together, the growth equations for these two climatic seasons (summer and winter) are given in Table 5. In all cases the slope ( b ) of the linear regression compar- ing the observed and calculated data was near 1, and the same was true for R2: winter cohort b: 1.01, R2: 0.96; spring cohort b: 0.97, R2: 0.99; summer cohort b: 1.01, R2: 0.95; and autumn cohort b: 1.03, R2: 0.99; indicating a good adjustment of data to the pattern of calculated growth. The maximum LWL growth percentage indicates that all the cohorts reach at least 76% of their growth in the first year of life (Fig. 4). The winter and autumn cohorts reach 85% of their LWL in the first year. After two years, all the cohorts reach 94% of their LWL, with the winter and autumn cohorts reaching the highest percentage in the two years (97%). DISCUSSION Each of the molluscs has a specific distribution pattern. Potamolithus spp. inhabit mainly the pool and to a lesser extent the water trapped among the rocks. Succinea sp. inhabits higher places where rocks begin to be colonized by vegetation. Acrorbis petricola and Chilina megastoma maybe found along the whole height of the waterfall; the former lives on the vertical rocks of the wall, while C. megastoma dwells generally in the crevices and in areas more strongly hit by water. Figure 3. Means (dots) and standard deviation (bars) of the normal curves fitted to each monthly shell-size frequency distribution, and growth curves for Chilina megastoma in Arrechea Falls. Lines repre- sent theoretical growth curves according to von Bertalanffy models: black lines represent winter cohorts; dark gray represents summer cohorts; light gray represents autumn cohort; dashed line represents spring cohort. The decrease in density of Chilina megastoma and Acrorbis petricola beginning in February of 2005 was due to the drought recorded in the Iguazu area from December 2004 to May 2005 (Center of Subtropical Ecological Investigations-CIES, unpubl. data) (Table 2). The two species inhabit rocky and humid sec- tors of the waterfall, which, because of the low water level of the river, remained dry. In samples in February and June 2005 (those most affected by drought) there were no speci- mens of the smaller size classes. There may have been insuffi- cient humidity to allow oviposition or keep hatchlings viable. 164 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Table 4. Exponents for the growth equations for each cohort, k, growth rate constant; tg, hypothetical time in which length = 0. Cohort k t0 Cohort I 1.93 0.63 Cohort II 1.46 0.77 Cohort III 1.88 1.10 Cohort IV 1.96 1.42 Cohort V 1.82 1.62 Cohort VI 1.52 1.97 Table 5. Exponents for the growth equations for each seasonal cohort. k , growth rate constant; t0, hypothetical time in which length = 0. Seasonal cohort k t0 winter 1.88 0.63 spring 1.46 0.77 summer 1.73 0.05 autumn 1.96 0.40 Populations of Potamolithus were not affected by the drought as they were mainly in the pools, which did not dry. During the last two samples (September and December 2005), the popula- tion of C. megastoma began recovering with an increase in density, and a predominance of juvenile specimens. Hydrology of the Iguazu River and its tributaries are not only affected by rainfall but also are regulated by 100 -i hydroelectric projects located upstream, as mentioned above. Chilina megastoma turned out to be the most vulnerable spe- cies because of its high humidity requirements, because the species does not migrate towards the water pool zone, and because of the long hatching period (25 days; pers. obs.). Oviposition by Chilina megastoma occurred throughout the year, and averaged between 1 1 and 14.5 per square meter. Chilina megastoma may reproduce continually because of the low temperature variation along the year in the Iguazu National Park. Species of Chilinidae in cold areas reproduce once a year. Chilina gibbosa, in the Ramos Mexla reservoir (provinces of Rio Negro and Neuquen, Argentina) and in Lake Pellegrini (Rio Negro province), reproduces during the summer (Bosnia et al. 1990, Gutierrez Gregoric et al. 2004, respectively). Chilina fluminea, from the “La Balandra” beach, Berisso (Buenos Aires province, Argentina), reproduces only in winter (Gutierrez Gregoric 2008). Chilina ovalis, in south- ern Chile, does so in spring (October) (Quijon and Jaramillo 1999, Quijon et al. 2001). Lor C. ovalis and C. gibbosa from Lake Pellegrini, the estimated lifespan for each species is about three years. Lor C. fluminea, Gutierrez Gregoric (2008) estimated a lifespan of between two and two and a half years, while in C. megastoma, longevity is approx, two years (Pig. 4). These two different trends can be due to factors such as tem- perature. While INP shows little variations throughout the year, in the other environments the seasons are very marked. Lor C. megastoma, its best reproductive period is when tem- peratures are around 15 °C (Table 1), similar to C. fluminea in “La Balandra” beach, where temperatures are near 12 °C. The growth constants for Chilina ovalis and C. gibbosa were similar and lower than those of C. fluminea and C. megastoma. This can be explained by the temperature drop recorded in win- ter (10 °C for C. ovalis in Chile) and the greater longevity in these species. Other families of gastropods from subtropical climates can have several recruitments per year, but never in winter (Ituarte 1989, 1994, Rumi et al. 2007). In Biomphalaria occidentalis Para- ense, 1981 (Planorbidae) (Corrientes province, Argentina), the growth per- centage of the first year of life was always greater than 80%. Biomphalaria straminea (Dunker, 1848) and Biompha- laria tenagophila (d’Orbigny, 1835) from Artigas, Uruguay reached 80% of maxi- mum length at nine and a half and eight months respectively (Ituarte 1989, 1994). Chilinidae in subtropical environments thus possess slower growth rates than Time (years) Figure 4. Growth of Chilina megastoma expressed as percentage of maximum length of the last whorl. WATERFALL GASTROPOD POPULATIONS 165 Planorbidae, but reproduces continually, while the Planorbi- dae cease reproduction in winter. ACKNOWLEDGMENTS The authors wish to thank the staff at Centro de Investiga- ciones Ecologicas Subtropicales (CIES) in Iguazu National Park, and N. Ferrando for their support during fieldwork. We specially thank M. Griffin for help with English. LITERATURE CITED Baluku, L. and M. Loreau. 1989. Etude comparative de la dynamique des populations de Biomphalaria pfeifferi (Gastropoda, Planor- bidae) dans deux cours d'eau du Zaire oriental. Journal African Zoology 103: 311-325 [In French], Basso, N. G. and A. I. Kehr. 1991. Postmetamorphic growth and popu- lation structure of the frog Leptodactylus latinasus (Anura: Lepto- dactylidae). Studies Neotropical Fauna and Environment 26: 39-44. Bosnia, A. S., F. J. Kaisin, and A. Tablado. 1990. Population dy- namics and production of the freshwater snail Chilina gibbosa Sowerby 1841 (Chilinidae, Pulmonata) in a North-Patagonian reservoir. Hydrobiologia 190: 97-110. Castellanos, Z. A. de and M. C. Gaillard. 1981. Mollusca Gasteropo- da: Chilinidae. Fauna deAgua Duke de la Republica Argentina, PROFADU (CONICET), Buenos Aires 15: 23-51 [In Spanish], Castellanos, Z. A. de and A. E. Miquel. 1980. Notas complemen- tarias al genero Chilina Gray (Mollusca Pulmonata). Neotro- pica 26: 171-178 [In Spanish], Estebenet, A. L., N. J. Cazzaniga, and N. V. Pizani. 2002. The natural diet of the Argentinean endemic snail Chilina parchappii (Ba- sommatophora: Chilinidae) and two other coexisting pulmo- nate gastropods. The Veliger 45: 71-78. Gloer, P., C. Albrecht, and T. Wilke. 2007. Enigmatic distribution patterns of the Bithyniidae in the Balkan Region (Gastropoda: Rissooidea). Mollusca 25: 13-27. Gutierrez Gregoric, D. E. 2008. Estudios morfoanatomicos y tenden- cias poblacionales en especies de lafamilia Chilinidae Dali 1870 (Mollusca: Gastropoda) en la cuenca Del Plata. Ph.D. Disserta- tion, Facultad de Ciencias Naturales y Museo, Universidad Na- cional de La Plata, La Plata, Argentina [In Spanish]. Gutierrez Gregoric, D. E. and A. Rumi. 2008. Chilina iguazuensis (Gastropoda: Chilinidae), new species from Iguazu Nacional Park, Argentina. Malacologia 50: 321-300. Gutierrez Gregoric, D. E., M. A. Roche, A. Rumi, and M. Maggioni. 2004. Crecimiento individual en Chilina gibbosa (Gastropoda: Chilinidae) en el lago Pellegrini, Rio Negro, Argentina. In: B. Cavaganaro, S. Claver, L. Marone, P. Villagra, and R. Villalba, eds., Book of Abstracts of the II Reunion Binacional de Ecologia. Universidad Nacional de Cuyo, Mendoza, Argentina. P. 434 [In Spanish]. Ituarte, C. F. 1989. Growth dynamics in a natural population of Biomphalaria straminea (Dunker, 1848) from Bella Union, Artigas, Uruguay. Studies Neotropical Fauna and Environment 24: 35-40. Ituarte, C. F. 1994. Temporal variation in age structure of a natural population of Biomphalaria tenagophila (Gastropoda: Planor- bidae) from a rice field irrigation channel system at Artigas, Uruguay. Malacological Review 27: 13-21. Loreau, M. and L. Baluku. 1987. Growth and demography of popu- lations of Biomphalaria pfeifferi (Gastropoda, Planorbidae) in the laboratory. Journal ofMolluscan Studies 53: 171-177. Miquel, S. E. 1986. El ciclo de vida y la evolucion gonadal de Chilina fluminea fluminea (Maton, 1809) (Gastropoda; Basommato- phora; Chilinidae). Neotropica 32: 23-34 [In Spanish]. Ponder, W. F. 1982. Hydrobiidae of Lord Howe Island (Mollusca: Gastropoda: Prosobranchia). Australian Journal of Marine and Freshwater Research 33: 89-159. Quijon, P. and E. Jaramillo. 1999. Gastropods and intertidal soft- sediments: The case of Chilina ovalis Sowerby (Pulmonata: Ba- sommatophora) in South-Central Chile. The Veliger 42: 72-84. Quintana, M. G. and A. Mercado Laczko. 1997. Biodiversidad en peligro. Caracoles de los rapidos en Yacyreta. Ciencia Hoy 7: 22-31 [In Spanish], Quijon, P., H. Contreras, and E. Jaramillo. 2001. Population biol- ogy of the intertidal snail Chilina ovalis Sowerby (Pulmonata) in the Queule river estuary, South-Central Chile. Estuaries 24: 69-77. Rumi, A. 1986. Estudio morfoldgico, taxondmico y bio-ecologico de los planorbidos argentinos. Ph.D. Dissertation, Facultad Ciencias Naturales y Museo, Universidad Nacional de La Plata, La Plata, Argentina [In Spanish], Rumi, A., D. E. Gutierrez Gregoric, and M. A. Roche. 2007. Growth rates fitting using the von Bertalanffy model: An analysis in natural populations of Drepanotrema spp. (Gastropoda: Plan- orbidae). Revista de Biologia Tropical 55: 559-567. Rumi, A., D. E. Gutierrez Gregoric, V. Nunez, I. I. Cesar, M. A. Roche, M. P. Tassara, S. M. Martin, and M. F. Lopez Armengol. 2006. Freshwater Gastropoda from Argentina: Species richness, distribution patterns, and an evaluation of endangered species. Malacologia 49: 189-208. Von Bertalanffy, L. 1938. A quantitative theory of organic growth. Human Biology 10: 181-213. Walford, L. A. 1946. A new graphic method of describing the growth of animals. Biological Bulletin 90 : 141-147. Submitted: 1 May 2009; accepted: 13 October 2009; final revisions received: 26 October 2009 Amer. Malac. Bull. 28: 167-181 (2010) Subtropical sacoglossans in Okinawa — at “special risk” or “predictably rare”? Cynthia D. Trowbridge1’*, Yayoi M. Hirano2, Yoshiaki J. Hirano2’3, Kosuke Sudo2’3, Yoichi Shimadu2’3, Tomohiro Watanabe2’3, Makiko Yorifuji2’3, Taro Maeda2’3, Yuki Anetai2’3, and Kanako Kumagai2,3 1 Oregon Institute of Marine Biology, University of Oregon, Charleston, Oregon 97420, U.S.A. 2 Marine Biosystems Research Center, Chiba University, Japan department of Biology, Graduate School of Science, Chiba University, Japan Corresponding author: cdt@uoregon.edu On low intertidal and shallow subtidal shores on the west coast of Okinawa, Japan, we investigated the trophic associations of sacoglossan opisthobranchs associated with Bryopsidalean green algae. During 1 1 short research visits (55 days total) from 2002 to 2008, we recorded almost 500 specimens of 11 species. These sacoglossans include a new record for Japan ( Caliphylla A. Costa, 1867), a recent record for Japan [Placida daguilarensis Jensen, 1990), two undescribed species ( Placida Trinchese, 1876 and Elysia Risso, 1818), one unnamed (but well- described) species ( Placida sp. sensu Baba 1986), and six other Indo-Pacifk species. Not only did we record more sacoglossan species but also we found higher slug abundances than other colleagues in Okinawa or the Indo-Pacifk region. Quantitative population attributes and feeding preferences are described for these sacoglossans. In contrast to temperate geographic regions, several of these Japanese sacoglossans specialized on a single algal genus rather than two or more genera in different families. This specificity is consistent with narrower host-plant associations in high-diversity communities; yet monophagy has not yet been demonstrated in this guild of Okinawan sacoglossans. Given the broad geographic ranges, restricted host ranges, often predictable populations, and high frequency of life cycles with planktotrophic larvae, western Pacific subtropical sacoglossans should be considered “predictably rare” ( sensu Rabinowitz 1981) rather than at “special risk” ( sensu Clark 1994). Keywords: Sacoglossa, Japan, herbivory, Codium, Bryopsis Western Pacific tropical and subtropical shores are char- acterized by the high species richness of the marine fauna and flora. The taxonomic and ecological diversity may reflect the cumulative effects of at least two biogeographic patterns: (1) Western Pacific shores are more diverse than eastern Pacific shores of comparable latitude. (2) Species richness increases along a latitudinal gradient from polar to tropical areas. Geo- graphic patterns of marine specialist herbivores, particularly the sacoglossan opisthobranchs, parallel these general eco- logical patterns. Many areas of the subtropical and tropical Pacific and Indo-Pacific are considered to be biodiversity “hotspots” for many taxonomic groups from coastal and pelagic fishes to seaweeds and invertebrates. Recent studies by Gosliner (1992), Ono (1999, 2004), Carlson and Hoff (2003), Jensen (2007), and Gosliner et al. (2008) exemplify this pattern. In contrast to these natural patterns, anthropogenic activities may reduce biodiversity by habitat destruction, introduced species’ effects, eutrophication, and other forms of pollution. Because anthropogenic activities are dispropor- tionately acute on tropical and subtropical shores affected by tourism and/or harvesting, coastal communities are often degraded or biologically compromised. Clark and DeFreese (1987) and Clark (1994) discussed the problems as they relate to sacoglossan occurrence, with Clark suggesting that the spe- cialists are at “special risk” of extinction because of their host specificity and high frequency of direct development. He pre- sented data on the decline of Key West populations of saco- glossans. In most other cases, the importance of natural vs. anthropogenic effects has often been asserted, rather than quantitatively demonstrated, for tropical and subtropical shores. As part of a large-scale project on Japanese sacoglossan assemblages (Hirano et al. 2003, 2005, 2006a, 2006b, 2006c, 2007a, 2007b, 2007c, 2008, Shimadu 2004, Trowbridge et al. 2005, 2006, 2007, 2008a, 2008b, 2009a, 2009b, Shimadu et al. 2006, Kumagai 2009), we quantified spatial, temporal, and taxonomic patterns of sacoglossan opisthobranchs and their macroalgal hosts on the subtropical shores of western Oki- nawa. Anthropogenic effects on Okinawa shores include habitat destruction, nutrient loading, sedimentation, heavy metal accumulation, human trampling, over-harvesting (fishes, invertebrates, and seaweeds), coral reef bleaching, etc. Although we did not document anthropogenic effects * Mailing address: P.O. 1995, Newport, Oregon 97365, U.S.A. 167 168 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 * 2010 during this study, one of our sites was severely disturbed by a shoreline modification project that replaced the natural shoreline with coral blocks and added a breakwater during our study (Eig. 1). Our other sites were heavily used recre- ational sites for scuba divers, swimmers, and collectors. Despite these anthropogenic effects, we recorded impres- sively high species richness of sacoglossan gastropods. The regional diversity of sacoglossans in Fiji, Guam, Thailand, Hong Kong, and Japan is well known to be extremely high (Carlson and Hoff 2003, Jensen 2007, Trowbridge et al. 2007, unpubl. ms, Gosliner et al. 2008). However, the local alpha- and beta-diversity ( e.g ., within individual sites or among closely adjacent sites) and the population abundance of sacoglossans in the Ryukyu Islands of Japan, particularly on the main island of Okinawa, have traditionally been consid- ered low (see Bolland’s Okinawa Slug Site at http://www. rfbolland.com/okislugs/). We quantitatively demonstrate that A. Figure 1. Anthropogenic disturbance at Sobe, Okinawa during our study: coastal modification to protect shore from storm waves. A, Replacement of vegetation and shoreline with coral blocks and sand; downward movement of coarse sand smothered intertidal and sub- tidal organisms. B, Addition of breakwater that changed the hydrol- ogy of site, including the shifting of the seagrass meadows and other marine macrophytes. local-scale assemblages of sacoglossans are not only diverse but also reliably found. Okinawan shores are clearly a “hot- spot” of sacoglossan biodiversity despite intense anthropo- genic effects. In this account, we focus on sacoglossan gastropods associated with hosts belonging to the green algal order Bry- opsidales (taxonomy based on Guiry and Guiry 2009). Spe- cifically, we investigated slugs on Codium (F. Codiaceae), Bryopsis (F. Bryopsidaceae), and Derbesia (F. Derbesiaceae). On NW and NE Atlantic shores, NE Pacific shores, S Pacific shores, and perhaps elsewhere, sacoglossans feed on 2-3 of these genera and, thus, are feeding on species in 2-3 algal families. Notable examples include Placida dendritica (Alder and Hancock, 1843), Elysia viridis (Montagu, 1804), Elysia hedgpethi Er. Marcus, 1961, and Placida sp. ( sensu Baba 1986). However, in Florida Placida kingstoni Thompson, 1977 appears to feed on Bryopsis but not Codium. Furthermore, Elysia trisinuata Baba, 1949 in Hong Kong and Japan appar- ently feeds exclusively on Codium (Jensen 2003, Trowbridge et al. 2008a). The ecological and conservation implications of diet breadth are far-reaching. In particular, species with a greater suite of alternate hosts would be more buffered from disturbance or habitat loss than monophagous species. Understanding the feeding specificity and population abun- dances of specialist gastropods will enable us to determine whether they are “special risk” as suggested by Clark (1994) or merely “predictably rare” ( sensu Rabinowitz 1981). MATERIALS AND METHODS Study region The Ryukyu Archipelago extends from Kyushu to Tai- wan and has several major island groups, each composed of numerous smaller islands. In the Ryukyu Shoto (the south- ern half of the islands), the Okinawa Islands are considered the central group or Ryukyu proper (Wikipedia 2009). The largest island in this group, Okinawa (also known as the Okinawan mainland or Okinawajima), is 1201 km2 in area. We examined sacoglossans on the west coast of Okinawa, facing the East China Sea. The climate of Okinawa is subtropical with mean air temperature of 23 °C, mean seawater temperature of 25 °C, and annual rainfall of >2 m. The tidal range is ca. 2.2 m on the western shore of Okinawa. Spring low tides expose coral reef platforms with diverse and abundant siphonous green algae (e.g., Codium, Bryopsis, Caulerpa, Halimeda, and allies). Algal populations also occur in shallow subtidal habitats (e.g., shallow fringing lagoons, reef crest, and reef slope). The siphonous green algal communities are flourishing despite intense trampling, marine species’ collection, nutrient enhance- ment, shoreline modification, and water sports. OKINAWAN SACOGLOSSANS 169 Surveys We visited the main island of Okinawa 11 times from July 2002 to December 2008 (Table 1). We surveyed the shore for 4-6 days per trip over a 6-y period including a variety of seasons. We concentrated our surveys at three primary sites between Naha and Zanpa-misaki on the west coast. From north to south, these sites were Zanpa (26°26'15"N, 127°42'48"E), Sobe (26°23'12"N, 127°43'24"E), and Sunabe (26°19'51"N, 127°44'36"E). We surveyed the low intertidal and shallow subtidal area of the shore where seaweeds grow profusely. This habitat has been understudied, as the majority of searches for opisthobranchs have involved scuba diving. We snorkeled over the intertidal zone during mid-tide, and the shallow subtidal area (<0.5 m) during low tide. During each survey, we collected Bryopsidalean green algae ( Codium , Bryopsis, and Derbesia) and subsequently investigated these in shallow trays on the shore. Slugs used in the feeding exper- iments were collected from the trays (for small species) or directly from submerged algal hosts in the field (for larger species). Slug abundances were determined on an algal bio- mass basis (#/kg wet weight of algae). Seaweeds of the NW Pacific are challengingly diverse. For algal identifications, we relied on Yoshida (1998) and Yoshida et al. (2005) as well as phycological expertise of colleagues in Okinawa (Prof. S. Kamura), mainland Japan (Prof. T. Kobara), and Taiwan (Dr. Jui-Sheng Chang). Codium repens — long reported for Okinawa— is an Atlantic species; Pacific specimens reported by that name belong to a morphologically complex series of species recently reviewed by Chang et al. (2002); Dr. Chang kindly identified our speci- mens as Codium geppiorum. Species documentation For all the species, we took high-resolution digital images to document coloration of living individuals. Voucher speci- mens were measured to the nearest millimeter, relaxed in Table 1. List of survey dates for sacoglossan surveys in Okinawa, Japan. Number of actual Year Month survey i 2002 July 7-12 6 2003 November 4-10 6 2004 March 8-11; July 1-6; November 22-25 4; 4; 4 2005 March 10-15; November 27-December 2 6; 6 2006 March 28-April 1; December 18-24 5; 5 2007 March 4-8 4 2008 December 10-16 5 * Inclement weather or sea conditions prevented us from surveying every day. 3.8% MgCl.,, and then preserved in 5% buffered formalin; vouchers were deposited in the extensive Hirano opistho- branch collection at Chiba University. Our initial identifi- cations were based on external morphological characters; subsequent dissections of radulae and reproductive systems were made to confirm species identifications relative to type descriptions. Feeding experiments Sacoglossans were used for two types of feeding experi- ments when available. In 16 pairwise-choice experiments (Tables 2-4), individual slugs were placed in separate con- tainers with pairwise choices of different green algae known to be host plants of sacoglossans. Containers (ca. 150 ml) were filled with freshly collected seawater and held at room temperature. Periodic observations were made over a 1 or 2-d period, monitoring the location of each individual relative to the algal choices. At the end of each experiment, grazing dam- age was noted. Such experiments are crucial to demonstrate what a sacoglossan prefers to consume, but not whether it could or would consume the less-attractive choice. We also conducted six no-choice experiments to see if specimens would feed on other potential hosts; feeding was determined by recording any visible grazing damage to the algae and/or freshly acquired chloroplasts in the sacoglossan diverticulae (readily visible in most sacoglossans). Because the literature presents confusing and indirect information about one sacoglossan’s algal hosts, we con- ducted eight feeding trials with the undescribed species of Placida Trinchese, 1876. Despite superficial similarities with Placida dendritica, Placida daguilarensis Jensen, 1990, and Placida sp. ( sensu Baba 1986), there are numerous external and internal differences between the undescribed Okinawan Placida sp. and these other three, well-described Placida spe- cies. Published records or photographs of the sacoglossan on Caulerpa sertularioides, Caulerpa lentillifera , and congeners ( e.g ., Ichikawa 1993, Ono 1999) prompted us to investigate whether Placida sp. could or would consume Caulerpa. We also evaluated whether this species would eat Bryopsis, Derbe- sia, Valonia, or the septate Rhizoclonium (Table 2). With Elysia trisinuata, we tested whether the species exhibited a preference between coexisting Codium species and whether the slug would eat Bryopsis in a no-choice situa- tion (Table 3). With Stiliger spp., we conducted four pair- wise-choice and two no-choice experiments to evaluate saco- glossans’ preference and capacity to feed on the sympatric Codium geppiorum, C. arabicum, and Bryopsis harveyana. For Elysia ornata (Swainson, 1840), we conducted four experiments to test whether the species would consume the non-host Derbesia or Codium as well as whether the species would consume sympatric Bryopsis spp. Finally, for Placida cremoniana (Trinchese, 1893), we were constrained by small 170 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Table 2. Feeding preference experiments conducted with Placida sp. (undescribed) in Okinawa between 2002 and 2006. Date Experimental design Result July 2002 • pairwise choice of Codium geppiorum • slugs very strongly preferred Codium; none fed on (host) and Caulerpa sertularioides Caulerpa although a few specimens explored the alga • N =22 slugs • overall 65% on Codium and 7% on Caulerpa • 86% on Codium and 0% on Caulerpa at 24 h July 2002 • pairwise choice of Bryopsis harveyana and • slugs preferred Bryopsis to Caulerpa but did not like Caulerpa sertularioides either choice (few explored either alga); no algae eaten • N= 21 slugs • overall 28% on Bryopsis and 12% on Caulerpa November 2003 • pairwise choice of Codium geppiorum (host) • slugs strongly preferred Codium geppiorum; only 1 even and C. arabicum explored C. arabicum • N- 4 slugs • overall 53% on C. geppiorum and 9% on C. arabicum November 2003 • pairwise choice of Bryopsis harveyana • slugs preferred associating with Derbesia; no evidence and Derbesia sp. of feeding on either alga • N = 2 slugs • overall 64% on Derbesia and 9% on Bryopsis March 2004 • pairwise choice of Codium geppiorum • slugs strongly preferred Codium; did not eat Valonia (host) and Valonia aegagropila • overall 56% on Codium and 0% on Valonia • N = 45 slugs November 2005 • no-choice design of Bryopsis harveyana • slug on alga but did not or could not feed • N = 1 slug • overall 92% on Bryopsis March 2006 • pairwise choice of Codium geppiorum • slugs strongly preferred Codium; did not eat Bryopsis (host) and Bryopsis harveyana • overall 54% on Codium and 15% on Bryopsis • N= 18 slugs March 2006 • pairwise choice of Bryopsis harveyana • slugs preferred to explore Bryopsis to Rhizoclonium and Rhizoclonium grande but did not like either choice; no alga eaten • N = 18 slugs • overall 18% on Bryopsis and 8% on Rhizoclonium sample sizes (N = 3) so conducted just two preference experiments: between Codium spp. and between Derbesia and Bryopsis. RESULTS Environmental conditions In July, the mean seawater temperature during each survey averaged 30-32 °C. In November and December, the mean seawater temperature ranged from 20.5 to 25.6 °C. Finally, during March surveys, mean seawater tem- perature varied from 20.0 to 22.0 °C. Thus, in nearshore habitats, the annual temperature range was at least 12 °C. This pronounced seasonality was associated with the strongly seasonal changes in macroalgal and sacoglossan assemblages. Sacoglossans The species recorded on Bryopsidalean green algae dur- ing this 6-y survey included seven cerata-bearing species and four elysiids (Figs. 2-3). These species were ( 1 ) an undescribed Placida Trinchese, 1876, (2) the well-described Placida sp. ( sensu Baba 1986), (3) Placida cremoniana (Trinchese, 1893), (4) Placida daguilarensis Jensen, 1990, (5) Stiliger ornatus Ehrenberg, 1828, (6) Stiliger aureomarginatus Jensen, 1993, (7) a species of Caliphylla A. Costa, 1867, (8) Elysia trisinuata Baba, 1949, (9) Elysia ornata ( sensu lato) (Swainson, 1840), (10) Elysia rufescens (Pease, 1871), and (11) an undescribed Elysia Risso, 1818. Codium associations Five sacoglossan species that feed on and associate with the perennial green algal host Codium were present during our surveys (Fig. 2). In particular, the small (< 5 mm), unde- scribed Placida sp. (Fig. 2A) was reliably found on the green macroalga Codium geppiorum and the sympatric encrusting congener Codium arabicum at Sobe and, to a lesser degree, at the other two sites. The species was more common on thalli on the seaward side of the reef crest than on the landward side. For example, in March 2005, the species was 2.3 times more common (based on number per kg wet weight of Codium ) on the seaward than the landward side of the fring- ing reef. We collected >320 specimens of Placida sp. (with comparatively little effort) between 2002 and 2008. Peak abundances were recorded in March surveys (Figs. 4-5) when we found 57 to 87 specimens per trip. The species was signifi- cantly more abundant in March than other survey times OKINAWAN SACOGLOSSANS 171 Table 3. Feeding preference experiments conducted with three other sacoglossan species from Codium spp. Sacoglossan species Date Experimental design Results Elysia trisinuata November 2003 • pairwise choice of Codium • strongly preferred Codium geppiorum; only geppiorum (host) and C. arabicum 1 slug even briefly explored C. arabicum • N = 6 slugs • overall 77% on C. geppiorum and 7% on C. arabicum Elysia trisinuata November 2005 • no-choice design of Bryopsis harveyana • 4 slugs tried to feed but no cell sap removed; • N = 8 slugs 2 slugs laid eggs on alga; other 2 slugs avoided alga Stiliger aureo- March 2005 • no-choice design of Bryopsis harveyana • only 1 slug even explored Bryopsis; no feeding marginatus • N= 15 slugs • overall 3% on Bryopsis Stiliger aureo- March 2005 • pairwise choice of Codium arabicum • slugs strongly preferred Codium; only 2 slugs marginatus and Bryopsis harveyana even explored Bryopsis but did not feed on it • N- 15 slugs • overall 53% on Codium and 8% on Bryopsis Stiliger aureo- March 2005 • pairwise choice of Codium geppiorum • fed on both Codium species irrespective of host source marginatus (host of 9) and C. arabicum (host of 2) • overall 41% on C. geppiorum and 43% on C. arabicum • N = 11 slugs Stiliger ornatus November 2003 • pairwise choice of Codium geppiorum • strongly preferred Codium geppiorum; only 1 slug (host) and C. arabicum even briefly explored C. arabicum • N = 2 slugs • overall 45% on C. geppiorum and 5% on C. arabicum Stiliger ornatus March 2005 • pairwise choice of Codium geppiorum • slug preferred Codium geppiorum (host) and C. arabicum • overall 55% on C. geppiorum and 9% on C. arabicum • N = 1 slug Stiliger ornatus March 2006 • no-choice design of Bryopsis harveyana • only 2 slugs even explored Bryopsis; they • N = 12 slugs did not feed on the alga • overall 8% on Bryopsis (Kruskal-Wallis, H = 10.7, P = 0.005). The mean March abundance was ca. 40 individuals per kg Codium (wet weight) (Fig 5A). Based on pairwise-choice experiments (Table 2), Placida sp. from Codium geppiorum (1) strongly preferred C. geppiorum to Caulerpa sertularioides and Valonia aega- gropila, (2) preferred Codium to Bryopsis harveyana , (3) preferred C. geppiorum to sympatric congener C. arabicum, and (4) preferred Derbesia sp. to B. harveyana. These results and the field associations indicate that Codium spp. are pri- mary hosts to Placida sp. Although Placida sp. did explore Derbesia and Bryopsis in experiments, we observed no actual evidence of feeding and documented no field asso- ciations; the algae do not appear to be hosts. Furthermore, based on the lack of these slugs on Caulerpa and Valonia, these algae are clearly not hosts: there was no direct or indi- rect evidence of Placida ever feeding on the delicate Caul- erpa sertularioides or occurring on the alga in the field dur- ing our study. Coexisting sacoglossans on Codium spp. included Placida sp. ( sensu Baba 1986) (Fig. 2B), Stiliger aureomarginatus (Figs. 2D-E), Stiliger ornatus (Fig. 2F), and Elysia trisinuata (Fig. 2G). The well-described but unnamed Placida sp. was observed in 3 of the 1 1 surveys (March 2005, 2006, and 2007); two authors (Shimadu and Kumagai) conducted their thesis research on this species in eastern Honshu, thus aiding in our recognition of the typically more northern species. The small (< 5 mm), brightly colored sacoglossan Stiliger aureomarginatus was seen on Codium geppiorum primarily in March surveys (Figs. 4A, 5C); this report constitutes the first formal record of this species for Okinawa. Peak densi- ties were recorded on 7 March 2007 with 23 individuals per 545 g C. geppiorum. This sacoglossan species is easily con- fused with the small (< 5 mm) Stiliger ornatus, particularly with juvenile slugs; the key distinguishing feature is the color pattern on the cerata: dark cerata with orange tips (Figs. 2D-E). In contrast, S. ornatus specimens have cerata with a yellow proximal area, then a dark blue to black encircling band, followed by a yellow band, and then a dark tip (Fig. 2F). Stiliger ornatus was most abundant in July (Figs. 4A, 5D) although not significantly so (Kruskal-Wallis test, H = 2.0, P = 0.360) due to the large variation among July collec- tions; for both species, grazing damage to Codium geppio- rum was highly visible. In feeding experiments (Table 3), the two species strongly preferred Codium to Bryopsis and would not consume the latter in no-choice trials. Stiliger aureomar- ginatus exhibited no preference between congeneric Codium spp. Although S. ornatus preferred its host (C. geppiorum ) to the coexisting C. arabicum, this may reflect the small sample size (N = 2). 172 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Table 4. Feeding preference experiments conducted with Elysia ornata from Bryopsis and Placida cremoniana from Derbesia in Okinawa between 2002 and 2008. Sacoglossan species Date Experimental design Results Elysia ornata November 2003 • pairwise choice of Bryopsis harveyana • slugs strongly preferred Bryopsis; few (host) and Derbesia sp. slugs explored Derbesia and covered it with • N =7 slugs • mucus; no damage to Derbesia overall 81% on Bryopsis and 5% on Derbesia Elysia ornata November 2003 • no-choice design of either Bryopsis • all Bryopsis was consumed harveyana (host) or Derbesia sp. • no Derbesia was consumed; slugs laid eggs • N = 8 slugs • on Derbesia or sat on it but did not eat it overall 48% on Bryopsis and 31% on Derbesia Elysia ornata March 2005 • no-choice design of Codium arabicum • N = 6 slugs • 3 slugs briefly explored alga but did not feed Elysia ornata March 2005 • pairwise choice of Bryopsis harveyana • slugs explored and consumed both and Bryopsis sp. (host unknown) Bryopsis species • N = 4 slugs • overall 18% on Bryopsis Placida cremoniana November 2003 • pairwise choice of Codium geppiorum • strongly disliked both Codium spp. and C. arabicum • overall 10% on C. geppiorum • N — 3 slugs and 0% on C. arabicum Placida cremoniana November 2003 • pairwise choice of Bryopsis harveyana • 1 small juvenile fed on Bryopsis and grew; and Derbesia sp. (host) other 2 larger slugs fed only on Derbesia • N = 3 slugs • overall 56% on Derbesia and 17% on Bryopsis Elysia trisinuata (Fig. 2G) was found in March, July, and November/December and was comparatively large (20-30 mm long). There was no significant seasonality in Elysia abundance ( H = 4.7, P = 0.093) although March collections tended to be the smallest. In feeding experiments, E. trisinu- ata from Codium geppiorum preferred its host to the coexist- ing but less common C. arabicum; the reciprocal experiment was not feasible but a few slugs were associated with C. arabi- cum on the shore. In a no-choice situation, this species would not eat Bryopsis. Bryopsis associations The delicately branching Bryopsis and Derbesia species were seasonal or ephemeral in Okinawa. Six species of sacoglossans were associated with the abundant Bryopsis harveyana and less common congeners: Elysia ornata ( sensu lato), Elysia rufescens, an undescribed Elysia sp., Placida daguilarensis, Placida sp. {sensu Baba 1986), and Caliphylla sp. Elysia ornata (Fig. 3A-C) was reliably associated with Bry- opsis, particularly in the November/December and March sur- veys when seawater temperatures were low and Bryopsis was abundant in the shallow subtidal areas, particularly near the reef edge. The sacoglossan species was large with specimens up to 52 mm long. We noted that specimens varied from pale body coloration to bright green, depending on the time since feeding on Bryopsis. More importantly, however, the rhino- phore color varied from bright orange or red (Figs. 3A-B) to green with only a faint tinge of red (Fig. 3C); the rhinophores lacked the black markings observed in the undescribed Elysia sp. (Fig. 3F). Black markings on the body surface of£. ornata, including inside and outside parapodia, varied from large (Fig. 3A) to small (Figs. 3B-C). In a pairwise-choice experiment (Table 4), E. ornata strongly preferred its host Bryopsis har- veyana to the morphologically similar Derbesia sp. In no- choice experiments, E. ornata specimens consumed all the Bryopsis but none of the Derbesia; they crawled all over Derbe- sia and spawned on it, but did not feed on it. Elysia rufescens also was frequent on Bryopsis and reached large sizes (42 mm). This species was less common than the large congener Elysia ornata (Fig. 4B). No feeding experiments were conducted with E. rufescens during this study; however, the species was not found in association with any algal host other than Bryopsis (we searched ca. 15 genera of green algae and many genera of red algal hosts during the large-scale study). In November 2004, an undescribed Elysia sp. (Fig. 3F) was found crawling on the benthos, not in association with any alga, but near Elysia ornata specimens which were in a habitat where small Bryopsis thalli grew in rock crevices and holes. However, the 50-mm sacoglossan readily consumed Bryopsis in the laboratory. This species, recognized by many colleagues as undescribed, may have been recorded on other OKINAWAN SACOGLOSSANS 173 dates. The primary distinguishing features of the undescribed Elysia sp. are the three pairs of parapodial thickenings that are tinged orange and the rhinophores with black markings. In E. ornata , the orange and white parapodial lines are typically continuous and the rhinophore coloration is distinct (Figs. 3A-C). The other had black dots and larger, more diffuse white markings inside and outside the parapodia. Detailed comparisons of the internal anatomy of the undescribed spe- cies and E. ornata ( sensu lato) have not yet been made by our research group (or others). A small Placida species (to 8 mm long) was recorded from Bryopsis (Fig. 3E). At first, the animal was lumped with Placida sp. ( sensu Baba 1986). After investigating the internal anatomy of the former, the species was positively identified as Placida daguilarensis Jensen, 1990 and reported as a new record for Japan (see Hirano et al. 2006b, 2006c). Because of our photographic technique and detailed notes, we were able to retroactively distinguish the species in our early records. The differences between P. daguilarensis (Fig. 3E) and the other Placida species include external morphology such as the shape of the cerata and the pericardium (Figs. 2A-C, 3E), internal morphology (reproductive system), branching pat- terns of digestive diverticulae (based on squash mounts of cerata), and algal diet. Placida daguilarensis, originally described from Hong Kong, was not particularly common during our Okinawan surveys but we found many additional specimens in Honshu. Placida sp. ( sensu Baba 1986) in Okinawa was primarily found on Codium. However, on 7 March 2007, we found one individual (8 mm) on Bryopsis harveyana (Fig. 2C). Consistent with our work on Honshu, specimens on Bry- opsis had longer cerata and sparser digestive diverticulae on the rhinophores, head, and dorsum than conspecifics on Codium (Fig. 2B). Finally, another new species record for Japan is the cerata-bearing Caliphylla sp. To date, we have found two specimens of Caliphylla (Fig. 3G) in December 2006 at Sobe (12 and 17 mm). In December 2008, we did not find the species despite intensive searching; however, Bryopsis pop- ulations were not as lush as normal due to prolonged per- sistence of warm-water in autumn. After investigating the internal anatomy of this species, examining the literature, and reviewing our previous photographic and specimen- based records, we also found Caliphylla sp. from Sagami Bay on Honshu in November 2002. Derbesia associations The rather flamboyant Placida cremoniana (Fig. 3H) was found in association with the green alga Derbesia sp. that was entangled around other seaweeds ( e.g ., among the cal- careous fronds of branching coralline red algae). This was the first apparent record of the sporophyte stage of Derbesia in Okinawa. In a pairwise-choice trial (Table 4), P. cremoniana preferred its algal host ( Derbesia ) to Bryopsis but we need more Okinawan specimens to test this more rigorously as one small juvenile fed on Bryopsis and grew. DISCUSSION Species richness Our surveys have yielded several important results about the species richness, trophic associations, and population abundances of sacoglossan opisthobranchs. From 2002 to 2008, we recorded 1 1 species of sacoglossans associated with Bryopsidalean green algae in Okinawa. This value was slightly higher than the 9 species recorded by Gosliner et al. (2008) for the Indo-Pacific region. The only Indo-Pacific species in this feeding guild that we did not record was Elysia punctata Kelaart, 1858 associated with Bryopsis-, based on the species range reported by Gosliner et al. (2008), E. punctata has not yet been recorded in Japan and environs. We consider that our undescribed Elysia sp. that feeds on Bryopsis probably corresponds to Elysia sp. 1 1 of the Sea Slug Forum and Elysia sp. 3 of Ono (2004). Whether our species also corresponds to Elysia sp. 2 of Gosliner et al. (2008) is unclear as their speci- men had a distinctive purple color inside the parapodia that our specimens lacked. This paper reports the new record of Caliphylla in Japan (previously reported by Hirano et al. 2007a). The Okinawan record is not the first record of the genus in the western Pacific or Indo-Pacific region; for example, Carlson and Hoff (2003) reported Caliphylla mediterranea A. Costa, 1867. However, based on the reproductive system, our species does not appear to be the described C. mediterranea. Gos- liner et al. (2008) also reported an undescribed Caliphylla sp. from Hawaii. More detailed ecological, morphological, and molecular investigations need to be conducted before synonymizing the NW Pacific species with the Mediterra- nean one. Furthermore, the recent reports by Camacho- Garcla et al. (2005) and (posthumously) by Jim Lance for the Mexican Pacific (Hertz 2006, Steinberg 2007) and other reports of Caliphylla sp. for the tropical NE Pacific merit reexamination of the genus. Furthermore, the status of Elysia ornata as a single spe- cies or as an assemblage of cryptic species (including Elysia marginata (Pease, 1871) and Elysia grandifolia Kelaart, 1858) is unclear (Jensen 1992, 2001, Jensen and Padmakumar 1999, Rudman 1999, Carlson and Hoff 1978, 2003). We therefore refer to this “species” as E. ornata {sensu lato). Some of this variation is apparent when comparing Figs. 3A-C and numerous photographs in Ono (1999, 2004), Gosliner et al. (2008), and Sea Slug Forum (http://seaslugfo- rum.net/). 174 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2-2010 OKINAWAN SACOGLOSSANS 175 Figure 3. Sacoglossans collected from the green algae Bryopsis spp. (mostly B. harveyana) (A-G) or Derbesia sp. (H) between 2002 and 2008. A-C, Specimens of Elysia ornata ( sensu lato): A (19 mm, 20 December 2006, Sobe); B (35 mm, 5 July 2004, Sobe); and C (40 mm, 10 December 2008, Sobe). D, Elysia rufescens (32 mm, 10 December 2008, Sobe). E, Placida daguilarensis (8 mm, 2 December 2005, Sobe). F, Elysia sp. (50 mm, 22 November 2004, Sobe). G, Caliphylla sp. (12 mm, 23 December 2006, Sobe). H, Placida cremoniana (8 mm, 7 November 2003, Sobe). Figure 2. Sacoglossans collected from the green algae Codium geppiorum and/or Codium arabicum (except for C) during surveys between 2002 and 2008. A, Undescribed Placida sp. (4 mm, 10 December 2008, Sobe). B, Placida sp. ( sensu Baba 1986) (4 mm, 28 March 2006, Sobe). C, Placida sp. ( sensu Baba 1986) (8 mm, 7 March 2007, specimen from Bryopsis harveyana, Toguchi, Okinawa). D, Juvenile specimen of Stiliger aureomarginatus (3 mm, 28 March 2006, Sobe). E, Adult specimen of S. aureomarginatus (6 mm, 10 March 2005, Sobe). F, Stiliger ornatus (6 mm, 10 December 2008, Sobe). G, Elysia trisinuata (29 mm, 29 November 2005, Sobe). 176 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 4. Number of specimens collected in different seasons from 2002 to 2008. A, Sacoglossan species associated with Codium spp. B, Sacoglossans associated with Bryopsis (first four species) or Derbesia (fifth species). Trophic associations Overview We recorded field associations and/or laboratory diets of 1 1 species of Okinawan sacoglossans. Based on our data and published records of other malacologists, we calculated the trophic associations of each species: the frequent hosts as well as the minor associations (Fig. 6). Based on our 6-y study and the literature, ten species (91%) typically or frequently fed on and associated with algal hosts in a single family; one species (9%) fed on hosts in two families. In contrast, on NE Pacific, NW Atlantic, and NE Atlantic shores, sacoglossans typically feed on 2-3 families of Bryopsidalean algae. If we broadened our focus to include ah records of associa- tion and/or feeding across each species’ geographic range, the pattern differs (Fig. 6). Seven of these sacoglossan species (64%) fed on hosts within the same algal family whereas three species (27%) fed on two families of Bryopsidalean green algae and one species (9%) fed on three families. In ecological and evolution- ary terms, how significant are the infrequent or unusual host associations? On one hand, the broader diets do not reflect what the majority of members of a population or species do. On the other hand, the exceptions or the infrequent records may reflect what host-switches starving sacoglossans can and will do when they have depleted ephemeral hosts ( e.g ., Bryopsis). Most of the “infrequent” records are due to the occasional association of Bryopsis- feeding sacoglossans with Derbesia or Pedobesia (F. Derbesiaceae) or of Derbesia- feeding sacoglos- sans with Bryopsis or Pseudoderbesia (F. Bryopsidaceae). A. Undescribed Placida sp. 60 40 20 0 Kruskal-Wallis H= 10.7 P= 0.005 B. Ely si a trisinuata Mar Jul Nov/Dec Figure 5. Abundance of three genera of sacoglossans on Codium spp. (mostly C. geppiorum ) on Okinawajima shores, Japan. Details Many of the associations were expected (based on the published liter- ature and from internet reports). How- ever, we substantially improved the understanding of several species. For example, the host association of Placida cremoniana, a species described in 1893, has long eluded malacologists: we recorded herein the Okinawan speci- mens associated with and fed on Derbe- sia. Additional work in Sagami Bay, Honshu (Hirano et al. 2007c) con- firmed that P. cremoniana feeds on Derbesia as weh as on a related alga Pedobesia ryukyuensis (F. Derbesiaceae). Both genera have the sporophyte (dip- loid) stage as dominant in the life cycle. In contrast, P. cremoniana would not readily consume Bryopsis or Pseudo- derbesia which have the gametophyte (haploid) stage dominant. Although OKINAWAN SACOGLOSSANS 177 # of Sacoglossan Species Figure 6. Diet breadth of sacoglossans associated with Bryopsidalean algae. Bars indicate the number of Okinawan species to feed within a single Bryopsidalean family or between two or more families. Data based on this study and the literature. some previous studies have suggested that Bryopsidalean sporophytes have the polysaccharide mannan as a primary constituent of cell walls and gametophytes have the polysac- charide xylan in walls, phycologists emphasize that these pat- terns are based on only a few algal species and are not abso- lute: life-history stage and wall chemistry are not necessarily linked. Thus, we do not extrapolate about cell wall constitu- ents of Okinawan algae vs. sacoglossan tooth shape. However, we do note that P. cremoniana has strikingly small teeth com- pared to other sacoglossans that feed on Bryopsidalean algae. Malacologists have long recognized that Elysia ornata ( sensu lato) feeds on Bryopsis. However, Hirano etal. (2007c) recently reported that in Japan the species also feeds on Pseudoderbesia (F. Bryopsidaceae) and Pedobesia (F. Derbes- iaceae). Thus, E. ornata is not monophagous: it feeds on many species of Bryopsis and on three genera in two algal families. This result has considerable applied significance as E. ornata and the sympatric conspecific Elysia rufescens contain sec- ondary metabolites effective against certain types of cancer. The role of sacoglossan feeding history and sequestration of algal metabolites may influence the types and amounts of anti-cancer chemicals produced. For example, Rao et al. (2008) reported the absence of kahalalide B, a chemical previ- ously isolated in abundance from Hawaiian E. rufescens , from their sacoglossans despite sites and times identical with previ- ous collections. While the authors suggest potential differ- ences in microbial associates, another realistic explanation would be different algal hosts or past feeding histories. Other sacoglossan species such as Placida daguilarensis have not been studied sufficiently since description for a full understanding of the diet breadth to be possible. Our recent record of the species in Japan, from Okinawa to Hokkaido (Hirano et al. 2006b, 2006c), was a considerable extension of the species known geographic range (namely Hong Kong). Working on a Sagami Bay population of P. daguilarensis, Hirano et al. (2007c) also expanded the known details of its algal associations from just Bryopsis and Derbesia to also Pseudoderbesia and Pedobesia. However, the species is most commonly associated with Bryopsis and prefers it to Pedobesia. In contrast to these records of wider-than-expected diets, the degree of genus-level specificity for the Codium feeders in Okinawa was unexpected. Most Codium- feeding sacoglossans in other geographic regions will consume both Codium and Bryopsis-, these algal genera are in different families but are structurally similar to a sacoglossan (thin-walled coenocytic siphons). With the Okinawan sacoglossan species, there appeared to be little to no trophic overlap between the two genera. For example, this study and one in Sagami Bay (Trow- bridge et al. 2008a) supported earlier work by Jensen (2003) that Elysia trisinuata does not appear to feed on Bryopsis. The undescribed Placida sp. also appeared to feed just on Codium; we have no records of it on Bryopsis. In contrast, Elysia setoen- sis Hamatani, 1968 from Honshu does feed on both genera as well as several other genera (Trowbridge et al. 2008a). Fur- thermore, ecological work on Placida sp. ( sensu Baba 1986) also reported the species fed on Codium and Bryopsis on Hon- shu shores with infrequent records from Pedobesia (Shimadu 2004, Shimadu et al. 2006, Kumagai 2009, Hirano, unpubl. data). Are these differences in degree of feeding specificity ecologically or genetically determined? Intriguingly, there was no evidence of monophagy in three sacoglossan species feed- ing on Codium in Sagami Bay, (Trowbridge etal. 2009b); they ate many of the approx. 20 species of Codium in Japan. Cer- tainly, the sacoglossan species reported herein from Okinawa are reported on other Codium spp. elsewhere in their range. Population abundances The population abundances of the Codium-feeding saco- glossans (Figs. 5, 7) seasonally peak at an average of ca. 5 slugs per kg algae (for Elysia trisinuata ), ca. 15 (two Stiliger spp.), and ca. 40 (undescribed Placida sp.). The largest of these spe- cies ( Elysia ) has the smallest population densities. How do 178 AMERICAN MALACOLOGICAL BULLETIN 28 -1/2-2010 A. Mean values 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -2.5 l i Dry weight (range) l Wet weight Elysia Stiliger Placida Sacoglossan Genera B. Maximum values 1.0 0.5 0.0 -0.5 -1.0 -1.5 -2.0 -2.5 t t Dry weight (range) Wet weight t Elysia Stiliger Placida Sacoglossan Genera Figure 7. Comparison of population densities of sacoglossans based on the number per gram of algal wet weight or dry weight, using mean (A) or maximum (B) slug abundances. Because the wet to dry weight conversion was based on a published range (1 kg Codium wet weight would be ca. 40-70 g dry weight), a range of dry weight estimates is shown. The horizontal dotted lines indicate sacoglossan population density predicted for Okinawan latitude, based on the latitudinal trend of Atlantic species (Clark and DeFreese 1987). these values compare to sacoglossan abundances reported for other regions? Clark and DeFreese (1987) reported 32.4 slugs/g dry weight of Codium in Connecticut (41°N) (for Placida dendritica) and 24.2/g dry weight of Bryopsis at 25°N (for Placida kingstoni). Based on sacoglossan populations spanning 40° latitude in the N Atlantic, Clark and DeFreese (1987) reported the following linear regression: logI0 (den- sity) = 0.1 109 (latitude) - 2.9683. Based on our 23°N location at Okinawa, we would predict densities of ca. 0.4 specimens per g dry weight of algae (Fig. 7, note log10 scale). Our study reported slug abundances based on algal wet weights, as a drying oven was not available. If the dry to wet weight ratio for Codium fragile (reported by Trowbridge 1998) can be extrapolated to Codium geppiorum, then 1 kg of Codium wet weight would be ca. 40-70 g dry weight. Consequently, our mean sacoglossan abundances would be estimated to be ca. 0.1 slugs/g dry weight algae ( Elysia ) to ca. 0.6- 1.0 slugs/g ( Placida ) at 23°N, lower than predicted for Atlantic populations (Fig. 7A). However, the Atlantic regression was based on peak densities rather than mean densities in the peak season. If we calculate peak densities based on maximum abundance of a species in a given collection, our estimates would be 0.4-0. 7 ( Elysia in July 2004) to 2. 1-3.7 slugs/g dry weight ( Placida in March 2006). Thus, the NW Pacific values would then be comparable to or higher than those slug populations in the NW Atlantic Ocean (Fig. 7B). A more rigorous examination of Pacific sacoglossan species along a large latitude gradient would provide a more precise comparison. However, our preliminary comparison indicates that Pacific sacoglossan populations are not only highly diverse but also compara- tively abundant for their latitude. Furthermore, qualitative observations on websites or in guidebooks may contribute to misconceptions about species’ abundance, diversity, and seasonality unless the sampling biases are explicitly indicated. Bolland repeatedly states on his website that many of the sacoglossans are extremely unusual to rare; yet our research group reliably finds these species during most of our surveys. For example, Bolland reported the undescribed Placida sp. (which he called Placida dendritica) as “very rare in the Keramas” and “currently unknown from the waters of Okinawa’s main island” (http://rfbolland.com/okislugs/placdend.html) whereas we have found >320 specimens of this species on the main island. How do we explain these differences? The primary difference is probably one of methodology: we snorkeled in extremely shallow water where macroalgae proliferate, whereas Bolland and colleagues generally dive in deeper water, using scuba. Thus, the two research groups are inves- tigating slightly different habitats. Furthermore, we have OKINAWAN SACOGLOSSANS 179 taken a phytocentric approach, investigating large numbers of thalli of each potential algal host. Because Bolland’s sam- pling methodology has not been specifically reported, we presume he and other colleagues (generally oriented more to nudibranchs than to sacoglossans) may have a less phy- tocentric approach. Consistent surveys by the same investi- gators at the same sites throughout many years will enhance the understanding of assemblages of sacoglossan opistho- branchs. Also, as well stated by Clark (1994: 901): “Once food species have been identified, sacoglossans can be relo- cated repeatedly”. By reporting the algal associations in this paper, we hope other investigators can start reliably and quantitatively recording populations of subtropical saco- glossans. A comparable point was made by Bouchet et al. (2002: 427): “The existence of such species [very stenoe- cious] is revealed only if the hosts are appropriately sam- pled and scrutinized.” At special risk vs. predictably rare? Different types of rarity Are “sparse” or “uncommon” mollusc populations necessarily degraded or at “special risk” as Clark (1994), Bolland (web site), and others have suggested? Clark (1994) emphasized that one source of vulnerability of the Key West sacoglossans was the high incidence of direct devel- opment (and hence, low dispersal); this high incidence is unusual within the group. However, he also suggested that planktotrophic larvae may not necessarily disperse large distances, so all sacoglossans may be at “special risk” due to their host specificity and known habitat destruction or deg- radation. Ecological theory, however, indicates that “uncommon” to “rare” species are not necessarily a conservation risk; ecol- ogists have long realized that there are fundamentally differ- ent forms or types of “rarity” in species (Rabinowitz 1981). Species can be rare in different ways: by having a small geo- graphic range, by having extreme habitat specificity, and by having small local populations (Table 5). The 2x2x2 matrix of the three traits lead to 8 cases: 7 of these denote different types of rare species and 1 indicates common species. If we characterize geographic range, habitat specificity, and local population abundance of sacoglossans, we would have a much better understanding of the vulnerability or conserva- tion status of these species. Larval development and population size Most of the 100-150 recorded Japanese sacoglossan spe- cies have planktotrophic larvae (Trowbridge et al., unpubl. data) and large geographic distributions and, thus, are prob- ably not highly vulnerable to local disturbance or change. At least 8 of the 1 1 species discussed herein have planktotrophic larvae ( Caliphylla sp., undescribed Elysia sp., and Elysia rufe- scens have not yet been investigated). Some sacoglossans (such as Elysia setoensis in Japan) do have a broad algal-host range with many genera of algae used; the broad host range can function as a buffer. Furthermore, temperate sacoglos- san species often have high local population sizes (Clark and DeFreese 1987) and, hence, may often be considered eco- logically common. Elysia viridis is an excellent example of this case with wide distribution, broad suite of algal hosts, and large population sizes (see Trowbridge et al. 2009c). Sacoglossans that are more stenophagous in host asso- ciations, limited to one or just a few algal genera, could be considered (1) “predictably rare” ( e.g ., undescribed Placida sp. and Elysia trisinuata ) — such that they can be found if the specific habitat is investigated — or (2) “sparse” (e.g., Caliphylla sp.) — such that the local population size is small and the species cannot be consistently located. The latter type of species is vulnerable to habitat loss such as the local or regional elimination of a host species (to dis- ease, disturbance, pollution); the relative size of local pop- ulations and the capacity to disperse mediate the species’ susceptibility to extrinsic factors. Yet, the situation dis- cussed by Clark (1994) where specialists are “at special risk” of natural or anthropogenic change would occur pri- marily in species with restricted geographic distributions and/or limited dispersal due to direct development. Thus, Table 5. Different forms of rarity based on three species traits: size of geographic range, habitat specificity, and local population size [based on Rabinowitz (1981)]. Of the eight possible combinations, seven cases denote different types of rarity and one case is the common species. NW Pacific sacoglossans generally fall in the two categories indicated with boldface, capitalized font. Geographic distribution Wide Narrow Habitat specificity Broad Restricted Broad Restricted Local population Somewhere large Common Predictable Locally common Endemic size Everywhere small Sparse Sparse Sparse Sparse 180 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 most NW Pacific sacoglossans are “predictably rare” to “sparse” rather than at “special risk” of local extinction. ACKNOWLEDGMENTS This study was initiated with funding from the Women-In-Science Collaboration (WISC) program from AAAS/NSF (2002-2003) and the Japanese Society for the Promotion of Science (JSPS) Grant-in-Aid for Scientific Research C-15570073 (2003-2006). We thank our malaco- logical colleagues for extensive discussions about these sacoglossans, particularly W. Rudman, C. Carlson, P. J. Hoff, P. Krug, K. Jensen, and D. Behrens. Constructive comments by two reviewers and K. Brown significantly improved this paper. We thank numerous colleagues for their phycological expertise, particularly S. Kamura, T. Kobara, and J.-S. Chang. LITERATURE CITED Baba, K. 1986. Anatomical information on Placida sp. = Hermaea dendritica of Baba, 1937 and 1955, from Japan. Shells and Sea Life 18: 21-22. Bouchet, P„ P. Lozouet, P. Maestrati, and V. Heros. 2002. Assessing the magnitude of species richness in tropical marine environ- ments: Exceptionally high numbers of molluscs at a New Cale- donia site. Biological Journal of the Linnean Society 75: 421-436. Camacho-Garcia, Y., T. M. Gosliner, and A. Valdes. 2005. Field Guide to the Sea Slugs of the Tropical Eastern Pacific. California Academy of Sciences, San Francisco. Carlson, C. and P. J. Hoff. 2003. The opisthobranchs of the Mariana Islands. Micronesica 35-36: 272-295. Carlson, C. H. and P. J. Hoff. 1978. The identifiable Elysia from Guam (Elysiidae, Sacoglossa, Opisthobranchia). Micronesica 14: 89-113. Chang J.-S., C.-F. Dai, and J. Chang. 2002. A taxonomic and karyo- logical study of the Codium geppiorum complex (Chlorophy- ta) in southern Taiwan, including the description of Codium nanwanense sp. nov. Botanical Bulletin of Academia Sinica 43: 161-170. Clark K. B. 1994. Ascoglossan (=Sacoglossa) molluscs in the Flor- ida Keys: Rare marine invertebrates at special risk. Bulletin of Marine Science 54: 900-916. Clark, K. B. and D. DeFreese. 1987. Population ecology of Caribbean Ascoglossa (Mollusca: Opisthobranchia): A study of specialized algal herbivores. American Malacological Bulletin 5: 259-280. Gosliner, T. M. 1992. Biodiversity of tropical opisthobranch gastro- pod faunas. Proceedings of the Seventh International Coral Reef Symposium, Guam 2: 702-709. Gosliner, T. M„ D. W. Behrens, and A. Valdes. 2008. Indo-Pacific Nudi- branchs and Sea Slugs: A Field Guide to the World’s Most Diverse Fauna. Sea Challengers and California Academy of Sciences, Gig Harbor, Washington and San Francisco. Guiry, M. D. and G. M. Guiry. 2009. AlgaeBase. World-wide electronic publication, National University of Ireland, Gal- way. Available at: http://www.algaebase.org; accessed 27 March 2009. Hertz, C. 2006. In remembrance of James Robert Lance. TheFestivus 38: 109-110. Hirano, Y. J., Y. M. Hirano, and C. D. Trowbridge. 2005. Sacoglossan opisthobranch gastropods on west-coast shores of Okinawa- jima: Diversity and richness. 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Morisawa, ed., A Scientific Study to Estab- lish a Marine Reserve for Environmental Preservation of Sagami Bay. Final JSPS (Japan Society for the Promotion of Science) Project Report, National Diet Library, Kyoto. Pp. 181-204 [In Japanese] . Hirano, Y. M., C. D. Trowbridge, and Y. J. Hirano. 2007b. Unrecog- nized diversity of the genus Ercolania (Mollusca, Gastropoda, Sacoglossa) on Japanese shores. 43rd Annual Meeting of the Jap- anese Society of Systematic Zoology, Kitakyushu, Kyushu, 9-10 June 2007 [In Japanese], Hirano, Y. M„ C. D. Trowbridge, and Y. J. Hirano. 2008. Redis- covery of “Stiliger akkeshiensis” (Opisthobranchia, Sacoglossa) with a review of its systematic position. Umiushi-Tsushin No. 59: 6-7 [In Japanese]. Hirano, Y. M., C. D. Trowbridge, K. Sudo, and Y. J. Hirano. 2007c. Food preference of Placida cremoniana (Mollusca, Gastropo- da, Sacoglossa). 78th Annual Meeting of the Zoological Society of Japan, Hirosaki, Honshu, 20-22 September 2007 [In Japanese]. Ichikawa, M. 1993. Saccoglossa (Opisthobranchia) from the Ryukyu Islands. Publications of the Seto Marine Biological Laboratory 36: 119-139. Jensen, K. R. 1992. Anatomy of some Indo-Pacific Elysiidae (Opis- thobranchia: Sacoglossa (=Ascoglossa)), with a discussion of the generic division and phylogeny. Journal of Molluscan Studies 58: 257-296. Jensen, K. R. 2001. (Nov 26) Re: Elysia ornata and E. grandifolia. [Mes- sage in] Sea Slug Forum. Australian Museum, Sydney. Available from http://www.seaslugforum.net/find.cfm/icU5727; accessed 14 December 2009. Jensen, K. R. 2003. Distributions, diets and reproduction of Hong Kong Sacoglossa (Mollusca: Opisthobranchia): A summary of OKINAWAN SACOGLOSSANS 181 data, 1980-2001. In: B. Morton, ed., Perspectives on Marine En- vironmental Change in Hong Kong and Southern China, 1977- 2001. Proceedings of an International Workshop Reunion Con- ference, Hong Kong 21-26 October 2001. Hong Kong University Press, Hong Kong. Pp. 347-365. Jensen, K. R. 2007. Biogeography of the Sacoglossa (Mollusca, Opist- hobranchia). Bonner zoologische Beitrage 55: 255-281. Jensen, K. R. and K. Padmakumar. 1999. Description of three spe- cies of Elysia (Opisthobranchia, Sacoglossa) from Southern In- dia with a discussion of the identity of E. grandifolia Kelaart, 1858. Phuket Marine Biological Center Special Publication 19: 245-246. Kumagai, K. 2009. Feeding Ecology of Sacoglossa — Capacity to Switch Food and Variation of the Radular Morphology. M.Sc. Disserta- tion, Chiba University [In Japanese]. Ono, A. 1999. Opisthobranchs of Kerama Islands. TBS - Britannica Co., Ltd, Tokyo [In Japanese], Ono, A. 2004. Opisthobranchs of Ryukyu Islands. Rutles Inc., Tokyo [In Japanese], Rabinowitz, D. 1981. Seven forms of rarity. In: H. Synge, ed., The Biological Aspects of Rare Plant Conservation. John Wiley and Sons, Chichester, West Sussex and New York. Pp. 205-217. Rao, K. V., M.-K. Na, J. C. Cook, J. Peng, R. Matsumoto, and M. T. Hamann. 2008. Kahalalides V-Y isolated from a Hawaiian collection of the sacoglossan mollusk Elysia rufescens. Journal of Natural Products 71: 772-778. Rudman, W. B. 1999 (November 4) Elysia ornata (Swainson, 1840). [In] Sea Slug Forum. Australian Museum, Sydney. Available from http Wwww.seaslugforum. net/ factsheet.cfm?base=elysorna; accessed 14 December 2009. Shimadu, Y. 2004. Food Algae and Life History of Placida sp. (sensu Baba, 1986) (Sacoglossa). M.Sc. Dissertation, Chiba University [In Japanese]. Shimadu, Y., Y. M. Hirano, C. D. Trowbridge, and Y. J. Hira- no. 2006. Food algae and life history variation in Placida sp. ( sensu Baba, 1986). 53rd Annual Meeting of the Ecologi- cal Society of Japan, Niigata, Honshu, 24-28 March 2006 [In Japanese], Steinberg, J. E. 2007. Opisthobranchs from the unpublished materi- als of James R. Lance. Opisthobranch Newsletter 28: 1-5. Trowbridge, C. D. 1998. Ecology ofthe green macroalga Codium frag- ile (Suringar) Hariot 1889: Invasive and non-invasive subspecies. Oceanography and Marine Biology: an Annual Review 36: 1-64. Trowbridge, C. D., Y. J. Hirano, and Y. M. Hirano. 2005. Sacoglos- san opisthobranch gastropods on west-coast shores of Oki- nawajima: Green algal host associations. Venus 64: 80. Trowbridge, C. D., Y. M. Hirano, K. Sudo, and Y. J. Hirano. 2006. Elysia trisinuata and E. sugashimae — distinct species or intraspe- cific variation? Venus 65: 272 [In Japanese]. Trowbridge, C. D., Y. J. Hirano, and Y. M. Hirano. 2007. Inven- tory of Japanese sacoglossan opisthobranchs: Historical review, current records, and unresolved issues. Venus 66: 117. Trowbridge, C. D., Y. J. Hirano, and Y. M. Hirano. 2008a. Saco- glossan opisthobranchs associated with the green macroal- gae Codium spp. on Pacific rocky shores of Japan. Venus 66: 175-190. Trowbridge, C. D., Y. J. Hirano, and Y. M. Hirano. 2008b. Sacoglos- san opisthobranchs on NW Pacific shores: Stiliger berghi Baba, 1937, and Elysia sp. on filamentous red algae. Venus 67: 108. Trowbridge, C. D., Y. J. Hirano, and Y. M. Hirano. 2009a. Saco- glossan opisthobranchs on northwestern Pacific shores: Stiliger berghi Baba, 1937 and Elysia sp. on filamentous red algae. The Veliger 51: 43-62. Trowbridge, C. D., Y. M. Hirano, and Y. J. Hirano. 2009b. Interac- tion webs of marine specialist herbivores on Japanese shores. Journal of the Marine Biological Association of the United Kingdom 89: 277-286. Trowbridge, C. D., C. Little, P. Stirling, and W. F. Farnham. 2009c. Sacoglossan gastropods on native and introduced hosts in Lough Hyne, Ireland: Larval retention and population asyn- chrony? Journal ofthe Marine Biological Association of the Unit- ed Kingdom 88: 771-782. Wikipedia. 2009. Ryukyu Islands, from Wikipedia, the free encyclopedia. Available at: http://en.wikipedia.org/wiki/ Ryukyu_Islands; accessed 26 March 2009. Yoshida, T. 1998. Marine Algae of Japan. Uchida Rokakuho Publish- ing Co., Ltd., Tokyo [In Japanese], Yoshida T., S. Shimada, K. Yoshinaga, and Y. Nakajima. 2005. Checklist of marine algae of Japan (revised in 2005). Japanese Journal of Phycology 53: 179-228. Submitted: 27 March 2009; accepted: 16 June 2009; final revisions received: 15 December 2009 Amer. Malac. Bull. 28: 183-184 (2010) RESEARCH NOTE Self-adhesive wire markers for bivalve tag and recapture studies Lance W. Riley, Shirley M. Baker, and Edward J. Phlips Fisheries and Aquatic Sciences Program, School of Forestry Resources and Conservation, University of Florida, Gainesville, Florida 32653, U.S.A. Corresponding author: phlips@ufl.edu Abstract: Bivalves are good candidates for tag and recapture studies because the accrual of shell material provides a stable record of growth. Obtaining measurements for tagged individuals over time relies on the resilience of markings or tagging devices to environmental stress as well as the readability of identifying markings upon capture. Tagging devices should also be easy and quick to apply in order to minimize potential stress to the animal during extirpation from the water. A variety of methods to attach devices to the inside and outside of bivalve shells have been used in tagging studies. This paper describes a low cost, commercially available, self-adhesive numbered tag for application to clams and other bivalves. Tests of the tag demonstrate its resilience to severe conditions over a short time scale with a ninety- five percent recovery of individuals with tags remaining intact. Ninety-four percent recovery of individuals with tags intact was also achieved in long-term studies. No problems with legibility of tags recovered occurred in any of the tests. Key words: shell, numbering, clam, enumeration, individuals Tag and recapture studies are important methods for a wide range of physiological and ecological research applica- tions, such as accurate measures of growth and survival of indi- viduals within populations. Tagging methods for bivalves can be classified into two main categories, internal and external. Placement of internal tags within shell cavities, such as passive integrated transponder tags (Kurth et al. 2007), is much less common than external application. Internal tags are generally more costly, require some sort of locator/reader or destructive sampling for recapture, and can stress the animal more during device application than externally applied tagging devices. External tagging does not require relaxation to open shell valves for internal tag insertion, possible rejection of the tag by the animal, or specialized locating equipment, unlike internal tags that cannot be found by visual observation. External tag- ging of bivalves is practical because of the hardness and stable surface area of the shell. Many approaches to external tagging involve physical alteration of the shell surface, such as engrav- ing (McMahon and Williams 1986, Mattice and Wright 1986) or permanent dye markers and paints (Buttner and Heidinger 1980). Other methods require adhesives, such as numbered bee-type tags (Lemarie et al. 1995, Kurth et al. 2007), elec- tronic Passive Integrated Transponder (PIT) tags (Kurth et al. 2007), monofilament tags (Layzer and Heinricher 2004), and anchored tethers attached to the shell (Foe and Knight 1986). Other monofilament tags anchor into shell ligaments without the use of adhesives (Lim and Sakurai 1999). Problems with external tags include decreased tag legibility/recovery result- ing from abrasion with sediments and stress from handling due to drying times necessary for adhesives. To avoid tags altogether, cages with numbered slots for individual clams have been used (Foe and Knight 1986, Cataldo et al. 2001). In this study, we tested the use of self-adhesive markers (originally designed for marking electrical wire) for tagging bivalves. Thomas & Betts Brand EZ-Code® Wire Markers (Product #WM-0-90, Thomas & Betts Inc., USA) were cho- sen because of their small size, readability, self-adhesive strength, and low cost (pack of 455 tags for about US$32 = $0.07 per tag). These self-adhesive, vinyl-coated cloth tags are intended for identification of wires, breakers, and service panels in the electrical industry and are therefore widely avail- able through local and online electrical products suppliers. The tags have 4 mm-tall black numbers with a 5 mm2 white background and a thickness of less than 0.5 mm. There are a variety of numbers, letters, and combination number/ letter schemes available (see the Thomas & Betts internet- based catalog, www.tnb.com). The tags are applicable for bivalve studies, even for small-sized animals such as Corbicula fluminea (Muller, 1774) since the tags are small but are read- able without magnification (Fig. 1) and stand out in contrast to untagged clams and substrates. Because of the ease of application, the out of water time of the test organisms is minimal. The tag application also causes less shell damage than engraving and time out of water than paint and tradi- tional liquid adhesive/tag applications. Two tests were performed to evaluate the durability of the markers for bivalves: ( 1 ) exposing tagged individuals to short- term extreme physical stress conditions and (2) longer-term 183 184 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 1. Tag applied to clam shell. exposure of tagged bivalves to simulated natural environ- ments in stream-type flumes. The freshwater clam ( Corbicula sp.) was used as the test organism. Tags were applied to the middle of one shell valve of clams that had been towel-dried. Tags were allowed to cure for approx. 10 seconds to ensure adhesion, even though it was found that individuals could be submersed in water without the loss of the tags directly after application. No external coatings or adhesives were added to the tags prior to placement onto the shells. The legibility of the numbers on each tag was noted after each experiment. The first test of durability was performed on a sample of 20 live clams obtained from the Santa Fe River at The Rapids, near High Springs, Florida in December 2001. The animals were placed in a 1-L Nalgene® bottle along with 200 mL of coarse-grade sand and 200 mL of water. The mixture was capped and shaken vigorously by hand for 15 minutes, after which the clams were removed for inspection. Nineteen of 20 clams retained their tags for a 95% recovery with no loss of legibility. All clams exhibited chipping around shell margins with exposure of soft tissue, suggesting that the conditions in this test were harsher than those normally encountered in most natural environments. In long-term tag exposure experiments, 972 tagged clams were stocked in large, sand-bottom, outdoor aquaculture raceways. Water velocities in the raceways ranged from 0.013 to 0.020 m s'1, and water depths were maintained at 0.2 m. A total of 324 clams were stocked in each of 3 separate experi- ments, for time periods of 440 days, 290 days, and 210 days in June 2002, November 2002, and February 2003, respectively. Stocks were obtained from locations in the Santa Fe River at Sandy Point, Florida, Lake Dalhousie near Umatilla, Florida, and Lake George near Astor, Florida. Clams were tagged and distributed in a stratified random fashion in the input, mid- dle, and output sections of each raceway. Of the 972 tagged individuals, only 55 tags were not unaccounted for at the end of the experiments, or a 94% recovery. All of the tags attached to clam shells remained legible. The durability, longevity, and readability of tags in this study indicates the self-adhesive tags are an ideal method for assessing the dynamics and condition of bivalves in aquacul- ture or for natural populations. A large percentage of the tags remained intact on the surface of the clam shells and remained legible after exposure to both extreme conditions and long- term exposure. We suggest the Thomas & Betts EZ-Code Wire Markers are an acceptable alternative for use in tag and recapture studies of bivalves in natural and engineered systems. LITERATURE CITED Buttner, J. K. and R. C. Heidinger. 1980. Seasonal variations in growth of the Asiatic clam, Corbicula fluminea (Bivalvia: Corbiculidae) in southern Illinois fish pond. The Nautilus 94: 8-10. Cataldo, D. H., D. Boltovskoy, J. Stripekis, and M. Pose. 2001. Con- dition index and growth rates of field caged Corbicula fluminea (Bivalvia) as biomarkers of pollution gradients in the Parana River delta (Argentina). Aquatic Ecosystems Health and Man- agements. 187-201. Foe, C. and A. Knight. 1986. The effect of phytoplankton and sus- pended sediment on the growth of Corbicula fluminea (Bival- via). Hydrobiologia 127: 105-115. Kurth, J., C. Loftin, and J. Zydlewski. 2007. PIT tags increase effec- tiveness of freshwater mussel recaptures. Journal of the North American Benthological Society 26: 253-260. Layzer, J. B. and J. R. Heinricher. 2004. Coded wire tag retention in ebonyshell mussels Fusconaia ebena. North American Journal of Fisheries Management 24: 228-230. Lemarie, D. P., D. R. Smith, and R. F. Villella. 1995. Evaluation of tag types and adhesives for marking freshwater mussels. The Conservation and Management of Freshwater Mussels II: Initia- tives for the Future, 16-18 October 1995, St. Louis, Missouri. National Biological Service, Kearneysville, West Virginia. Lim, B. K. and N. Sakurai. 1999. Coded wire tagging of the short necked clam Ruditapes philippinarum. Fisheries Science Tokyo 65: 163-164. Mattice, J. S. and L. L. Wright. 1986. Aspects of growth of Corbicula fluminea. Proceedings of the Second International Corbicula Symposium. American Malacological Bulletin, Special Edition 2: 167-178. McMahon, R. F. and C. J. Williams. 1986. A reassessment of growth rate, life span, life cycles and population dynamics in a natural population and field caged individuals of Corbicula fluminea (Muller) (Bivalvia: Corbiculacea). Proceedings of the Second International Corbicula Symposium. American Malacological Bulletin, Special Edition 2: 156-166. Submitted: 23 September 2008; accepted: 30 July 2009; final revisions received: 19 October 2009 Amer. Malac. Bull. 28: 185-188 (2010) RESEARCH NOTE Occurrence of the red abalone Haliotis rufescens in British Columbia, Canada Alan Campbell, Ruth E. Withler, and K. Janine Supernault Department of Fisheries and Oceans Canada, Pacific Biological Station, Nanaimo, British Columbia V9T 6N7, Canada Corresponding author: Ruth.Withler@dfo-mpo.gc.ca Abstract: We document the first occurrence of a live red abalone, Haliotis rufescens Swainson, 1822, found on the central coast of British Columbia. The initial identification was based on morphological characteristics. Previously, the northern or “pinto” abalone Haliotis kamtschatkana kamtschatkana Jonas, 1845 has been the only abalone species considered to naturally occur in the coastal waters of British Columbia. Since hybridization of H. kamtschatkana with H. rufescens was known to occur, genetic analysis of tissue samples was undertaken to confirm the morphological identification as a purebred red abalone. Key words: genetics, VERL, hybridization, Haliotis kamtschatkana Along coastal regions of the eastern North Pacific Ocean, from Baja California to Alaska, there are seven abalone Hali- otis species, all of which coexist in the waters off California and northern Mexico (Geiger 2000, Geiger and Poppe 2000). One species, Haliotis kamtschatkana, has been recognized as two morphologically distinct subspecies. These are the nor- thern or “pinto” abalone Haliotis kamtschatkana kamts- chatkana Jonas, 1845 and the threaded abalone Haliotis kamtschatkana assimilis Dali, 1878. Abalone were once suffi- ciently abundant to support commercial and recreational fisheries in British Columbia (B.C.), Canada, coastal states of the U.S.A., and Baja California, Mexico. Abundances of all abalone species have declined and most abalone fisheries have been closed (Rogers-Bennett 2007). In B.C., the northern abalone was legally listed and protected in June 2003 as threatened under Schedule 1 of the Species at Risk Act. The northern abalone has been the only abalone species consid- ered to naturally occur in the coastal waters of B.C. based on surveys spanning 30 years (Sloan and Breen 1988, Adkins 1996, Campbell 2000, Hankewich et al. 2008). Herein, we document the identification of an unusual abalone that was discovered by a diver on the north end of Athabaskan Island, central coast of B.C. (52°3.26’N, 128°18.18>W) on 18 Decem- ber 1998. Vernier caliper measurements of the shell were length 210 mm, width 170 mm, and height 65 mm. The inside edge of the shell was red and the epipodium was black. The abalone, a female based on its green gonad, was tentatively identified as a red abalone, H. rufescens Swainson, 1822, according to the morphological characteristics of the shell and epipodium (Cox 1962, Mottet 1978, Haaker et al. 1986). The North American abalone species have overlapping species distributions in the southern part of their ranges, where at least one of the subspecies of Haliotis kamtschatkana has been sympatric with red abalone (Fig. 1). Close genetic relationships among the North American abalone species have been documented in a number of studies on mito- chondrial and nuclear DNA sequences, and the underlying evolutionary forces responsible for the relatively recent diver- gence of the species identified ( e.g ., Swanson and Vacquier 1998, Geiger 2000, Swanson et al. 2001, Galindo et al. 2002, 2003). All species recognized on the basis of morphological differences have been confirmed to be distinct on the basis of genetic sequences, with only two subspecies of H. kamtschat- kana indistinguishable on the basis of molecular analysis (Gruenthal and Burton 2005, Supernault et al. 2009). Hybridization of Haliotis kamtschatkana with Haliotis rufescens is known to occur (Owen et al. 1971, Talmage 1977). Analysis of nuclear DNA sequences, in which one copy of the DNA is inherited from each parent, is useful in distinguishing purebred organisms from first generation (F ) hybrids. Consequently we examined DNA sequences for the Vitelline Egg Receptor for Lysin (VERL) gene, which differ in nucleotide sequence and/or length among species of abalone (Galindo et al. 2003), from the abalone of unknown provenance. Amplification of the VERL sequence from genomic DNA of the unidentified abalone was carried out according to Super- nault et al. (2009) with the primers of Galindo et al. (2003). Cloned amplified products were sequenced in duplicate and the VERL amplification products from the unknown specimen were sized on an ABI 3730 Sequencer (Supernault et al. 2009). VERL sequence data were analyzed using Sequencher 4.5 (GeneCodes, Ann Arbor, Michigan) and MEGA 4.0 (Tamura et al. 2007) and aligned with abalone VERL sequences from GenBank for pink ( Haliotis corrugata Wood, 185 186 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 1. Distribution of the northern/threaded abalone Haliotis kamtschatkana (indicated by light shading) and the red abalone Haliotis rufescens (indicated by dark shading) along the Pacific coast of North America. The location of the red abalone found on Athabaskan Island, B.C., is shown with the circle indicated by an arrow. 1828); green (Haliotis fulgens Philippi, 1845); black (Haliotis cracherodii Leach, 1814); Haliotis discus hannai Ino, 1953; northern/threaded (Haliotis kamtschatkana kamtschatkana/ assimilis ); red (Haliotis rufescens ); flat (Haliotis wallalensis Stearns, 1899); and white abalone (Haliotis sorenseni Bartsch, 1940) (Galindo et al. 2003, Gruenthal and Burton 2005). MEGA was used to calculate pairwise nucleotide (p) dis- tances for 775 base pairs (bp) of the unknown sequence and known VERL sequences from northern and red abalone and to construct a multispecies neighbor-joining dendrogram of VERL based on pairwise p distances. The new VERL sequence was submitted to GenBank (accession number EU939887). We obtained 980 bp of sequence of the VERL gene from the unknown specimen, encompassing 5’ to 3’, the entire repeats 1 and 2, and 100 bp into repeat 3. The two sequences obtained from the unknown specimen were identical. The pairwise nucleotide (p) distance obtained from the compari- son of this VERL sequence with the red abalone VERL sequence was 0.004 (Table 1). In contrast, the pairwise p dis- tances between the unknown VERL sequence and northern/ threaded sequences were approximately ten times greater, ranging from 0.034 to 0.053 (Table 1). Alignment of the new sequence with GenBank sequences of all seven distinct North American and one common Asian abalone species of the North Pacific Ocean, Haliotis discus hannai, also confirmed the morphological identification of the specimen as a red abalone, Haliotis rufescens (Fig. 2). Comparison of species-specific sites of the VERL amplified product from the unknown individual with sequences from red and northern/threaded abalone provided a match with the red abalone (Table 2). Sizing of the VERL amplification products from genomic DNA extracted from the specimen revealed a single size of 980 bp. Thus, the individual was unlikely to be a hybrid between red and northern abalone. An Fj hybrid would be expected to possess a VERL sequence from each parental species, and the VERL sequence of the northern/threaded abalone is 1010 bp long. According to Cox (1962), the natural geographic dis- tribution of Haliotis rufescens ranges from Sunset Bay, Oregon to Bahia San Bartolome, Baja California, including the Farallon and Channel Islands. Haliotis kamtschatkana ranges from Sitka Sound, Alaska to Turtle Bay, Baja California, but the two sub- species may have disjunct distributions. The northern abalone is distributed from Alaska south to Point Conception in central California and the threaded abalone is distributed from central California to Turtle Bay in Baja California (Geiger 2000). An Table 1. Pairwise nucleotide p-distances between VERL sequences from the unknown abalone sample and known red and northern/threaded abalone. Values indicative of intraspecific differentiation are in bold. Species Red Unknown Northern 1 Northern 2 Threaded 1 Threaded 2 Red _ Unknown 0.004 - Northern 1 0.041 0.053 - Northern 2 0.036 0.036 0.004 - Thread 1 0.034 0.034 0.005 0.002 - Thread 2 0.051 0.053 0.004 0.005 0.007 - RED ABALONE IN BRITISH COLUMBIA 187 Pink 0.005 Figure 2. Neighbor-joining dendrogram of VERL sequences from abalone species of the North Pacific Ocean, including one sequence from an individual of unknown species identity. The bootstrap per- centage support levels for the tree nodes, based on 1000 iterations, are shown. overlap of distribution between the red and northern abalone was previously considered to occur only in the southern areas of the northern abalone range. How the red abalone occurred in B.C. is unknown. Fac- tors that may affect the geographic range of red abalone include climate change and or human intervention associated with aquaculture activities. Hobday and Tegner (2002) predicted from simulation modeling of red abalone pop- ulations that as water temperatures increase the real red abalone range may extend northward through larval trans- port. Rogers-Bennett (2007) also suggested northward (and southward) range shifts might be expected in the warmer water abalone species if ocean warming occurs. Importing an alien species into an area for aquaculture purposes could result in the accidental or intentional intro- duction of the species into the wild. The red abalone has been a species of interest for aquaculture in Chile (Godoy and Jerez 1998, Pereira et al. 2007) and South Africa (Griffiths 2000). Sloan and Breen (1988: 6) reported that “In northern Wash- ington State, Haliotis kamtschatkana and Haliotis rufescens hybrids were seen a few years after transplantation of Califor- nia H. rufescens (S. Olsen, Washington State Department of Fisheries, pers. comm.).” The introduction of red abalone into Washington State might have provided a convenient launch pad for their northern dispersal under suitable conditions. Invasions of different biological species into coastal marine habitats have become common in recent years (Ruiz et al. 2000, Robinson et al. 2005). The implications of an aba- lone species invasion into B.C. are unknown, but range exten- sions, based on temporary environmental alterations or human interventions, are often not permanent. On-going quantitative abalone surveys throughout B.C. (e.g., Hanke- wich and Lessard 2008) have not found red abalone, sug- gesting that this species is probably rare in B.C. There are several anecdotal reports of red abalone shells being observed in B.C. Abalone habitat (Lessard and Campbell 2007) is not considered to be a limiting factor for northern abalone in B.C. (Abalone Recovery Team 2002). Since the northern abalone coexists with other abalone species in Cal- ifornia, we assume that there probably is sufficient abalone habitat for a number of abalone species to coexist in B.C. However, natural competition with other marine species, Table 2. Partial nuclear VERL DNA sequences for red, northern, and threaded abalone compared with the sequence obtained from the unknown abalone found in British Columbian waters. Polymorphic DNA sites are indicated by numbers using the numbering system of Gruenthal and Burton (2005). Letters indicate the nucleotide found in each polymorphic DNA site; periods indicate no change from the top sequence. VERL sequence sites Species rVCCC5MUll number 122 207 215 230 278 336 363 375 380 415 423 443 445 498 550 551 Red AF453553 A G T G C A C G C T C C A G T G Unknown EU939887 Northern AY8 17705 T A G A G G A C T A A A C A C A Northern AY8 17704 T A G A G G A C T A A A C A C A Threaded AY8 17697 T A G A G G A C T A A A C A C A 188 AMERICAN M ALACOLOGICAL BULLETIN 28-1/2-2010 predation, and reduced fertilization success due to low aba- lone abundance and density would be expected to keep red abalone scarce in B.C. ACKNOWLEDGMENTS We thank S. Hutchings, I. Winther, D. Brouwer, and L. Stenhouse for technical assistance and J. Lessard, D. L. Geiger, and an unknown reviewer for helpful suggestions to improve this manuscript. LITERATURE CITED Abalone Recovery Team. 2002. National Recovery Strategy for the Northern Abalone ( Haliotis kamtschatkana ) in Canada. Available at: http://www-comm.pac.dfompo.gc.ca/pages/ consultations/fisheriesmgmt/abalone/ documents/ 04 Abalone_ RS.pdf ; accessed fanuary 2008. Adkins, B. E. 1996. Abalone surveys in south coast areas during 1982, 1985 and 1986. Canadian Technical Report of Fisheries and Aquatic Sciences 2089: 72-96. Campbell, A. 2000. Review of northern abalone, Haliotis kamtscha- tkana, stock status in British Columbia. Canadian Special Publication of Fisheries and Aquatic Sciences 130: 41-50. Cox, K. W. 1962. California abalones, family Haliotidae. California Fish and Game Bulletin 118: 1-133. Galindo, B. E., G. W. Moy, W. J. Swanson, and V. D. Vacquier. 2002. Full-length sequence of VERL, the egg vitelline envelope receptor for abalone sperm lysin. Gene 288: 111-117. Galindo, B. E., G. W. Moy, W. J. Swanson, and V. D. Vacquier. 2003. Positive selection in the egg receptor for abalone sperm lysin. Proceedings of the National Academy of Science U.S.A. 100: 4639-4643. Geiger, D. L. 2000. Distribution and biogeography of the Recent Haliotidae (Gastropoda: Vetigastropoda) worldwide. Bolletino Malocologico 35: 57-120. Geiger, D. and G. Poppe. 2000. A Conchological Iconography. The Family Haliotidae. Conch Books, New York. Godoy, C. and G. Jerez. 1998. The introduction of abalone in Chile: Ten years later. Journal of Shellfish Research 17: 603-605. Griffiths, C. L. 2000. Overview on current problems and future risks. In: G. Preston, G. Brown, and E. van Wyk, eds., Best Management Practices for Preventing and Controlling Invasive Alien Species. The Working for Water Programme, Cape Town, South Africa. Pp. 235-241. Gruenthal, K. M. and R. S. Burton. 2005. Genetic diversity and species identification in the endangered white abalone ( Haliotis sorenseni). Conservation Genetics 6: 929-939. Haaker, P. L., K. C. Henderson, and D. O. Parker. 1986. California Abalone. State of California Marine Resources Leaflet No. 11. Hankewich, S., J. Lessard, and E. Grebeldinger. 2008. Resurvey of northern abalone, Haliotis kamtschatkana, populations in southeast Queen Charlotte Islands, British Columbia, May 2007. Canadian Manuscript Report of Fisheries and Aquatic Sciences 2839: vii + 39 pp. Hankewich, S. and J. Lessard. 2008. Resurvey of northern abalone, Haliotis kamtschatkana, populations along the central coast of British Columbia, May 2006. Canadian Manuscript Report of Fisheries and Aquatic Sciences 2838: vi + 41 pp. Hobday, A. J. and M. J. Tegner. 2002. The warm and cold: Influence of temperature and fishing on local population dynamics of red abalone. California Cooperative Oceanic Fisheries Investigations Report 43: 74-96. Lessard, J. and A. Campbell. 2007. Describing northern abalone, Haliotis kamtschatkana, habitat: Focusing rebuilding efforts in British Columbia, Canada. Journal of Shellfish Research 26: 677-686. Mottet, M. G. 1978. A review of the fishery biology of abalone. State of Washington, Department of Fisheries Technical Report 37: 1-81. Owen, B., J. H. McLean, and R. J. Meyer. 1971. Hybridization in the Eastern Pacific abalones (Haliotis). Bulletin of the Los Angeles City Museum of Natural History Science 9: 1-37. Pereira, L„ J. Lagos, and F. Raya. 2007. Evaluation of three methods for transporting larvae of the red abalone Haliotis rufescens Swainson for use in remote settlement. Journal of Shellfish Research 26: 777-781. Robinson, T. B., C. L. Griffiths, C. D. McQuaid, and M. Rius. 2005. Marine alien species of South Africa - status and impacts. African Journal of Marine Science 27: 297-306. Rogers-Bennett, L. 2007. Is climate change contributing to range reductions and localized extinctions in northern ( Haliotis kamtschatkana) and flat (H. walallensis) abalones? Bulletin of Marine Science 81: 283-296. Ruiz, G. M., P. W. Fofonoff, J. T. Carlton, M. J. Wonham, and A. H. Hines. 2000. Invasion of coastal marine communities in North America: Apparent patterns, processes, and biases. Annual Review of Ecology and Systematics 31: 481-531. Sloan, N. A. and P. A. Breen. 1988. Northern abalone, Haliotis kamtschatkana, in British Columbia: Fisheries and synopsis of life history information. Canadian Special Publication of Fisheries and Aquatic Sciences 103: 46 pp. Supernault, K. J., A. Demsky, A. Campbell, T. J. Ming, K. M. Miller, and R. E. Withler. 2009. Forensic genetic identification of abalone (Haliotis spp.) of the northeastern Pacific Ocean. Conservation Genetics. DOI 10.1007/sl0592-009-9925-x. Swanson, W. J. and V. D. Vacquier. 1998. Concerted evolution in an egg receptor for a rapidly evolving abalone sperm protein. Science 281: 710-712. Swanson, W. J., C. F. Aquadro, and V. D. Vacquier. 2001. Polymorphism in abalone fertilization proteins is consistent with the neutral evolution of the egg’s receptor for lysin (VERL) and positive Darwinian selection of sperm lysin. Molecular Biology and Evolution 18: 376-383. Talmadge, R. R. 1977. Notes on a California hybrid Haliotis (Gastropoda: Haliotidae). The Veliger 20: 37-38. Tamura K., J. Dudley, M. Nei, and S. Kumar. 2007. MEGA4: Molecular Evolutionary Genetics Analysis (MEGA) software version 4.0. Molecular Biology and Evolution 24: 1596-1599. Submitted: 17 May 2009; accepted: 6 October 2009; final revisions received: 23 October 2009 INDEX TO VOLUME 28 Anetai,Y. 28: 167 Baker, S. M. 28: 183 Bogan, A. E. 28: 151 Bringolf, R. B. 28: 151 Campbell, A. 28: 185 Coles, B. F. 28: 29 Coppolino, M. L. 28: 97 Eads, C. B. 28: 151 Evans, R. R. 28: 135 Fahy, N. E. 28: 59 Garner, J. T. 28: 121 Gerber, J. 28: 15 Greiner, R. D. 28: 151 Gutierrez Gregoric, D. E. Haggerty, T. M. 28: 121 Hirano, Y. J. 28: 167 AUTHOR INDEX Hirano, Y. M. 28: 167 Riley, L. W. 28: 183 Kohn, A. J. 28:215 Rodrigues, L. 28: 127 Kumagai, K. 28: 167 Rumi, A. 28: 159 Levine, J. F. 28: 151 Rundell, R. J. 28: 81 Maeda, T. 28: 167 Shimadu, Y. 28: 167 Mikkelsen, P. M. 28: 191 Silveira, M. J. 28: 127 Minton, R. L. 28: 91 Sinclair, C. S. 28: 105 Mormul, R. P. 28: 127 Myers Flaute, C. J. 28: 1 Naranjo-Garcia, E. 28: 59 Sudo, K. 28: 167 Supernault, K. J. 28: 185 Nekola, J. C. 28: 29 Thomaz, S. M. 28: 127 Nunez, V. 28: 159 Trowbridge, C. D. 28: 167 Orstan, A. 28: 113 Watanabe, T. 28: 167 Perez, K. E. 28: 13, 91 Watters, G. T. 28: 1 Phlips, E. J. 28: 183 Withler, R. E. 28: 185 Ray, S. J. 28: 135 Yorifuji, M. 28: 167 189 Amer. Malac. Bull. 28: 191-213 (2010) Seventy-five years of molluscs: A history of the American Malacological Society on the occasion of its 75th annual meeting Paula M. Mikkelsen Paleontological Research Institution, 1259 Trumansburg Road, Ithaca, New York 14850, U.S.A. Corresponding author: pmm37@cornell.edu Abstract: The American Malacological Union (now Society), founded in 193 1 as a national organization of collectors, students, professionals, and others interested in the holistic study of molluscs, is now an international society mainly of professionals. Although diminished in size, it continues to attract and fund students, publish a respected peer-reviewed journal, and host annual meetings featuring world-class symposia. In recognition of the society’s 75th annual meeting in 2009, 1 provide a detailed account of the founding, meetings, membership, publications, governance, and societal identity of AMS, gleaned from meeting programs, newsletters, scrapbooks, correspondence, and the memories of Past Presidents and other members. Anniversaries are times of celebration, remembering, reflection, and summarizing. The 75th annual meeting1 of the American Malacological Union (AMU; now Society, hereaf- ter AMS) held in Ithaca, New York, in the summer of 2009, was no exception. Today’s AMS is fraught with problems: diminishing membership, increasing costs, decreasing return on investments, and fewer and fewer members willing to hold office. Nevertheless, it thrives, by virtue of its respected peer- reviewed journal, continuing student interest, worthwhile projects to benefit students, conservation, and members, and a quorum of loyal members willing to dedicate uncompen- sated hours to assure its future. Now, as ever, knowing where we came from will help us guide the future. FOUNDING The American Malacological Union was founded in 1931 mainly as the brain-child of and through the organizing efforts of Norman W. Lermond (1861-1944), a New Eng- lander described by one biographer2 as a self-promoter, uto- pian, idealist, socialist, natural leader and organizer, editor and writer, naturalist, and collector, who founded and oper- ated the Knox Academy of Arts and Sciences Museum and Arboretum (and bird sanctuary) in Thomaston, Maine. He -was also a frustrated politician, unsuccessfully running for 1 2009 was actually the 79th anniversary of the American Malacologi- cal Society; no annual meetings were held in 1942-1945. 2 According to Scott M. Martin, in a presentation at the 2009 AMS meeting in Ithaca, New York. Lermond’s shell collection was dis- persed to Colby College in Waterville, Maine, then from there to the Museum of Comparative Zoology (Harvard University) and the Delaware Museum of Natural History. Congress as the Populist Party candidate in 1898, and for governor of Maine in 1900 as the Socialist Party candidate (Martin 1995, Murray 1999, see additional references on Lermond cited by Coan et al. 2009). Lermond’s museum included Indian artifacts, rocks and minerals, herbarium specimens, stuffed mammals and birds, bird eggs and nests, pinned insects, and the largest shell collection in the state of Maine (of ca. 100,000 shells)2. By the 1930s, the desire for a national organization of malacologists had long been discussed. In 1890, the American Association of Conchologists (AAC) was founded by John C. Campbell of Philadelphia. Its mem- bership roster listed 29 members, including such notables as Henry A. Pilsbry, William H. Dali, Frank C. Baker, and Josiah Keep, but the organization disbanded a few years later, likely due to the declining health and death of Campbell in 1897. Nearly 40 years later, on January 8, 1931, Lermond dis- cussed the merits of such a national organization with two acquaintances, Dan L. Emery and Charles C. Allen, while wintering in St. Petersburg, Florida.3 Soon thereafter, he (or Emery) sent letters to an estimated 200 students and col- leagues (including amateurs and professionals, neontologists and paleontologists) who were in any way interested in mol- luscs or their shells. That letter read: 3 This account is from Lermond’s handwritten report of the first meet- ing, in his capacity as Provisional Secretary, preserved in the 1931- 1951 scrapbook in the AMS Archives. William J. Clench of Harvard University remembered it differently, recalling that the idea for AMU was conceived over a bowl of chop suey shared between him and his good friend Lermond in Yung Lee’s Restaurant off of Harvard Square (Annual Reports for 1952and 1981;Teskey 1981); this was followed by the meeting between Allen, Emery, and Leonard in St. Petersburg. 191 192 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 St. Petersburg, Fla., January 21, 1931 To North American Conchologists Greeting: A small group of Conchologists (Active Shell Collec- tors), here in the East, have been talking amongst them- selves, for several years past, about the desirability of and need for an Association of American Conchologists, organized on a similar plan to that of the A. O. U. [Amer- ican Ornithologists Union], We have decided that the time has arrived for action, for organization:- First, by enrolling a goodly number of “Foundation” or “Charter” members; Second, by calling a meeting, in the near future, to perfect organization. In response to the urgent request of the others, I have consented to serve as Provisional Secretary until such time as an election of officers can be held. When sending in for enrollment, we will ask each to contribute 25 cents towards the initial expense for postage, statio- nery, typewriting and printing. Having waited a sufficient time for all the two to three hundred shell collectors and conchologists of the U. S., Canada, Cuba and Canal Zone, who are interested, to reply, a printed list of members, with their Post Office Addresses, what they collect and what they wish to exchange, will be mailed to each. With this list will also go a call for election of officers. The whole territory will be divided into some eight sections, and a vice president elected for each. Annual meetings should be held in rotation in each section, the first, quite likely, in Philadelphia (the Pioneer Center of American Conchology), say May 1st to 3rd, inclu- sive; at which time and place the organization will be com- pleted by the adoption of a Constitution and By-laws. The Nautilus, of course will be the Association’s “Official Organ”, for reports, etc. Two Veteran Conchologists of the U. S. - Chas. T. Simpson and L. S. Frierson - have already sent in their names for enrollment. Mr. Simpson writes: “I heartily welcome anything that will help to simplify the awful mud- dle of our biological nomenclature. I am as glad as you are that the Bolton rubbish is to be discarded.”4 4 “Bolton” here is a misspelling for Bolten, more properly cited as Roding (1798). The work was in fact not discarded as Lermond apparently hoped. William Healy Dali (USNM) had earlier recog- nized it as the original source of numerous taxonomic names and published an index in 1915, and the work was ultimately placed on the Official List of Titles of Works in Zoology by the International Commission on Zoological Nomenclature. A photographic fac- simile of Museum Boltenianum was reprinted by AMU in 1986. Trusting to hear from one and all, I remain, Sincerely and Conchologically yours, Norman W. Lermond, Acting Secretary of A. A. C. It is interesting to note the emphasis in this letter on col- lecting and exchanging specimens (now largely the aegis of our sister organization, the Conchologists of America), and on an apparent crisis in nomenclatural problems that needed addressing. Two additional letters calling for Charter Mem- bers followed a month later (note that letter-mail postage was only two cents in 1931!), each quoting testimonials from respondents who had already joined. There was mention of one disapproving response, but besides that, the reaction was nothing less than glowing. Bohumil Shimek, a botany professor at the State University of Iowa (who also amassed a shell collec- tion of 2.4 million specimens, ultimately transferred to the Smithsonian), was quoted as writing “Hooray! Your circular let- ter comes like manna to the hungry, or water to the thirst-cursed wanderer on the desert, to one who has never lost his interest in the field.” Others were less eloquent, but equally supportive, and many sent in more than the required twenty-five cent fee. The response was very good - 169 persons were enrolled as Charter Members. In a letter on March 21, 1931, Lermond called for nominations for officers and for a vote on five sug- gested names: Association of American Conchologists (his original suggestion), American Association of Conchologists (name of the earlier, failed society, to which Lermond did not belong), American Conchological Society, Malacological Society of America, and Conchological Society of America. The last name garnered the most votes and appeared on the March 1931 listing of Charter Members (Abbott 1956). The first meeting was set for April 30 to May 2, 1931, at the Academy of Natural Sciences in Philadelphia, to be hosted by ANSP’s Henry Pilsbry. Paul Bartsch, curator at the United States National Museum (now National Museum of Natural History), shared the enthusiasm for the new society, but in a letter to Pilsbry dated just before the meeting (23 April 1931) explained part of his fervor: “My dear Pilsbry ... as to the meeting: I thoroughly agree with you in everything that you say. I think it is a good thing to have an organization of this kind. It sort of serves to re-awaken an interest in our pets. In these days of the microtomist we must do all we can to keep the pendu- lum from casting us off altogether in its extreme swing. While I understand the importance of knowing all about the cross-section of an elephant’s hair, I can’t help but believe that we should not altogether lose an interest in the elephant as a whole. And the work that you and a handful of us have been doing now for a lifetime has been to continue the interest in the entire animal. And this group which is to meet at your place will do a lot to A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 193 retain that interest and to again eventually give system- atic zoology that basic status which it deserves. However, a group of well meaning enthusiasts, inexperienced in the real basic side of the work, can do a lot of harm, and this is where I agree with you fully that we must quietly and diplomatically serve as a steering combination that will keep the ship on an even keel while its passengers are enjoying the picnic.” We detect in these words, even in those early days, hints at tensions that we still occasionally face today in meeting venues, e.g., between whole-organism biologists and micro- analysts (including today’s molecular biologists), and between those who work and those who merely attend (the “well meaning enthusiast ... [at] the picnic”). By meeting time, membership had swelled to 191. Twenty- nine members from 12 states attended that first meeting, at which a constitution was drafted and adopted. That very brief, eight-paragraph Constitution set the name, officers, and goals of the society, limited membership to the Americas and Hawaii, set annual dues at $1.00, and named the existing publication The Nautilus as the society’s official serial.5 Inter- estingly, and without explanation that I can find, the name American Malacological Union was adopted at that meeting although it had not even appeared among the choices that Lermond had circulated earlier. It is also not clear why “Con- chological Society of America,” which had earned the most votes the previous spring, did not persist. Although the word “union” in the society’s original name might sound odd to modern readers, in 1931, the choice was appropriate because the organization was conceived as a union of professionals, amateurs, and shell clubs; one of Lermond’s socialist organizations, the Brotherhood of the Cooperative Commonwealth (with which he co-founded the utopian community of Equality in W ashington State in 1 898 ) , used the term “union” in this way. It was also the epithet of other prominent societies of the time, e.g., the American Ornithologists Union, with which Lermond had drawn com- parison in his initial letter. Yet Lermond originally proposed “Association,” not “Union,” so the origin of American Mala- cological Union remains unresolved. Murray (1999) wrote that William Clench attributed the name to Henry Pilsbry, saying that “whatever Pilsbry wanted, Pilsbry got,” in respect for the latter’s eminence during this era of American malacol- ogy. Regardless of its origin, the word “union” later gradually evolved to refer especially to trade and labor groups, present- The account of the first meeting, a list of its Charter Members, and the text of the first Constitution were published in The Nautilus, 45: 1-5, July 1931. ing problems in recent years (unforeseen by the founders) for AMS Treasurers in establishing banking arrangements and for AMS Presidents seeking funding for annual meetings and various other functions. According to Murray (1999), there were at least five attempts to change the name from “Union” to “Society,” the first appearing in the meeting minutes of 1952. Each time it was defeated largely in deference to tradi- tion. The name change was strongly supported by respon- dents to a society-wide questionnaire in 1997, went through the lengthy procedure for a constitutional change, and was finally made effective in 1998, bringing the American Mala- cological Society more in line with comparable groups and better expressing its focus, goals, and activities. The stated purpose of the society according to the 1931 Constitution was “the promotion of the science of malacol- ogy by holding meetings for reading and discussion of papers, and for furthering the interests of students and collectors of shells by facilitating acquaintance and co-operation among the members.” Following the call for nominations, Henry Pilsbry was elected as President (by one account, after Lermond declined). Lermond was chosen as Secretary-Treasurer, but was inactive, and his title was reduced to Corresponding Secretary somewhat later. Mrs. Harold R. (Imogene C.) Robertson6 of the Buffalo Museum of Science was made Financial Secretary. Mrs. Robertson, either by design or default, wrote a detailed account of the meeting. She effec- tively served in this capacity, that is, informally as Secretary- Treasurer, with Lermond still officially as Corresponding Secretary, until after Lermond’s death in 1944 and the War Years of 1944-1946. Although the call for nominations also selected eight Vice Presidents representing regional sections of the country7, this system was not activated (because the sections apparently never organized) and Paul Bartsch was elected the sole Vice President at the second meeting. Four Council Members (which we now call Councilors-at-Large) 6 In accordance with convention at the time, Imogene Robertson was often listed as Mrs. Harold R. Robertson, so I here combine the two in the interest of clarity and completeness. 7 Section 1 (Eastern and Atlantic Coast States), Paul Bartsch, United States National Museum; Section 2 (Western and Pacific Coast States), Ida S. Oldroyd, Stanford University; Section 3 (Central States), L. C. Glenn, Vanderbilt University, Nashville; Section 4 (Gulf States), T. H. Aldrich, Alabama Museum of Natural History, Birmingham; Section 5 (Canada), F. R. Latchford, Toronto, and Aurele La Rocque, National Museum of Canada, Ottawa; Section 6 (Cuba and the West Indies), Carlos de la Torre, Universidad de la Habana; Section 7 (Canal Zone, Central America, and Mexico), James Zetek, Institute for Research in Tropical America, Balboa, Canal Zone; Section 8 (Hawaiian Islands and the Philippines), D. Thaanum, Honolulu. Members were listed within these sections in Lermond’s first membership list in March 1931. 194 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 1. Officers and council members of the newly formed American Malacological Union at the second annual meeting in Washington, D. C., in May 1932. Left to right: Councilor William J. Clench, Councilor Junius B. Henderson, Councilor Ida S. Oldroyd, Vice President Paul Bartsch, President Henry A. Pilsbry, Secretary-Treasurer Norman Lermond, Financial Secretary Imogene C. Robertson, and Councilor Calvin Goodrich. were also chosen in 1932: William J. Clench, Harvard University; Junius B. Henderson, University of Colorado; Ida S. Oldroyd, Stanford University; and Calvin Goodrich, University of Michigan (Fig. 1). The very first paper presented at the first AMU meeting was on South American unionoids: “ Ruganodontites , a new subgenus of Anodontites,” by William B. Marshall, Assistant Curator in the Division of Mollusks at the USNM. Ten addi- tional papers followed at that first meeting. In her annual report, Imogene Robertson wrote an extensive summary of each presentation (there being no formal abstracts) and of the discussion following each paper; this became the standard format for the next 25 years. ANNUAL MEETINGS The AMS has traditionally met in the summer every year since its founding (see Appendix 1). The series was broken only once, by the four years of World War II (1941-1945). During this hiatus, by carrying on voluminous correspon- dence, Secretary Imogene Robertson was able to compile and issue Annual Reports for 1943 and 1944-1945, including member and affiliated club news (which carried on for many years in the various iterations of the annual reports and newsletter). In the last Annual Report during these “off’ years, the death at age 83 of founder Norman Lermond was reported. The 1946 meeting (the twelfth) in Washington, D.C. was recorded as an especially joyous occasion, as old friends and new ones gathered for the first time since the outbreak of World War to exchange news and exp- eriences. Subsequent meetings have taken place throughout the United States, as well as once in Havana, Cuba (1938), three times in Canada (Toronto 1939, Montreal 1960, Ottawa 1967), and once officially in the Bahamas (although actually aboard the cruise ship Nordic Empress, out of Miami, in 1993). The Cuban meeting was unique in at least one aspect. President Carlos de la Torre (also President of the Uni- versidad de la Habana, Director of its Museum, Curator of Mollusks, and Cuba’s Minister of Education) - by far the highest ranking malacological sys- tematist - arranged to have Fulgencio Batista’s government send the Cuba (a Cuban Navy vessel) to Key West to transport the delegates to and from Havana, most of the 49 delegates hav- ing taken the train from Miami to Key West (A. Kabat, pers. comm., 8 January 2010). Although AMS members are now accustomed to the meeting venue being held at the institu- tion (or at least in the city) of the President, this has actually been a rather recent custom and the minority case through- out our history; only 33% (25 of the 75 annual meetings) have been “local.” Council and members have occasionally discussed alter- nate meeting schedules. A questionnaire in 1997 posed this question, and there was considerable support for meeting less often, particularly in not meeting every third year when Unitas Malacologica holds its international meeting. This has never been put into effect although our society has met jointly at three of the last four World Congresses of Malacol- ogy. The first WCM (Washington, D. C., 1998) was so named principally because UM and AMU (along with Western Society of Malacologists) agreed to meet jointly. The 1997 questionnaire also confirmed that most members supported occasionally meeting jointly with another organization. AMS has met jointly most often with the Western Society of Malacologists. The first joint meeting of AMU and WSM was held at San Diego State University in 1975. Today, WSM and AMS are on friendly terms, and the two have met jointly ten times at western sites (1975, 1979, 1983, 1986, 1989, 1991, 1997, 2000, 2005, 2006), with plans again in 2010 in San Diego. Discussions about meeting jointly with other organi- zations ( e.g ., Society for Integrative and Comparative Biology, National Shellfisheries Association, Conchologists of America) have not yet met with success. A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 195 Figure 2. Membership and attendance at annual meetings of the American Malaco- logical Union/Society, 1931-2009. Numbers were taken from annual reports or were physically counted in membership directories, however, it is likely that membership numbers have been calculated in different ways over the years (e.g., whether or not sub- scribers were included; they were excluded here whenever specified). Attendance not available for 1942-1945 (no meetings during World War II), 1998, and 2000. Organization of the meeting has always been the job of the President8 although some- times with the help of a local meeting orga- nizer, particularly when the meeting took place away from the President’s home institu- tion. Dolores S. “Dee” Dundee served in this capacity twice in New Orleans, once for John Q. Burch of Los Angeles in 1964, and a second time for Louise Russert-Kraemer of University of Arkansas in 1982. Shell clubs also assisted on local organizing committees, serving as official hosts of the meeting beginning in 1955 (New York Shell Club at Staten Island, New York) through 1986 (Monterey Peninsula Shell Club at Monterey, California). Even after the tradition had passed, local shell clubs continued to sponsor evening or social events at the annual meeting into the 1990s and as late as 2006. Presidents have also invariably needed (and found) many additional hands to assist. Past President William G. Lyons recalled, “There were times at the 1987 Key West meeting when I felt like I was doing more than my share of the work, but at the end I had more than 40 members to thank for their substantial help as regis- trars, drivers, tour guides, symposia organizers, audio-visual techs, T-shirt artists, etc. Without all of us working together, it would have been a disaster. It seems that it has always been thus, which in great part is why we still have an organization today” (W. G. Lyons, pers. comm., 12 January 2010). Attendance at the annual meetings has varied greatly over the years (Fig. 2). Past Secretary Margaret Teskey (1981) noted that attendance at annual meetings averaged 25% of total membership, but records gathered during the writing of this paper indicate a mean of only 13%. Mean meeting atten- dance, taken over the 69 meetings for which attendance fig- ures are available, is 127 persons. A low of only 17 attendees was tallied in 1953 at the meeting at University of Kansas, Lawrence, despite 476 members in the organization at that time (or only 3.6% attending). No explanation was put on record for the low attendance, and 13 papers were neverthe- less presented (Fig. 3). The maximum attendance recorded so far at a meeting was in 1970 when 268 of 764 members (35.1%) attended the meeting in Key West, Florida. Perhaps there is a lesson here for future presidents: location, location, 8 In 1996, Council discussed separating the duties of running the society and organizing the meeting into two offices - Immediate Past Presi- dent for the former and President for the latter. This idea received strong support (the President is usually too busy organizing the meet- ing to attend to anything else), but was never put into effect. location! Woods Hole in 1990 was a close second with 261 attending, thanks to a highly successful cephalopod sympo- sium. Three years later, only 83 people attended, but that was the “Bahamas” meeting aboard the Nordic Empress, and many otherwise loyal AMUers were unable to justify a business meeting aboard a cruise ship. Attendance immediately rebounded to 170 at the 1994 meeting in Houston. Figure 2 also shows that in years when there is a World Congress of Malacology (Washington, D.C., 1998; Vienna 2001, Perth 2004; and Antwerp 2007), meeting attendance is depressed irrespective of whether AMS meets with the WCM (2001, 2007; meeting attendance unavailable for 1998) or separately (2004) (D. O Foighil, pers. comm., 11 January 2010); AMS will meet separately again in San Diego in 2010 in lieu of the WCM in Phuket, Thailand. Annual meetings were initially informal, three or four days long, with no time limit placed on presentations; early “calls for papers” asked only how long a presenter needed for his or her talk. In 1959, the first concurrent sessions were organized, separating scientific papers from talks on more popular subjects, such as collecting trips. Since 1979, concur- rent sessions, organized by topic, have been the norm. The first posters were presented in New Orleans in 1982, orga- nized by Clement Counts III; the Woods Hole meeting in 1990 still holds the record for the most posters (32) at a sin- gle meeting (excluding World Congresses of Malacology). Printed programs have existed from the beginnings of AMU, at first listing only titles and presenters in addition to other meeting events. Abstracts of presentations were 196 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 3. The 1953 annual meeting of the American Malacological Union at the University of Kansas, Lawrence (top), organized by President A. Byron Leonard of that institution, holds the record for the lowest attendance at any meeting, with only 17 registrants (left to right: first row: Juan Jose Parodiz, Albert R. Mead, R. Tucker Abbott, Jeanne S. Schwengel, A. Byron Leonard, Mrs. Leonard, Fritz Haas, Mrs. Berry; second row: Dorothea Franzen, Margaret C. Teskey, Joseph C. Bequaert, A. Myra Keen, Joseph P. E. Morrison, Elmer G. Berry). The Key West meeting in 1970 (bottom), organized by President Alan Solem of the Field Museum of Natural History (Chicago), attracted the most attendees (excluding World Congresses of Malacology), with 268 registrants. not included in the meeting program until the Key West meeting in 1970, under President Alan Solem. In recent years, the Program and Abstracts volume of the meeting is a publica- tion available both in print at the meeting, sometimes mailed to non-attending members, and is now often provided online. Social networking is the new term for it, but AMU/AMS meetings, like any organization, have provided at least as much social as scientific opportunity. Coffee breaks and field trips (either at the end of the meeting, or in the middle as a “break”) became the best places to see and be seen, to introduce oneself to the community, to start new collabora- tions, and to make new friends (Fig. 4). Early meeting reports noted Mrs. Frank R. [Jeanne] Schwengel’s annual cocktail party as a regular event (the eleventh was in 1953), looked forward to by many attendees. Apparently, Jeanne’s hus- band, a retired Brigadier General, was a senior executive at Seagram and Sons, once the largest distiller of alcoholic beverages in the world. Every year, “The General” shipped several crates of Sea- gram’s products to the AMU meeting so that his wife could entertain like roy- alty (A. Rabat, pers. comm., 8 January 2010). Texan members banded together for a similar event beginning in 1972; by 1974, the annual “Texas Party” was already a tradition, and continued through 1980. Philadelphia members threw a similar event at the Corpus Christi, Texas, meeting in 1979. Other forms of entertainment often included multimedia presenta- tions. At the meeting in Rockland, Maine (1941), Henry Russell (Museum of Comparative Zoology) brought a home movie camera and produced the first-ever (?) film of attendees at a meet- ing, primarily during social events. At the next meeting (in 1946, following the World War II hiatus), he showed “highlights” from Rockland, and the audience roared when they saw his clip — surreptitiously filmed — of H. Burrington Baker and Miss Bernadine Barker, then unwed, wandering off into the woods, hand-in-hand. (Later in 1941, Miss Barker apparently became Mrs. Baker.) Unfortunately, Russell’s films do not survive (A. Rabat, pers. comm., 8 January 2010). In 1957, at her meeting in New Haven, President Ruth Turner arranged to have a party in an outdoor courtyard near the Yale residence hall where they were staying, and as part of the “entertainment” A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 197 Figure 4. The social nature of AMS meetings is at least as important as the scientific offerings. Field trips and the annual auction provide informal occasions to spend time with colleagues, new and old. Top left: North McLean (left) shares an interesting find with a colleague, while Paul Bartsch (right) autographs a piece of driftwood riddled with shipworms, on the beach at Plum Point, Maryland, 1932. Top right: Rebecca Rundell and then-President Warren Allmon search for Devonian fossil molluscs near Ithaca, New York, 2009. Bottom left: then-President Paula Mikkelsen and Auctioneer Paul Callomon at the AMS auction during the World Con- gress of Malacology in Antwerp, Belgium, 2007. Bottom right: Gerhard Haszprunar and Rudiger Bieler share a malacological moment while waiting for the auction to start, Antwerp, Belgium, 2007. (i.e., a conversation piece) had a slide projector that continu- ously ran with various molluscan scenes, broadcast on a large outdoor screen, which was well received (A. Rabat, pers. comm., 8 January 2010). More recently, William E. “Bill” Old Jr. (American Museum of Natural History) was notable in the late 1970s in offering slide shows of previous meetings at the President’s Reception held on the first night of the meeting. This past summer, the tradition revived for the 75th anniver- sary, with a presentation (assembled by the present author) of all group photographs taken at past meetings. AMU Pacific Division In 1948, prompted by the expense and difficulties that west coast members experienced in attending east coast meet- ings, a Pacific Division was organized to convene separate annual meetings in western states. Andrew Sorenson of Pacific Grove, California, is credited as the founder and was made an Honorary Life Member in 1956. The organizational meeting of “AMUP” (also called AMUPD or simply “the PD”) was held at Allan Hancock Foun- dation of the University of California, Los Angeles, 10-11 April 1948. Ruth E. Coats of Tillamook, Oregon, was elected Chairman (that term was used equally for men and women at the time), with Vice Chairman John Q. Burch (Los Angeles), Secretary-Treasurer Leo G. Hertlein (California Academy of Sci- ences), and Councilors Joshua L. Baily, Jr. (San Diego), S. Stillman Berry (Red- lands, California), Wendell O. Gregg (Los Angeles), and A. Myra Keen (Stan- ford University). Initially, all Past Chair- persons served as Councilors although only four were ever listed officially. There were approximately 40 attendees at that first meeting, and nine papers were presented. Meeting format was more-or-less identical to that of an AMU meeting, including a shell club host, similar number of days, group photograph, presentations, field trips, committees, and occasional symposia. One major difference was shell exhibits, which were regular features of AMUP meetings; a shell auction with Rudolf Stohler as auctioneer was held in 1960 at AMUP specifically to raise funds for cabinets to be used for shell exhibits at annual meetings. Beginning in 1949, the Chairman of AMUP served as Second Vice President on AMU Council. Meeting attendance was good, usually mirroring or exceeding that at the national meeting, and peaking at 1 10 in 1961 at University of California at Santa Barbara in Goleta, California. The first (and only) joint meeting of AMU and AMUP was held at the Hotel Lafayette in San Diego, California, in 1956. Bylaws were adopted in 1958. AMUP was close to having a “headquarters,” holding seven of its 22 annual meetings at Asilomar Conference Grounds in Pacific Grove, California. Past attendees remem- ber this as an ideal meeting site, with the dunes, salt air, rocks, cozy lodge and lodgings, and Stanford University’s lab nearby (AMS returned there in 2005, with similar effect). Dues were activated in 1960, set at 50 cents added to AMU dues; mem- bership peaked at 190 in 1964 (the same year as the AMU membership maximum). Still the PD felt in many respects like a poor step-child to the national organization. Friction began in the early 1960s when AMUP wanted to bestow various 198 AMERICAN MALACOLOGICAL BULLETIN 28-1/2*2010 members with life membership, but this recognition was denied at the AMU level. Awards of Honor were given in rec- ognition of service to malacology for several years running, but these never carried any official status in the national orga- nization. Past President (and Past Chairman of AMUP) Alan Kohn recalled hearing an unverified rumor that “there was a dispute involving the sum of 50 cents. . . . Mr. A.J. Ostheimer, who then lived in Hawaii, was charged 50 cents AMUPD addi- tional dues, and he refused to pay it. This set off the dispute that resulted in the demise of the PD” (A. J. Kohn, pers. comm., 15 January 2010). Regardless of the cause(s), the end was in sight. In 1968, PD members held only a business meet- ing at Asilomar during the inaugural meeting of the Western Society of Malacologists (WSM). The 1969 meeting was tech- nically a joint meeting with WSM, again at Asilomar. In 1970, a Committee on AMU East-West Organization, with Albert Mead (then University of California) as Chair, presented a report to acknowledge the dissolution of AMUP and to pro- pose reorganizing the national society to hold a Congress every three years supplemented by regional annual meetings (East- ern, Western, Hawaiian, Foreign). The fate of this proposal is unknown, except that it was not accepted. AMUP held no fur- ther meetings and was formally dissolved in 1972, effectively replaced by the independent organization WSM. See Appen- dix 2 for a full list of AMUP meetings and chairmen. World Congresses of Malacology In 1998, the AMU met jointly in Washington, D. C., with the international society Unitas Malacologica and the West- ern Society of Malacologists, forming the first World Con- gress of Malacology; this was also the first time that Unitas had met outside of Europe, so it was truly a unique event. AMU President Robert Hershler organized the congress, together with UM President Rudiger Bieler, and attracted ca. 400 attendees (the number of AMU members was not recorded). AMS has convened its annual meetings at subsequent World Congresses in Vienna, Austria (2001), and Antwerp, Belgium (2007). Although such meetings were very well attended, the actual numbers of AMS members in attendance was generally lower than at national meetings. Nevertheless, participation in the triennial WCM increases the stature of the society in the international community, and most recent Presidents have supported such joint ventures. Unlike the first WCM in Washington in 1998, in which AMU was a full joint organizer, sharing in the expenses and profits, subsequent WCM participa- tion has been limited to attendance at a Unitas-hosted congress, with specific AMS-sponsored events ( e.g ., auction, symposia). Symposia The first symposium on record at a meeting was in Rockland, Maine (under President Harald Rehder), in 1941. The seven papers comprising “Methods of Collecting and Preserving Mollusca,” organized by Blenn R. Bales, an Ohio physician and amateur, was published in the AMU Bulletin for 1941. This was a handy reference source and was so much in demand that in 1955 the symposium papers were incorporat- ed into the first printing of the popular AMU booklet, How to Collect Shells (Abbott et al. 1955; see Special Publications below). Following this successful beginning, three symposia were held in 1957 (The Distribution of New World Mollusca, organized by Thomas E. Pulley; Some Aspects of Medical Malacology, organized by Edward H. Michelson; Research work in the U.S. Fisheries Laboratory, organized by Victor L. Loosanoff), but symposia did not become annual events until 1968. In 1978, Council specifically mandated symposia “of national prominence” at the next two meetings, to increase membership and visibility of the society. Since then, sympo- sia, special topic sessions, and workshops have become an integral part of each meeting, to highlight key issues in mala- cology and attract participants who might not otherwise attend. The first plea for donations specifically for symposium support was issued by President Clyde Roper in 1980. The 1981 symposia in Ft. Lauderdale, Florida, were partly financed by the shell and book auction and by 50th Anniversary com- memorative cards and cancelled envelopes sold by member Richard E. Petit (Fig. 5). In 1982, a Symposium Endowment Fund was established by then-President Louise Russert Kraemer to provide annual funding to presidents and sympo- sium organizers to support travel for symposium speakers and other associated costs of these special sessions. Auction pro- ceeds and individual donations added to the fund, and a goal was set in 1985 of $30,000 (achieved in 1988). AMS-funded symposia are now mandated to be of “world-class” caliber (by a Council motion in 1996, also requiring only one symposium Figure 5. AMU’s 50th anniversary souvenir envelope in 1981 fea- tured a commemorative postmark from the meeting venue (Ft. Lauderdale, Florida) as well as the AMU logo Io fluvialis drawn by Anthony D’Attilio in 1960. Richard E. Petit produced the covers and sold them for a few dollars each to raise funds for symposia support. The Symposium Endowment Fund was formally established the fol- lowing year. A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 199 per year to receive society funding), and are required to submit proceedings for possible publication in the American Malacological Bulletin ( AMB reserves right of first refusal for all AMS-funded symposia). This action does not preclude other non-funded symposia, workshops, or special sessions designed to bring in additional cadres of participants, with the result that modern meetings often have three or more “special events” to strengthen the attractiveness of attending. Sales at meetings Meetings have frequently included space for book dealers and various other businesses offering goods of interest to attendees. Early meetings often included shell sales and “swaps.” One of the most controversial measures ever taken by AMS was the motion to prohibit the sale of specimen shells at annual meetings, in the spirit of molluscan conservation. It began in 1976, with a motion approved by Council to forbid commer- cial sales and the exhibit of shells at meetings. The motion did not pass, but sensitivity to the subject prompted special per- mission being asked of Council to allow shell sales in the auc- tion in Ft. Lauderdale in 1981 (specifically restricted to marine, non-endangered species). There was another motion to ban shell sales proposed by Council in 1983, and several years of shifting positions followed; at the time, a vote of members present at the Annual Business Meeting was required for ratifi- cation, and the membership always reversed Council’s deci- sion. One particularly confusing year was 1984, which had an Exhibits Committee to organize vendor tables at the meeting. That committee was charged not to allow shell sales, but shells were allowed in the auction, so the exhibits ban was rescinded. Shell exhibits and/or shells in the auction were present in 1985, 1987-1990, 1992, and 1994. The sale of shells continued to be controversial until another ban was hotly debated by Council in 1994-1995, tabled, and ultimately defeated within Council. After the defeat, a motion to ban shell sales was made and passed at the Annual Business Meeting. Although this vote had no control over Council, it expressed the strong opinion of the membership, and Council passed the ban in 1996: “The Amer- ican Malacological Union does not allow selling, buying, or trading of shells or shell products at its annual meetings.” Another motion to overturn was proposed in 2005, but was defeated. This policy remains in place today. Annual meetings have usually included a fundraising auction since 1980. The first such event was at AMUP at Asilomar in 1960, to raise funds to buy shell cabinets to be used at meetings. The next on record was in 1966 at the national meeting in North Carolina, to help cover meeting expenses. A shell auction was a special event in 1975 in San Diego during the joint meeting with AMUP. Then in 1976, the first “Liter- ature Auction and Book Bazaar,” with Morris “Karl” Jacobson as auctioneer, raised $983.53. When held at joint meetings with Western Society of Malacologists, proceeds were often split fifty-fifty. In 1983, the auction was called the William E. Old Jr. Memorial Auction, in honor of one of our first “regu- lar” auctioneers, Bill Old, who died suddenly on New Year’s Eve the previous year. Proceeds of the 1981-1998 auctions went to the Symposium Endowment Fund. In 1999, the deci- sion was made to redirect auction proceeds to the Student Research Endowment Fund, and the auction continues to support student programs today. Still one of the most enjoy- able events at the annual meeting, the auction offers a wide variety of books and “shell paraphernalia” (T-shirts, toys, ceramics, quilts, etc.) auctioned by some of most vibrant per- sonalities, most notably Morris “Karl” Jacobson (1966, 1976, 1980), William E. Old Jr. (1980, ??-1982), Richard E. Petit (1983-2003, jointly with Hank Chaney or Carole Hertz dur- ing west coast meetings), Chris Garvie (2004), Paul Scott (2004), and Paul Callomon (2004-2009). Sales of T-shirts bearing the annual meeting logo have also become annual (for more about logos, see Branding and Societal Memory, below). The earliest mention of such a T-shirt that I have been able to locate was for the Los Angeles meeting in 1989, however then-President James H. McLean recalls this being a tradition long before the L.A. meeting (pers. comm., 8 January 2010). MEMBERSHIP Membership in AMU grew steadily from 191 at its found- ing to a peak of 838 in 1964 (Fig. 2), a growth period of just a little over 30 years. The second thirty years (1960-1990) saw membership wavering in the 700-800 member level. However, the last 15 years has shown a nearly consistent decline, and society records indicate concern beginning much earlier. In 1973, Council discussed a perceived drop in younger mem- bers joining and continuing their membership (although I could find no mention of any enacted remedy). In 1978, Recording Secretary Connie Boone calculated that AMU was losing members at the rate of approximately 10% per year, prompting measures aimed at increasing membership and meeting attendance, including a mandate for symposia “of national prominence.” The curve in Figure 2 indicates that such measures had a positive effect and reversed the decline. Lastly in 1993, Janet Voight, Chair of the Membership Com- mittee, authored an article in AMU News about declining membership, citing a 33% drop below 1987 level. Today, the total is approaching founding level again, hovering for the last three years at approximately 280 persons. What happened? There has been considerable discussion in AMU/AMS Council about attracting and maintaining members, and a combination of issues are clearly involved. Certainly, the rise of Conchologists of America (COA; which boasted a mem- bership of 1,500 members in 1998) for amateurs, as well as 200 AMERICAN MALACOLOGICAL BULLETIN 28-1/2-2010 special interest malacological organizations (e.g., Freshwater Mollusk Conservation Society, Cephalopod International Advisory Council), have decreased AMS membership in recent years. Other factors no doubt include decreased aca- demic funding for memberships and travel, a generally reduced interest in membership organizations (not restricted to malacology; see Putnam (2000) for an interesting analysis of this social phenomenon), and the ready availability of email, listservers, and other online sources of information. Recent attempts to increase student benefits have resulted in a slight increase in student participation at the latest meet- ings, which is heartening to all and so important to the future of malacology. Nevertheless, the future remains uncertain. Amid the curve in Figure 2, three sharp declines (in 1971, 1993, and 2001) warrant further discussion. At least two of these are clearly artifacts of intentional corrections to the mem- bership roster. It is a well acknowledged fact that during some years, officers in charge of membership were lax in removing unpaid members, thus artificially inflating the membership total. Other societies have admitted similar problems, so AMS is certainly not unique in this regard, and many membership organizations intentionally carry unpaid members on their rolls as “prospects” to be regained; so the problem is actually one of improper reporting. The first misleading decline occurred in 1971 when 192 members were dropped because of unpaid dues. Concern was expressed and a flurry of letter writing by outgo- ing Recording Secretary Marion Hubbard followed. Member- ship increased by 43 people in 1976, so that activity obviously had a positive effect. A similar correction occurred in 2001, when nearly 200 members who had not renewed their mem- bership were dropped from the roster. No similar mention was made in 1993 reports, but the same correction could conceiv- ably have occurred then as well. The extent of membership over- inflation across the curve in Figure 2 cannot be traced with accuracy, perhaps lending limited usefulness to the pre- sented data, and one wonders what paid membership actually was during the days of 600-800 members. Nevertheless, it is clear that membership now is much less than it used to be, but that the three sudden and drastic dips in the curve were proba- bly not responses to something that AMU/AMS “did,” or sud- den reversals of interest on the part of our community. Today’s officers are taking greater care of the membership database, if only in recognition of the high cost of unwarranted mailings, so we can probably trust at least the most recent totals. That said, the undoubtedly genuine general decline in membership over recent years continues to occupy AMS Council, which seeks new and better ways to attract and keep members. Special categories AMU has had special membership categories since 1932. Originally limiting membership to persons living in the Americas, Corresponding Membership was created for those in other countries; there were two Corresponding Members in 1932 (Professor Shintaro Hirase, Zoological Institute of Tokyo, and Dr. Sohtsu G. King, Pekin Laboratory of Natural History, Peiping, China). By the 1960s, AMS had achieved a recognized place among the scientific societies of the world, and included corresponding members from Japan, the Philippines, Australia, Saudi Arabia, Yap, and the Netherlands. By 1981 in its fiftieth year, Germany, France, Oman, New Zealand, South Africa, Brazil, Belgium, Ireland, Taiwan, Thai- land, Hong Kong, Austria, and the Arabian Gulf were added to the roster. Other organizations were originally listed as regular members in the annual rosters, but in 1967, 42 Affiliated Shell Clubs were listed separately for the first time. Although Affili- ated Membership is still offered (at the same rate as an indi- vidual membership), Affiliated Members last appeared as a special category in the AMU/AMS Directory in 1996. Other special price memberships have long been offered (though exact years are not available) in various categories. AMS currently recognizes Regular Members, Student Mem- bers (at reduced fees), Sustaining Members (for those who contribute above regular dues), Affiliated Members (see above), and Life Members (who pay a lump sum and thus avoid all subsequent annual fees; currently 15 to 30 times Regular membership depending upon age). Honorary Membership was also created in 1932, “for those that have contributed in an outstanding way to American Con- chology.” Charles Torrey Simpson (USNM, emeritus), Bryant Walker (Detroit, Michigan), and Victor Sterki (Carnegie Muse- um of Natural History) were the first elected. Dr. Thomas Barbour, Director of the Museum of Comparative Zoology, was elected to honorary membership in 1933. Later, Honorary Membership was awarded in recognition for service to malacol- ogy. Honorary Life Membership was first awarded in 1952 to Imogene C. Robertson, in recognition for her 20 years of service as AMU Secretary when she retired from that position. Many others have been so honored through the years; this title (now codified in the Constitution) is now available to a maximum of ten persons at any one time. Honorary Presidency was conferred upon Ida S. Oldroyd (Stanford University, when the meeting was held there) in 1934 and Henry Pilsbry in 1937, but that category seems to have been abandoned in favor of Honorary Life President, reserved for only one person at a time. Those so honored have been Paul Bartsch (1959), S. Stillman Beriy (1960-1984), Harald A. Rehder (1985-1996), Ruth D. Turner (1997-2000), and Alan J. Kohn (2008-present). A full list of honorary appointments awarded by the society appears in Appendix 3. Students AMS has always strived to support students of malacology as the future of our science, but student benefits were not always available. The first student paper award was initiated by A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 201 President William Old at his 1979 meeting in Corpus Christi, Texas. Robert S. Prezant (University of Delaware; Prezant would become President in 1999) won the $100 award for his paper on mantle glands of lyonsiid bivalves. Thereafter, student paper awards were offered annually, usually by virtue of indi- vidual donations from members. In the 1980s, donations were solicited to help subsidize student papers in the Bulletin when it first became a peer-reviewed journal. The first standing com- mittee for student programs (the Student Paper Award Com- mittee) was established in 1983 and has continued ever since. Today, AMS offers lower membership dues and lower registra- tion and event fees at annual meetings to enrolled and newly graduated students; some meetings ( e.g ., Woods Hole 1990, Sarasota 1992, Chicago 1996) have also provided reduced or free housing for students. Various presidents have been suc- cessful in raising funds to support student travel and attendance at the meetings (notably for Pittsburgh 1999 [where 25% of attendees were students], Vienna 2001, and Antwerp 2007). The two annual AMS student grants honor Past Presidents for their support of student programs: since 2000, the Constance Boone Award for the Best Student Presentation, given for the best student paper or poster presented at the annual meeting, and the Melbourne R. Carriker Student Research Awards in Malacology (created in 1997, so named since 2007). 9 A standing committee now manages student programs and evaluates pro- posals. Since 1999, proceeds of the annual auction have been added to the Student Research Endowment Fund (originally proposed in 1990, established in 1998 with a goal of $50,000). Since 1999, a student member serves as one of the four Council- ors-at-Large, and students often gather separately at annual meetings to discuss issues pertinent to their special status. Amateurs Amateur conchologists have played an active and inte- gral role in AMS throughout its existence. Founder Norman 9 In 1985, two student paper awards honored two past presidents who were recently deceased: the William J. Clench Prize (awarded to Janice Voltzow [Duke University], for “Functional Morphology of the foot of the Lightning Whelk, Busycon contrarium ”; Janice was President of AMS in 2001) and the Joseph Rosewater Prize (awarded to Silvard Kool [George Washington University] for “Systematic Revision of the Thaidid Genera Based on Anatomy”). In 1987, the student paper award was named the Maude N. Meyer Award for that year only, in recognition of a generous bequest; John B. Wise (Grice Marine Biological Laboratory) was the winner for his presentation “Contributions to the Biology of Boonea im- pressa (Say) (Gastropoda: Pyramidellidae).” The Carriker Award was preceded by the Joseph C. Bequaert Award for Field Studies of Land or Freshwater Mollusks, established in 1982, at $400 annu- ally, but seems to have only lasted through 1983. Lermond himself was an amateur, as have been many Presi- dents and other officers. Shell clubs have held Affiliated mem- berships, and for many years served as co-hosts of annual meetings. When Arthur Clarke held his meeting in Corpus Christi, Texas, in 1968, six Texas shell clubs united to play host in lieu of university auspices. “Shell Club Night” was an annual evening event beginning in 1964, and most years through 1988, with shell exchanges, slide shows of past meet- ings, and reports by shell club representatives. Shell Club Representatives became official in 1978, when they were allowed a reduced registration fee. I personally remember the pride felt by many (myself included) who attended the meet- ings in this capacity. Shell club representatives continued at most meetings through 1997 although Shell Club Nights ended a decade before. The 1980 meeting established work- shops aimed at amateurs and hobbyists; these continued through 1984. During the last several decades, however, ama- teur participation has declined, and AMS today is largely a professional organization. The reasons for this are controver- sial and laden with emotion. Certainly, the growing promi- nence of the COA, founded in 1971, which perhaps provides more of the type of programming sought by non-professionals, played a strong role in this shift. The professionalization of the annual AMS meeting program, the prominence of con- tributed papers and symposia using academic methodologies (e.g., electrophoresis, electron microscopy, biochemistry, sequencing), the upgrade to a peer-reviewed publication, and the prohibition of shell sales have also been perceived as limiting (or even excluding) by many amateurs. It is well documented that AMS Council never discussed or sought to eliminate amateurs, and in fact often discussed how to enhance programs to retain such members (Murray 1999). The loss of the participation of shell clubs, which once played such a prominent role in annual meetings, places an increased burden on presidents and local organizing committees. Today, only a small number of amateurs still actively partici- pate in AMS, but all continue to be welcomed as members. PUBLICATIONS Annual Reports From 1931 to 1933, reports of the annual meeting were provided in the molluscan specialty journal The Nautilus. The report of the second meeting was colorfully entitled “Mrs. Imogene C. Robertson’s Rambling Notes on the Second Annual Meeting of the AMU in Washington DC, May 26-28, 1932.” From 1934 to 1970, Annual Reports of AMU were issued (but still in the same size and format of The Nautilus) that included meeting abstracts, some extended papers, reports on the business meeting, a group photograph (often with each attendee identified, a tradition that continued through 202 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 198 1 10), and a membership list. The first 26 of these were printed on the Buffalo Museum of Science press, where Imo- gene Robertson (AMU Secretary) was Curator, and also large- ly the author of the Annual Reports. Authors did not submit their own abstracts of their papers until 1948; prior to that, Mrs. Robertson wrote everything. In the revised Constitution of 1953, the Annual Reports series was listed instead of The Nautilus as the official publication of the society. The first full financial report appeared in Annual Reports in 1961. The 1966 Annual Reports included the first extended abstracts, written by the authors. In 1971, the annual publication was enlarged and renamed Bulletin of the American Malacological Union, with contributed papers occupying a more substantial frac- tion of the content. A comprehensive index to the annual reports and bulletins for 1934-1974, compiled by long-time Secretary-Treasurer Margaret C. Teskey (and informally called the “Teskey Index”) was published in 1975 and separately dis- tributed (preceded by a shorter version, Anonymous 1966). American Malacological Bulletin The Bulletin metamorphosed in 1982, becoming a full- fledged, peer-reviewed scientific journal called the American Malacological Bulletin ( AMB ), with Robert S. Prezant (then at University of Southern Mississippi) as its first editor. Prezant explained in the first issue, “The persistently high quality research reported in AMU Bulletins deserves an accentuated and expanded AMU journal.” Five Associate Editors and a Board of Reviewers of 35 persons were listed in the first issue. A Managing Editor, Ronald B. Toll, was added in 1986. Annual issues were published in 1983 and 1984, after which AMB became biannual (although some recent years have produced single, technically combined, issues). Volume 1 in July 1983 was somewhat transitional between the old and new, including peer-reviewed “outside” papers plus most of the traditional Annual Report contents (minutes of the busi- ness meeting, abstracts, and membership list). Page charges supported the new format, and the dues notice included a plea for donations to assist student publication in AMB. Three Special Editions have also been published (all fully underwritten): no. I, Perspectives in Malacology: a Symposium to Honor Melbourne Carriker (at the 1985 AMU meeting in Rhode Island, organized by Robert S. Prezant and Clement L. 10 There is no record of how attendees in the group photograph were identified in the early years, but in the 1970s and 1980s, I personal- ly recall the method. Immediately after the photograph was taken, clipboards were passed down each row for each person to inscribe their name in order. The last time this occurred was at the 1981 Ft. Lauderdale meeting - over 200 attendees waited for the clipboards, not-so-patiently, on the beach in the sweltering Florida July sun. We never passed a clipboard again! Counts III, and supported by the University of Delaware, 1985); no. 2, Proceedings of the Second International Corbicula Symposium (held in 1983 in Little Rock, Arkansas, organized by Louise Russert Kraemer and others, and supported by the U. S. Nuclear Regulatory Commission and the Electric Power Research Institute, 1986); and no. 3, Entrainment of Larval Oysters (proceedings of a 1985 workshop in Lewes, Delaware, organized by Robert S. Prezant and others, and supported by the Baltimore Army Corps of Engineers and the Waterways Experiment Station, Vicksburg, Mississippi, 1986). The report of the annual business meeting, list of Executive Council members, group photograph, membership list, and abstracts of papers presented at the annual meeting were omitted from AMB in 1987, after which they became available only in the newsletter or meeting program provided to attendees. An index to the first six volumes and three special editions was published in 1988 (Counts 1988). Symposia presented at annual meetings have made up a substantial part of the papers published in the AMB since its inception. AMB reserves the right of first refusal for symposia at annual meetings that have been financially supported by AMS. After the original team of Editor Prezant and Managing Editor Toll, the Bulletin has been produced by Toll and Paula M. Mikkelsen, Toll and Timothy A. Pearce, Janice Voltzow and Angel Valdes, and Kenneth M. Brown and Cynthia D. Trowbridge. Details about the annual meetings of AMU/AMS and resulting publications were summarized by Coan and Kabat (2009). In 2009, the AMB joined the e-generation by becoming part of BioOne (www.bioone.org), the electronic aggregation of bioscience research journals. A full collation of the Annual Reports, Bul- letins, and AMB through 2007 was published by Coan and Kabat (2007). Membership newsletters The first American Malacological Union Newsletter was produced in September 1968. It consisted of just two pages and is marked “NYSC Notes No. 144” but does not appear to have ever been printed in the New York Shell Club Notes or perhaps even distributed widely (Eugene V. Coan, pers. comm., 9 January 2010). It announces plans for the newslet- ter, has a paragraph on “Why join the American Malacologi- cal Union?,” an announcement about the 1968 Corpus Christi meeting, results of the last business meeting in terms of elect- ed officers, and a membership application form. The follow- ing January, vol. 1, no. 2 appeared as part of the New York Shell Club Notes no. 148, with a history of the AMU Annual Reports by Morris. K. “Karl” Jacobson (American Museum of Natural History, hence the connection with the New York Shell Club; Jacobson was also the newsletter editor), and “Teskey Testers,” amusing anecdotes from the correspon- dence received by Secretary Margaret C. Teskey. In fall 1970, under Jacobson’s editorship, an independent AMU newsletter A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 203 began what would become a long-term, useful feature of summarizing news from shell dubs, museums, members, and research institutions. The newsletter was issued twice per year and also included book reviews, news of conservation issues, and in spring 1972, a conservation questionnaire. In follow- ing years, Jeanne Whiteside, Dorothy E. Beetle, Paula M. Mikkelsen, Raymond E. Neck, M. Bowie Kotrla, Donna D. Turgeon, S. Dawne Hard, Paul Callomon, and Christine Parent have served as Newsletter Editor. In 1984, responsibility for the newsletter was added to the duties of the Correspond- ing Secretary; it was separated out again when the two offices of secretary plus treasurer were combined in 1989. In 1980 (before email, Facebook, and personal or institutional web pages), the newsletter began focusing on shell club news in the fall issue, and on research reports by professionals and institutions in the spring issue. Member and institutional news dwindled in the 1980s, but affiliate member (shell club) news continued through 1992. In 1987, the newsletter took over disseminating annual reports of the officers, the group photograph, and the general membership list, formerly included in the AMB and its predecessors. A third annual issue of AMU News, printed on archival-quality paper, was authorized that year to accommodate these additions but continued only through 1990. The annual financial report continues to be printed in AMB as a requirement for perma- nence of public record (R. S. Prezant, pers. comm., 11 Janu- ary 2010). The membership directory was separated from the newsletter in 1997 and since 2003 has been circulated elec- tronically to membership as an independent document. In 2001, the newsletter “went electronic” and ceased hard-copy distribution except by request. Issues back to 2000 are archived and available for download on the AMS website. AMS website In 1997, almost overnight, email became the dominant method of communication among Council members. That same year, the AMU website (now at http://www.malacological. org/index.php) was launched, managed successively by Deborah Wills, Daniel L. Graf, Liath Appleton, and Brian Gollands. The AMS Secretary now routinely emails informa- tion and documents to the entire membership. The website has expanded to include a continually increasing range of reports, meeting information, electronic publications, imag- es, and other resources for members, potential members, and students. Perhaps the most widely used electronic publica- tion is the often updated “2,400 Years of Malacology,” which provides information on biographical and bibliographical publications on approx. 10,000 malacologists, conchologists, paleontologists, and others in the history of our science (Coan et al. 2009). This year, the website will begin to include pass- worded tools for officers ( e.g ., an online membership data- base) to facilitate the operations of the society. Common Names List One external publication project was (and continues to be) strongly sanctioned by AMS over the course of many years. The American Fisheries Society’s Common and Scientific Names of Aquatic Invertebrates from the United States and Canada: Mollusks, otherwise known as the “Common Names List,” cov- ers those species occurring on the American continent north of Mexico and/or within 200 miles of its shores (to 200 meters deep). For the sake of completeness, the list also includes land snails and freshwater molluscs. It provides a complete checklist plus an “official” common name for use by conservation legis- lation and other non-academic projects that are uncomfort- able with taxonomic nomenclature. The project began in 1977, when President Carol Stein established a committee “to pre- pare a list of common names of molluscs of medical impor- tance, commercial importance, of major food supply, and selected molluscs as determined by the committee.” David Stansbery took the chair of what would soon be called the Committee on Common Names of Mollusks. By 1981, Stansbery was seeking funds to publish the work. In 1983, Stansbery stepped down and Donna Turgeon took over the chair, and responsibility for the project was transferred from AMU to the Council of Systematic Malacologists. In 1985, Turgeon announced that the list of 4,700 species had been compiled by 100 contributors and reviewers. The work would ultimately be published by the American Fisheries Society as part of a new initiative to document all aquatic invertebrates of the United States. Funds to support the publication were received from Shell Oil Company, and an agreement with AFS awarded AMU a percentage (15%) of the profits after costs were recov- ered. The first edition was published in 1988 (Turgeon et al. 1988), and the second (with 6,270 species and a CD-ROM), which documented additions and changes since the first edi- tion, appeared in 1998 (Turgeon et al. 1998). Plans for a third edition are now underway, which will again track changes plus expand by adding state indicators for each species plus Hawaii and all U.S. territories {e.g., Guam, Puerto Rico, and the U.S. Virgin Islands). AFS has since published additional inverte- brate lists (in addition to its fish list, now in its 6th edition, 2004), including Cnidaria and Ctenophora (2nd edition, 2003), and Crustacea (2005, superseding Decapoda in 1989). Special publications In 1939, a Publication Committee was established to con- sider publishing a periodical helpful to beginners. In 1941, the symposium “Methods of Collecting and Preserving Mollusks” was presented at the annual meeting, which ultimately evolved into the needed publication How to Collect Shells (later How to Study and Collect Shells ) that, in the days before the internet, was extremely popular. This booklet was revised three times (La Rocque 1961, Abbott et al. 1966, Jacobson 1974, as How to Study and Collect Shells). This last title change emphasized 204 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 observation, prompted by the Conservation Committee, and was reprinted several more times, and sold for a modest $ 1 .00 per copy for most of that time. Following a workshop at the 1999 meeting in Pittsburgh (“Malacology Curation for Ama- teurs,” organized by Charles Sturm by invitation from then- President Robert S. Prezant), the “how to collect” concept was expanded in an effort to bring amateurs and professionals together (R. S. Prezant, pers. comm., 9 January 2009). The result, after several years of recruiting, writing, and editing, was the more comprehensive revision, The Mollusks: A Guide to their Study, Collection, and Preservation (Sturm etal. 2006). A few other additional official AMU publications were produced as fundraisers. Scientific Contributions Made from 1882 to 1939 was published in 1940, honoring AMU’s first president Henry A. Pilsbry on the occasion of the society’s tenth anniversary. This was a Pilsbry bibliography, 63 pages long, compiled by H. Burrington Baker (who also did anatomical work for Pilsbry; R. Robertson, pers. comm., 12 January 2010) and published by AMU, listing the 986 titles produced “in first 58 years” of Pilsbry’s career. It sold for $1.00 per copy. In the early 1980s, member and shell book dealer Richard E. Petit investigated reprinting out-of-print classic molluscan works and selling them on behalf of AMU. Roding’s Museum Boltenianum (1798) was produced in 1986 (from a facsimile originally assembled from the British Muse- um copy by C. Davies Sherborn and E. R. Sykes in 1906) and sold for $20.00 per copy with a 25% discount to members. Despite additional ideas, no further issues were produced. A gift of reprints of Binney and Tryon’s Complete Conchological Writings ofC. S. Rafinesque was also sold by AMU, as were the separates of S. Stillman Berry, Dee Dundee, and Brantley Branson, through the office of Corresponding Secretary. Most recently, Harold Murray (1999) produced a bound manuscript of a June 1979 series of ten interviews with William J. Clench at his home in Dorchester, Massachusetts, in an attempt to capture some of the oral history of malacol- ogy (as well as some of its personality - Bill was telling jokes in Chapter 10!). The volume was sold at the 1999 Pittsburg meeting, with the proceeds going to the student research fund (R. S. Prezant, pers. comm., 9-12 January 2009). GOVERNANCE AMU created its first Constitution at its first meeting in 1931. Bylaws (which govern the day-to-day operations of the society, such as the amount charged for annual dues, and require less procedure to modify) were created in 1953. The present Constitution and Bylaws have been revised nine times since 1985 (last in June 2008), each time becoming more com- plex, but at the same time staying accurate and becoming more precise in defining how AMS operates. Another important document for serving officers is the “Motions of Council and Annual Business Meetings,” which lists, by category, all of the motions “on the books” since 1982; technically all such motions are binding unless rescinded by a subsequent motion. AMU began to investigate the process of incorporation as a non-profit organization in 1961, with R. Tucker Abbott, Joseph P. E. Morrison, and Harald Rehder leading the com- mittee. Three years later, in early 1964, AMU was incorpo- rated in the State of California (because that is where the then-Treasurer, Jean Cate, resided). The society voted to maintain the incorporation in California in 1974, and in 1999, “Restated Articles of Incorporation” were filed to reflect the name change to American Malacological Society and to ensure compliance with current California Non-Profit Cor- porations law (E. V. Coan, pers. comm., 8 January 2010). AMS is governed by a Council. The first meeting created only a President, Secretary, and Treasurer (then called Finan- cial Secretary). A Vice President (who would assume the Presi- dency the following year) and four Councilors were added in 1932. A Publications Editor was added in 1954, with George M. Moore (Durham, New Hampshire) elected to fill it, succeeded in 1962 by Morris K. “Karl” Jacobson (Rockaway Beach, New York), then by Arthur Clarke (National Museum of Canada) in 1972. President Elect was added in 1971, allowing two full years for a presumptive president to learn the ropes before assuming office. Except for the offices of Presidential succession, there were no term limits at first, with each officer simply continuing as long as he or she consented to serve. Term limits were put in place in the 1970s; two years for Councilors-at-Large, three years for Corresponding Secretary, Recording Secretary, and Treasurer, and five years for Publications Editor. Secretary, Treasurer, and Publications Editor now each serve five years, and Councilors hold staggered two-year terms, with two new Councilors elected each year. The offices of Secretary and Treasurer have existed in numerous combinations. Although Norman Lermond held the office of Corresponding Secretary during the early years of the society, Imogene Robertson essentially performed his duties plus those of her own office of Financial Secretary (= Treasurer). At the 1946 meeting, the office of Treasurer was formally created, with Harold Robertson elected (he had informally and jointly occupied the effective office of Secretary-Treasurer with his wife Imogene since 1932). The two offices were recombined after the death of Harold Robertson and retirement of Imogene in 1951. Margaret Teskey occupied the office of Secretary-Treasurer until 1961, then was Secretary after a separate office of Treasurer was again created; she served in this capacity until 1969. In 1970-1971, the office of Secretary was further split into Corresponding Secretary and Recording Secretary, prompted by the work- load. By 1972, the Corresponding Secretary was receiving “about a letter a day” and answered most of them with one A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 205 of several form letters or informational lists. Constance Boone of Houston, Texas, served as Recording Secretary for nearly the entire span of its existence, from 1974 until 1989 when the three offices of Corresponding Secretary, Record- ing Secretary, and Treasurer were once more combined into one office, this time in an effort to increase efficiency.11 That attempt failed, again because of workload, and the offices were again separated into Secretary and Treasurer in 1994, as they remain today. Past Presidents were first added to Council in 1938, and from then on, all Past Presidents served on Council, with the result that by the 1970s, at least 25 Past Presidents could con- ceivably attend and vote at annual Council meetings. The number of Past Presidents actively serving on Council was reduced in 1987 (by a Bylaws amendment proposed in 1985, after a long series of debates) to the immediate three Past Pres- idents, two who had served as President 4-10 years earlier, plus two who had served as President 1 1 or more years earlier. This complement was further reduced in 2008 to only one from each of those categories, largely because of difficulties in find- ing Past Presidents willing to serve. Regrettably, the restriction of Past Presidents on Council disheartened many, who have seldom been seen at meetings after a few years past their own. The absence of the “organizational memory” inherent in our most respected and senior members is acutely felt by many members and by the officers of the AMS Council. AMS Council now consists of a President, President- Elect, Vice-President, Secretary, Treasurer, AMB Editor, three Past Presidents, and four Councilors-at-Large, one of which is a student member. Non-voting members of Council include the webmaster, the AMB Managing Editor, and the Newslet- ter Editor. The Council confers throughout the year via email and other media, and meets during the annual meeting. Committees of AMU/AMS have been difficult to trace. Early committees had specific short-term charges. In 1936, a “Checklist Committee” was established to compile a check- list of molluscs of North America north of Mexico. Then in 1939, a Publication Committee was charged with creating a periodical helpful to beginners, which ultimately resulted in the booklet How to Study and Collect Shells. A Committee 1 1 In the days before email, telephone and more often written cor- respondence between the offices of Recording Secretary, Cor- responding Secretary (by the 1980s, also Newsletter Editor), and Treasurer were required on an almost weekly basis. Most of this concerned payment of dues and changes to the membership list. For instance, the Treasurer received the dues payments, often with address changes, but the membership list was maintained by the Recording Secretary. The Corresponding Secretary often also received address changes. Today, this problem continues, but is more efficiently handled through electronic means. on Nomenclature was created in 1949 to communicate nomenclatural problems to the International Commission on Zoological Nomenclature. A Legal Committee in 1971 was charged to consider the limits of AMU activities in influ- encing legislation regarding endangered species, and to con- sider the implications of dropping the non-profit status of the society. Standing committees first appeared in the minutes spo- radically (and I have little confidence that these are the first years of these units): Constitutional Revision (1952), Public Relations (1960), Finance (1960), Auditing (1964), Nomi- nating (1964, although it surely must have existed earlier), Annual Meeting Site (1971, again probably earlier), History and Archives (1972), Membership (1972), Publications (1972), and Awards (1973). The Conservation Committee was created in 1969 and became very active politically through the 1970s and 1980s (reference duties of Legal Committee, above, in 1971), soliciting letters of support from the society in support of various actions, e.g., the Gatun Lake Resolution, supporting the maintenance of the existing freshwater barrier in the Panama Canal in 1973. A Code of Ethics was approved by the committee in 1973, and a Policy on Biological Conser- vation in 1997. The Council of Systematic Malacologists first appeared in 1973, to consider inter-institutional problems not suitable for AMU consideration. CSM operated as an independent society associated with AMU until 1997, when it disbanded; it was concurrently replaced by the standing Sys- tematics (sometimes Systematics and Collections) Commit- tee. The Institute of Malacology (the body that publishes the journal Malacologia) has also often met at AMU/AMS (from 1983 to present) although it is an independent organization. Most of the standing committees exist in some form today, appointed by the President annually to conduct society busi- ness. The current roster is Publications, Student Awards, Conservation, Constitution and Bylaws, Membership, Sys- tematics, Nominating, Endowment Review, Auditing and Budget, and Resolutions and Recognition. SOCIETAL IDENTITY In 1960, after thirty years of wildly variable covers of the Annual Reports, a juvenile freshwater Spiny Riversnail, Io fluvialis Say, 1825, chosen as “a typically American shell” and drawn by Anthony D’Attilio (with assistance from Happy Robertson), first appeared on the cover of an Annual Report. It remained there until creation of the peer-reviewed Ameri- can Malacological Bulletin in 1983. In 1964, Io fluvialis, as the “recognized symbol of AMU,” was chosen as the logo of the annual meeting in New Orleans. Io has since served as the official logo of the society, and the logo of four additional meetings (Naples 1977, Ft. Lauderdale 1981, New Orleans 206 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 1982, Charleston 2002) although most meetings since 1961 have born some sort of original meeting logo of their own. Interestingly, the first two meeting logos recognizable as such were fossil gastropods, Ecphora quadricostata (Say, 1824) (Washington, D.C., 1961) and Vasum horridum Heilprin, 1886 (St. Petersburg, Florida, 1962). Meeting logo shells have often reflected either the locale of the meeting [e.g., Scotch Bonnet, Phalium granulatum (Born, 1778), state shell of North Carolina in 1966 at the University of North Carolina; Amaea mitchelli (Dali, 1896) a prized regional beach find for Corpus Christi, Texas, in 1968] or the research interest of the President [e.g., the pyramidellid snail Fargoa bartschi (Winkley, 1909) with a spermatophore, for President Robert Robertson in 1984; a stylized snail of the family Mathildidae for President Rudiger Bieler in 1996]. Many have been original works of art [e.g., Busycon carica (Gmelin, 1791) by Anthony D’Attilio, 1965; Cancellaria gladiator R. E. Petit, 1976, by Sue Stephens, 1988]. These logos have graced program covers, T-shirts, coffee mugs, bottle openers, and reusable water bot- tles, and have become collectors’ items in their own right for many loyal society members. Charter Member Imogene Robertson began keeping AMU scrapbooks in 1931, continued in 1952 by Margaret Teskey until 1972. The “AMU Library,” consisting of scrapbooks, Annual Reports, shell club publications, and “a small but increasing collection of memorabilia and pictures of shell collectors,” had its own dedicated room at the Delaware Museum of Natural History in 1973, during R. Tucker Abbott’s curatorship. By 1975, these records were called “archives” and saw regular donations (the 1976 Annual Report noted receipt of photographic slides of past meetings and Imogene Robertson’s correspondence). In 1978, the archives were moved to the Department of Malacology, Acad- emy of Natural Sciences of Philadelphia, where they reside today. The wall of the ANSP Malacology Department displays the framed 50th Anniversary congratulatory scroll received from the Malacological Society of London by President Richard S. “Joe” Houbrick in 1981 (Fig. 6). THE FUTURE OF AMS Membership is down and declining, to the lowest totals since the 1930s. Most of the amateurs have left for CO A, and many professionals (especially freshwater and cephalopod specialists) often choose another annual meeting (sometimes more specialized, often for wider or just different exposure) due to reduced travel budgets throughout academia. There are too many specialty journals for the relatively small science of malacology, and American Malacological Bulletin was reduced recently to a single annual issue for budgetary rea- sons, so is it really important enough to keep publishing? Figure 6. The archives of the American Malacological Society reside today at the Academy of Natural Sciences in Philadelphia. Part of this includes a congratulatory scroll presented to President Richard S. “Joe” Houbrick by the Malacological Society of London on the occasion of the society’s 50th anniversary. The citation reads: “The Officers and Council of the Malacological Society of London, on behalf of its members, are pleased to send greetings and congratula- tions to the American Malacological Union on the occasion of the Fiftieth Anniversary Meeting at Fort Lauderdale, Florida 19-25 July 1991. With best wishes for your continued success.” It is signed by E. R. Trueman, President, and A. Bebbington, Secretary. [Ed. Comment: The editors spend a tremendous amount of time on the Bulletin, and would appreciate feedback on whether their efforts are appreciated .] Professors do not encourage their students to join AMS, and often they themselves do not attend. Does AMS really need to meet every year? These are bleak statements, but ones that some of us hear at nearly every meeting. Today’s Presidents must be A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 207 more and more creative in making meetings attractive, and in fundraising to make meetings self-sustaining with- out outrageous costs to attendees. Yet despite the grum- blings and difficulties, AMS meetings are attracting more and more students. Recent meetings have been praised because of their small size — as better opportunities to meet people and engage in meaningful conversations without the hustle and bustle of a gigantic meeting venue. AMS enjoys the respect of national and international organiza- tions and malacologists around the globe. Would the foun- ders be pleased? Yes, I think so. True, we must continually strive to make membership and meeting attendance mean- ingful to today’s malacologists; such is the task of all orga- nizations. Yet our society survives, evolves with the times, and in at least most respects, is thriving. In the words of Paul Bartsch in 1931, “it is a good thing to have an organi- zation of this kind.” In the eyes of at least this Past Presi- dent, it still is. ACKNOWLEDGMENTS I have had some excellent resources available to assist me with this task. Most especially, this included the AMU scrapbooks (1931-1976) in the society archives at the Acad- emy of Natural Sciences of Philadelphia, initially compiled by Imogene C. Robertson, the society’s first Financial and Recording Secretary (1931-1951), later continued by her successor Margaret Teskey (1952-1972). I am indebted to Paul Callomon, “keeper” of the archives at ANSP for allow- ing me to access this resource. I also had three previous his- torical versions to assist: the brief summary history on the AMS website (http://www.malacological.org), two articles from the Bulletin of the American Malacological Union for 1981 (on AMS’s 50th anniversary; Keen 1981, Teskey 1981), and an unpublished history of the society by Harold Murray (1991). Scott Martin (of Columbus, Ohio) provided addi- tional insight on and information about Norman Lermond, which he presented at the 2009 AMS meeting. Finally, I have been a member of AMS since 1977 (and thus now qualify as a “long-term member”), and have served as Corresponding Secretary, Newsletter Editor, Managing Editor of the AMB, and the three Presidential officers; this experience and my indelible habit of accumulating and filing away my meeting notes served me very well in this enjoyable endeavor. Alan Kabat and Gene Coan provided many interesting addenda that were unknown and unrecorded elsewhere. Thanks also to Past Presidents Rudiger Bieler, Gene Coan, Alan Kohn, Bill Lyons, Jim McLean, Diarmaid O Foighil, Bob Prezant, and Robert Robertson for editing and providing details about their annual meetings and other items under their charge. LITERATURE CITED Abbott, R.T. 1956. Shell clubs in America. Proceedings of the Phila- delphia Shell Club 1: 1-5. Abbott, R. T., G. M. Moore, J. S. Schwengel, and M. C. Teskey, eds. 1955. How to Collect Shells (a Symposium). American Malaco- logical Union, Buffalo, New York. Abbott, R. T., M. K. Jacobson, and M. C. Teskey, eds. 1966. How to Collect Shells (a Symposium), 3rd Edition. American Malaco- logical Union, Marinette, Wisconsin. Anonymous. 1966. Abstract index (1949-1965) [and] author index, abstracts, 1949-1965. American Malacological Union, Annual Reports for 1965: 105-122. Coan, E. V. and A. R. Kabat. 2007. The publications of the American Malacological Union/Society. American Malacological Bulletin 23: 1-10. Coan, E. V. and A. R. Kabat. 2009. Annotated Catalog of Malacological Meetings, Including Symposia and Workshops in Malacology. http://www.malacological.org/pdfs/catalog_of_symposia_ %28june_2009%29.pdf; last accessed on 9 January 2010. Coan, E. V., A. R. Kabat, and R. E. Petit. 2009. 2,400 Years ofMalacology, 6th ed. http://www.malacological.org/pdfs/2400years/2400yrs_ of_Malacology_Intro.pdf; last accessed on 9 January 2010. Counts, C. L., III. 1988. Index to the American Malacological Bul- letin: 1983 to 1988. Volumes 1 through 6, Special Edition num- bers 1-3. American Malacological Bulletin 6: 219-305 [see also anonymous correction in 7: 89, 1989], Dali, W. H. 1915. An index to the Museum Boltenianum. Smithsonian Institution, Washington, D.C. Jacobson, M. K., ed. 1974. How to Study and Collect Shells (a Sym- posium). American Malacological Union, Wrightsville Beach, North Carolina. Keen, A. M. 1981. A footnote to the history of malacology in the United States. Bulletin of the American Malacological Union for 1981, pp. iv-v. La Rocque, A., ed. 1961. How to Collect Shells (a Symposium), 2nd ed. American Malacological Union, Marinette, Wisconsin. Martin, S. M. 1995. Maine’s early malacological history. Maine Nat- uralist, 3: 1-34. Murray, H. D. 1991. History (Evolution) of the American Malacologi- cal Union (Society). Unpublished report prepared for the 1999 annual meeting, Pittsburgh, Pennsylvania. Murray, H. D. 1999. The Clench Tapes: A Transcription of an Interview of William J. Clench (1897-1984). Interview from 4-11 June 1979 by Harold D. Murray, Department of Biolo- gy, Trinity University, San Antonio, Texas, USA. Prepared for the July 1999 American Malacological Society Annual Meeting, Pittsburgh, Pennsylvania. Putnam, R. D. 2000. Bowling Alone: The Collapse and Revival of American Community. Simon 8c Schuster, New York. Roding, P. F. 1798. Museum Boltenianum, sive, Catalogus Cimelio- rum e Tribus Regnis Naturae quae olim Collegerat Joa. Fried Bol- ten. Pars Secunda continens Conchylia sive Testacea Univalvia, Bivalvia & Multivalvia. Hamburg, Germany [facsimiles pub- lished by Sherborne 8c Sykes, British Museum (Natural His- tory), 1906, and AMS, 1986]. 208 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Sturm, C. F., Jr., T. A. Pearce, and A. Valdes, eds. 2006. TheMollusks: A Guide to their Study, Collection, and Preservation. Universal Publishers, Boca Raton, Florida. Teskey, M. C. 1981. Half-century of AMU 1931-1981. Bulletin of the American Malacological Union for 1981, pp. v-vii. Turgeon, D. D., A. E. Bogan, E. V. Coan, W. K. Emerson, W. G. Lyons, W. L. Pratt, C. F. E. Roper, A. Scheltema, F. G. Thompson, and J. D. Williams. 1988. Common and Scientific Names of Aquatic In- vertebrates from the United States and Canada: Mollusks. American Fisheries Society Special Publication 16, Bethesda, Maryland. Turgeon, D. D., J. F. Quinn, Jr., A. E. Bogan, E. V. Coan, F. G. Hochberg, W. G. Lyons, P. M. Mikkelsen, R. J. Neves, C. F. E. Roper, G. Rosenberg, B. Roth, A. Scheltema, F. G. Thompson, M. Vecchione, and J. D. Williams. 1998. Common and Scientific Names of Aquatic Invertebrates from the United States and Can- ada: Mollusks, 2nd Edition. American Fisheries Society, Special Publication 26, Bethesda, Maryland. Submitted: 18 January 2010; accepted: 22 January 2010; final revisions received: 26 January 2010 A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 209 Appendix 1. American Malacological Union/Society annual meeting years, venues, and presidents. Year Meeting venue President Henry A. Pilsbry, Academy of Natural Sciences of Philadelphia 1931 Academy of Natural Sciences of Philadelphia, Philadelphia, Pennsylvania 1932 United States National Museum, Washington, D.C. 1933 Biological Institute, Harvard University, Cambridge, Massachusetts 1934 Geological Building, Stanford University, Stanford, California 1935 Buffalo Museum of Science, Buffalo, New York 1936 Detroit Hotel, St. Petersburg, Florida 1937 Museum of Zoology, University of Michigan, Ann Arbor, Michigan 1938 Escuela de Ciencias, Universidad de la Habana, Havana, Cuba 1939 Royal Ontario Museum of Zoology, Toronto, Canada 1940 Academy of Natural Sciences of Philadelphia, Philadelphia, Pennsylvania 1941 Knox Academy of Arts and Sciences, Thomaston, Maine, and Crescent Beach Inn, Owl’s Head, Rockland, Maine 1942 [no meeting, World War II] 1943 [no meeting, World War II] 1944 [no meeting, World War II] 1945 [no meeting, World War II] 1946 United States National Museum, Washington, D.C. 1947 Asilomar Hotel and Conference Grounds, Pacific Grove, California 1948 Carnegie Museum, Pittsburgh, Pennsylvania 1949 University of Miami, Coral Gables, Florida 1950 Chicago Museum of Natural History, Chicago, Illinois 1951 Buffalo Museum of Science, Buffalo, New York 1952 Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 1953 Memorial Union Building, University of Kansas, Lawrence, Kansas 1954 Nesmith Hall, University of New Hampshire, Durham, New Hampshire 1955 Wagner College, Grymes Hill, Staten Island, New York 1956 Hotel Lafayette, San Diego, California (Joint meeting with AMU Pacific Division) 1957 Peabody Museum, Yale University, New Haven, Connecticut 1958 South Quadrangle, University of Michigan, Ann Arbor, Michigan 1959 Roberts Hall, Haverford College, Haverford, Pennsylvania, and Academy of Natural Sciences of Philadelphia, Pennsylvania 1960 Redpath Museum, McGill University, Montreal, Canada 1961 Natural History Museum, United States National Museum, Washington, D.C. 1962 Florida Presbyterian College, St. Petersburg, Florida Henry A. Pilsbry, Academy of Natural Sciences of Philadelphia Paul Bartsch, United States National Museum Junius Henderson, University of Colorado Museum William J. Clench, Museum of Comparative Zoology, Harvard University Calvin Goodrich, Museum of Zoology, University of Michigan Joshua L. Baily Jr., San Diego Museum of Natural History Carlos de la Torre, Universidad de la Habana Maxwell Smith, Florida H. Burrington Baker, University of Pennsylvania Harald A. Rehder, United States National Museum Frank Collins Baker, Museum of Natural History, University of Illinois at Urbana Louise M. Perry, Asheville, North Carolina Henry van der Schalie, Museum of Zoology, University of Michigan Henry van der Schalie, Museum of Zoology, University of Michigan Henry van der Schalie, Museum of Zoology, University of Michigan Henry van der Schalie, Museum of Zoology, University of Michigan A. Myra Keen, Stanford University Elmer G. Berry, Museum of Zoology, University of Michigan Fritz Haas, Chicago Museum of Natural History Joseph P. E. Morrison, United States National Museum Jeanne S. Schwengel, Scarsdale, New York A. Byron Leonard, University of Kansas Joseph C. Bequaert, Museum of Comparative Zoology, Harvard University Morris K. “Karl” Jacobson, American Museum of Natural History Allyn G. Smith, California Academy of Sciences Ruth D. Turner, Museum of Comparative Zoology, Harvard University Aurele La Rocque, Ohio State University R. Tucker Abbott, Academy of Natural Sciences of Philadelphia Katherine Van Winkle Palmer, Paleontological Research Institution Thomas E. Pulley, Museum of Natural History, Houston William K. Emerson, American Museum of Natural History 210 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Appendix 1. (Continued) Year Meeting venue President 1963 Buffalo Museum of Science, Buffalo, New York 1964 Sheraton-Charles Hotel, New Orleans, Louisiana 1965 Wagner College, Staten Island, New York 1966 University of North Carolina, Chapel Hill, North Carolina 1967 Carleton University and National Museum of Canada, Ottawa, Canada 1968 Robert Driscoll Motor Hotel, Corpus Christi, Texas 1969 Science Hall, Marinette County Campus, University of Wisconsin-Green Bay, Marinette, Wisconsin 1970 Key Wester Motor Inn and Villas, Key West, Florida 1971 Atlantis Convention Center, Cocoa Beach, Florida 1972 Galvez Hotel, Galveston, Texas 1973 Clayton Convention Center, University of Delaware, Newark, Delaware, and Delaware Museum of Natural History, Greenville, Delaware 1974 Museum of Science, Springfield, Massachusetts 1975 San Diego State University, San Diego, California (Joint meeting with the Western Society of Malacologists) 1976 Ohio State University, Columbus, Ohio 1977 Beach Club Hotel, Naples, Florida 1978 University of North Carolina, Wilmington, North Carolina 1979 La Quinta Royale Motor Inn, Corpus Christi, Texas (Joint meeting with the Western Society of Malacologists) 1980 Executive Inn East, Louisville, Kentucky 1981 Galt Ocean Mile Hotel, Ft. Lauderdale, Florida 1982 Fountain Bay Club Hotel, New Orleans, Louisiana 1983 McCarthy Hall, University of Washington, Seattle, Washington (Joint meeting with the Western Society of Malacologists) 1984 Holiday Inn - Waterside, Norfolk, Virginia 1985 Chafee Hall, University of Rhode Island, Kingston, Rhode Island 1986 Monterey Sheraton, Monterey, California (Joint meeting with the Western Society of Malacologists) 1987 Marriott’s Casa Marina Resort, Key West, Florida 1988 Radisson Francis Marion Hotel, Charleston, South Carolina 1989 Davidson Conference Center, University of Southern California, and Natural History Museum of Los Angeles County, Los Angeles, California (Joint meeting with the Western Society of Malacologists) 1990 Swope Conference Center, Marine Biological Laboratory, Woods Hole, Massachusetts 1991 Clark Kerr Campus, University of California, Berkeley, California (Joint meeting with the Western Society of Malacologists) 1992 Hyatt Sarasota, Sarasota, Florida 1993 Nordic Empress Cruise, Miami, Florida, to the Bahamas 1994 Hyatt Regency Downtown, Houston, Texas 1995 University of Hawai’i, Hilo, Hawaii 1996 Field Museum of Natural History, Chicago, Illinois Albert R. Mead, University of Arizona John Q. Burch, Los Angeles, California Juan J. Parodiz, Carnegie Museum Ralph W. Dexter, Kent State University Leo G. Hertlein, California Academy of Sciences Arthur H. Clarke Jr., National Museum of Canada Joseph Rosewater, United States National Museum Alan G. Solem, Field Museum of Natural History David H. Stansbery, Museum of Zoology, Ohio State University Arthur S. Merrill, National Marine Fisheries Service, Oxford, Maryland Dolores S. “Dee” Dundee, Louisiana State University Harold D. Murray, Trinity University, Texas Donald R. Moore, Institute of Marine Sciences, Miami Dorothea S. Franzen, Illinois Wesleyan University George M. Davis, Academy of Natural Sciences of Philadelphia Carol B. Stein, Ohio State University William E. Old, Jr., American Museum of Natural History Clyde F. E. Roper, National Museum of Natural History Richard S. “Joe” Houbrick, National Museum of Natural History Louise Russert Kraemer, University of Arkansas Alan J. Kohn, University of Washington Robert Robertson, Academy of Natural Sciences of Philadelphia Melbourne R. Carriker, University of Delaware James Nybakken, Moss Landing Marine Laboratory William G. Lyons, Florida Department of Natural Resources Richard E. Petit, Charleston, South Carolina James H. McLean, Natural History Museum of Los Angeles County Roger T. Hanlon, University of Texas Medical Branch, Galveston Carole S. Hickman, University of California at Berkeley Robert C. Bullock, University of Rhode Island Fred G. Thompson, Florida Museum of Natural History Constance E. Boone, Houston Museum of Natural Science E. Alison Kay, University of Hawai’i at Hilo Rudiger Bieler, Field Museum of Natural History A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 211 Appendix 1. (Continued) Year Meeting venue President 1997 Radisson Hotel, Santa Barbara, California (Joint meeting with the Western Society of Malacologists) 1 998 S. Dillon Ripley Conference Center, Smithsonian Institution, Washington, D.C. (at World Congress of Malacology) 1 999 Sheraton Hotel Station Square, Pittsburgh, Pennsylvania 2000 Seven Hills Conference Center, San Francisco State University, San Francisco, California (Joint meeting with the Western Society of Malacologists) 2001 Institute of Zoology, University of Vienna, Vienna, Austria (at World Congress of Malacology) 2002 Lightsey Conference Center, College of Charleston, Charleston, South Carolina 2003 University of Michigan, Ann Arbor, Michigan 2004 Sundial Beach Resort, Sanibel Island, Florida 2005 Asilomar Conference Grounds, Pacific Grove, California (Joint meeting with the Western Society of Malacologists) 2006 University of Washington, Seattle, Washington (Joint meeting with the Western Society of Malacologists) 2007 Groenenborger Campus, University of Antwerp, Antwerp, Belgium (at World Congress of Malacology) 2008 Student Center, Southern Illinois University, Carbondale, Illinois 2009 Cornell University, Ithaca, New York Eugene V. Coan, Santa Barbara Museum of Natural History Robert Hershler, National Museum of Natural History Robert S. Prezant, Queen’s College, New York Terrence M. Gosliner, California Academy of Sciences Janice Voltzow, University of Scranton Robert T. Dillon, Jr., College of Charleston Diarmaid M. O Foighil, University of Michigan Jose H. Leal, Bailey-Matthews Shell Museum, Sanibel Island Dianna K. Padilla, State University of New York at Stony Brook Roland C. Anderson, Seattle Aquarium Paula M. Mikkelsen, Paleontological Research Institution Frank “Andy” Anderson, Southern Illinois University Warren D. Allmon, Paleontological Research Institution Appendix 2. American Malacological Union Pacific Division annual meeting years, venues, and chainnen (then applied to both men and women). Year Meeting venue Chairman 1948 Hancock Foundation, University of Southern California, Los Angeles, California Ruth E. Coats, Tillamook, Oregon 1949 Municipal Auditorium, Long Beach, California [not available] 1950 The Barbara Hotel, Santa Barbara, California John Q. Burch, Los Angeles, California 1951 Mills College, Oakland, California Leo G. Hertlein, California Academy of Sciences 1952 Founders Hall, University of Southern California, Los Angeles, California Wendell O. Gregg, Los Angeles, California 1953 Asilomar Hotel and Conference Grounds, Pacific Grove, California Allyn G. Smith, California Academy of Sciences 1954 Founders Hall, University of Southern California, Los Angeles, California Elsie M. Chace, Lomita, California 1955 Department of Geology, Stanford University, Stanford, California Ralph O. Fox, California Academy of Sciences 1956 Hotel Lafayette, San Diego, California (Joint meeting with AMU) Edward P. Baker, Downey, California 1957 Mar Monte Hotel, Santa Barbara, California Edward P. Baker, Downey, California 1958 Life Sciences Building, University of California, Berkeley, California Albert R. Mead, University of Arizona 1959 University of Redlands, Redlands, California John E. Fitch, California State Fisheries Laboratory 1960 Asilomar Conference Grounds, Pacific Grove, California Rudolf Stohler, University of California at Berkeley 212 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Appendix 2. (Continued) Year Meeting venue Chairman 1961 University of California at Santa Barbara, Goleta, California Howard R. Hill, Natural History Museum of Los Angeles County (deceased during year), succeeded by Robert W. Talmadge, California Academy of Sciences 1962 Asilomar Conference Grounds, Pacific Grove, California Robert W. Talmadge, California Academy of Sciences 1963 University of California at Santa Barbara, Goleta, California Crawford N. Cate, Los Angeles, California 1964 Asilomar Conference Grounds, Pacific Grove, California A. Myra Keen, Stanford University 1965 University Lodge, Point Loma Campus, California Western University, San Diego, California Edwin C. Allison, La Jolla, California 1966 Department of Zoology, University of Washington and Pacific Science Center, Seattle, Washington Alan J. Kohn, University of Washington 1967 Merrill Hall, Asilomar Conference Grounds, Pacific Grove, California Gale G. Sphon, Jr., Santa Barbara Museum of Natural History 1968 Asilomar Conference Grounds, Pacific Grove, California Fay Wolfson, La Jolla, California 1969 Asilomar Conference Grounds, Pacific Grove, California (Joint meeting with Western Society of Malacologists) G. Bruce Campbell, Lynwood, California Appendix 3. Honorary appointments of the American Malacological Union/Society. Honorary Members (1932-1952): Thomas Barbour, 1933-1946 Charles Torrey Simpson, 1932 Victor Sterki, 1932-1933 Bryant Walker, 1932-1936 Honorary Life Members (1952-present): R. Tucker Abbott, 1981-1995 Ralph Arnold, 1959-1960 H. Burrington Baker, 1958-1971 Paul Bartsch, 1953-1958, thereafter Honorary Life President Joseph C. Bequaert, 1958-1982 S. Stillman Berry, 1953-1960, thereafter Honorary Life President Melbourne R. Carriker, 1997-2007 Emery P. Chace, 1973-1980 William J. Clench, 1961-1984 Eugene V. Coan, 2008-present William K. Emerson, 1987-present Dorothea S. Franzen, 2006-2008 Julia Gardner, 1959-1960 Fritz Haas, 1960-1969 Leo G. Hertlein, 1970-1972 Morris K. “Karl” Jacobson, 1980 A. Myra Keen, 1968-1985 Alan J. Kohn, 2004-2008, thereafter Honorary Life President James H. McLean, 2004-present Joseph P. E. Morrison, 1978-1983 Harold D. Murray, 1999-present A HISTORY OF THE AMERICAN MALACOLOGICAL SOCIETY 213 Katherine Van Winkle Palmer, 1961-1982 Richard E. Petit, 1997-present Henry A. Pilsbry, 1953-1957 Harald A. Rehder, 1978-1985, thereafter Honorary Life President Imogene C. Robertson, 1952-1953 Robert Robertson, 1997-present Margaret C. Teskey, 1967-1996 Ruth D. Turner, 1981-1997, thereafter Honorary Life President Henry van der Schalie, 1982-1986 J. Z. Young, 1990-1997 Honorary Life Members (Pacific Division): Andrew Sorenson, 1956-1962 Honorary Presidents (1932-1958): IdaS. Oldroyd, 1934-1940 Henry A. Pilsbry, 1937-1957 Imogene C. Robertson, 1940-1953 Honorary Life Presidents (1959-present, reserved for only one person at any one time): Paul Bartsch, 1959 S. Stillman Berry, 1960-1984 Harald A. Rehder, 1985-1996 Ruth D. Turner, 1997-2000 Alan J. Kohn, 2008-present Amer. Malac. Bull. 28: 215-217 (2010) James W. Nybakken: September 16, 1936 - June 20, 2009 An Appreciation Alan J. Kohn Department of Biology, Box 351800, University of Washington, Seattle, Washington 98195-1800, U.S.A. Corresponding author: kohn@u.washington.edu A long-time member of AMU/AMS, President in 1986, and distinguished teacher, marine biologist, and researcher, James W. Nybakken (Fig. 1) died of leukemia June 20, 2009 at the age of 72. Jim did his graduate work at the University of Wiscon- sin, completing his Ph.D. in 1965 with a dissertation on the ecology of Three Saints Bay on Kodiak Island, Alaska. Shortly after his study, an earthquake elevated much of the bay’s intertidal zone several meters, and Jim’s thesis, published by the University of Alaska (1969) provided invaluable baseline data to assess the ecological effects of the earthquake. In Sep- tember 1965, Jim joined the faculty of California State Uni- versity Hayward (now CSU East Bay). In January 1966, he began teaching at the CSU system’s Moss Landing Marine Laboratories (MLML), where he was a member of the found- ing faculty and where he spent the rest of his career. He served as acting director of the Laboratories in the mid-1990s. “Throughout his tenure at MLML he witnessed the transfor- mation of a small field station cobbled together in an old cannery building to a modern marine institution with an international reputation for excellence in marine science.” (www.mlml. calstate.edu/ announcements/news/6-24-09). I first met Jim when he was a graduate student participat- ing in a three-month International Indian Ocean Expedition (IIOE) cruise on the then Stanford University teaching and research vessel Te Vega in 1963. This ship facilitated inshore and coral reef studies in the western Indian Ocean, from Singapore through the Strait of Malacca and Malaysia to northern Thailand, and then through the islands off the south coast of Sumatra, ending rather ignominiously with an irrevocably broken drive shaft off Padang. The scientific personnel consisted of six faculty members and museum curators and six graduate students from diverse universities. Jim and I worked together on com- parative ecology of the important reef- associated gastropod Conus, discovering in the process the most species-rich assem- blages known at the time. Pig. 2 shows Jim, as well as Joe Rosewater, who also served as AMU president, in 1969, aboard the Te Vega during the IIOE cruise. In 1968-69, Jim, together with his wife Bette and their two small sons, Kent and Scott, spent a year’s leave of absence from MLML in Seattle, where Jim and I analyzed the field data from the IIOE cruise and worked on writing up the results. We were able to obtain ecological data on 48 species, and we found up to 27 of them on a single reef. Co-occurring Conus species typically specialize differentially on prey spe- cies, but those in more diverse assemblages proved not to be more specialized. Rather, they consumed a wider array of prey taxa, and they specialized more on different microhabi- tats than did those in habitats that support fewer species. We also demonstrated size- selective predation, that only the larg- est individuals of prey species may be large enough to repay foraging effort, and that Conus individuals shift their diets to larger prey species as they grow (Kohn and Nybakken 1975). During that year, we also collaborated on the first scan- ning electron microscopic morphological study of the harpoon- like Conus radular tooth (Kohn etal. 1972), and Jim embarked on studies of the ecology and radular morphology of Eastern Pacific Conus (Nybakken 1970a, 1970b, 1971, 1979a, 1979b). While Jim’s biological interests were broad, most but not all of his research focused on molluscs. His other notable con- tribution to the biology of Conus was the first study ever made of ontogeny of radular teeth (Nybakken and Perron 1988, Figure 1. James W. Nybakken (1936-2009) in 2007. 215 216 AMERICAN MALACOLOGICAL BULLETIN 28 • 1/2 • 2010 Figure 2. Two future AMU presidents aboard Te Vega off Suma- tra during the International Indian Ocean Expedition in December 1963. Left to right: Klaus Rutzler (Smithsonian Institution), Jim Nybakken (AMU President, 1986), Joseph Rosewater (AMU President, 1969), and Llewellya Colinvaux (Ohio State University). Nybakken 1990), and he honored the genus on his car’s license plate (Fig. 3). He also carried out a number of studies of nudi- branch ecology along the central California coast {e.g., Nybakken 1978). In the 1990s, Jim turned his attention to the deep sea in a 2-year study of the ecology of communities at the base of the continental slope off California (Nybakken et al. 1998). However, Jim is really best known for his five books, especially Marine Biology: An Ecological Perspective. This widely used text has gone through six editions, the first five Figure 3. Jim Nybakken’s personalized “CONUS” California license plate in place on his vehicle. Figure 4. Upper figure: Alan Kohn’s graduate students and visiting researchers at end-of-the year party, Edmonds, Washington in June 1969. Back row, left to right: Robert Kaufman, Richard Strathmann, Andrea Hammond, James Nybakken, Charles Galt, Margaret Lloyd, and Daniel Luchtel. Front row: Pamela Roe, Susanne Lawrenz Miller, Alan Kohn, and Sarah Ann Woodin. Lower figure: Seven from the upper group at retirement party, in the same positions as in upper figure. Denver, Colorado in January 1999. self-authored and the most recent (2004) co-authored with Mark Bertness of Brown University. Jim’s other books include Diversity of the Invertebrates: A Laboratory Manual (1996), Guide to the Nudibranchs of California (1980), and Readings in Marine Ecology (1986). When Jim presided over AMU in 1986, he organized the Annual Meeting at Monterey. The last meeting he attended was the 2005 meeting at Asilomar, California. Jim retired in 1998 but remained an active participant in MLML activities as well as pursuing his diverse recreational activities, including orchid culture — as a semi-professional with a 6-greenhouse nursery, wine making, Norwegian cul- ture, and world travel. In 1999, I was honored that Jim joined a party that my former students and associates arranged at the Society for AN APPRECIATION 217 Integrative and Comparative Biology meeting in Denver on the occasion of my retirement. We took advantage of the occasion to place those who had also attended the 1969 labo- ratory group party at our home in the same positions for a photograph (Fig. 4). Clearly, we had not changed very much in 30 years! Jim Nybakken will be greatly missed, but his legacy lives on in his dozens of scientific papers, his widely used books, the students he influenced during his career marked by edu- cational leadership, and the students who have been and will be supported through the James W. Nybakken Scholarship administered by the Friends of Moss Landing Marine Labora- tories (friends@mlml.calstate.edu). CITED PUBLICATIONS OF JAMES W. NYBAKKEN Kohn, A. J. and J. W. Nybakken. 1975. Ecology of Conus on eastern Indian Ocean fringing reefs: Diversity of species and resource utilization. Marine Biology 29: 211-234. Kohn, A. J., J. W. Nybakken, and J.-J. Van Mol. 1972. Radula tooth structure of the gastropod Conus imperialis elucidated by scan- ning electron microscopy. Science 176: 49-51. McDonald, G. R. and J. W. Nybakken. 1980. Guide to the Nudibranchs of California. American Malacologists, Melbourne, Florida. Nybakken, J. W. 1969. Pre-earthquake intertidal ecology of Three Saints Bay, Kodiak Island, Alaska. Biological Papers of the Uni- versity of Alaska, No. 9: 117 pp. Nybakken, J. W. 1970a. Correlation of radula tooth structure and food habits of three vermivorous species of Conus. The Veliger 12:316-318. Nybakken, J. W. 1970b. Radular anatomy and systematics of the West American Conidae (Mollusca, Gastropoda). American Museum Novitates No. 2414, 29 pp. Nybakken, J. W. 1971. The Conidae of the Pillsbury Expedition to the Gulf of Panama. Bulletin of Marine Science 21: 93-110. Nybakken, J. W. 1978. Abundance, diversity and temporal variabil- ity in a California intertidal nudibranch assemblage. Marine Biology 45: 129-146. Nybakken, J. W. 1979a. Population characteristics and food resource utilization of Conus in the Sea of Cortez and West Mexico. Journal ofMolluscan Studies 45: 82-97. Nybakken, J. W. 1979b. Population characteristics and food re- source utilization of Conus in the Galapagos Islands. Pacific Sci- ence 32: 271-280. Nybakken, J. W. 1986. Readings in Marine Ecology, 2nd Edition. Harper and Row, New York. Nybakken, J. W. 1990. Ontogenetic change in the Conus radula, its form, distribution among the radula types, and significance in systematics and ecology. Malacologia 32: 35-54. Nybakken, J. W. 1996. Diversity of the Invertebrates: A Labora- tory Manual; Pacific Coast Version. Wm. C. Brown Publishers, Dubuque, Iowa. Nybakken, J. W., S. Craig, L. Smith-Beasley, G. Moreno, A. Summers, and I. Weetman. 1998. Distribution density and relative abundance of benthic invertebrate megafauna from three sites at the base of the continental slope off central California as determined by camera sled and beam trawl. Deep-Sea Research, Part II 45: 1753-1780. Nybakken, J. W. and F. Perron. 1988. Ontogenetic change in the radula of Conus magus (Gastropoda). Marine Biology 98: 239-242. Nybakken, J. W. and M. D. Bertness. 2004. Marine Biology: An Eco- logical Perspective, 6th Edition. Pearson, Benjamin Cummings, San Francisco. Submitted: 9 January 2010; accepted: 9 January 2010; final revisions received: 22 January 2010 INFORMATION FOR CONTRIBUTORS Scope. The American Malacological Bulletin is the scientific publication of the American Malacological Society and serves as an outlet for reporting notable contributions in malacological research. Manuscripts concerning any aspect of original, unpublished research, important short reports, and detailed reviews dealing with molluscs will be consid- ered for publication. Format. Manuscripts and illustrations should be submitted electronically (in a Word document or pdf file with embed- ded figures). Text must be typed in 12 pt font on 8.5 X 11 inch (letter-sized) paper, double-spaced, with all pages numbered consecutively. Leave ample margins on all sides, and left-justify the text. Final submission of accepted, revised manuscripts must include the text, tables, etc. as a man- datory MS Word file on a CD, DVD, or e-mail attach- ment, along with high resolution TIFF files of all figures. 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The full generic name along with specific epithet should be written out the first time that taxon is referred to in each paragraph. The generic name can be abbreviated in the remainder of the paragraph as follows: C. virginica. The taxonomic authorities of generic names must be provided if species names are not included. Please refer to recent issues for examples. Text References. Literature citations should be cited within text as follows: Hillis (1989) or (Hillis 1989). Dual author- ship should be cited as follows: Yonge and Thompson (1976) or (Yonge and Thompson 1976). Multiple authors of a single article should be cited as follows: Beattie et al. (1980) or (Beattie et al. 1980). Literature Cited Section. References must also be typed double-spaced. All authors must be fully identified and listed alphabetically; journal titles must be unabbreviated. When more than one publication with the same first author is cited, please arrange citations as follows: (a) single author, according to publication dates; (b) same author and one co-author in alphabetical, then chronological order; (c) same author and more than one co-author. Citation Format. Beattie, J. H., K. K. Chew, and W. K. Hershberger. 1980. Differential survival of selected strains of Pacific oysters ( Crassostrea gigas) during summer mortality. Proceedings of the National Shellfisheries Association 70: 184-189. Hillis, D. M. 1989. Genetic consequences of partial self fer- tilization on populations of Liguus fasciatus (Mollusca: Pulmonata: Bulimulidae). American Malacological Bulle- tin 7: 7-12. Seed, R. 1980. Shell growth and form in the Bivalvia. In: D. C. Rhoads and R. A. Lutz, eds., Skeletal Growth of Aquatic Organisms. Plenum Press, New York, New York. Pp. 23-67. Yonge, C. M. and T. E. Thompson. 1976. Living Marine Molluscs. William Collins Son and Co., Ltd., London. For more detailed examples of journal series, supplements, graduate theses, governmental reports, Internet citations, non-English citations, or other categories of references, please check the following AMS web page under “Submission”: http:// malacological.org/publications/ authors.html. Resources. The manuscript should follow the style rules out- lined in The CSE Manual for Authors, Editors, and Publishers (7th edition, June 2006). This can be purchased from the CSE at 12100 Sunset Hills Rd., Suite 130, Reston, Virginia 20190, USA or at the following web site: http://www. councilscienceeditors.org/publications/style.cfm. Spelling should follow American English as listed in Merriam- Webster Online (http://www.m-w.com/) or recent hardcopy editions. Punctuation. Punctuation should follow that shown in the most recent issues of AMB. Specific common examples include: • Italics: e.g., i.e., sensu, per se, et al. • Non-italicized text: pers. comm., pers. obs., and unpubl. data • Font: please use Times New Roman 12 (if possible). 219 Figures. Authors are strongly encouraged to submit elec- tronic copies of the figures on CDs or DVDs (hardcopy submissions may incur additional costs). Illustrations should be clearly detailed and readily reproducible. Fine patterns and screens do not reproduce well. All figure panels must be marked with capitalized letters (A, B, C, etc . . .) and ad- equately labeled with sufficiently large labels to remain read- able with reduction by one half. Magnification bars must appear on the figure (except for graphs), or the caption must read horizontal field width = x mm or x pm. All measure- ments must be in metric units. When creating figures, use font sizes and line weights that will reproduce clearly and accurately when figures are sized to the appropriate column width. Please do not include figure legends in a graphic file. Explanations of abbreviations used in a figure should occur in the legend. Please see recent journal issues for cor- rect format. Indicate in text margins the appropriate loca- tion in which figures should appear. Color illustrations can be included at extra cost to the author (currently about $650-700/figure). Figure Format. For final revisions of papers submitted to the editor, all electronic figure files should be in TIFF for- mats, preferably with LZW image compression. Each individ- ual figure or graphic must be supplied as a separate, stand- alone file (not as an embedded object). Figure files must be named with their respective numbers and graphic type such as Figl.tif, Figure2.tif, etc. Figure Resolution. AMB quality reproduction will require grayscale and color files at resolutions yielding approxi- mately 350 dpi. Bitmapped line art should be submitted at resolutions yielding 600-1200 dpi. These resolutions refer to the output size of the file (85 mm for single column or 170 mm for double column); if you anticipate that your images will be enlarged or reduced, resolutions should be adjusted accordingly. Please check your output resolu- tion prior to manuscript submission. Directions on how to check are available at the following AMS web page under “Submission”: http://malacological.org/publications/authors. html. The production of high-resolution figures is the re- sponsibility of the author(s). Mandatory AMB Style. Any manuscript not conforming to AMB format will be returned to the author for revision before publication. Manuscripts are accepted contingent upon authors making final AMB stylistic revisions. New Taxa. The Bulletin welcomes complete descriptions of new molluscan taxa. Establishment of new taxa must con- form with the International Code of Zoological Nomencla- ture (1999). Descriptions of new species-level taxa must in- clude the following information in the order as given: higher taxon designation as needed for clarity; family name with author and date; generic name with author and date; Genus species author sp. nov. followed by numeration of all figures and tables; complete synonymy (if any); listing of type material with holotype and any other type material clearly designated along with complete museum catalogue or acces- sion information; listing of all additional non-type material also with full museum deposition information; type locality; diagnosis and full description of material done in telegraphic style including measurements and zoogeographic distribu- tion as necessary; discussion; etymology. Descriptions of new supraspecific taxa should include type species (for new genus) or type genus (for new family), diagnosis and full description done in telegraphic style, and list of included taxa. Proofs. Page proofs will be sent to the author and must be checked for printer’s errors and returned to the Managing Editor within a 1-week period. Changes in text, other than printer errors, will result in publishing charges that will be billed to the author. Charges. There are no mandatory page costs to authors lacking financial support. Authors with institutional, grant, or other research support will be billed for page charges. The current rate is $50 per printed page. Acceptance and ulti- mate publication is in no way based on ability to pay page costs. However, changes at the proof stage are mandatory costs set by the publisher. Reprints. Order forms and reprint cost information will be sent with page proofs. The author receiving the order form is responsible for insuring that orders for any co-authors are also placed at that time. Electronic e-reprints (with unlim- ited distribution) are available from BioOne (www.bioone. org/loi/malb) after the electronic version of the volume is released. Submission. Submit all manuscripts to: Dr. Kenneth M. Brown, Editor-In-Chief, Department of Biological Sciences, Louisiana State University, Baton Rouge, Louisiana 70803, USA. Please refer to the AMS web page for more detailed information prior to submission, http://malacological.org/ publications/authors.html. Subscription Costs. Institutional subscriptions are available at a cost of $75 per volume. Membership in the American Malacological Society, which includes personal subscriptions to the Bulletin, is available for $60 ($20 for students, $60 for affiliated clubs). All prices quoted are in U.S. funds. Outside the U.S. postal zones, add $5 airmail per volume (within North America) or $10 airmail per volume (other locations). For membership information contact Dr. Dawn Dittman, Treasurer, Tunison Laboratory of Aquatic Science, 3075 Gracie Rd„ Cortland, New York 13045-9357, USA. For in- stitutional subscription and back-issue information contact Dr. Kenneth M. Brown, Editor-In-Chief, Department of Bio- logical Sciences, Louisiana State University, Baton Rouge, Louisiana 70803, USA. Complete information is also avail- able at the AMS website: http://www.malacological.org. 220 THE AMERICAN MALACOLOGICAL SOCIETY NAME Title MRS. AMANDA S. LAWLESS, Secretary Department of Malacology Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA 19103-1195, U.S.A. MEMBERSHIP RENEWAL 2010 CALENDAR YEAR (January 1-December 31). Please complete form and mail with dues payment to the Treasurer at the address above. First Name Middle Initial Last Name MAILING INFORMATION: Please list my e-mail in the AMS Directory and contact me via e-mail to save time and money. I I Please do not list my e-mail in the Directory. I I Please use snail-mail when corresponding with me. My mailing address and e-mail address are correct in the AMS Directory. EH Please correct my mailing address and/or e-mail address as follows: Members outside the US: please note that there is a postage & handling fee for the Bulletin. MEMBERSHIP CATEGORY (please check box and circle amount paid): □ Regular Member - One year dues (2010) $ 60.00 □ Regular Member - Two years (2010 & 201 1) $105.00 I I Regular Member - Three years (2010-2012) $145.00 I I Each additional family member per year $ 1 .00 I I Student Member $ 20.00 I I Sustaining Member - dues plus $25.00 $ 85.00 I I Affiliate Membership (Shell Clubs & other organizations) $ 60.00 I I Membership reinstatement/back issues $60 Regular/$20 Student for 2009 $ . POSTAGE EH Canada & Mexico $5.00 per year I I All other foreign addresses $10.00 per year $ . TAX DEDUCTIBLE GIFT I I To Symposium Endowment Fund $ . EH To Student Research Grant Endowment Fund $ . TOTAL ENCLOSED $ . Payment can be made by check on a U.S. bank, by International Money Order, or by MasterCard or Visa. Make checks payable to the AMERICAN MALACOLOGICAL SOCIETY. AMS does not issue receipts or confirm membership status unless a request is sent to ddittman@usgs.gov If you wish to make payment via VISA or MasterCard, please complete the following: EHVisa EHmC Card# Expires 3- or 4-digit security code Signature of cardholder Thank you for your continued support of AMS! AMERICAN MALACOLOGICAL SOCIETY, INC. FINANCIAL REPORT General Accounts 2007 Income and Expenses TOTAL ASSETS (January 1, 2007) $193,587.74 INCOME Membership Dues 14,455.83 Membership Dues (2006) 1,112.00 Membership Dues (2007) 1 10,503.58 Membership Dues (2008) 1,656.68 Membership Dues (2009) 1,135.23 Membership Dues (2010) 48.34 Publications Income 13,730.88 AMB Subscriptions 3,425.00 AMB Page Charges 5,907.00 AMB Back Issues 270.00 AMB and Book Royalties 4,068.88 AMB Postage & Misc. Income 60.00 Donations 5,719.65 Student Travel Fund 1,200.00 Student Endowment Fund 545.00 Student Fund from Auction 3,974.65 Interest and Dividends from Endowment 8,691.68 Life Membership Fund 1,120.14 Symposium & Student Fund 7,571.54 $42,598.04 EXPENSES $30,661.19 Treasurer and Secretary Office Expenses 66.40 Affiliate Memberships 173.00 Banking & Credit Card Fees 794.35 Refunds 1,445.00 Incorporation & Registration Fees 45.00 Insurance/Bond Fees 282.00 Website Expenses 110.35 Annual Meeting & Symposium Expenses 4,445.89 Publication Expenses AMB (22 1/2) 12,449.05 Reprints 1,170.45 Managing Editor Travel etc. 556.81 Misc. postage etc 1,002.52 Student Research Grants 3,000.00 Student Travel Awards 3,000.00 Travel Expenses for Officers 1,620.33 Student Paper Awards 500.00 Net Gain in 2007 $11,936.85 TOTAL ASSETS (December 3 1 , 2007) $205,524.59 ** Includes capital gains and losses in endowment portfolios which fluctu- ate with the market. 222 The American Malacological Society and the Western Society of Malacologists 76th (AMS) and 43rd (WSM) • 26 June - 1 July 2010 Joint Annual Meetings San Diego, California As the current President of AMS, it is my pleasure to announce the upcoming joint 76th Annual Meeting of the AMS and 43rd Annual Meeting of the Western Society of Malacologists. The meeting will kick off in style with a welcome reception on the evening of Saturday, June 26th, with scientific sessions held from Sunday (June 27th) to noon on Wednesday (June 30th). Stay the following day, Thursday, July 1st, for organized or informal excursions in the San Diego area. Venue. - We are fortunate to have reserved our meeting site at San Diego State University. The new SDSU convention center is attractive and easily accessible by car or via public transportation. SDSU has both new and somewhat less expensive older dorms, but both are affordable. Alternatively, there are many hotels close by. Meeting attendees, whether staying in the dorms or not, can choose between an excellent optional meal plan or forage on their own in nearby university district or convention center restaurants. Public transit options include a new trolley stop, which is adjacent to the convention center or a brief walk across a footbridge from the dorms. The trolley connects directly to the downtown train station and airport (search in Google maps for SDSU Transit Center, San Diego), and provides easy access to the outstanding selection of restaurants and bars in famous Old Town, San Diego about 29 minutes by trolley. The meeting will be affordable for students and close to world-class ocean beaches and other well-known attractions that make San Diego such a popular destination to visit. Sessions. - Scientific sessions are historically great at joint AMS/WSM West Coast meetings. • Dr. Peter Marko ( Assistant Professor, Clemson University) and Dr. Alan Kohn ( Professor Emeritus, University of Washington) are organizing an AMS-sponsored symposium on “Biogeography of the Pacific.” An impressive line-up of speakers has agreed to present their research on Pacific molluscan biogeography on June 29. This symposium and associated contributed paper and poster sessions are expected to be a memorable highlight of the meeting. • Dr. Jennifer Burnaford {Assistant Professor, Cal State Fullerton) is organizing a complementary special session on invasive molluscs. • Dr. Eric Gonzales ( Postdoctoral Researcher, UC Berkeley) will present a timely Student Workshop, “Genomic Tools for Molluscan Ecology and Evolution.” • Further details on the meeting website: http://www.malacological.org/meetings/ Other events include ever-popular auction, banquet, reprint sales, and other fun activities. Please plan to attend and spread the word. Contact me if you would like to ship donations for the auction. Help us to make this joint meeting a success by planning to attend, responding when the call for contributed talks and posters is announced, getting involved, and especially encouraging students and colleagues, including those from Latin American countries, to participate. If you have any questions, please feel free to contact the meeting organizers, myself (deernisse@fullerton.edu) or WSM President, Dr. George Kennedy (gkennedy@bfsa-ca.com). See you in San Diego! Voticf'EermA$&/ Professor of Biology, Cal State Fullerton 223 ' Junes. 26-30 Y 2010 ^ at San Diego State University m&fm «U.WBV w JtiSYM wesmem sow®'®; lfflKwar jBVJHI Featured Symposium on Pacific iicigeograpltf Special Session on Invasive Molluscs Student Workshop: “Genomic Tools for Molluscan Ecology and Evolution” iacologicai.org/meefings/ 224 Strategies for collecting land snails and their impact on conservation planning. MARLA L. COPPOLINO 97 Surfing snails: Population genetics of the land snail Ventridens ligera (Stylommatophora: Zonitidae) in the Potomac Gorge. COLLEEN S. SINCLAIR 105 Reproductive biology and the annual population cycle of Oxyloma retusum (Pulmonata: Succineidae). AYDIN ORSTAN 113 Independent Papers Distribution, density, and population dynamics of the Anthony Riversnail (Athearnia anthonyi) in Limestone Creek, Limestone County, Alabama. JEFFREY T. GARNER and THOMAS M. HAGGERTY 121 Epiphyton or macrophyte: Which primary producer attracts the snail Hebetancylus moricandii ROGER PAULO MORMUL, SIDINEI MAGELA THOMAZ, MARCIO JOSE DA SILVEIRA, and LILIANA RODRIGUES 127 Distribution and environmental influences on freshwater gastropods from lotic systems and springs in Pennsylvania, USA, with conservation recommendations. RYAN R. EVANS and SALLY J. RAY 135 Fish hosts of the Carolina heelsplitter ( Lasmigona decorata), a federally endangered freshwater mussel (Bivalvia: Unionidae). CHRIS B. EADS, ROBERT B. BRINGOLF, RENAE D. GREINER, ARTHUR E. BOGAN, and JAY F. LEVINE 151 Population studies of an endemic gastropod from waterfall environments. DIEGO E. GUTIERREZ GREGORIC, VERONICA NUNEZ, and ALEJANDRA RUMI 159 Subtropical sacoglossans in Okinawa — at “special risk” or “predictably rare”? CYNTHIA D. TROWBRIDGE, YAYOI M. HIRANO, YOSHIAKI J. HIRANO, KOSUKE SUDO, YOICHI SHIMADU, TOMOHIRO WATANABE, MAKIKO YORIFUJI, TARO MAEDA, YUKI ANETAI, and KANAKO KUMAGAI 167 Research Notes Self-adhesive wire markers for bivalve tag and recapture studies. LANCE W. RILEY, SHIRLEY M. BAKER, and EDWARD J. PHLIPS 183 Occurrence of the red abalone Haliotis rufescens in British Columbia, Canada. ALAN CAMPBELL, RUTH E. WITHLER, and K. JANINE SUPERNAULT 185 Index to Vol. 28 189 Society Business Seventy-five years of molluscs: A history of the American Malacological Society on the occasion of its 75th annual meeting PAULA M. MIKKELSEN 191 James W. Nybakken: September 16, 1936 - June 20, 2009 An Appreciation ALAN J. KOHN 215 Information for Contributors 2010 219 Membership Form 2010 221 Financial Report 2007 222 AMS/WSM Meeting Announcement 223