1VIBL/WH0I THE NAUTILUS Volume 110 1996-1997 AUTHOR INDEX Ahlstedt, S. a 97 Kkausk, M. K 31, 42 Arnold. VV. S 31. 42 Lavzek. J. B 102 Beis, 1 1 Lyons, W. G 31. 42 DEL Rio. C. J 77 Marelll D. C 31. 42 Dillon. R. T. ]R 97 Petuch. E. J 17, 122 E\'ERsoLE. A. G 107 Sabei.i.i. B 94 Graf, D. L 45 Sysoev, A. V. 22 Harasewtch. M. G 55 Tamam. M 94 HouART, R 12 Theler, J. L 94 Kaloyianni, M 1 Wise, J. B 76 Kirk, S. G 102 Xii\i\i,i, R 1 NEWTAXA PROPOSED IN N'OLUME 110 (199fi-!997) GASTROPODA Astriwa (Astraliiiin ) polkcnsis Petiicli, 1997, new .specie.s (Tnrbinidae) 125 Pijrazishms ki'itdrcui Petuch, 1997. new species (Potiimididae) 126 Ci'stiiinccrithiniu Pefiicli, 1997, new genus (Cerithiitlae) 126 Prismaccrithium Petuch, 1997, new genus (Geritliiidae) 127 Prismacerithium prisma Petuch, 1997, new species (Cerithiidae) 127 Pachijcroinmium dalli Petuch, 1997, new species (Naticidae) 127 Pachi/rrommiiim mansfieldi Petuch, 1997, new species (Naticidae) 127 Cahisacyprnca Petuch. 1997. new species (Cvpraeidae) 17 Cahisdciiprncn liriani Petuch. 1997. new species (Cvpraeidae) 18 Cnhisiuiipniea cliierri Petuch. 1997. new species (Cvpraeidae) 19 C.nlitsdnipraca sdraaotiwnsis Petuch. 1997. new species (C\praeidae) 19 Ciiprarorhis kenilmii Petucli. 1997. new species (Cvpraeidae) 129 Iphitus rohertsi .SabeHi & Taviani, 1997, new species (Epitoniidae) 94 Ergalatax obscura Houart. 1996. new species ( Muricidae ) 13 Spiniful', School of Science, Aristotle University of Tliessaloniki. Thessaloniki 54006, Greece ABSTRACT Anaerobic energy metabolism \\ as investigated in foot, mantle and hepatopancreas of the subtidal gastropod Hexaplex trim- ciilu-s (Linne, 1758) {=Murex tnineulu.s). C^hanges in the le\els of adein lates, substrates and end products were monitored over a time course of 16 hours of anoxia. It is suggested that unlike intertidal species, the succinate pathway is a metabolic route of minor importance during anoxia in subtidal gastropods. The existence of an alternative electron acceptor, apart from Oj and fumarate, in the tissues of the subtidal gastropods under anaerobic conditions is discussed. Key M'ords: metabolism, anaerobiosis, subtidal, gastropods, foot, mantle, hepatopancreas, Hexaplex trunculus. INTRODUCTION It is generally accepted tfiat within the animal kingdom the ability to sustain periods of oxygen deprivation varies among the different phyla. MoUusks and particularly bivalves present well developed capacities for the sur- vival of environmental anoxia and for this reason have been extensively studied in terms of their anaerobic en- ergy metabolism (de Zvvaan, 19 « o E :i w ra ti. o c ra a o +j n a X 300 Duration ofanaerobiosis (h) Figs 1-3. Changes in the concentrations of aspartate, glycogen and glucose in the hepatopancreas of Hexaplex tninculus dur- ing various periods of anaerobiosis. 1) aspartate, 2) glycogen, ■3) glucose. * Significantly different from the corresponding aerobic condition (t-test P<0.05). ** Significantly different from the corresponding aerobic condition (t-test P<0.00.5) ing the use of the succinate pathway by Htxuplcx tnin- culus tissues during anaerobiosis. MATERIALS AND METHODS Animals and Chemicals: Specimens of Hexaplex trun- culus were obtained from the shores of Thessaloniki through a local fisherman. They were kept in circulating seawater at about 2()°C and were used two to three days after arrival. Aerobic animals were sampled directly from the seawater tank. To impose anoxia the animals were placed in a vessel of seawater with an air tight cover, containig 1 L of seawater that had been previously de- o.xygeiiated by bubbling a steady stream of nitrogen gas through it for 1 h to give zero oxygen tension as moni- tored w ith an ox\ gen electrode (Digital Oxygen System, model 10, Rank Brothers Manufacture). Animals were kept under anoxic conditions for 2, 4, 8, 12 and 16 h at 20°C. At the end of the anoxic incubation the shells were quickly opened and the tissues were rapidly excised, blotted and immediately frozen in liquid nitrogen. Tis- sues were stored at -80°C until assayed. Substrates, enzymes and coenzymes were purchased from Sigma Chemicals Co. (St Louis, L^SA). All other chemicals were purchased from Serva (Heidelberg, Ger- many). Preparations of Tissue Extracts: Samples of frozen tissue were ground to a powder under liquid nitrogen using a mortar and a pestle. The powdered tissue was extracted by adding three volumes of cold perchloric acid (HCIO4, 10% w/v). The precipitated protein was removed b\ centrifugation at 4000 x g lor 10 minutes and the su- pernatants were neutralized with 3M potassium bicar- bonate (KHC03).The precipitated potassium perchlorate was removed by centrifugation as above and the super- natants were taken for determination of metabolite con- centrations. Determination of Metabolites: Glycogen was deter- mined by the method of Keppler and Decker (1974), aspartate by the method of Bergmeyer et al. (1974a) and glucose by the method of Bergmeyer et al. (1974b). Al- anine was assayed according to Williamson D.H.(1974) and succinate according to Williamson JR. (1974). L-Lactate was measured according to Gutmann and Wahlefeld (1974) and octopine according to Storey et al. (1979). ATP was quantified bv the method of Lam- precht and Trautschold (1974), ADP and AMP by the method of Jaworek et al. (1974). Statistics: Results are presented as means ± SEM (n=5). Tests for significant differences between groups used the Student's t-test. RESULTS Hepatopancreas: Figures 1-3 show the changes in as- partate, gl\ cogen and glucose content of the hepatopan- creas of Hexaplex trunculus during 16h of anaerobiosis. Aspartate content decreased throughout all the periods of anoxia tested. Aspartate levels measured in the tissue were lowest alter 12h of anaerol)iosis. (iKcogen and glu- cose levels tended to increase throughout anaerobiosis. (ilucose concentration increased 10-told during 16h of anoxia. Figures 4-7 illustrate the time course of the changes R. Xomali et al., 1996 Page 3 0 501 (A (A re E 0) "35 a» o E It X c o o CO Duration of anaerobiosis (h) Figs 4-7. Changes in the concentrations of lactate, octopine, alannie and succinate in the hepatopancreas of Hexaplex trunculiis during various periods of anaerobiosis. 4) lactate, 5) octopine, 6) alanine, 7) succinate. * SignificantK different from the corresponding aerobic condition (t-test P<0.05). ** Significantly different from the corresponding aerobic condition (t-test P<0.00.5) in levels of potential end products of Hexaplex trunculus hepatopancreas during anoxia. Alanine accumulated during the first hours of anoxia. Thereafter its levels started to fall, but remained significantK increased com- pared to the aerobic state. Succinate accumulated after the 4th hour of anoxia. The maximum alanine accu- mulation (2.9 ^mol/g wet mass) observed during the first two hours of anoxia exceeded that of succinate (0.8 ^mol/ g wet mass) observed after 16 h of anaerobiosis. Octopine accumulated significantly after 12h of anoxia, while no significant lactate accumulation was observed through- out the 16h anaerobiosis. From the data shown in figures 8-11 a sharp decline in ATP content can be detected, while levels of ADP and AMP are not significantly changed. The energy charge shifts from 0.61 to 0.49 after 4h of anoxia. There- after, it slightly increases to 0.5.3 after 16h of anoxia. Foot Muscle: Figures 12-14 show changes in concentra- tions of aspartate, glycogen and glucose in the foot muscle of Hexaplex trunculus during 16h of anoxia. As in the hepatopancreas, aspartate levels had fallen significantly by the 4th hour of anoxia. At that time its levels are reduced by 5.5 ^mol/g wet mass. Thereafter, its con- centration remained almost constant. Contrary to the results in hepatopancreas, glycogen levels had fallen by the 4th hour of anoxia. After 8h of anaerobiosis a sharp reduction in glycogen content was observed (44.2 nvao\/ g wet mass). Glucose levels were significantly increased after 8 hours, after 12h of anaerobiosis its concentration was almost double that of the aerobic state. Changes in the concentrations of lactate, octopine, al- anine and succinate during 16h of anoxia in the foot muscle of Hexaplex trunculus are shown in figures 15- 18. Onh alanine levels ro.se significantly during anoxia. The maximum alanine accumulation was observed after 12h of anoxia and was about 3.7 ixmo\/g wet mass. No significant changes in octopine. lactate and succinate lev- els were obser\ed. In figures 19-22 the changes in adenylates and in the energy charge in the foot muscle during 16h of anoxia of Hexaplex trunculus are presented. A significant de- Page 4 THE NAUTILUS, Vol. 110, No. 1 CO (/) ro E ^^ O) To o E re OJ u. O c re a o ♦^ re a X 0 081 0 04 000 1 On S 05 < 00 1 0 o uj 0 5 12 16 11 K- 0 OH ' r 12 16 -1 ' 1 ' 1 — 8 12 16 Duration of anaerobiosis (h) Figs 8-11. Changes in the concentrations of ATP, ADP, .■\MP and energy charge in the hepatopancreas of Hcxaplex trunculus (hiring various periods of anaerobiosis. S) ATP, 9) ADP, 10) AMP, 1 1 ) energy charge. * Significantly different from the corresponding aeroljic condition (t-test P<0.0.5). ** Significantly different from the corresponding aerobic condition {t-test P<0.005) dine in ATP content together with a signiBcant increase in ADP level were observed after 4h of ano.xia. AMP levels remained almost unaltered. A decline in energy charge was also evident. Energy charge shifted from 0.87 to 0.72 during the first four hours of ano.xia. Thereafter, it remained almost constant. iMantlc: Figures 23-25 show changes in the concentration of aspartate, glycogen and glucose in the mantle of Hex- aplex trunctiluH during various [periods of anaerobiosis. Among the substrates measured only aspartate showed a significant decline, being reduced to 3.4 ^imo]/ g wet mass during the 8 first hours of anoxia. Glycogen levels remained almost constant throughout anoxia. Glucose concentration in the mantle increased and showed a sig- nificant rise after 12 and 16h of anoxia exposure, com- pared to the aerobic state. Among the metabolites presented in figures 26-29, a significant rise in lactate content was observed after 8h of an(jxia. During the first four hours alanine levels de- creased. Thereafter an apparent increase in alanine con- tent v\as observed. This increase was not significantly different from the value of the aerobic control, owing to high variation in the alanine content of different animals. There were no significant differences in octopine and succinate content among the difierent periods tested. Concerning the changes of the adenylates shown in figures 30-33, a significant decrease in ATP content was seen until the fourth hour of anoxia. Levels of ADP increasexl during the first eight hours, but declined there- after again, reaching control values. No significant change in AMP concentration was observed. Energy charge de- clined, mainly during the first eight hours, decreasing from 0.9 to 0.73. DISCUSSION The ability of in\ertel)rate anaerobes to sustain long pe- riods of anoxia is niainlv based on the metabolic adap- tations of their tissues. These species are able to reduce their metabolic rate and at the same time to maintain R. Xomali et al., 1996 Page 5 their energy reserves so as to confront the energetic needs of their tisssues. The ATP production is generally based on the anaerobic breakdown of glycogen. In contrast to mammalian systems, this anaerobic breakdown of gly- cogen does not lead to lactate production. Various end products accumulate during anaerobiosis depending on species, as well as on the duration of anaerobiosis. In intertidal bivalve mollusks, the main end product of the first stages of anaerobiosis is alanine, while at later stages succinate, propionate, acetate and fatty acids can be de- tected (de Zwaan, 1983). Hochachka and Mustafa (1972) proposed that amino acids should be an important an- aerobic fuel in intertidal bivalve tissues and they put forward a metabolic map integrating the metabolism of glucose, aspartate and glutamate during ano.xia. The metabolic behaviour of subtidal species during anaero- biosis is almost unknown. The results of the present study revealed that foot muscle, mantle and hepatopancreas of Hexaplex trun- ciilits catabolize aspartate and accumulate alanine in re- sponse to 16h of ano.xia. The pattern of metabolism differs among tissues. In the hepatopancreas, aspartate declines during the earlv stages of anaerobiosis and is accompa- nied b\ a simultaneous rise in alanine content (figs, 1 and 6). After this initial response, aspartate declines more slowly, while alanine accumulates up to the 16th hour of anaerobiosis, compared to the aerobic state. Gluta- mate, a possible end product of anaerobiosis in intertidal species, is not accumulated in the hepatopancreas of Hex- aplex trtinculus during anoxia (results not shown). A significant rise in glucose content was observed, starting from the very beginning of oxygen deprivation and rose by a factor of ten till the 16th hour of anaerobiosis (fig. 3). It could be suggested that Hexaplex trtinculus he- patopancreas responds rapidly to anoxia b\ catabolising mainh aspartate and accumulating initialK alanine and subsequently succinate (fig. 1,6,7). Elevation of tissue glu- cose could not be attributed to glycogen degradation since glycogen levels showed no signs of decreasing throughout anoxia. Similarly no significant decrease in glycogen content was found in hepatopancreas, foot, and phasic adductor muscle of the cherrystone clam Mer- cenaria mereenaria (Korycan & Storey, 1983), No change in the percentage of phosphorvlase in the active a form was found in the adductor muscle of Mytilus edulis over a time course of up to 18h of anoxia (Ebberink & Sali- mans, 1982). In the foot muscle of Hexaplex Iruneulus, the response to anoxic stress is less rapid than in the hepatopancreas. This agrees with the findings of Whitman and Storey (1989), who suggested that the processes involved in an- oxia induced modification of givcolv tic enzymes are more rapidh initiated in non muscular than muscular tissues. A sharp decline in aspartate content is obvious after four hours of anoxia, while alanine accumulation is maximal after twelve hours of anoxia (fig. 12 and 17). The low levels of succinate did not change throughout anoxia (fig. 18). This could either suggest that the succinate pathway is not functioning in the foot muscle of Hexaplex trnn- nj E *-* 0) « O E o O o 100 Duralion of anaerobiosis (h) Figs 12-14. Changes in the concentrations of aspartate, gly- cogen and glucose in the foot muscle of Hexaplex trunculus during various periods of anaerobiosis. 12) aspartate, 13) gi\- cogen, 14) glucose. * SigiiificaiitK different from the corre- sponding aerobic condition (t-test P<0.05) cuius, or that succinate is rapidly metabolized to another end product (i.e. propionate or acetate). Similarly, it was reported that in the subtidal marine gastropod Nassarius reticulatu.s only alanine and no succinate accumulated throughout anoxia (Wieser, f980). The Mediterranean bivalve Scapharca cf. cornea Reeve, 1843 also produced no succinate during 72h of anoxia (Carpene et al., 1981). However, in the foot muscle of the subtidal whelk Nassa Page 6 THE NAUTILUS, Vol. 110, No. 1 (/) (/) (Q E 0) o o E zi oT o (/) 3 O o 0 10 w c o 005 u 3 CO 0 00 18 — r- 4 — r- 8 — I — 12 — I — 16 Duration of anaerobiosis (h) Figs 15-18. Changes in the concentrations of lactate, octopine, alanine and succinate in the foot muscle of Hexaplex trunculus during various periods of anaerobiosis. 15) lactate, 16) octopine, 17) alanine, 18) succinate. * SignificantK different from the corresponding aerobic condition (t-test P<0.05). mutalnlis succinate was produced, but only in low con- centrations (Gade et ai, 1984). In contrast to hepato- pancreas, glycogen content falls sharply after four hours of anoxia (fig. 13). The observed glycogen degradation in the foot muscle could be correlated with the contin- uous effort of the animal to move away from areas de- prived of oxygen. The elevated glucose levels, probably derived from glycogen catabolism, could be the main metabolic substrate used to support muscular work under anaerobic conditions. No lactate and octopine accumu- lation was found {fig. 15, 16). Our results concur with the data presented by Kluytmans et ai (1983) and Gade et al. (1984). It seems, therefore, that the foot muscle of Hexaplex truncitlus shows no signs of energ\' stress dur- ing the first two hours of anoxia. Later, it responds to anoxia by catabolizing aspartate and glycogen and ac- cumulating mainly alanine. Due to the declining levels of aspartate in the mantle during anaerobiosis, aspartate appears to be the main source of energy production during anoxia. Alanine ac- cumulation presented a somewhat different pattern of accumulation compared to the two other tissues exam- ined (fig. 28). There was an initial decline in its level, followed by a subsequent increase up to the 16th hour of anoxic stress. However, owing to great variation in mantle alanine content among different individuals, the values did not differ from the control. Similarly, alanine did not accumulate in the mantle tissue of Mytilus ediilis at reduced oxygen tensions, but it accumulated in the posterior adductor muscle under the same conditions (Livingstone & Ba\iie, 1976). Glucose levels increased during anoxia in all three tissues examined. Glucose levels in the hepatopancreas rose from the very begining of anoxia, reaching its max- imum value after 16h of oxvgen deprivation where a tenfold increase in its content could be observed. How- ever, glucose increases in loot muscle only after 8h of anoxia and in mantle ti.ssue after 12h of anoxia. This delayed increase could be correlated w ith the time need- ed for glucose residues to be mobilized from the hepa- topancreas and transferred to the foot muscle and the mantle via the hemolv mph. While glucose levels in the R. Xomali et al., 1996 Page 7 w 0) (0 E o W 0) o E zi. o" u er, H. U., E Bernt, H Miillering and G Phleiderer. 1974a. L-aspartate. In: Bergine>er, H. L'.(ed.). Methods of enz\matic anahsis .\cademic Press, New York, 4:1696- 1700. ' Bergmeyer, H. I , E. Bernt, F, Sclimidt and H Stork 1974b, D-glucose, /;i. Bergmeyer, H. U.(ed.). Methods of enzy- matic analysis, .Academic Press, New York. 3:1196-1201. Carpene, E., W. Cortesi and VV. Zurburg. 1981. Anaerobic energy metabolism in the bivalves Scap/iarca inaeqtiivalvis (Bruguiere) and \'(';h« gallina L. .•Abstracts of the Third ESC^PB Congress. Pergamon Press, Oxford, pp. 118-119. de Zv\aan, A, 1983. Carbonh\drate catabolism in bivalves. In: Hochachka, P. W (ed ). The Moliu.sca, .Academic Press, New York. 1:137-175. Ebberink, R. H. M. and M. Salimans. 1982. Control of gly- cogen phosphorylase activit\ in the posterior adductor muscle of the .sea nuissel Mytilus cclulis. Journal of Com- parative Physiolog) 148:27-33. Eberlee. J. C. and J .M Storey. 1988. Tissue-specific bio- chemical responses tluring anoxia and reco\er\ in the channelled whelk. Journal ol Experimental Marine Biol- og\ and Ecology 121:l(i5-171 Eberlee, J. C, J. M. Storey and K. B. Store). 1983. .Anaero- biosis, recovery from anoxia, and the role of strombine and alanopine in the oyster Crassustrea virginica. Cana- dian Journal of Zoology 61: 2682-2687, Glide, G, 1983, Energ\ metalxilism (}f arthropods and mol- lusks during en\ionniental and lunctioiial anaeroljiosis Journal oi Experimental Zoolog\ 228415-429 Gade, G.. K II Carlsson ami (.', Mcinardus, 1984 Energy metabniisni ui the loot of the marine gastropod \assa R. Xomali et al., 1996 Page 9 V> (0 E w « o E o c (0 0) o 0 75 0 50 025 0 00 0 50n 01 c ■q. o o O 0 25 27 0 00 Duration of anaerobiosis (h) Figs 26-29. Changes in the concentrations of lactate, octopine, alanine and succinate in the mantle of Hcxaplex ininculus during various periods of anaerobiosis. 26) lactate, 27) octopine, 2S) alanine, 29) succinate. * Significantly different from the corresponding aerobic condition (t-test P<0.05). ** Significantly different from the corresponding aerobic condition (t-test P<0.05). nmtabilis during environmental and functional anaero- biosis. Marine Biolog> 80:49-56. Gutmann, I. and W. Wahlefeld. 1974. L- Lactate. In: Berg- meyer, H. U.(ed.). Methods of enzymatic analysis. .Aca- demic Press, New York. .3:1464-1468. Henmiinga, M. A., J. J. Maaskant, \\ Koomen and J Joosse. 1985a. Neuroendocrine control of gKcogen mobilization in the freshwater snail Lymnea stagnalis. General Com- parative Endocrinology 57:117-123. Hemminga, M. A., J. J. Maaskant, J. van der Plas and P.A. Gabbott. 1985b. The hyperglycemic factor of the CNS of the freshwater snail Lymnea stagnalis: interaction with glucose stimulation of glycogen s> nthesis and evidence for its release during anaerobiosis. General Comparative En- docrinology 59:301-307. Hochachka. P. W and T. Mustafa 1972 Invertebrate Fac- ultative .Anaerobiosis. Science 178:1056-1060. Holwerda DA. and A. de Zwaan. 1979. Fumarate reductase of Mijtilus edulis L. Marine Biological Letters 1:33-40 Holwerda, D. A. and A. de Zwaan. 1980. On the role of fumarate reductase in anaerobic carbohydrate catabolism of Mytilus ednlis L. Comparative Biochemistrv and Phys- iology 67B:447-453. Jaworek, D., \V. Gruber and H U. Bergmeyer. 1974. Aden- osine-5-diphosphate and Adenosine-5-monophosphate. In: Bergmeyer, H. U.(ed.). Methods of enzymatic analysis. Academic Press, New York. 4:2127-213l' Keppler, D. and K Decker 1974. Glycogen. In: Bergmeyer, H. U.(ed.). Methods of enzvmatic anaKsis. .Academic Press, New York. 3:1127-1131. Kluytmans, J. H., A. M. T. de Bont, J. Janus and T. C. M. Wijsman. 1977. Time dependent changes and tissue specificities in the accumulation of anerobic fermentation products in the sea mussel Mytilus edulis. Comparative Biochemistry and Physiolog) 588:81-87. Kluytmans, J H.] A. M. T. de Bont, E. C. J. Kruitwagen, H J. L. Ravenstein and P. R. V'eenhof. 1983. Anaerobic ca- pacities and anaerobic energy production of some medi- terranean bivalves. Comparati\e Biochemistrv and Phys- iology. 75B:171-179. Kohler, P 1976. Hydrogen transport in the muscle mito- chondria of Ascaris suum. In: Bossche, H. V. D. (ed.). Biochemistry of Parasites and Host-Parasite Relationships. North-Holland, Amsterdam, pp. 125-132. Kohler, P. 1980. The function of mitochondrial enzsmes in parasitic helminths. /;i: \itale, L. and \'. Simeon (eds). Page 10 THE NAUTILUS, Vol. 110, No. 1 (0 re E ■«-> o E zl c re S 0,030 0015 0000 u uJ 05- 00 12 16 -I— 8 12 16 Duration of anaerobiosis (h) Figs 30-33. Changes in the concentrations of ATF, .ADP, AMP anil energy charge in the foot muscle of Hcxaplcx tninculus during various periods of anaerobiosis. 30) ATP, 31) ADP, 32) AMP, .33) energy charge. * SignificantK- different from the corresponding aerobic condition (t-test P<0.05). Proceedings of the FEBS Special Meeting on Enzymes: Industrial and Clinical Enzvmologv. Pergamon Press, Ox- ford. 61:24.3-256. Korycan, S. A. and K. B Storey, 1983, Organ specific me- tabolism during ano.xia and recovery from anoxia in the cherrystone clam, Mcrcenaria mercenaria. Canadian journal of Zoology 61: 267-4-2681, Lamprechi, VV. and 1. Trautschold. 1974. Adenosine-5-tri- phosphate. In: Bergmeyer, H. U.(ed.). Methods of enzy- matic analysis. Academic Press, New York. 4:2101-2110. Livingstone, D R. and B. L. Bayne. 1976. Responses of My- lilus cdnlis L to low oxygen tension: Anaerobic metabolism of the posterior adductor mu.scle and mantle tissues Jour- nal of Comparative Physiology 114:143-15.5 Meinardus, G. and G. V.Mv. 1981. Anaerobic luelabolisni ol the common cockle Cardium edule — IV. Time dependent changes of metabolites in the foot and gill ti.ssue induced by anoxia and electrical stimulation Comparative Bio- chemistry and Physiology 70B: 271-277, Michaelidis, B and I. Beis, 1990 Studies on the anaerobic energy metabolism in the loot muscle of marine gastropod Patella cacrulea L. Coniparati\'e Biochemistr\ and Phvs- iology 95B:493-500. Storey, K. B. 1988. Mechanisms of glycolytic control during facultati\e anaerobiosis in a marine mollusc: tissue-specific analysis of glycogen phosphorylase and fructose- 2,6-bis- phosphate, Canadian Journal of Zoolog) 66:1767-1771. Store) , K B,, J. M. Storey, J. Kjell and P. \V. Hochachka. 1979, Octopine metabolism in Sepia officinalis, effect of h\ poxia and metabolite loads on the blood levels of octopine and related compounds C'anadian |ournal of Zoologv 57:2331- 2336 Storey, K, B,, D, A, Kelly, J. A. Duncan and J. M Storey 1990, ,\naerobiosis and organ specific regulation of glycoKsis in a rji.ii iiir u liclk C>anadian Journal of Zoology 68:974-980. \\ luljiian, K, E and K. B, Storey, 1989. Organ-specific reg- ulation of phosphofructokinase during facultative anaero- biosis in tile marine whelk Busycotijpu.s canaliciilatiiin. (Canadian Journal of Zoolog\ 6970-75. \\ ieser, VV. 1980. Metabolic end products in three species of marine gastropoils Journal oi Marine 15iological Associa- liciii nl the I riUril Kuii;d(.iii 60 175-1.80, R. Xomali et al., 1996 Page 11 Wijman, T. C. M , J J Maaskant, P Balm and J FClijnstra 1988. H\ pergKcaeniia in relation to anoxia and stress in the freshwater snail Lynuiat'a stagnalis. 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THE NAUTILUS 110(1):12-16, 1996 Page 12 On the Identity of Morula martensi Dall, 1923 and Description of a New Species of Ergalatax from the Red Sea (Gastropoda: Muricidae: Ergalataxinae) Roland Houart Research associate Institut Royal des Sciences Natiirelles de Belgique Departement des Invertebres Recents Rue \'autier, 29 B-1000 Brussels, Belgium ABSTRACT Morula martensi Dall, 1923 differs from the similar species Ergalatax margariticula (Broderip, 1833) by having a smaller, narrower and white shell The species erroneously identified as Ergalatax or Cronia martensi (Dall, 1923) in recent literature is named here as Ergalatax obscura n.sp. Key Words. Gastropoda, Muricidae, Morula martensi, new species INTRODUCTION Martens (1874:95, pi. 5, fig. 49) identified and illustrated a shell from the Hausknecht collection as Purpura (Ri- cinula) siderea Reeve, 1846. Examination of the type materia! of Ricinula siderea (3 syntypes BMNH 1968478, type locality: Philippines, e.\ coll. H. Cuming), revealed that it is a buccinid, and not the species illustrated by Martens, which Dall (1923:304) subsequently named Morula martensi. The seven specimens examined by Martens, all from the Persian CJulf, are stored in the Museum fiir Xaturkunde, Berlin. Martens' original figure is here reproduced (fig. 2), and the actual specimen fig- ured (figs. 12, 13). Martens' description, figure, and comparisons with Purpura iindata (Chemnitz, 1795:pl. 192, fig. 1852) [ = Ergalatax margariticola (Broderip, 1833)], my re-e.\- amination of the specimens of the Hausknecht collection and comparison with specimens from Qatar (coll. R. Houart), reveal that this species is similar to E. margari- ticola. The shells from Oman and the Persian Gulf are white with black blotches on the spiral cords (figs. 12- 15), and differ from E. margariticula. which has a broad- er, more darkly pigmented shell. Specimens from Oman were illustrated by Boscii and Bosch (1982:95, 1989:60) as Cronia konkanensis (Melvill, 1893), which, however, is probably a distinct species, more closely related to Thais muricina (Blainville, 1832) and T. marginatra (Blainviile, 1832), both classified in Rapaninae. II the populations from the Persian Gulf prove to be only an ecological form of £. margariticola. then E. martensi (Dall, 1923) will become a junior synonym of tliat species. Ergalatax n^artensi was reported to be a s\non\ m of E. contracta (Reeve, 1846) by Houart (1995:251). It is here considered a distinct species. Three distinct species with similar gross morpholog\ have been confused in the recent literature. Ergalatax martensi (Dall, 1923), from Oman and the Persian Gulf, is closely related to but probably distinct from E. mar- gariticola (Broderip, 1833); Ergalatax obscura, new spe- cies, from the Gulf of Aden, the Red Sea, the Gulf of .Acjaba and the Mediterranean coast of Turke\ , is a dis- tinct species identified incorrectly as Cronia (or Erga- latax) martensi (Dall) in recent literature; and Thais konkanensis (Melvill, 1893), from the Indo-West Pacific, which is closely related to T. muricina (Blainville, 1832) and T. marginata (Blainville, 1832). My researchs were incomplete when I received pho- tographs and material for identification from Turkey, from G. Buzurro and VV. Engl. .\t the time, I provision- ally identified the new Mediterranean immigrant as Er- galatax martensi, and it was reported as such by Buzurro, Engl and Tiimtiirk (1995) and by Engl (1995). It is now- evident that E. martensi is confined to Oman and the Persian Gull, and tiiat the Turkish species is £. obscura n.sp. SYSTEMATICS Order Neogastro()oda Thiele, 1929 SuperfamiK Muricoidea Rafinesque, 1815 Family Muricidae Rafinesque, 1815 SubfamiK Ergalataxinae Kuroda & Habe, 1971 (Jenus Ergalatax Iredale, 1931 Ergalatax Iredale, 1931:231. T\pe species (by original R. Houart, 1996 Page 13 designation): Ergalatax rcciinens Iredale, 1931; oft Syd- ne\' Harbour, New South Wales, Australia. Ergalatax martensi (Dall, 1923) (figs. 2, 12-15) Morula nujrtcnsi Dall, 1923:304, new name for Morula aiderea von Martens,1874, not Reeve, 1846. Cronia konkanauis—V.O'iCH & BOSCH, 1982:95, te.xt fig.; SMYTHE, 1982:60, pi 1, fig i: B0SC:H & BOSCH, 1989: 60, text fig. (not Ricinula hjiikanensis Melvill, 1893). NOT Cronia mar(e;!si— SHARABATI, 1984:pl. 19, fig, 9; SINGER & MIENIS, 1991b:58, fig. 19 (in part); COU- LOMBEL, 1994:73, text figs {==Ergalatax olxcura n.sp). NOT Ergalatax martensi BUZl'RRO, ENGL & TiiMTiiRK, 1995: 17, text fig,; ENGL,1995: 46, fig. 10 (= Ergalatax obscura n.sp). Description: Shell up to 26.5 mm in length at maturity, slender, squamous. Spire high with 6 weakly convex, shouldered teleoconch whorls. Suture adpressed. Proto- conch eroded in all specimens examined. Axial sculpture of teleoconch whorls consisting of high, strong, nodose ribs, producing blunt knobs at intersection with spiral cords. 9 or 10 ribs on first and second whorls, 8-10 on third, 9-12 on fourth and fifth, 7-10 on last whorl. Ribs more strongK developed at shoulder. Other axial sculp- ture of numerous growth lamellae. Spiral sculpture of low, squamous cords of various strength, lisually 13 or 14 cords on last whorl, and 6 additional, smaller cords on shoulder. Aperture narrow, ovate. Columellar lip smooth, occasionalK with 1 or 3 weak folds abapicalK . Lip adherent. Anal notch narrow, deep. Outer lip cren- ulate, with 6-8 weak, elongate denticles within. Siphonal canal short, broad, open. White, with primary spiral cords, or knobs only, topped with dark brown or black. Aperture glossv white. Operculum dark brown with subterminal nucleus, l^adula unknow n. Type material: Lectotxpe (here selected), the specimen figured by Martens (1874) as Purpura (Ricinula) siderea, not Reeve, 1846, and 6 paralectotypes ZMB 21596. Type locality: Persian Gulf, here designated. Other material examined: QATAR: Doha, landing stage, coll. R. Houart (10). Distribution: Oman and the Persian Gulf. Remarks: .\s stated above, the shell is close to Ergalatax margariticola (Broderip, 1833), but differs in being usu- ally smaller relative to the number of teleoconch whorls, narrower, and in having a white ground color and a white, glossy aperture, instead of cream or light brown in E. margariticola. Ergalatax obscura n.sp. (Figs 1, 3-8) Cronia mar/fri.si— SHARABATI. 1984; pi 19, fig. 9, 9a, 9b; 49 Figures 1.2. 1. Protoconch of Ergalatax obscura n.sp. (scale bar: 0.5 mm), 2. Fig. 49 of Martens (1874), as Purpura siderea (Reeve). SINGER & MIENIS, 1991b: 58, fig. 19; COULOMBEL, 1994: 73, text figs) (not Morula martensi Dall, 1923). Drupella rugosa SINGER & MIENIS, 1991a: 18, fig. 6 (not Murex rugosus Born, 1778), Cronia cf konkanensis—GlVfpe (4. 5) and paralectot\ pe (9. 10). 6. Cymnobela (Theta) chrysopelex. holot>pe. 7. 8. Typhlonmngelia polythele, lectotype (7) and paralectotype (8). Four weak, narrow, widely spaced cords between sub- sutural cord and row of peripheral gemmules, 3 distinct cords below gemmuiate periphery on last whorls. Shell base and canal covered by 17 narrow, widely spaced cords, which gradually weaken towards the canal end. Peripheral gemmules on spire whorls mostly eroded, but preserved ones strong and acute, becoming progressively weaker, longitudinally elongate, arcuate towards the body whorl. They are numerous, irregularly placed on last half of the body whorl, almost obsolete near aperture. Pen- ultimate whorl bears 20 gemmules, the body about 33. Paralectotypes have less angulate whorls with less prominent peripheral gemmules. Subsutural ramp may be concave, especially on last half of the body whorl. Page 24 THE NAUTILUS, Vol. 110, No. 1 Siphonal canal varies in length and nia\ be almost straight or turned to left, as in iectot\ pe. The pattern of sculpture is not similar to that shown in Barnard s drawing (Bar- nard, 1963:£ig. 2a), i.e., with very strong and closely set bilobed tubercles forming the subsutural cord. Remarks: This species was also recorded in the area from 33°26'S to 34°37'S and from 16°23.5' E to 17°03'E, at 2378-3036 m (Barnard, 1963). The shells found in the John Murray E.xpedition, sta. 118, off Zanzibar, 1789 m, and identified as G. teschi (Powell, 1964) (Sysoev, in press, figs. 68-70), are G. lobatopsis. Family CONIDAE Fleming, 1822 Subfamily CLATHURELLINAE H. & A. Adams, 1858 Typhlomangelia polijthele Barnard, 1963 (figures 7-8) Typhlomangelia (?) polylhele Barnard, 196.'3:422, fig. .Se, i. Material examined: Two s\ ntypes (SAM A9802). Type locality: "Africana 11", sta. A317, 33°50'S, 16''30'E, 1480-1660 fms (2707-3036 m). Description: Largest (12.3 x 5.4 mm) and better pre- served of two syntypes here designated as lectotype. Its shell rather thick, consisting of 5,5 remaining whorls. Protoconch missing, first teleoconch whorl eroded. Whorls angulate at periphery. Subsutural ramp slightly concave, with narrow subsutural cord and numerous, regular, nar- row, arcuate folds formed by thickened growth lines. Axial sculpture represented by numerous, very short ribs below subsutural ramp. Ribs rapidly weaken, do not reach lower suture or peripherv of body whorl. Ribs number about 22 on penultimate whorl (its dorsum broken, ap- parently during extraction of soft body) and 26 on body. Spiral cords (7 on penultimate whorl and 32 on body whorl) are narrow, separated by wide interspaces (2-4 times wider than cords). Spiral cords are more closely set on siphonal canal. Canal short, straight. Svmmetrical anal sinus occupies entire subsutural ramp, being deepest in its middle. Paralectotype (11.5 x 5. l mm at 5+ pre- served whorls) has the same conchological characters. Remarks: Barnard provisionalK placed this species in Typhlomangelia. \\ hich seems to be appropriate for the species, despite its small and broad shell lacking spiral sculpture on the subsutural ramp. Typhlomangelia ad- enica Sysoev, a very similar species both conchologically and in radular characters, is known from the Gulf of Aden, John Murray Expedition sta. 26 and 185, 200- 2312 m (Svsoev, in press). Abyssothauma Sysoev, 1996, new genus Type species: Moniliopsis psilarosis Barnard, 1963. Diagnosis: Shell large, to 60 mm, lusiiorm, w ith rounded and vseakly convex whorls, covered b\ thick olivaceous periostracum. Spire more than 30% of shell height. Earlv' v\li()rLs wcaklv angulate. Axial sculpture of numerous oblique narrow folds below shoulder in early whorls, or absent. Spiral sculpture of low , rather wide cords belov\ subsutural ramp. Shell base weakly convex, not diff^er- entiated from canal. Aperture narrowly oval, siphonal canal wide, long. Anal sinus moderately deep, rounded, occupv ing the subsutural ramp, with deepest point in its middle. Operculum large, leaf-shaped, with terminal nu- cleus. Radular teeth hollow, long, slightK curved, not barbed, with rounded base and terminal opening of tooth cavity. Remarks: The new genus resembles the genera Typhlo- syrinx Thiele, 1925 and Belaturricula Powell, 1951. From the former genus it differs in having a stronger shell with better developed spiral sculpture, thick dark-colored periostracum, different form of axial sculpture on early whorls, and presence of operculum. This new genus dif- fers from Belaturricula in having a narrower shell with stronger sculpture, pronounced siphonal canal and deep anal sinus. The new genus includes the type-species and a new species from the Tasman Sea, 3610-3688 m that will be described elsewhere. Pontiothauma ergata Hed- ley, 1916, resembles A. psilarosis in general shell outline, predominance of spiral sculpture, structure of radular teeth (see Egorova, 1982, figs. 63a. b), presence of oper- culum with terminal nucleus, but differs in having a shallow anal sinus (holotype illustrated b\ Dell, 1990: figs. 418-419) and very thin light-colored periostracum. The taxonomic position of P. ergata is uncertain, but it obviously does not belong to Pontiothauma, differing in the shell form, sculpture, the presence of operculum and different radular teeth indicating the placement in the Clathurellinae, not Raphitominae. Probably P. ergata belongs to a separate genus, closely related to Abysso- thauma. Abyssothauma psilarosis (Barnard, 19(i3) (figures 4-5, 9-10) Moniliopsis psilarosis Barnard, 1963:421-422, fig. 4a. IVIaterial examined: Two specimens (SAM A9789) in- dicated by Barnard (1963) as "Types". Type locality: Barnard (1963) did not designate the type localitv nor indicate which of his two samples contained 'tvpes '. However, judging from the indicated number and cjualitv of specimens from '.\fricana 11 sta. A316, 34°42'S, 16°54'E, 1725-1780 fms (3155-3255 m), cor- responding to specimens from the type series, the t\pe localitv is restricted to these coordinates. Description: Largest of two tvpe specimens is here des- ignated as lectotvpe. Dimensions: 47.2 x 16.0 at 6.5 preserved whorls (lectotype) and 40.8 x 14.4 at 6 pre- served whorls (paralectotype). Lectotype shell strong, solid, covered by thick dark-olivaceous periostracum. Protoconch and at least one teleoconch whorl missing. Upper whorls distinctiv angulate above peripherv, an- gulation weakening in subsequent whorls, becoming ob- -solete in penultimate whorl. Subsutural ramp weakly A. V. Sysoev, 1996 Page 25 concave. Upper whorls with strong, obhque axial ribs below subsutural ramp U8 ribs on first well-preserved whorl). Ribs form small knobs in upper part. Ribs mark- edly weaken towards lower suture, becoming shorter in successive whorls. Last 3.5 whorls devoid of a.\ial sculp- ture e.xcept for indistinct, irregular folds formed b> growth lines. Spiral sculpture of initial w horls consists of wide, very low cords separated by narrow grooves. Cords be- come stronger and more widely spaced on two last whorls, but interspaces do not exceed cord width. Subsutural ramp lacks spiral sculpture except of 3-4 feeble striae on lower part. Aperture narrow, inner lip evenly curved, covered by a smooth callus. Siphonal canal moderately long, narrow. Paralectotype smaller, with only 2.5 whorls, lacking axial ribs. Spiral cords slightly weaker than those of lectoty pe. Outer lip intact, evenly projecting forward below anal sinus. Sinus rounded, moderately deep, slight- ly asymmetrical, with its deepest point just above middle of subsutural ramp. Remarks: Barnard (1963) placed his species in Moni- liopsis Conrad, 1865. However, this Eocene genus con- taining few species from southern USA and Peru, is char- acterized by a strong sculpture of very wide spiral cords cut into subrectangular nodes that are present on the subsutural ramp. In Barnard's time, Moniliopsis was used to include some Recent West American species which are now included in the genus Ophioclennella Bartsch, 1944. This genus, in turn, cannot accomodate the Bar- nard's species either, because it is characterized by a much lower position and different type of anal sinus. Subfamily RAPHITOMINAE Bellardi, 1875 Gymnobcla glaucocreas (Barnard, 1963) (figures 15, 19) Cijthara (.'■") glaucocreas Barnard, 1963:422-42.3, fig, 2cl. Material examined: One specimen (SAM A9821) indi- cated by Barnard (1963) as "Type ' (holotype). Type locality: "Africana 11", sta. A318, 33°52'S, 16°51'E, 1380-1520 fms (2524-2780 m). Description: Shell 26 0 x 11.8 mm, with 7.5 preserved whorls, greyish-white, covered by thin periostracum. Protoconch whorls eroded, probabK partK missing. Ear- ly teleoconch whorls with only a slight angulation at periphery. Towards body whorl, whorls become more angulate, subsutural ramp becomes concave. Sutures shallowl) channeled. Axial ribs begin in lower part of subsutural ramp, numerous (27 on body and penultimate whorls), narrow, oblique, with sharp crests. Two ribs may rarely merge together near whorl shoulder. Subsutural ramp covered by thin arcuate folds following traces of anal sinus. Folds are not always continuations of axial ribs, and are more numerous than latter (41 on bod\ whorl, plus several similar short folds that do not reach lower part of subsutural ramp). Spiral sculpture of nar- row, flattened, widely spaced cords covering entire shell surface. Cords on subsutural ramp more closely spaced. Interspaces between cords bear thinner secondary thread. Remarks: This species has all the characteristics of Gym- nobcla \errill, 1884. It is most similar to the North At- lantic G. engonia N'errill, 1884 (t\ pe species of the genus) and differs from the latter mainly in having less angulate whorls, less oblique axial ribs, and fainter spiral sculpture. The radula of G. glaucocreas. described by Barnard as similar to that of G. verecunda (Barnard, 1963), also has much in common with that of G. engonia (see Bouchet & Waren, 1980, fig. 26). Gymnobcla dagama (Barnard, 1963) (figures 11-14) Cythara ('.■') dagama Barnard, 1963:423, fig. 2g, h. Material examined: Four specimens (SAM A9731) out of 5 indicated by Barnard as "Types ', Type locality: "Africana 11 ", sta. A191, 33°36'S, 16°15'E, 1520-1570 fms (2780-2871 m). Description: Largest synt\pe (38.4 x 16.3 mm) with heavily eroded surface. Second largest specimen (32.8 x 13,8 mm) with well preserved shell (fig. 12) is here des- ignated as lectotype. Shell of 6.5 whorls. Protoconch miss- ing, upper whorls eroded. Sutures deeply channeled. Whorls almost evenly convex, with only slight angulation at about 1/4 of whorl height, Subsutural ramp narrow, slightK concave. Axial ribs weak, narrow, with rounded or flattened crests, numerous (19 on penultimate whorl and 27 on bod\ whorl) and somewhat irregular in strength, especialK' on bod\ whorl. Ribs begin at border of sub- sutural ramp and extend to lower shell base, Subsutural ramp devoid of sculpture. Spiral sculpture on spire whorls appears to consist of wide, flattened cords separated by narrow grooves due to the erosion of surface. Cords well preserved and are narrow and widely spaced (interspaces 1.5-2 times wider than cord) only in some shell areas, particularly on body whorl. Aperture oval, with short, wide siphonal canal poorly difterentiated from aperture. Judging from growth lines, outer lip very slightK projects forward below anal sinus. Anal sinus narrow, with its deepest point at middle of subsutural ramp. Two smaller paralectot\pes (31.3 x 1,3.1 mm and 23.6 x 11.5 mm at 7-1- and 6-1- whorls, respectively) have more angulate whorls with more concave subsutural ramp, especially in smallest, best preserved specimen. This specimen has weak spiral cords on lower part of subsutural ramp, cov- ered In numerous thin arcuate folds formed by thickened growth lines. Remarks: The radula (Barnard, 1963, fig. 2h) is not char- acteristic of Gymnobcla and more closeK resembles that of Bcnthomangclia Thiele, 1925. However, the concho- logical characters indicate the assignment of this species to Gymnobcla. Besides the t\pe locality, the species was found in the area from 33°36' to 34°37'S and from 16°15' to 17°03'E, at 2707-3219 m (Barnard, 1963). Page 26 THE NAUTILUS, Vol. 110, No. 1 Figures 11-20. \l-l'i. Cymnobcla dagama, Iw-totvpo (12) ami paraU-dotypes ( 1 1, 13, 14; .38.4, ;31.;3, ami 2.3,6 mm, respectively). 1.5. 19. Gyrmwhcla glaucocrcas. hololype. 16-1«, 20. Xanlhodaphnc pyrropelex, lectotype (17-18) and paralectotypes {16. 20; 49.3 and 44 7 mm, respectively). A. V. Sysoev, 1996 Page 27 Gi/tnnobcla verccunda (Barnard, 1963) (figures 21-24) Daphnella {?} verecunda Barnard, 1963:427-428, fig. 2e. Material examined: Three syntypes (SAM A9697). Type locality: "Africana 11". sta. A189, 33°50'S, 17°21'E, 600 1ms (1097 m). Description: Largest (shell 22.4 x 10.8 mm) ol three syntypes best preserved, and here designated as lectotype (figs. 21-22). Sliell white, consisting of 7 preserved whorls (uppermost broken). Protoconch missing. Spire whorls angulate at periphery, subsutural ramp slightly concave. Axial ribs begin in lower part of subsutural ramp, very rapidly become prominent at shoulder, and gradually weaken below, not reaching canal. Ribs strong, oblique, narrow, 15 ribs on penultimate whorl, 16 on body whorl. Some growth lines on subsutural ramp thickened near upper suture. Spiral sculpture on subsutural ramp con- sisting of weak, narrow threads, which appear at some distance from upper suture and become stronger towards shoulder. Spiral sculpture on subsutural ramp not always visible due to shell erosion. Spiral cords below the shoul- der rather wide, flattened, sometimes with a narrower secondary cord in interspaces. Siphonal canal short, slightly twisted. Anal sinus, judging from growth lines, is asymmetrical, with its deepest point in lower half of subsutural ramp. Paralectotypes are 19.5 x 9.5 and 16.6 X 8.3 mm in size. Larger paralectotype better preserved, with 15 axial ribs on body whorl and wider spiral cords than in lectotype. All three shells have a characteristic light-orange staining on columella. Remarks: Besides the type localitv , the species was also found off northern Namibia (19°45'S), at 1600 m, R/V "Professor Shtokman", sta. 1563 (Sysoev, unpublished data). Gijmnohela (Theta) chrysopelex (Barnard, 1963) (figure 6) Typhlusyrinx chrysopelex Barnard, 1963:425, fig, 3g, h. Material examined: One specimen (SAM A9857) (ho- lotype). Type locality: "Africana 11", sta. A322, 34°36'S, 17°00'E, 1500-1760 fms (2743-3219 m). Description: Shell 18.8 x 8.6 mm, white, with glossy surface. Protoconch of three preserved whorls (at least 1-1.5 initial whorls missing), diameter 1.25 mm. Pro- toconch sculpture not well preserved, consisting of ar- cuate axial riblets passing from suture to suture. Riblets on lower half of whorl diagonalK crossed by oblique riblets begiiming at lower suture. Teleoconch of 4.75 whorls. First whorl angulate slightly above periphery, penultimate whorl at about 1/3 of whorl height. Oblique axial ribs beginning just above whorl angulation, slightly weaken below, and reaching lower suture and upper shell base. Ribs 15 on penultimate whorl, 18 on body whorl. Spiral sculpture of obscure striations below whorl an- gulation, feeble threads on lower part of shell base and canal. Based on growth lines, anal sinus very shallow as compared to congeners. Remarks: This species agrees well with the subgenus Theta Clarke, 1959 in the shell outline and sculpture. The protoconch is very similar to that of G. (T.) chariessa (Watson, 1881) (see Bouchet & Waren, 1980: figs. 254, 255), but larger than in that species. The radular teeth (Barnard, 1963, fig. 3h) are typical of Gymnobela. Gijm- nohela (T.) chrysopelex is closely related to G. (T.) char- iessa. but differs in having a shallower anal sinus, larger protoconch, and less obliciue axial ribs. Xanthodaphne pyrropelex (Barnard, 1963) (figures 16-18, 20) Pletirotoma (Surcida)dissimili.s (nun Watson) — Barnard, 1958: 147-148, fig. 23a, Typhlusyrinx pyrropelex Barnard. 1963:423-425, fig, 2c. Material examined: Four specimens (SAM A1643, SAM A1644, SAM A1645)out of 5 indicated by Barnard (1963) as "Types". Type locality: Barnard did not designate the type lo- cality, and the type series came from three different localities described as "Cape Point N. 77°E. 650-700 fathoms; NE. x E. 1/2 E. 43 miles, 900 fathoms; NE. X E 3/4 E. 38 miles, 750-800 fathoms". The syntypes were collected by the Cape Goverment trawler s.s. "Pie- ter Faure , but the respective station numbers were not indicated in the publications (Barnard, 1958, 1963) or on labels. Assuming that Barnard (1963: 424) listed lo- calities and museum numbers in the same order, the lectotype was found in the second of above-mentioned localities. Description: Largest syntype (SAM A1643, 49.3 x 18.5 mm at 7 teleoconch, 1.5 preserved protoconch whorls) with slightly broken canal, eroded parts of body whorl. Another syntype (SAM A1645, 45.0 x 15.3 mm) strongly eroded, with broken aperture. Largest of two remaining syntypes designated here as lectotype (figs. 17-18). Shell 48.2 X 17.2 mm, white, thin, fragile. Protoconch of 1.5 preserved brown whorls w ith thin oblique arcuate axial riblets. Ribs more crowded and irregular near the tran- sition to teleoconch, less inclined, occasionally bifurcat- ing in lower part. Teleoconch whorls (6.8 in number) almost evenl\- rounded, with well-marked subsutural ramp, slightK concave in earl> whorls, almost flat in last ones. Border of subsutural ramp defined by shallow groove. Some growth lines on subsutural ramp thickened below suture, which is more pronounced in early whorls. Subsutural ramp devoid of spiral sculpture except for indistinct striation on its lower part. Spiral sculpture be- low shoulder consisting of very low, flattened, wide, slightly wavy cords with serrated edges, separated by narrow grooves. Cords becoming stronger, narrower be- low bod\' whorl periphery. Based on growth lines, the Page 28 THE NAUTILUS, Vol. 110, No. 1 Figures 21-29. 21-2J-. C.ymmjhcki icmiinda. Ii-etot) pi' (21-22) and paralectotypes (23, 24-, 195 and 16.6 mm. respectively). 25-27. XaulIuHlaphnr aubruscu. lectotype (25-26) and paraleetolvpe (27). 28-29. Fanwiica hilntdis. lectotvpe (28) and par- alectotype (29). anal sinus subsutural, wide, deep. Its margin almost ver- tical below suture, nearly horizontal at border of sub- sutural ramp. Outer lip very strongly projecting forward below sinus. Paralectotypes very similar to lectotype, varying insignificantly in details of spiral sculpture. None have more intact protoconch than on lectotype. Remarks: The species has all the essential characters of XanlluHlapInw Powell, 1942. Protoconch sculpture is sinular to that oi the type species oi the genus, Plcuro- toma membranacea Watson, 1886. However, the shell of X. pyrropclcx is more slender than that of most other repre.sentati\es of the genus. Xanthodaphne pyrropclcx A. V. Sysoev, 1996 Page 29 species was also recorded in the area from 33°36'S to 34°36'S and from 16°15'E to 17°00'E, at 2524-3219 m (Barnard, 1963). Xanthodaphne suhrosea (Barnard. 1963) (figures 25-27) Typhlosijrinx subrosea Barnard. 1963:425-426 , fig.3a-d. Material examined: Two syntypes (SAM A9822). Type locality: "Africana II", sta. A318, 33°52'S, 16°51'E, 1380-1520 fms (2524-2780 m). Description: Larger of two syntypes (36.0 x 16.2 mm) designated here as lectotype. Shell white, semi-transpar- ent, fragile. Protoconch of at least 3.5 brown whorls (upper whorls eroded) sculptured b\ numerous arcuate a.xial riblets on upper half, diagonally cancellated on lower half. Protoconch diameter 1.45 mm. Teleoconch of 4.75 evenly convex, rapidh' expanding whorls. Whorls smooth initially, then obscure spiral striations appear, turning into rather strong and flattened cords (about 60 on body whorl) co\ering the entire whorl except for narrow area just below upper suture. Strongest, most widely spaced cords situated on whorl shoulder. These cords appear first on shell. Interspaces between cords usually do not exceed the cord width at whorl periphery. Siphonal canal rather long, slightly twisted. Inner lip without callus. Based on growth lines, anal sinus subsu- tural, outer lip greatK projecting forward below sinus. Paralectotype with much smaller, proportionally broader shell (20.3 X 11.0 mm), its protoconch consisting of about 4.5 whorls. Upper whorls eroded, others with sculpture similar to that of lectotype. Protoconch diameter 1.35 mm. Spiral sculpture similar to that of respective whorls of lectotype. Sculpture irregular on most of body whorl, due to scar of heavy damage to shell. Inner lip covered by very thin callus. Remarks: The species corresponds well to the diagnosis of Xanthodaphne, and is quite similar to its type-species, A', memhranacea. The only difference is that the pro- toconch sculpture in X. memhranacea is mainly axial. However, the protoconch sculpture of X. subrosea falls within the range of variability characteristic of the genus in general (see Bouchet & Waren, 1980: figs. 234-237). Famelica bitrudis (Barnard, 1963) (figures 28-29) Daphnella (.^) bitrudis Barnard, 1963: 428. fig. lb. Material examined: Two s\nt\pes (SAM A9754). Type locality: "Africana 11", sta. A193, 33°49'S, 16°30'E, 1500 fms (2743 m). Description: Larger s>nt\pe (15.2 x 3.4 mm) with bro- ken aperture, partly broken penultimate whorl. Smaller synt\pe (13.6 x 3.3 mm) here designated as lectotype. Shell very narrow, semi-transparent, white, very fragile, consisting of 7.5 whorls, uppermost vvhorl(s) missing. Up- per 3 w horls with eroded surface, differing in their pro- portions from subsecjuent whorls, apparentK' represent- ing a protoconch. Teleoconch w horls rather evenly con- vex, with clearly defined subsutural ramp, bordered by spiral cord corresponding to lower, almost horizontal branch of anal sinus. Spiral sculpture developed below subsutural ramp, consisting of weak narrow cords sep- arated by wide (2-3 times wider than cords) interspaces. Cords weaken on shell base, with smooth band at tran- sition of shell base to canal. Sculpture resuming on canal, but with very weak cords. Siphonal canal long, narrow, twisted. Anal sinus subsutural, not very broad. Outer lip projects forward below anal sinus. Paralectotype differs onl\ in having slightly more numerous and more closely spaced spiral cords (12 on the penultimate whorl vs. 11 in the lectotype). Remarks: In their original description of Famelica, Bouchet and Waren (1980), the authors included Daph- nella bitrudis in their new genus. ACKNOWLEDGEMENTS The author is greatly indebted to Mrs. Michelle G. van der Merwe, Collection Manager, South African Museum, for the loan of the material. LITERATURE CITED Barnard, K. H. 1958 Contributionsto the knowledge of South African marine Moliusca. Part I. Gastropoda: Prosobran- chiata: Toxoglossa. Annals of the South .African Museum 44:73-163, Barnard, K. H. 1963, Deep sea Moliusca from west of Cape Point, South Africa. Annals of the South African Museum 46:407-452. Bouchet, P. and A. Waren. 1980. Revision of the North-East Atlantic bathyal and abyssal Turridae (Moliusca, Gastrop- oda), Journal of Molluscan Studies, supplement 8:1-119, Dell, R, K, 1990, Antarctic Moliusca: with special reference to the fauna of the Ross Sea, Bulletin of the Royal Societ\ of New Zealand 27:1-311, Egorova, E. N. 1982. Biological results of the Soviet Antarctic expeditions, 7. MoUusks of the Davis Sea. Issledovaniya fauny morei 26(34):3-142 (in Russian). Kilburn, R. N. 1983. Turridae (Moliusca: Gastropoda) of southern Africa and Mozambique, Part 1, SubfamiK Tur- rinae. Annals of the Natal Museum 25(2):549-585. Sysoev, A. \'. 1996. 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Box 7279 Silver Spring, MD 20907-7279 THEt^NAUTILUS CONTENTS Volume 110, Number 2 Fehmanj 19, 1997 ISSN 0028-1344 Dan C. Marelli Maureen K. Krause William S. Arnold William G. L^•ons Systematic Relationsliips Among Florida Populations of Aroopcctcn irmdidiis (Lamarck. 1819) (Bi\al\ia: Fectinidae) 31 Subspecific Status oi Araopccten irradiaiis coiiroithcns (Say, 1S22) and of the Bay Scallops of Florida 42 Dan C. Marelli William G. Lyons William S. Arnold Maureen K. Krause Daniel L. Graf Distribution ol Uuionoid (Bi\al\'ia) Faunas in Minnesota, USA 45 M. G. Harasewych The Life and Malacological Contributions ol H. Tucker Abbott ( 1919-1995) 55 John B. Wise FctitiUa. New Name for PetitcIIa Wise, 1996, a Preoccupied Name (Mollusca: Gastropoda: Pyramiilt'llidae) 76 STATEMENT OF OWNERSHIP, MANAGEMENT AND CIRCULATION 1. Publication Title, THE NAUTILUS 2. Publication No,, 0028-1344 3. Filing Date, October 14, 1996 4. Issue Frequenc), Quarterh 5. No. of Issues Published .■VnnualK , Four 6. 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Paid Circulation 1. Sales through dealers, carriers etc. 2. Mai! Subscriptions C. Total paid and requested circulation D. Free Distribution b> Mail E. Free Distribution outside the mail F. Total free distribution G. Total distribution H. Copies not distributed (1) Office use, leftovers, spoiled (2) Return from Newsagents I. Total Percent Paind and/or Requested Circulation 93^ 95'" Average Single 12 months Issue 550 550 0 0 440 440 440 440 23 23 10 2 33 25 473 465 77 85 0 0 550 550 THE NAUTILUS 110(2):31-41, 1997 Page 31 Systematic Relationships Among Florida Populations of Argopecten irradians (Lamarck, 1819) (Bivalvia: Pectinidae) Dan C. Marelli Maureen K. Krause' - William S. Arnold William G. Lyons Florida Department of Environmental Protection Florida Marine Research Institute 100 8th Avenue SE St. Petersburg, FL 33701-5095 USA ABSTRACT Morphometric and genetic examinations (using allozyme elec- trophoresis) were conducted on two Florida populations of bay scallops, Argopecten irradians (Lamarck, 1819), to investigate the status of the subspecies A. i. taijlorae Petuch, 1987. One other Florida population [A. i. coiicentricus (Say, 1822)] was examined morphometrically. Morphometric examinations em- phasized mensural and meristic characters used in previous systematic diagnoses, Morphometric data were analyzed using anal) sis of variance and principal component analysis. Scallops taken from Florida Bay (putatively A. i. taijlorae Petuch, 1987) were smaller but otherwise not morphologically distinct from populations of A. i. concentricus (Say, 1822) from Pine Island Sound and Homosassa Bay, Florida. Mean Nei s modified ge- netic distance value shows a close relationship between Florida Bay and Homosassa Ba\ scallops. Neither morphometric nor genetic e\idence supports the proposed status of A. i. taijlorae as distinct from Florida populations of A. i. concentricus. Key words: Pectinidae, Argopecten irradians. morphometries, svstematics. INTRODUCTION Scallops are often distributed in patchy or contagious patterns, and, although disjunct local populations may be joined by variable degrees of lar\ai exchange to form metapopulations (Levins, 1970; Andrewartha k Birch, 1984; Roughgarden et ai, 1985; Roughgarden & Iwasa, 1986; Simberloff, 1988; Hanski, 1989; Orensanz et a!., 1991), local populations along a species' range are prob- ably self-sustaining (Sinclair ct ai. 1985). ExpectedK', ' Department of Ecology and Evolution, State University of New York at Stony Brook, Stony Brook, NY 1179-4-5245 USA. ^ Current address: Southampton College, Long Island Univer- sity, Southampton, NY 11968 USA. reproductive isolation may contribute to genetic drift that can produce locally distinctive morphologies. The bay scallop, Argopecten irradians (Lamarck, 1819), which inhabits semi-enclosed coastal bays, sounds, and estuaries from Cape Cod, Vlassachusetts, to Tampico, Mexico (Clarke, 1965; Abbott, 1974), exemplifies the results of such isolation (Clarke, 1965; Waller, 1969; Kraeuter et ai, 1984). Three extant subspecies ot A. irradians have custom- arily been recognized: Argopecten irradians irradians (Lamarck, 1819), distributed from Cape Cod to New Jersey; A. i. concentricus (Sa\-, 1822), distributed from New Jersey to South Carolina and from Palm Beach Inlet, Florida, to Louisiana; and A. i. aniplicostatus (Dall, 1898), distributed from Texas into Mexico (Clarke, 1965; Wal- ler, 1969, 1991; Abbott, 1974). A fourth subspecies, A. i. sablensis (Clarke, 1965), described from shells collected in Nova Scotia, apparently became extinct during the Holocene (Waller, 1969). These subspecies have been distinguished morphologically by plical number and shape, valve size and shape, and valve coloration (Clarke, 1965; Waller, 1969; Abbott, 1974). Bricelj et at. (1987), however, cautioned that subspecific distinctions within A. irradians have not yet been substantiated by bio- chemical techniques. Recently Petuch (1987) described an additional sub- species of ba\' scallop, Argopecten irradians taijlorae, based upon specimens from Rabbit Key Basin, Florida, and reported it to be restricted to Florida Bay and the western sides of the middle and upper Florida Keys (figure 1). Petuch (1987) reported that plicae of A. i. taylorae were more numerous than those of the other subspecies and that plicae were lower, wider, and more flattened than in A. i. concentricus. The valves of A. i. taylorae were also said to be much smaller, more fragile, and much more colorful than those of A. i. concentricus (table 1). To emphasize the geographic isolation of Argopecten irradians taylorae, Petuch (1987) gave the distributional Page 32 THE NAUTILUS, Vol. 110, No. 2 ST. JOSEPH BAY GULF OF MEXICO .% JOHNSON o .KEY-? ;:.^----%;:**''\..^;: ::;*;/■■■ i RABbir'"^ /} KEY BASIN s and additional localities relevant to bay scallop populations. Inset shows location of collection uilliin Hahhil Ke\ Dasin. range of .A. i. concent ricus as "New Jerse\ to Georgia, and. . .Tampa, Florida to Louisiana." In iloing so, lie overlooked earlier Florida records of bay scallops at Boca Grande and Sanibel Island (Clarke, 1965); at Whitewater Bay, Sandy Ke) Basin, and south of I'laniiiigo in Florida Bay (Tabb & Manning, 1961); and at several locations from Vaca Key in the middle Keys northward through Biseayne Bay to Palm Beach Inlet on the Florida east coast (Waller, 1969). Clarke (1965) and Waller (1969) had pre\iousl\ reported specimens from all of these lo- cations as A. i. concent ricus. but at least some of the specimens came from within the range of A. i. tatjlorae. An investigation of the s\ stematic status of Argopecten irradians taylorae was prompted b\ the need to address resource management issues. Ba\ scallops have histori- cally supported small-scale commercial fisheries at var- ious locations between Pensacola Ba\ and Pine Island Souiul along Florida s west coast. .Although the com- mercial fishery has decreased substantially in recent years, the species continues to be the target of an important recreational fisherv throughout much of that region (Ar- nold, 1990), The presence of an isolated subspecies of ba> scallop in Florida Ba\ , as reported b\ Petuch (1987), could have ramifications for resource management. The Dan C. Marelli et a I., 1997 Page 33 Table 1. Characters of subspecies of Argopeclen irradians reported by previous authors. Subspecies Clarke, 19te Waller, 1969 Abbott, 1974 Petuch, 1987 Argopecten irradians irradians Argopecten irradians concenlricus Argopecten irradians amplicostatus Argopecten irradians taylorae 16-20 (usuall) 16-lS ribs; ribs rounded; valve length and height nearl) equal, L/H ratio approx. 1.06; W/H ratio ap- pro.\. 0.21 (left) and 0.24 (right); right valve color as in Ab- bott (1974) but com- monly white (or nearly so), 17-23 ribs; ribs round- ed or with flattened tops; L/H ratio ap- prox. 1.03 (1) and 1.02 (r); valves some- what inflated, W/H ratio approx. 0.26 (1) and 0.32 (r); 75- 100'^ of specimens with v\ hite or nearly white right valves, color otherwise like left valve except lighter. 12-18 (usually 14-16) ribs; ribs rounded to slightly flattened; L/H ratio approx. 1.0(1) and 1.01 (r); shell more inflated than other 2 subspe- cies, W/H ratio ap- prox. 0.29 (1) and 0.34 (r); color of right valves usually white, sometimes with slight color tinge. 15-20 ribs; ribs low and 17-18 ribs; ribs low and rounded; L = H; valves thinner and flatter and ribs lower than A. i. concentri- cus. 15-22 ribs; ribs high, sharplv rounded, be- coming semi-hexago- nal distally; differs from A. i. irradians by having thicker and more convex valves and higher ribs; differs from A. i amplicostatus by having thinner and less convex valves. 13-17 ribs; ribs high, sharply rounded proximally, becoming low and trapezoidal to semi-hexagonal distally; differs from A. I. concentricus by having thicker, more convex valves with fewer ribs. roundish; valves most compressed of the 3 subspecies; right valve color only slightlv lighter than left. 19-21 ribs; ribs squar- ish; right valve much more convex than left; right valve light- est in color of the subspecies, common- Iv all white. 12-17 ribs; right valve with high, squarish ribs; shell more gib- bose than other sub- species; right valve commonly white. 23-25 ribs; ribs flatter and wider when compared with A. i. concentricus. shell smaller, more fragile and more colorful than A. i. concentri- cus; right valve gen- erally yellow rather than white, with more brown mot- tlings than A. i. con- centricus. population might require separate management because of the increasingly popular recreational fishery or might require protection as a rare and isolated subspecies. We used both allozyme electrophoresis and morpho- metric analyses to investigate the relationship of the Flor- ida Bay population of Argopecten irradians taylorae to the Homosassa Bay population of A. i. concentricus. Valve characters were analyzed morphometrically for two Florida populations of A. i. concentricus and one of A. i. taylorae. as well as for the t\ pe series of A. i. taylorae. Page 34 THE NAUTILUS. Vol. 110, No. 2 Although subspecific morphometric variation within A. irradians has been investigated in other studies (Clarke, 1965; Waller, 1969), no effort had been made to char- acterize corresponding genetic variation. This study is the first to examine both morphological and genetic vari- ation within a putative subspecies of Argopecten irra- dians. MATERIALS AND METHODS Bay scallops (n = 66) were collected from Rabbit Key Basin, the type locality of Argopecten irradians taylorae, on 29 August 1990. The scallops were located in the southeastern portion of the basin, in a turtle grass (Thal- assia testudinum Banks e.\ Konig) bed, at a depth of appro.ximately 2 m. At that time we also collected empty valves (n of right valves = 13) of A. i. taylorae from that locality. Scallops were returned to the Florida Depart- ment of Environmental Protection s Keys Marine Lab- oratory on Long Key, where sections of adductor muscle and mantle tissue were dissected from live animals and stored in liquid nitrogen for subsequent genetic analysis. Shells were disarticulated, numbered, and returned to the Florida Marine Research Institute at St. Petersburg. Additional samples of bay scallops were collected from Homosassa Bay (n = 60), just west of the mouth of the Homosassa River, on 24 July 1990 and from Pine Island Sound (n = 56) on 28 June 1991. The specimens from Homosassa Bay and Pine Island Sound were also collected from shallow (z < 2 m) T. testudinum beds; Homosassa Bay scallops were also examined both genetically and morphometrically, but Pine Island Sound scallops were examined only morphometrically. Electrophoresis. Tissue samples of scallops from Hom- osassa Bay and Rabbit Key Basin were analyzed for ge- netic composition using methods reported in Bricelj and Krause (1992). Eight polymorphic loci (frequency of the most common allele < 0.99) were examined, represent- ing the following enzymes: phosphoglucomutase (PGM, EC 2.7.5.1), glucose-6-phosphate isomerase (GPI, EC 5.3.1.9), octopine dehydrogenase (ODH, EC 1.5.1.11), superoxide dismutase (SOD-1, EC 1.15.1.1), a-amino acyl peptide hydrolase (LAP, EC 3.4.1.-), alanyl aminopep- tidase (AAP, EC 3.4.1.-), dihydrolipoamide dehydroge- nase (DHLD, EC 1.8.1.4), and nonspecific aminopepti- dase (AP, EC 3.4.1,-). SOD-1 was only weakly poly- morphic (three alleles, H = 0.03), and DHLD was in- consistently resolved; thus these two loci were not in- cluded in the data analysis. The following enzymes examined were monomorphic: superoxide dismutase (SOD-2), /i-galactosida.se (/i-GAL, EC 3.2.1.23), isocitrate dehydrogenase (two loci, IDH- 1 and IDH-2, EC 1.1.1.42), mannose phosphate isom- erase (MPI, EC 5.3.1.8), 6-phosphogluconate dehydro- genase (PGD, EC 1.1.1.44), catalase (CAT, EC 1.11.1.6), glycerol-3-phosphate dehydrogenase (GPDH, EC l.i.1.8), trio.se phosphate isomerase (TPl, EC 5.3.1.1), esterase (two isozymes, EST-1 and EST-2, EC 3.1.-.-), arginine kinase (ARK, EC 2.7.3.3), L-iditol dehydroge- nase (IDDH, EC 1.1.1.14), and malic dehydrogenase iso- zymes (MDH-1 and MDH-2, EC 1.1.1.37). Alleles were designated using standard notation: the most common allele at each locus was assigned a value of 100 and other alleles were defined based on mobility relative to the most common allele. Observed genotypic frequencies were compared with those expected under Hard> -Weinberg equilibrium us- ing the G-test and, when necessary, Williams correction for small sample size (Sokal & Rohlf, 1981). Rare alleles were pooled with the electrophoretically closest common allele to obtain genotypic class frequencies >5. Hetero- zygote deficit or excess was determined from the D sta- tistic where D is the percentage deficit (Selander, 1970); negative D values indicate a deficit of heterozygous ge- notypes. Allele frequencies among populations were compared using the R x C test of independence and the G-test (Sokal & Rohlf, 1981). Nei's modified genetic dis- tance (Nei, 1978) was calculated for each population pair surveyed. Morphometries. Morphological characters of scallops in populations from Homosassa Bay, Pine Island Sound, and Rabbit Key Basin and of the holotype (USNM 859901) and paratypes (USNM 859902) of Argopecten irradians taylorae were examined following the methods of Waller (1969). We also collected and examined a sample of recently emptied valves of scallops (putatively A. i. tay- lorae) from Rabbit Key Basin that appeared morpholog- ically identical to living scallops from that area. For each scallop we counted right valve plicae and measured plical height, plical width, and 13 other valve characters di- mension that have been applied in morphological com- parisons of bay scallops (Clarke, 1965; Waller, 1969; Ab- bott, 1974; Petuch, 1987) (Appendix 1). Mensural characters used by Petuch (1987) in his di- agnosis were examined using analysis of variance (ANO- VA) for their relationship to valve size in the same four populations. Data from the type series of A. i. taylorae from Rabbit Key Basin were deleted from the analyses of population effects because of the small sample size (n = 9) and the resultant undue influence on the statistical models. Principal component analysis (PCA) was performed using only mensural variables. Marcus (1990) suggested that measurement error can unduly influence variability in the PCA model, so standard errors of measurement were calculated for each mensural character to estimate measurement error. Each character was measured 10 times on each of six specimens, two from each population, and standard errors were calculated for each specimen- by-character combination. Standard errors were then ex- pressed as a percentage of the mean. L'sing all fifteen variables we log transformed Uogio) the \alues and per- formed a standard PCA (PRINCOMP procedure, SAS Institute, Inc., 1985). Burnab\ s (1966) method of size- corrected PCA was performed separateK on all data [size-corrected PCA (N. Macleod, personal communi- cation)]. Burnaby s method removes most of the influence of size on the PCA by constructing variables orthogonal Dan C. Marelli et a/., 1997 Page 35 Table 2. Allele frequencies for polymorphic enzymes from populations of Argopecten irradians from Rabbit Key Basin and Homosassa Bay, Florida, n = number of alleles sampled (2 X number of individuals). N^ = effective number of alleles (after Hartl & Clark, 1989). Locus AP AAP LAP ODH PGM GPI Allele 108 104 100 96 94 n Ne 102 100 98 97 96 94 n N. 100 96 94 n N, 106 104 102 100 96 94 90 n N. 98 96 95 94 92 91 90 n N. 150 106 104 100 99 98 n Frequency RKB HOM 0 0,38 0.53 0.08 0.01 132 2.32 0 0.35 0.17 0.13 0.19 0.15 132 4.37 0.62 0.37 0,01 132 1,92 0.01 0.08 0 0.77 0.07 0.01 0.05 132 1.63 0.02 0.49 0.1-4 0.13 0.11 0.03 0.07 132 3.36 0 0.06 0.61 0.30 0.02 0.01 132 2.14 0.02 0,23 0.51 0.24 0 118 2.70 0.03 0.40 0.24 0.04 0.17 0.13 112 3.79 0.85 0.15 0 118 1,35 0.02* 0,13 0.01 0.80 0 0.04 0 118 1.52 0.01* 0.56 0.18 0.19 0.01 0 0.05 lis 2.61 0,01 0.02 0.62 0.33 0.01 0 118 2()2 * Genotype frequencies not in Hardy-Weinberg equilibrium. to a variable that is considered to represent size (Rohlf & Bookstein, 1987). The relative contribution of each variable to the variation represented by PC2 and PC3 was determined, and Cattell s (,1966) scree test was used to identify and eliminate variables that failed to add 60- ? £50- E3 E3 1- I O ^40- UJ >i ^30- 20- ■' E I i ■ ■ RKB-USNM RKB-FMRI PISL LOCALITY HOM Figure 2. Valve heights of Florida Gulf coast Argopecten ir- radians populations. Plots show mean, range, and ± 1 standard error. RKB-USNM: Rabbit Key Basin, type seriesoi Argopecten irradians taylorae. USNM 859901 & 859902; RKB-FMRL Rab- bit Ke\ Basin, collection made for this study, FSBC I 41441; PISL: Pine Island Sound, FSBC 1 44439; HONI: Homosassa Bay, FSBC 1 40337. CD s 3 Z -I < o 26- 25- 24- 23- j E E3 22- ' E 3 21- 20- 19- - I RKB-USNM RKB-FMRI PISL LOCALITY HOM Figure 3. Numbers of right valve plicae for Florida Gulf coast Argopecten irradians populations. Plots show mean, range, and ± 1 standard error RKB-L SNM: Rabbit Ke\ Basin, type series of Argopecten irradians taylorae. USNM 859901 & 859902; RKB-FMRL Rabbit Key Basin, collection made for this study, FSBC I 41441; PISL: Pine Island Sound, FSBC I 44439; HOM: Homosassa Bav, FSBC I 40337. substantially to the variance in the analysis, leaving 9 variables: AM, AD, DG. AK, GP, BJ, IW, ad, and ce (see Appendix 1 for explanations). Using only these variables, data were reanalyzed, and size-adjusted shape variation among populations was examined using PC2 and PC3 generated b\ the Burnaby technique. Page 36 THE NAUTILUS, Vol. 110, No, 2 Figures 4-7. Left valves of Argopeclen irradians from Florida. 4. A. i. concentricus. height 60.0 mm. Rabbit Ke> Basin, FSBC I 41441. 5. A. i. taylorae. holotype, height 44.0 mm. Rabbit Key Basin, USNM 859901. 6. A. i. concentricus. height 60.1 mm, Homosassa Bay, FSBC I 40337. 7. A. i. concentricus, height 57.9 mm. Pine Island Sound, FSBC I 44439. Scale bar = 10 mm Growth relationships of \alve characters within and between populations were examined using the method of Jolicoeur (1963). Eigenvectors (coefficients) of PCI from the standard PCAs using log-transformed data for all populations combined and separately on populations from the three localities were compared with (l/p)"^, where p = the number of variables and therefore (1/ p)0o = 0.333. Coefficients for valve characters that varied above or below 0.333 represented factors that reflected positive or negative allometry. Valve color and pattern were examined following the criteria of Elek and Adamkewicz (1990), with the ex- ception that the relationship between color pattern and population was examined using the nonparametric Krus- kal-W'allis procedure. Disposition of specimens: Except for the t\pe material of Argopectcn irradians tatjlorac, valves of all specimens examined have been deposited in the Florida Marine Research Institute Invertebrate Collection, lot numbers FSBC I 40337 (Homosassa Bay), FSBC I 41441 (Rabbit Key Basin), and FSBC I 44439 (Pine Island Sound). RESULTS Electrophoresis. Among the poK morphic loci no sig- nificant differences were found bet\veen populations for the loci GPI, ODH, AAP, or PGM. Allele frequencies between Homosassa Bay and Rabbit Key Basin popu- lations were significantly different for amino peptidase (AP, p < 0.05) and leucine amino peptidase (LAP, p < 0.001) (table 2). The mean percentages of heteroz) gous loci per indi- vidual (MLH) for Homosassa Bay and Rabbit Key Basin populations were 43^7 and 44*7 using the six polymorphic loci. Homo.sassa Bay scallops fit Hardv -\\ einberg expec- tations for heterozygosit) at all loci, but the Rabbit Key Basin population had heterozvgote deficits at the PGM (p < 0.05) and ODH (p < 0.01) loci. Dan C. Marelli et a/„ 1997 Page 37 Table 3. Growth relationships for valve characters from Rab- bit Key Basin (RKB), Pine Island Sound (PISL), and Homosassa Bay (HOM), Florida Argopecten irradians populations ana- lyzed using principal component analysis. Numbers are eigen- vectors of principal component 1 and indicate positive or neg- ative allometric growth where eigenvectors var>' above or below l/Vp. where p = the total number of characters, following Jolicoeur (1963). Popu ation All Char- popula- acter tions RKB PISL HOM AM 0.3132 0.3157 0.3265 0.1672 AD 0.3037 0.3095 0.3385 0.1447 DG 0.3087 0.3340 0.2589 0.2415 AK 0.3063 0.6240 0.3139 0.2051 GP 0.3302 0.3212 0.4259 0.1548 BJ 0.2635 0.3348 0.3178 0.2051 \\\ 0.0796 0.2789 0.3457 0.5943 ad 0.3696 0.3301 0.3739 0.4236 ce 0.5469 0.4313 0.2675 0.5015 The mean genetic distance between the Homosassa Bay and Rabbit Key Basin populations was 0.035. Morphometries. Standard error estimates for measure- ments of morphometric characters were universally low, averaging 0.64% of the mean for all variables, suggesting that measurement error did not overly influence the vari- ance in subsequent morphometric analyses. Scallops from the Pine Island Sound collection were significantly larger than those from Homosassa Bay, and both Pine Island Sound and Homosassa Bay scallops were much larger than those from our Rabbit Key Basin col- lection (AXOVA, all p < 0.05) (figures 2-3). The type specimens of ArgopfTfen irradians fay/oraf were slightly larger than the live specimens from our Rabbit Key Basin collection and similar in size to the empty valves we collected from Rabbit Key Basin, although two pairs of empty valves exceeded 50 mm in height. Representative valves from the sample populations are illustrated in figures 4-7. Numbers of right-valve plicae were similar for all sam- ple populations that we examined morphometrically, and plical number was not significantly related to valve height (0.05 < p < 0.06). However, the effect of population on plical number was significant (p < 0.0001), and right- valve plicae were significantly more numerous on Hom- osassa Bay scallops (p < 0.05) than on Rabbit Key Basin and Pine Island Sound scallops. Plical numbers were not significantly different between Rabbit Key and Pine Is- land Sound populations. Width of plicae relative to valve size was positively related to valve height (p < 0.0002). Patterns of growth for individual mensural characters varied greatly and inconsistently among populations (ta- ble 3). Although some characters varied isometrically, there was no consistent trend among populations. Coloration of valves from all populations was similar; more than 90% of specimens from each population ex- hibited the color pattern of t\ pe E of Elek and Adamke- wicz (1990). Variations in color pattern were not signif- icantly different among populations. More than 95% of the valves from each scallop population had white back- grounds, and 100% of the specimens had mottled left valves (table 4). Excluding one yellow and two orange scallops from Rabbit Key Basin and one orange scallop from Pine Island Sound, right valves in all populations were consistently all white except in the early juvenile region of the shell. The first principal component (PCI) from the standard PCA of all data accounted for 83.3% of the total variance, and the second and third principal components account- ed for 7.7%' and 4.1%, respectively, of the variance. Bur- naby size-corrected PCA showed that principal com- ponents 1 through 3 accounted for 59.2%-, 14.9%, and 9.8% of total variance, respectively. Two variables in the Burnaby PCA accounted for most of the variance in PC2 and PC3: width of plical interspaces (IW) represented 28.5% of PC2 variance and 69.5% of PC3 variance, and ce (length of ligament insertion) accounted for 33.6% of PC2 variance and 20.4% of PC3 variance (table 5). Plots of PC2 and PC3 from the size-corrected PCA of the data from all 3 populations and including the separate col- lection of empty valves from Rabbit Key Basin (figure 8) demonstrate that scatterplots from each population overlap with those from all other populations. Particu- larly extensive overlap occurs between the Rabbit Key Basin and Homosassa Bay populations, and less overlap is seen in plots representing the Pine Island Sound pop- ulation and those of all other populations. DISCUSSION The genetic distance (D = 0.035) between the Homosassa Bay and Rabbit Key Basin populations of Argopecten Table 4. Summar\' of Island Sound (PISL, n = color and color pattern on valves of Argopecten irradians from Rabbit Key Basin (RKB, n — 66), = 56), and Humusassa Ba\ iHOM, n = 60), Florida. Numbers expressed as percentages. Pine Popula- tion Bac ■kg round color Rays present Mottling Ba iding White Yellow Orange Left Right Left Right Left Right RKB PISL HOM MEAN 95.5 98.2 100 97.8 1.5 0 0 0.5 3.0 l.S 0 1.6 90.9 96.4 98.3 95.1 30.3 12.5 6.7 17.0 100 100 100 100 80.3 85.7 93.3 86.3 100 41.1 98.3 81.3 57.6 58.9 90.0 68.7 Page 38 THE NAUTILUS, Vol. 110, No. 2 Table 5. Percentage of variance in principal components 2 and 3 from Burnaby PCA attributable to individual valve char- acters for Argopecten irradian^ populations from Rabbit Key Basin, Pine Island Sound, and Homosassa Bay. Character ab- breviations are defined in the appendi.x. 0.4 Principal Principal Character component 2 component 3 AM 4.78 1.13 AD 3.60 1.73 DC 2.09 0.72 AK 6.86 2.28 GP 9.18 3,09 BJ 8.13 0.20 I NX- 28.50 69.50 ad 3.28 0.89 ce 33.57 20.43 irradians s.l. is on the order of distances between local races (Nei, 1976, 1987). This value is comparable to those between disjunct but apparently recently separated pop- ulations of the Baltic clam Macoma balthica (Linnaeus, 1758) (Nei's D = 0.058) (Meehan et ai, 1989) and be- tween Great Barrier Reef and Enewetak Atoll popula- tions of the giant clam Tridacna maxima (Nei's D = 0.033) (Ayala, 1975). Electrophoretic data from five ad- ditional populations of Argopecten irradians s.l. exam- ined by Krause (1992) (Martha's Vineyard, Massachu- setts; Niantic River, Connecticut; Orient Harbor, Long Island, New York; Core Banks, North Carolina; and St. Joseph Bay, Florida), treated identicalK , were used along with data from the Rabbit Ke) Basin and Homosassa Bay populations to create a genetic distance matrix. A dendrogram was constructed from the genetic distance matrix using the unweighted pair-group method with arithmetic mean (UPGMA Sneath & Sokal, 1973), pro- vided by the computer program NTSYS (Applied Bio- statistics, Inc.) (figure 9). Standard errors of tree branch- ing points were estimated using the procedure of Nei et al. (1985) who point out that, when using electrophoretic data and less than 30 loci, the topology of a reconstructed tree is subject to a large stochastic error. The size of the errors allows us little confidence in the dendrogram, but two factors may contribute to an inflation of the esti- mated standard errors. When genetic distance values are very low, less than 0.105, the value of 1 in the equation D = -log,, I exceeds 0.9. When I >0.9 and average het- erozygositv (VILH) is >0.2 serious overestimation of the variance may occur (Nei et ai, 1985). Twelve of 21 pairwise I values in the scallop genetic distance matrix exceed 0.9 and MLH = 0.44 p. Despite the size of the standard errors, the genetic distance data indicate a close relationship between the Rabbit Key Basin and Homo- sassa Bay scallops within Argopecten irradians. The small but significant differences in allele fre- quency between Homosassa Bay and Rabbit Key Basin populations at the AP and L.'\P loci might be evidence for localized selection among genotypes between these sites (McMillen et ai, 1994), for reduced gene flow be- 0.3 CO H u 0.2 Z o a o u < a. O 0.1 z oc a. -0.1 o D O DO • D '^ 0 d'^d • ncP • °o^°*°o^ ° • ^ 0 °o \ rfto a D D i '^ a ^ a A A - A i^ • a a a a a8 a ^A A ^ ■ ■« . ■ , & ^^ ■ ■ ■ ■ *■ • * 1 ■* .- ■ ■ 1 1 1 1.3 1.4 1.5 1.6 1.7 PRINCIPAL COMPONENT 2 1.8 1.9 Figure 8. Bivariate scatterplot of principal component 2 and 3 scores generated using Burnabv 's (1966) size-corrected prin- cipal component analvsis on morpliometric measurements of Argopecten irradians from Rabbit Key Basin (D), Pine Island Sound (■). and Homosassa Bav (A) populations. Population indicated by (•) represents empty valves collected from Rabbit Kev Basin localitv , tween the populations (Beaumont & Zouros, 1991). or some combination of these processes. The overall mag- nitude of genetic variation among the Florida popula- tions is onl\ slightK greater than that found among Krause's (1992) northern populations of A. i. irradians. Therefore A. i. taylorae appears, based on genetic evi- dence, not to differ at the subspecific level from other Florida populations examined. X'alues for the mean percentage of heterozygous loci per individual (MLH) in the Homosassa Ba\ (43?c) and Rabbit Ke\ Basin (44%) populations are relativeK high but similar to the 45.3% MLH reported by Bricelj and Krause (1992) for Argopecten irradians irradians from the Niantic River estuary, Connecticut. These values in- dicate that high MLH values may be characteristic of Argopecten irradians s.l., but Wall et al. (1976) reported a MLH of 11.6% for A. irradians from Bogue Sound, North Carolina. Values of MLH reported for 5 other pectinids range from 9.4% to 32.1% (Nikiforov & Dol- gonov , 1982; Beaumont & Beveridge, 1984). Color in .\rgopecteri irradians is genetically controlled Dan C. Marelli et al.. 1997 Page 39 (0.026) (0.014) [(oToos) — (0.04) (0.019) (0.021) MASS CONN LINY CBNC SJFL HOM RKB 0.15 0.125 0.1 0.075 0.05 GENETIC DISTANCE 0.025 -I 0 Figure 9. I'PGMA tree constructed trom tlie Nei (1972) genetic distance matrix for populations of Argopecten irradians. Population abbreviations are MASS: Martha's Vineyard, Massachusetts; CONN: Niantic River, Connecticut; LINY: Orient Harbor, Long Island, New York; CBNC: Core Banks, North Carolina; SJFL: St. Joseph Bay, Florida; HOM: Homosassa Bay, Florida; RKB: Rabbit Key Basin, Florida. Numbers in parentheses are standard errors of branching points estimated using the procedure of Nei et al. (1985). and may sometimes be useful in distinguishing individ- uals and populations (Kraeuter et al.. 1984; Adamkewicz & Castagna, 1988). Nevertheless, we found no substantial differences in valve coloration or color pattern between the Homosassa Bay, Pine Island Sound, and Rabbit Key Basin scallop populations to support Petuch's (1987) con- tention that shells of A. i. taylorae are much more colorful than those of A. i. concent ricii.s. Most scallops in each of the populations had a rayed pattern on the left valve, and additional color was generalK expressed as mottling and banding, somewhat obscuring the rayed pattern. Variations in the intensity of color were affected by the nature and e.xtent of fouling on the left \alve and possibly by ontogenetic change. Petuch (1987) identified the small size of scallops from the Rabbit Key Basin population as an important char- acter that distinguishes Argopecten irradians taylorae from A. i. concentricus. Although heights of live scallops from our Rabbit Key Basin collection were significantly smaller than those in all other collections, the empty valves that we collected from the same locality were comparable in size to those in Petuch's type series. This suggests that the scallops in Rabbit Ke\' Basin may indeed be distinctively smaller than other Florida scallops, al- though the close genetic similarity between the Rabbit Key Basin and Homosassa Ba\ populations indicates that scallop size is readily influenced by local conditions. The small size of our live Rabbit Key Basin collection coupled with the heterozygote deficits at the loci ODH and PGM that occurred among the scallops from this collection [similar to deficits at these loci that have been reported for juveniles of other species of Pectinidae (Volckaert & Zouros, 1989; Bricelj & Krause, 1992)] may indicate that the scallops we collected from Rabbit Key Basin were juveniles. Discontinuities in plical number occur between pop- ulations of Argopecten irradiana along the geographic range of the species, and these differences have been emphasized ta.xonomically (Clarke, 1965). The number of plicae ranges from 12 to 25 in the described subspecies (Clarke, 1965; Waller, 1969; Abbott, 1974; Petuch, 1987) and is reportedly under genetic control (Kraeuter et al., 1984). We substantiate the elevated plical numbers that have been reported for Florida A. i. concentricus. but we did not find even greater plical counts in the Rabbit Key Basin population as reported b> Petuch (1987). In fact, the mean plical number for our sample of scallops from Rabbit Key Basin was statistically indistinguishable from that of scallops from Pine Island Sound. Higher counts of plicae for the type series reported by Petuch (1987) suggest that he may have misidentified as plicae some of the "riblets" that occur on the disk flanks of this species; such confusion has been previously reported by Clarke (1965) and Waller (1969). Our analysis of right valve plical numbers rejects the conclusion that scallops from Rabbit Key Basin represent a separate and unique subspecies. Width of the plicae at the ventral margin was iden- tified b\ Petuch (1987) as a character that can be used to separate Argopecten irradians taylorae from A. i. concentricus. However, because plical width relative to valve size is positively correlated with valve height, its use in a univariate comparison of scallops of different sizes without accounting for allometry is invalidated. Moreover, our data indicate that Rabbit Key Basin scal- lops have relatively narrower plicae than do scallops from Homosassa Bay or Pine Island Sound — not wider as re- ported by Petuch. Inconsistencies in the relationship between growth and valve characters between populations indicate that mor- phometric characters do not vary with growth in a simple manner; variability due to small sample sizes and mea- Page 40 THE NAUTILUS, Vol. 110, No. 2 surement error may also affect the allometric coefficients (Marcus, 1990). Regardless of the source, the inconstancy of shape variables and the presence of allometric rela- tionships among mensural characters, morphometric ra- tios, and size cast doubt upon the use of univariate char- acters in describing morphometric differences between scallop populations, adding empirical emphasis to warn- ings b\ previous authors (summarized in Humphries et ai. IQ'sI). Burnab)' size-corrected PCA failed to distinctly sep- arate the Rabbit Key Basin, Pine Island Sound, and Hom- osassa Bay scallop populations. Although morphometric overlap among populations is e\ ident in the plots of PC2 and PCS, scatterplots of the individual populations form distinct clusters and indicate that the more geographi- calK distant populations (Rabbit Ke> Basin and Hom- osassa Bay) in our samples were morphometrically more similar. Variation in shape, and hence some separation in the plots of PC2 and PC3, may be influenced by ontogeny of individuals, because the Burnaby technique does not completeK remove the effect of size (Humphries etal., 1981), but separationon the basis of shape variables indicates that scallop shape is heavily influenced by local conditions and may simply reflect ecophenotvpic vari- ation. We assessed alleged differences between typical Ar- gopecten irradians taylorae and representatives of other bay scallops from peninsular Florida. Based on the results of our electrophoretic and morphometric examinations, we refute those differences and conclude that bay scal- lops from Rabbit Key Basin do not represent a subspecific taxon distinct from Florida populations of A. i. concen- tricus. ACKNOWLEDGEMENTS Clarita Lund, Brenda Hedin, Richard Darden, Yantian Lu, Don Hesselman, and James Seagle assisted in col- lecting scallops, and C. Lund also performed data entry. Robert McWilliams and Charlotte LaTorre measured valve characters. Catherine Bray assisted in data analysis and figure preparation. Ll\ n French prepared figures. Dr. Theresa Bert and Hector Cruz-Lopez helped with interpretation of electrophoretic data. Everglades Na- tional Park biologist Daniel Foxen facilitated our Florida Bay collection. This project was partially supported b\ funds created by the Florida Saltwater Products License and the Florida Saltwater Fishing License. Electropho- retic anaKses were supported by a National Science Foundation dissertation improvement grant (BSR- 9015991) to MKK LITERATURE CITED Abbott, R. T. 1974. .\merican Seashells, 2nd edition. Van Nostrand Reinhold, New York, 663 pp. Adamkewicz, L. and M. Castagna. 1988. Genetics of shell color and pattern in the bay scallop Argopecleu irradians. Journal of Heredity 79:14-17. Andrewartha, H. G. and L. C. Birch. 1984. The ecological web Universit) of Chicago Press. 500 pp Arnold, \V. S. 1990. .-^ review of the biology of the bay scallop, Argopecten irradians, in Florida waters. Unpublished re- port to the Florida Marine Fisheries Commission, Florida Department of Natural Resources, Tallahassee, Florida, 59 p, Ayala, F. J. 1975. Genetic differentiation during the specia- tion process. In: Dobzhansky, T., M. Hecht and W. C. Steere (eds. ). Evolutionary Biology, Vol. II. Plenum Press, New York, pp. 1-78. Beaumont, A. R. and C. M. Beveridge. 1984. 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Developments in aquaculture and fisheries science 21. Elsevier Science Publishers, New Y'ork, pp. 625-713. Petuch, E. J. 1987. New Caribbean molluscan faunas. The Coastal Education and Research Foundation, Charlottes- ville, N'irginia, 158 pp. Rohlf, F. J. and F. L. Bookstein. 1987. as a method for "size correction. 356-367. Roughgarden, J. and Y". Iwasa. 1986 D\namics of a meta- population with space-limited subpopulations. Theoretical Population Biology 29: 235-261. Roughgarden, J, Y" Iwasa and C. Baxter 1985. Demographic theor> for an open marine population with space-limited recruitment. Ecology 66:54-67. SAS Institute, Inc. 1985. SAS user's guide: statistics. Version 5 edition. SAS Institute, Inc., Gary, North Carolina, 956 pp. A comment on shearing S\stematic Zoology 36: Sa} , T. 1822. .'\n account of some of the marine shells of the United States. Journal of the Academy of Natural Sciences of Philadelphia 2(2):257-276. Selander, R K 1970. Behasiour and genetic variation in natural populations. American Zoologist 10:53-66. Simberloff , D. 1988 The contribution of population and com- munity biology to conservation science. Annual Review of Ecology and Systematics 19:473-512. Sinclair, M.,'r. K. Mohn, G. Probert and D. L. Roddick. 1985. Considerations for the effective management of Atlantic scallops. Canadian Technical Report of Fisheries and Aquatic Sciences 1382, 97 pp. Sneath, P. H. A. and R. R. Sokal. 1973. Numerical taxonomy. Freeman, San Francisco. 573 pp. Sokal, R R and F. J Rohlf. 1981. Biometry, 2nd edition. W. H. Freeman, San Francisco, 859 pp. Tabb, D. C. and R. B. Manning. 1961. A checklist of the flora and fauna of northern Florida Bay and adjacent brackish waters of the Florida mainland collected during the period July, 1957 through September, 1960. Bulletin of Marine Science of the Gulf and Caribbean ll(4):552-649. Volckaert, F. and E. Zouros. 1989. Allozyme and physiolog- ical \ariation in the scallop Placopecten magellanicus and a general model for the effects of heteroz\ gosity on fitness in marine molluscs, Mamie Biology 103:51-61. Wall, J. R, S. R. Wall and M. Castagna' 1976. Enzymes [sic] polymorphisms and genetic variation in the bay scallop, Argopecten irradians. Genetics 83(3, part 1, Suppl.):81. Waller, T. R. 1969. The evolution of the Argopecten gibbus stock (Mollusca: Bivalvia), with emphasis on the Tertiary and Quaternary species of eastern North America. Journal of Paleontology 43:1-125. Waller, T. R. 1991. Evolutionary relationships among com- mercial scallops (Mollusca: Bivalvia: Pectinidae). In: Shumway, S. E. (ed.). Scallops: Biology, Ecology and Aqua- culture. Developments in aquaculture and fisheries science 21. Elsevier Science Publishers, New York, p 1-73. APPENDIX I Table of morphological characters and abbreviations used in this study, following Waller (1969). Abbreviations re- fer to either right or left-valve; apostrophes are added in te.xt to indicate left valve measurements. AM AD DG AK GP DF CD EI BJ DE LO ad ce PW IW Valve height Length of posterior portion of valve Length of anterior portion of valve Height of posterodorsal portion of valve Height of anterodorsal portion of valve Length of anterior ligament Length of posterior ligament Height of anterior auricle Height of posterior auricle Length from valve midline to ventral insertion of anterior auricle Valve width Height of resilial insertion Length of resilial insertion Width of medial plicae at ventral margin Width of medial interplical spaces at ventral margin THE NAUTILUS 110(2):42-44, 1997 Page 42 Subspecific Status of Argopecten irradians concentricus (Say, 1822) and of the Bay Scallops of Florida Dan C. Marelli William G. Lyons William S. Arnold P'lorida Department of Environmental Protection Florida Marine Research Institute 100 8th Avenue SE St. Petersburg, FL 33701-5095 USA Maureen K. Krause' Department of Ecology and Evolution State University of New York at Stonv Brook Stony Brook, NY 11794-5245 USA ABSTRACT Nei genetic distances between Florida and North Carolina pop- ulations of ba\' scallops, all putatively Argopecten irradians concentricus, are greater than distances between the North Carolina population and populations of A. i. irradians from New York to Massachusetts. The subspecies A. i. concentricus is probably untenable. Evidence for a subspecies, A. i. taylorae, in Florida and the eastern Gulf of Mexico is discussed. Key words: Argopecten. bay scallops, genetics, Pectinidae, subspecies. INTRODUCTION Bay scallops, Argopecten irradians (Lamarck, 1819), range from New England to Florida, Texas, and north- eastern Mexico. Northern, southern, and western popu- lations have been accorded subspecific rank as A. i. ir- radians, A. i. concentricus (Say, 1822), and A. i. ampli- costatus (Dall, 1898) (Clarke, 1965; Waller, 1969). We have found no morphometric or genetic characters to separate scallops of a recently proposed fourth subspe- cies, A. i. taylorae Petuch, 1987, from other Florida populations (Marelli et al., 1996) customarily assigned to A. i. concentricus. However, our work shows that Nei (1972) genetic distances between Florida populations and a North Carolina population of Argopecten irradians, all putativeK of A. i. concentricus, are greater than dis- tances between that North Carolina population and pop- ulations of A. I. irradians from Massachusetts, Connect- icut, and New York (figure 1), prompting us to review and reassess the subspecific status of A. i. concentricus. Say (1822) described Pecten concentricus based upon ' Current address: Southampton College, Long Island Univer- sity, Southampton, NY 11968 USA. specimens from the coast of New Jersey and also rec- ognized as distinct a variety from New England that he named Pecten borealis-. the latter name is now known to be a junior synonym of A. i. irradians. Because Say's specimens of P. concentricus are lost, Clarke (1965) se- lected a neotype from Great Egg Harbor near Atlantic City in southeastern New Jersey, thereby fixing the type locality. Clarke also restricted the type locality of P. irradians to Waquoit Bay near Falmouth, Massachusetts. Clarke (1965) evaluated the status of subspecific units among bay scallops by comparing four "primary char- acters" [plical count; width/length; height/length; and color (% white)] of shells from 23 populations throughout the range of what he called "the Argopecten irradians superspecies. Clarke's results, derived from relatively few specimens (.\ = 13) of disparate sizes (height range: 17.7 to 80.6 mm), must be viewed with uncertainty be- cause of our findings of allometric \ariabilit\ among individual populations (Marelli et at., 1996). Neverthe- less, Clarke s four samples from the "New Jersev coast "; Atlantic City, New Jersey; Sinepuxent Bay, Maryland; and Hog Island, Virginia are instructi\ e in understanding his concept of A. i. concentricus. The New Jersey coast sample (depicted as northern New Jersey but actually unlocalized) had characters consistent with four other samples from Massachusetts and Rhode Island (i.e., A. i. irradians): the Atlantic Cit\ and Sinepuxent Ba\' samples shared characters of both the more northern and more southern (North Carolina) samples; and the Hog Island sample grouped with two samples from North Carolina, which in turn most resembled five samples from the eastern Gulf of Mexico between Sanibel Island, Florida, and Ft. Morgan, Alabama. A specimen from Chandeleur Island, Louisiana, was depicted as intermediate between those of eastern and western Guli (Matagorda, Texas, to Tampico, Mexico; i.e., A. i. aniplicostatus) populations, but its plical count was clearly assignable to the eastern group. Clarke interpreted his results to indicate that northern. Dan C. Marelli et ai, 1997 Page 43 (0.026) (0.014) [(oToos) — (0.04) (0.019) (0.021) MASS CONN LINY CBNC SJFL HOM RKB 0.15 0.125 0.1 0.075 0.05 0.025 0 GENETIC DISTANCE Figure 1. UPGMA tree constructed from the Nei (1972) genetic distance matrix for populations of Argopecten irradians. See Marelli et al. (1996) for methods. Abbreviations are MASS; Martha's Vineyard, Massachusetts; CONN; Niantic River, Connecticut; LINY; Orient Harbor, Long Island, New York; CBNC; Core Banks, North Carolina; SJFL; St. Joseph Bay, Florida; HOM; Homosassa Bay, Florida; RKB; Rabbit Key Basin, Florida. Numbers in parentheses are standard errors of branching points estimated using procedures of Nei et al. (1985). southern, and western populations of A. irradians are morphologically dissimilar throughout most of their ranges but that they intergrade between New Jersey and Virginia and again in Louisiana. Despite the transitional features of scallops in the t\ pe population, Clarke tried to maintain nomenclatural stability by retaining the name concent riciis for the economically important subspecies of the southern United States. In retrospect, Clarke s findings seem to have provided more support for the existence of clines than for the existence of subspecies. The subspecies concept is regarded by some as arbi- trary because it attempts to separate populations that, in fact, may show gradual transitions in morphology, phys- iology, and genetics over the range of a species (Wilson & Brown, 1953; Endler, 1977). The classification of Ar- gopecten irradians exemplifies difficulties inherent in attempts to distinguish such populations. The genetic distances we report for bay scallops from Florida to Mas- sachusetts could indicate a cline among populations be- tween which no clear boundaries exist, and that is prob- abl\' the case among the Atlantic coast populations north of Florida. The transitional nature of morphological characters of the type population of A. i. concent ricus seems to support that explanation, as do genetic similar- ities we found among populations between North Car- olina and Massachusetts. Thus, the concept of A. i. con- centricus as a subspecies seems untenable. The subspecies concept gains strength, however, w hen it is applied to populations that are separated by distri- butional gaps or barriers that constrain contact {sensii Mayr, 1969). Waller (1969) described a distributional gap of approximately 764 km (475 miles) between the east- ernmost population of A. !. aniplicostatus and the west- ernmost population he considered to represent A. i. con- centricus. and he proposed that the Mississippi Delta acts as a barrier to genetic interchange between those stocks. Thus, subspecific designations for those populations seem appropriate. A distributional gap of similar magnitude in the south- eastern United States has received less attention, perhaps because it contains no obvious barrier. The northernmost population of bay scallops along eastern Florida is at Palm Beach Inlet (Lake Worth Inlet) (Waller, 1969) near latitude 26°45'\, and the next documented occurrence of the species seems to be at Lockwood Folly Inlet near Cape Fear, North Carolina (Porter, 1974), near SS'SO'N; these populations are separated by a minimum distance of nearly 800 km. Even if reports of uncommon occur- rence in South Carolina (Clarke, 1965; Shoemaker et ah, 1978) and an unsubstantiated report from Georgia (Pe- tuch, 1987) are considered, no population of bay scallops is known in the region between Palm Beach Inlet and the Florida-Georgia border at 30°45'N, a gap of nearly 450 km. Thus, despite reported similarities in shell mor- phology, the genetic gap between the North Carolina and Florida stocks is not a simple expression of distant populations separated by other, more contiguous popu- lations distributed along a cline. Instead, the bay scallop stocks of Florida (and probably those extending to eastern Louisiana) seem to comprise an aggregation of popula- tions isolated b\ large distances from other aggregations to the north and to the west, i.e., a subspecies, sensu Mayr (1969). If A. i. concentricus is not appropriate as a name for bay scallop stocks of Florida and the eastern Gulf of Mexico, and if those stocks merit subspecific recognition, then A. i. taijlorae Petuch, 1987, is the first available name. Another possibility, Pecten circularis Sowerby, 1835, was shown by Waller (1995) to be a junior synonym of Argopecten irradians concentricus. Dr. Waller in- Page 44 THE NAUTILUS, Vol. 110, No. 2 formed us (in litt.. 1/30/96) that the holot\pe of Pecten circiilaris "compares favorabl\ w ith North Carohna con- centricus in overall shape, auricular shape, rib shape, and rib count, and it differs in these features from Gulf populations." We advocate caution to those who might immediately appl\- the name Argopccten irradians taijlorac for the Florida and eastern Gulf stocks. Our study (Marelli et al., 1996) was intended only to assess differences between t\pical A. 1. taylorae and other bay scallops of Florida. Those differences were refuted using analytical tech- niques involving both genetics and morphometries. Sim- ilar work is needed to elucidate the genetic compositions of populations in the transition zone between New Jersey and \'irginia, in South Carolina and Georgia (if such populations exist), in the region between northwestern Florida and eastern Louisiana, and in the western Gulf of Me.xico (A. !. amplicostatus) to provide perspective for evaluating genetic distances among all of the popu- lations before the question of subspecies is resolved. ACKNOWLEDGEMENTS Dr. Arthur Bogan, then at the Academy of Natural Sci- ences of Philadelphia, provided a copy of Say (1822). Dr. Thomas R. Waller, National Museum of Natural History, Washington, D. C, provided additional infor- mation on the holotype of Pecten circiilaris. This project was partially supported by funds created by the Florida Saltwater Products and the Florida Saltwater Fishing License. LITERATURE CITED Clarke, A. H, Jr. 1965. The scallop superspecies Aequtpec/en irradians (Lamarck). Malacoiogia 2:161-188, Dall, W. H. 1898. Contributions to the Tertiary fauna of Florida, with especial reference to the Miocene silex beds of Tampa and the Pliocene beds of the Caloosahatchie River; Wagner Free Institute of Science Transactions, Vol. 3, Part 4, 377 p. Endler, J. A. 1977. Geographic variation, speciation, and clines. Princeton Universitv Press, Princeton, New Jersey, 246 p. Lamarck, J. B P. A de VI de. 1819. Histoire natiirelle des animaux sans vertebres. Paris, Vol. 6, Part 1, 343 p. Marelli, D. C, M. K. Krause, W. S. Arnold and W. G. Lyons. 1 997. Systematic relationships among Florida populations of Argopccten irradians scnsu lata (Lamarck. 1819) (Bi- valvia: Pectinidae), The Nautilus 110(2):31-41. Mayr, E. 1969. Principles of svstematic zoologv. McGraw- Hill, New York, 428 p. Nei, M. 1972. Genetic distance between populations. The American Naturalist 106: 283-292. Nei, M., J. C. Stephens and N. Saitou. 1985, Methods for com- puting the standard errors of branching points in an evo- lutionary tree and their application to molecular data from humans and apes. Molecular Biolog\ and E\ olution 2: 66- 85, Petuch. E, J, 1987 New Caribbean molluscan faunas. The Coastal Education and Research Foundation, Charlottes- ville, Virginia, 158 p. Porter, H, J. 1974, The North Carolina marine and estuarine Mollusca — an atlas of occurrence. University of North Car- olina Institute of Marine Sciences, Morehead City. North Carolina, 351 p. Say, T, 1822. .\u account of some of the marine shells of the United States, Journal of the Academs' of Natural Sciences of Philadelphia 2(2):257-276, Shoemaker, A, H,, H, J, Porter. B. Boothe, R. E. Petit and L. S, Eyster, 1978, Marine mollusks. In: R, G, Zingmark (ed.) An annotated checklist of the biota of the coastal zone of South Carolina. University of South Carolina Press, Co- lumbia, SC. p. 123-135. Waller, T. R. 1969. The evolution of the Argopecten gibhus stock (Mollusca: Bivalvia), with emphasis on the Tertiary and Quaternar\ species of eastern North .America, Journal of Paleontology 43:1-125 Waller. T. R 1995. The misidentified holot\ pe of Argopecten circularis iBi\aK ia: Pectinidae), The \eliger 38: 298-303, Wilson, E. O. and W. L. Brown. 1953. The subspecies concept and its taxonomic applications. Svstematic Zoologv 2:97- 111. THE NAUTILUS 110(2):45-54, 1997 Page 45 Distribution of Unionoid (Bivalvia) Faunas in Minnesota, USA Daniel L. Graf Department of Biology- Northeastern University Boston, MA 02115 USA and Department of Mollusks Museum of Comparative Zoology Cambridge, MA 02138 ABSTRACT The freshwaters of Minnesota are significant because they form the conjunction of three major drainage s\ stems: the Mississippi, the St, Lawrence, and Hudson Bay; 46 species of freshwater mussels (Bivalvia: Unionoidea) inhabit these rivers and lakes. Of these, 31 are found only in the Mississippi River and its tributaries below Minneapolis. Nine species have distributions that include all of the state's drainage systems; 5 species have been reliably reported from only the Red River of the North and Lower Mississippi River drainage systems; and Elliptio complanata is found only in the Lake Superior Basin. Prob- lematical records within the state are addressed. Key words: Biogeography, Unionidae, Margaritiferidae, fresh- water mussels. INTRODUCTION No equal area on earth has such a diversity of Naiad life or such magnificent shells. Here are found the largest species in the world; here are forms with knobs, pustules, angles, lobes, and concentric sculpture. The nacre of many of them is wonderfully rich in tints ot silver, pink, purple, salmon or red, and it is equaled in beauty by the elegant patterns of e.xternal painting, in stripes and mot- tlings and delicate hair lines. Charles T. Simpson (1896) on the Lhiionoidea of the Mississippi Valley The distributions of the 46 species of freshwater mussels (Bivalvia; Unionoidea) that inhabit Minnesota were pre- sented by Charlotte Webster Davvley in her doctoral thesis (1944) and a widely cited paper (Dawley, 1947). However, additional data have accumulated in the near- ly half century since then. This paper seeks to update the literature record and establish the known distribu- tions of these mollusks. The life-cy cle of a freshwater mussel involves the use of one or more species of fish or, in one case, an am- phibian as a host for the parasitic larva or glochidium. In general, the major phase of dispersal occurs via par- asitism of their host. Thus, barriers that inhibit the va- gility of fish (drainage divides, waterfalls, etc.) also block the dispersal of the Unionoidea. Hydrology of Minnesota. Minnesota's surface area of 218,500 km- is drained by three divergent watersheds (Figure 1 ). The Lake Superior Sy stem, the smallest, drains to the St. Lawrence River via Lake Superior. It is defined as the lake itself and all its Minnesota tributaries. This system drains about 15,300 km- or 7% of the state. The next largest watershed, draining 76,500 km- or 35% of the state, runs to Hudson Bay and is subdivided into the Red River System (the Red River of the North and all its Minnesota tributaries) and the Lake of the Woods System (the Rainy Ri\ er and all Minnesota waters draining to Lake of the Woods). The two systems are confluent at Lake Winnipeg, Manitoba, but because they have different mussel faunas, they are considered sepa- rately. The remaining 126,700 km- (58''() drains to the Gulf of Mexico via the Mississippi River. The basin is subdi- vided into the Upper and Lower Mississippi River Sys- tems (following L'nderhill, 1989) based on their divergent histories and \astly disparate mussel faunas. The Upper Mississippi River System is that portion of the Mississippi River and all of its tributaries above the Falls of St. Anthony at Minneapolis. The Lower Mississippi River System is the river and all of its Minnesota tributaries below the Falls of St. Anthony. This includes not only those watersheds draining directly into the Mississippi River in Minnesota (the St. Croix, Minnesota, Zumbro, Cannon, and Root River Watersheds), but also those wa- ters in southwestern Minnesota draining to the Missouri and Des Moines Rivers. For the purposes of this study, state boundaries formed by streams extend to the opposite bank to relax possible political tensions among mussels on opposing sides of the same river. These five drainage systems were in place following the final northward retreat of Glacial Lake Agassiz. Be- fore that time, the enormous volumes of meltwater left Page 46 THE NAUTILUS, Vol. 110, No. 2 Figure 1. The Drainage Systems of Minnesota. The Red River System (RRS) and Lake of the Woods System (LVVS) are con- fluent at Lake Winnipeg, eventually emptying to Hudson Bay. The Lake Superior System (LSS) drains to the Great Lakes. The remainder of the state drains via the Mississippi and is comprised of the Upper Mississippi River System (UMRS) and Lower Mississippi River, Minnesota River, and St. Croi.x River subsystems (LMRS, MRS, and SCRS, respectively). See te.xt for e.xplanation of the state s drainage systems. in the wake of wasting glaciers over-ran present drainage divides and connected now dissociated basins. At such times, unionoids were free to migrate up the Mississippi River from southern refugia (Johnson, 1980) and into adjacent basins until water levels receded. MATERIALS AND METHODS Species have been assigned to watersheds using not only specimens housed in the University of Minnesota's James Ford Bell Museum of Natural History (JFB) Invertebrate Collection but al.so from an extensive literature survey. In the Appendi.x, for each species, a single lot is listed for each basin when available, generally those used by earlier authors. The following literature references where utilized to establish the distributions of the Unionoidea in each of the above defined Minnesota drainage systems. No ar- cheological or fossil literature has been included. LakeSuperiorSystem;Dall(1905), Dawley (1944, 1947), Goodrich and van der Schalie (1932: Lake Superior in general), Moyle (1947), Walker (1913). Lake of the Woods System: Baker (1929, 1935), Clarke (1973), Dawley (1944, 1947). Red River System: Baker (1929), Clarke (1973), Coker and Southall (1915), Cvancara (1966, 1967, 1970, 1977, 1979, 1983), Cvancara et al. (1981), Daniels (1909), Dawley (1944, 1947), Grant (1885), MN DNR (1984, 1986), Radke (1992), Sargent (1895). Wilson and Danglade (1914). Upper Mississippi River System: Baker (1929), Bright (1988), Dawley (1944, 1947), Grant (1885), Movie (1940), Sargent (1895), Wilson and Danglade (1914). Lower Mississippi River Subsystem: Baker (1928), Bright (1988), Bright ct al. (1989),' Coon et al. (1977), Cooper (1834), Davis (1990), Dawley (1944, 1947), Fuller (1978, 1980), Grant (1885), Grier (1922, 1926), Grier and Mueller (1922); Havlik (1981), Holzinger (1887), Hornbachef al. (1992), Johnson (1980), Mathiak (1979), Nachtrieb (unpublished), Southall (1925), Thiel (1981), van der Schalie and van der Schalie (1950), Wilson and Danglade (1914). Minnesota River Subsystem: Bright et al. (1990), Dawley (1944, 1947), Grant (1885). MN DNR (1985), Nach- trieb (unpublished), Wilson and Danglade (1914). St. Croix River Subsystem: Baker (1928), Bright (1988), Cooper (1834), Dawley (1944, 1947), Fuller (1978, 1980), Imlay (1972), Mathiak (1979), Stern (1983), Wilson and Danglade (1914). Wilson and Danglade (1912) and Ellis (1931), provided only vernacular names, but these have been reworked in a scientific context by Wilson and Danglade (1914) and van der Schalie and van der Schalie (1950), respecti\el\. Grant (1887) reported collecting unnamed Anodonta and Lampsilis from unspecified watersheds of the Arrowhead Region of northeastern Minnesota. Few data are avail- able on the Unionoidea of the Lake Superior System. Of the general Great Lakes literature reviewed (e.g., van der Schalie, 1961; Walker, 1913), most study has centered on the eastern lakes, emphasizing Lake Erie. The ref- erences cited aboxe provide almost no information on the mussels found in the streams of the North Sliore of Lake Superior (with the exception of the St. Louis River). Smith and Moyle (1944), however, sur\e\ed the fauna of these streams, including the macrobenthos, and re- ported no unionoids. Since Moyle, in his other reports (1940, 1947), has provided mussel data it can be assumed that no unionoids are present in these streams. For mussel distributions adjacent to Minnesota, the reader is referred to the follow ing references: North Da- kota: Cvancara (1966, 1967, 1970, 1975, 1976, 1983), Cvancara et al. (1966, 1972, 1976); South Dakota: Coker and Southall (1915), Over (1913); Canada: Clarke (1973, 1981); Wisconsin: Baker (1928), Havlik and Stansbery, (1977), Mathiak (1979), Stern (1983). RESULTS Figure 2 presents the distributions of the Unionoidea in Minnesota. For each species, the systematic assignment follows Williams et al. (1993), which claims to provide Daniel L. Graf, 1997 Page 47 w; ■r. ^ ._ y, __ y: 5 t^ — — L. ■J-. J-. y. 'S' L- i> r. J. X 3 ^ X c •^ L. u ii ^ J^ < = u Upper Mississippi River Faina — iMsinifioiui tiimplanaia (B;irnes, IS23) | Lasmi^ona contpri_'SSti (Lea. IS2'^)) | Liunpsih:! ciirdium (Rafinesque. IS20) | Pygdiioiioii grandis (Say. 1824) | Ugiimia recta (Lamarck. 1X19) | LumpsiUs siliquoidea (Barnes. 1823) | AnodonioiJesfenissMioints {Lea. 18.'14) | Slrophiuis iiiidiilcinis (Say. 1817) Q UilerlnuUa imbecillis (Say. 1829) Q D D D D Red River of the North Fai na Piikimiliis aliiliis (Say. 1 8 1 7 ) □ □ l.usinigonu coskira (Ra{\ni:f.quc. 1820) Q [31 FiiscoiHiici Jlava (Rafinesque. 1820) Q Q Ambleniii piuala (Say. 1817)0] CI yi/ Lake Superior Fauna omphiiuna (Lightfool. 1786) | D D nnnn _ -Specimen with LiteraUire Citalicni or VoLiclier only H Literature Citation only \T\ Problematical Record (see te.xt) □ Not Reported Lower Mississippi River Faina Sinipsontilas tunhigiui (Say. 1825) I'oHiiiidus capax (Green. 1832) Pli'iiiiibinui ciHiineum (Conrad, 1836) Anidens ciinpdgiisiis (Say, 1819) Elliplid cnissidtns (Lamarck. 1819) PUiliiihasus cyphyiis (Rafinesque. 1820) Eihptui dilataiiis (Rafinesque. 1820) Tnincillii do/Hicifoimis (Lea. 1828) FiiscdiHiui fheiHi (Lea. 1831 ) Vi'iuiskuonclui ellipsifdiniis (Conrad. 1836) Leplodcii Jrcigilis (Rafinesque. 1820) QiiadndajragDSci (Conrad. 1836) Liiinpsilis higgiiisi (Lea. 1857) Airniiiiiiiuis ligamenlina (Lamarck. 1819) Ellip.stiriii llneiiliila (Rafinesque. 1820) Alasiiiulanki iihiri^iiuiki Say 1819 QiuidntUi mekinevw (Rafinesque. 1820) CiimherUindia nuinodonui (Say. 1829) Mfgalamuas nenosii (Rafinesque. 1820) Qiiiidnilu nitdiilaki (Rafinesque. 1820) Ptiranuliis (ihieiisis (Rafinesque. 1820) I Ihiivcinci olivtirui (Rafinesque, 1820) Ttixitltisina purvtis (Banies, 1823) Qiiadruhi pustuUtsci (Lea. 1831) Oblicpuiriu njh'xii Rafinesque. 1820 Anddiinia subdibuuUiki Say 1831 Lanipsitis teres (Rafinesque, 1820) Epiiiblasmu irkptetra (Rafinesque, 1820) TrunciUii Inincuia (Rafinesque. 1820) Cyclimaias iiibcriiiUita (Rafinesque. 1820) Tntiifiiiiiia tcrnuiisti (Rafinesque. 1820) = :/: '^- i; E c o 4> lis! O & .£ .i ^ > t_ X ■' fi " '^ o j: L. 3 4h > "=<*;- C/D o ^ u u S i; 01 4* -^ a 5 = ^ n « O 5" ^ s -j; n n D DDHB n n n nnnn n n n niBB n n n nam n n n nBHD D n n nann n n n naaa n n D naaa n n n naaa D D D nana n n f"i naaa D n D naan D n D naaa n n D naaa n n D naaa n D D naaa n D D naaa n n D nana D n n naaa n n D nnaa n n D naaa D D D naaa n n D naaa n n D naaa D n D naaa n n n nnnn D D D naan n n D nnna n n n naaa D D D nana n n D naaa Figure 2. The Distribution of L'nionoid P^aunas in Minnesota. the same list as Turgeon et al. (1988) but with spelling and other such errors corrected. Figures depicting these mollusks can be found in Cummings and Mayer (1992) and Fuller (1985). The only exception, Elliptio com- planala (Lightfoot, 1786), is illustrated in Clarke (1973: 56-57, plate 3). The species list includes only those spe- cies verified to occur in Minnesota; distributions pre- sented exclude erroneous records (see Problematical Re- cords below). The Lower Mississippi Drainage System is divided into three subsystems: the Lower Mississippi River Subsystem, excluding the Minnesota and St, Croix River watersheds; the Minnesota River Subsystem, w hich includes the main stem of the river and all its tributaries; and the St. Croix River Subsystem, also including the main stem of that river and its tributaries. Questionable and Problematical Records. Problemati- cal records, both from literature reports and JFB speci- mens, may lead to false hy potheses; those identified by the author are summarized and dealt with below . Questionable Literature Records. Four species reported from the Red River System are supported by only a single literature citation each: Qtiadrula pustulosa by Coker and Southall (1915), Pleiirobetna coccineum by Wilson and Danglade (1914), Elliptio dilatata by Daniels (1909), and Obliquaha reflexa h\ Dawley (1944, 1947). How- ever, no corroborating voucher specimens exist at the JFB and none of these species were collected by Cvancara (1966, 1967, 1970, 1975, 1976, 1983; Cvancara et al, 1966; 1972; 1976) or others (e.g. Clarke, 1973) who have extensively surveyed basin. Qiiadrula pustulosa, P. coc- Page 48 THE NALTILLS. Vol. 110. No. 2 cineum, and E. dilatata are likely incorrectly identified Q. quadrula, Fusconaia flava. and Ligumia recta, re- spectively. In the case of O. reflexa, both Cvancara (1970) and Clarke (1973) reported personal communications with Da\vle\ in which she admitted her misidentification. The true distributions of these four species are probably lim- ited to onl\- the Lower Mississippi River System. Problematical Voucher Specimens. The presence of Tritogonia verrucosa in the Upper Mississippi River Sys- tem is indicated by an unpaired valve (JFB 3190) col- lected b> John Moyle (1940). Mo\le believed the valve to be evidence of Lower Mississippi species overcoming the Falls of St. Anthony but not surviving because of poor habitat. Dawley (1944, 1947) considered the pres- ence of Tritogonia to be an accident, and her opinion is likeK correct. Thus, the distribution of T. verrucosa is limited to only the Lower Mississippi River System. An unpaired valve of Amblema plicata (JFB 7567) was col- lected from the Cloquet River in the Lake Superior Sys- tem. However, since A. plicata has otherwise not been reported from that system in Minnesota or even from the Lake Superior Basin in general, this single specimen is probably not a remnant of a population and is assumed to be trash or the remains of some introduced individual or individuals. Amblema plicata has been reliably re- ported from only the Red and Lower Mississippi River Systems, and this presumably reflects the mussel's true distribution within the state. The problematical distribution of Acttnonaias liga- mentina in the Minnesota has been addressed by Clarke (1973). Dawley "s Red River specimen could not be lo- cated, but records of A. ligamentina reported by her (1944, 1947: JFB 2863), Moyle (1940: JFB 2872), and Radke (1992: JFB 7569) have been re-identified as Lamp- silis siliquoidea. Cvancara (1970) disregarded Dawley "s Red River voucher, reporting that in a personal com- munication, she agreed that her record was probably an error. Thus, Actinonaias is limited to only the Lower Mississippi River System. The single lot, JFB 2332 Lampsilis teres, labeled as coming from Leech Lake in the Upper Mississippi River System is likely the result of a cataloging error. Since Dawley (1944, 1947) apparently ignored it, the validity of this record is questionable. Other Problematical Data. Utterbackia irnbecillis has never been reported from any of the Lake Superior or Hudson Bay drainage systems. This makes L'. irnbecillis unique among species reliably reported from the Upper Mississippi Ri\er System. Also making V. irnbecillis dis- tinctive is the fact that it mav complete its reproductive cycle with or without a glochidial host (Dawley, 1944; Fuller, 1985). This would hamper its upstream vagility, and perhaps 17. imbecillis could not extend beyond the Upper and Lower Mississippi River Systems before drain- age connections were severed. The presence of Anodontoides ferussacianus in the Lake of the Woods S\stem is not supported by a specimen in the JFB, though both Baker (1935) and Dawley (1944, 1947) reported the species from that basin. Strophitus undulatus has never been reported from the Lake Superior System and its presence in the Lake of the Woods System is supported by only a single lit- erature record: Clarke (1973). Dawley (1944, citing Le- fevre & Curtis, 1911) noted that S. undulatus is another mussel that can complete its life-cycle independent of a glochidial host. However, the mussel is not obligately anomalous in its reproduction; its widespread distribu- tion indicates that it probably is dispersed by fish at least some of the time. Three of its published glochidial hosts (Hoggarth, 1992) are found in the that watershed as well as Minnesota's four other drainage systems (Underbill, 1989), providing a means of statewide distribution. The apparent absence of S. undulatus may actually be the result of the lack of an adequate surve> of the St. Louis River System. DISCUSSION There is a pattern to the distributions of the Unionoidea in Minnesota. Species with similar distributions can be grouped into faunas; members of each fauna are h\- pothesized to have shared modes and tempos of dispersal into the state s drainage systems. Ninety-eight percent of the species (45 of 46) are found in the Lower Missis- sippi River System, supporting the hypothesis that the freshwater mussel fauna oi Minnesota dispersed north- ward from southern refugia by way of the Mississippi River at the close of the Pleistocene (Johnson, 1980). The remaining species, Elliptio complanata, appears to have migrated into the state via Lake Superior from the St. Lawrence watershed. Of the 45 species of unionoids reported from the Low- er Mississippi River System, 30 have been reliably re- corded from onl\ that system; these mussels constitute the Lower Mississippi River Fauna (Figure 2). Much of this fauna is made up of mussels ecologically limited to larger rivers that were unable to extend their ranges beyond the Lower Mississippi and the lowest reaches of the Minnesota and St. Croix Ri\ ers (e.g., Fusconaia ebena, Ellipsaria lineolata). Others of the species characteristic of the Low er Mississippi River System ma\ be relatively recent arrivals to the state (e.g., Anodonta suborbiculata, Quadrula nodulata) since the> were not reported by earlier authors. Those freshwater mussels commonly found above the Falls of St. .^nthonx' are collectively know n as the Upper Mississippi River Fauna (Figure 2). The Upper Missis- sippi Ri\ er Fauna must ha\ e been present at a time w hen all ot the states watersheds were connected. Of the 9 species that belong to this assemblage, only Utterbackia imbecillis and Strophitus undulatus have not been re- ported from all five of Minnesota s drainage systems (dis- cussed above). Five mussel species are found oni\ in the Red and Lower Mississippi River Systems; these belong to the Red River of the North Fauna (Figure 2). .\t the close of the Daniel L. Graf. 1997 Page 49 Pleistocene, with the formation of Glacial Lake Agassiz, these t\\o drainage SNStems were connected more than once, and this allowed the dispersal of the Red Ri\er of the North Fauna from the Minnesota River basin into that of the Red River of the North. Elliptio complanata is the only unionoid of the Lake Superior Fauna, It is known oiiK' from the Lake Superior System. It has probabK migrated into Minnesota in the last 80 years or so from the eastern Great Lakes. The problematical species discussed above are difficult for exactly the reason that the\ don not fit neatly into any of the four described unionoid faunas. However, except for the case of Actinonaias ligarnentina. these problems are the result of single specimens. Further study of the unionoid faunas of the Lake of the Woods, Lake Superior, and Upper Mississippi River systems promises to advance our knowledge on the dis- tributions of freshw ater mussels of the Minnesota region. ACKNOWLEDGMENTS This paper is dedicated to my mentor and friend, the late Dr. Robert C. Bright of the James Ford Bell Museum of Natural History, University of Minnesota. Dr. Bright granted me the encouragement, resources, and freedom to completeK reorganize and computerize the museum's unionoid collection; this paper is the result. Professor James C. Underhill, also of the Bell Museum, provided invaluable intellectual and moral support. N'ersions of this manuscript were criticalK reviewed by J.C. Under- hill, E. Ruber of Northeastern University, and K.J. Boss and R.I. Johnson of the Museum of Comparative Zoology; all provided worthwhile input. Two anonymous review- ers suggested numerous improvements. Thanks also to C. Gatenby of Virginia Polytechnic Institute for iden- tifying those specimens that I was unable to acquire, T. Deneka of Macalester College for providing data on spec- imens he has collected, and all others w ho have assisted me directK and indirectly in completing this study. LITERATURE CITED Baker, F. C. 1903. Shell collecting on the Mississippi. Nautilus 16: 102-10.5. Baker, F. C. 1928. The fresh water Mollusca of Wisconsin: Part II. Pelecypoda. Bulletin of the Wisconsin Geological and Natural History Surve\ , vol. 70, No. 2. L'niversity of Wisconsin. 495 p. Baker, F. C. 1929. Mollusca from \'ermilion and Pelican Lakes, Minnesota, with the description of a new variety of Helisoma corpulenla. The Nautilus 42:95-97, 131-136. Baker, F. C. 1935. Land and freshwater Mollusca from North Star Lake and vicinity, Itasca Co., Minnesota: a systematic and ecological studv American Midland Naturalist 16: 257-274. Bright, R C. 1988. Mollusks In: Coffin, B., L. Pfannmuller (eds. ). Minnesota's Endangered Flora and Fauna. Univer- sity of Minnesota Press, Minneapolis, MN for the Natural Heritage and Non-Game Wildlife Programs of the Divi- sion of Fish and Wildlife, Minnesota Department of Nat- ural Resources, pp. 397-406. Bright, R. C, E, Plummer and D. Olson. 1989. A survey of the mussels of the Zumbro River Drainage, Southeastern Minnesota. Bell Museum of Natural History, University of Minnesota, St. Paul, Minnesota, pp. 1-27 -I- figures -i- plates -I- appendix. Bright, R., C. Gatenby, D. Olson and E. Plummer. 1990. A survey of the mussels of the Minnesota River, 1989. Bell Museum of Natural History, University of Minnesota, St. Paul, MN. pp. 1-36 -I- figures + plates + appendix. Clarke, A. H. 1973. The freshwater Molluscs of the Canadian Interior Basin. Malacologia 13:1-509. Clarke, A. H 1981. The Freshwater Molluscs of Canada. National Museum of Natural Sciences. National Museums of Canada, Ottawa, Canada. 446 p. Coker, R. E. and J.B. Southall 1915. Mussel resources in tributaries of the Upper Missouri River. Appendix IV to the Report of the U.S. Commissioner of Fisheries, 1914 (Bureau of Fisheries Document No. 812). 17 p. Coon,T. G.,J. W. EckbladandP M, Trygstad. 1977. Relative abundance and growth of mussels (Mollusca: Eulamelli- branchia) in pools 8, 9 and 10 of the Mississippi River. Freshwater Biolog\ 7:279-285. Cooper, W. 1834. List of shells collected b\ Mr. Schoolcraft in western and North Western Territory. Appendix to Narrative of an Expedition through the Upper Mississippi to Lake Itasca, etc. under direction of Henry B. School- craft. New York, Harper and Bros. pp. 153-156. Cummings, K. S. and C. A. Mayer. 1992. Field Guide to Freshwater Mussels of the Midwest. Illinois Natural His- tory Survey, Manual 5. 194 p. Cvancara, .\. M. 1966. Distribution and ecology of mussels in the Red River Valley, Grand Forks to Dayton. Pro- ceedings of the North Dakota Academy of Science 20: 156-157 (abstract) Cvancara, A. M. 1967. Mussels of the Red River Valley in North Dakota and Minnesota and their use in deciphering drainage history. In: Mayer-Oaks, W.J. (ed.). Life, Land and Water. University of Manitoba Press, Winnipeg, Man- itoba, pp. 187-196. Cvancara, A. M. 1970. Mussels (Unionidae) of the Red River \'allev in North Dakota and Minnesota, U.S..\ Malacologia 10:57-90. Cvancara, A. M. 1975. Aquatic malacogeography of North Dakota. Proceedings of the North Dakota Academy of Science 27:68-82. Cvancara, A. M. 1976. .Aquatic mollusks in North Dakota during the last 12 000 vears, Canadian Journal of Zoology 54:1688-1693. Cvancara, .\. M. 1977. Comparison of mussels (Bivalvia: Unionacea) collected during high and low water on the Red River of the North. Prairie Naturalist 9:54-56. Cvancara, A. M. 1979. Larger lake than stream mussels (Bi- valvia: Unionacea): an exception to the rule in north- western Minnesota. Proceedings of the North Dakota Academy of Science 33:23, Cvancara, A. M. 1983. Aquatic Mollusks of North Dakota. Report of Investigation No. 78. North Dakota Geological Survey. Kaye's, Inc., Fargo, ND 141 p. Cvancara, A. M., D. J. Brown, D K. Cudworth and T R. Klett. 1981, Mass mortality of mussels from slumping along the Red Lake River near Crookston. Minnesota, Prairie Nat- uralist 13: 13-14. Cvancara, .A. M., J.M. Eriksonand J, J, Delimata, 1972. Pres- Page 50 THE NAUTILUS, Vol. 110, No. 2 ent and past Mollusks of the Forest Ri\er. North Dakota. Proceedings of the North Dakota Academy of Science 25: 55-65. Cvancara, A. M., R. G. Heetderksand F. J Iljana. 1966. Local distribution of mussels. Turtle River, North Dakota. Pro- ceedings of the North Dakota Academy of Science 20; 149-155. Cvancara, A. M, R. D. Norby, and J.B. Van Alstine. 1976. Mollusks of the Sheyenne River, North Dakota, USA: pres- ent and past. Malacologicai Review 9:25-38. Dall, W, H. 1905. Land and freshwater mollusks of Alaska and adjoining regions Harriman Alaska, E.\pedition 13: 1-171. Daniels, L. E. 1909. Records of Minnesota Mollusks. The Nautilus 22:119-121. Davis, M. 1990. Freshwater mussels (Mollusca: Bivalvia: Unionidae) of the Cannon River drainage in Southeastern Minnesota. Minnesota Department of Natural Resources Non-Game Wildlife Report, pp. 1-22. Dawley, C. W. 1944. Distribution and growth studies of the Unionidae and aquatic Gastropoda found in Minnesota. Ph.D. Thesis, University of Minnesota 307 p. Dawley, C. VV. 1947. Distribution of aquatic mollusks in Minnesota. American Midland Naturalist 38:671-697. Ellis, M. M. 1931. Some factors affecting the replacement of commercial freshwater mussels. United States Bureau of Fisheries Circular 7:1-10, Fuller, S. L. H. 1978. Freshwater mussels (Mollusca: Bivalvia: Unionidae) of the Upper Mississippi River: Observations at selected sites within the 9-foot channel navigation pro- ject on behalf of the United States Army Corps of Engi- neers. The Academy of Natural Sciences of Philadelphia, Division of Limnologv and Ecology, Report No. 78-33:1- 401. Fuller, S. L. H. 1980. Freshwater mussels (Mollusca: Bivalvia: Unionidae) of the Upper Mississippi River: Observations at selected sites within the 9-foot channel navigation pro- ject for the St. Paul District, United States Army Corps of Engineers. The Academy of Natural Sciences of Phila- delphia, Division of Limnology and Ecology, Report No. 79-24F: 1-175 -I- Appendices.' Fuller, S. L. H. 1985. Freshwater Mussels of the Upper Mis- sissippi River. Wisconsin Department of Natural Re- sources, Madison, WI. Based on a poster by the same name published by the United States Fish and Wildlife Service and the United States Army Corps of Engineers. 63 p. Goodrich, C. H. and H. van der Schalie. 1932. I. On an increase in the Naiad fauna of Saginaw Bay, Michigan; 11. The Naiad species of the Great Lakes. Occasional Papers of the Museum of Zoology, University of Michigan 238: 1-14. Grant, U. S. 1885. Conchological Notes. Minnesota Geological and Natural History Survey Annual Report 14:114-124. Grant, V. S. 1887. Notes on the Molluscan fauna of Minnesota. Minnesota Geological and Natural Histors Survey Annual Report 16:481-484. Grier, N. M 1922 Final report on the stud> and appraisal of mussel resources in selected areas of the L'pper Missis- sippi River. American Midland Naturalist 8: 1-31. Grier, N. M. 1926. Report on the study and appraisal of the mussel resources in selected areas of the Upper Mississippi River, 1920-25. American Midland Naturalist 10:89-110, 113-134. Grier, N. M. and J. F. Mueller. 1922. Notes on the naiad fauna of the Upper Mississippi River: II. The naiads of the Upper Mississippi Drainage. The Nautilus 35:46-49, 96- 103. Havlik, M. E. 1981, The northern extension of the range of Anodonta suhorbiculata Say (Bivalvia: Unionidae). The Nautilus 95:89-90. Havlik, M. E. and D. H. Stansberv . 1977. The naiad mollusks of the Mississippi River in the vicinity of Prairie du Chien, Wisconsin Bulletin of the American Malacologicai Union, Inc., 1977:9-12. Hoggarth, M. A. 1992. An examination of the glochidia — host relationships reported in the literature for North American species of Unionacea (Mollusca: Bivalvia). Mal- acology Data Net 3:1-30. Holzinger, J. M. 1887. Notes on the Mollusca of Winona County. Minnesota Geological and Natural History Survey Annual Report 16:485-491. Hornbach, D. J., A. C. Miller and B. S. Payne. 1992. Species composition of the mussel assemblages in the upper Mis- sissippi River. Malacologicai Review 25:119-128. Imlay, M. J. 1972, Reproduction of Amhlema cosiata (Raf- inesque) of the Moose River, Minnesota, The Nautilus 85: 146, Johnson, R. I. 1980. Zoogeography of North .\merican Union- acea (Mollusca: Bivalvia) north of maximum Pleistocene glaciations. Bulletin of the Museum of Comparative Zo- ology 149:77-189. Lefevre, G. and W. C. Curtis. 1911. Metamorphosis without parasitism in the Unionidae. Science 13:893-865. Mathiak, H. .\. 1979. A River Survev of the I'nionid Mussels of Wisconsin, 1973-1977. Sand Shell Press, Horicon, WI. 75 p. Minnesota Department of Natural Resources. 1984. Biological survey of the Otter Tail River. Special Publication No. 137. Minnesota Department of Natural Resources, Divi- sion of Fish and Wildlife. Table 26. Minnesota Department of Natural Resources. 1985. Biological survey of the Minnesota River. Special Publication No. 139. Minnesota Department of Natural Resources, Divi- sion of Fish and Wildlife, p, 84, Minnesota Department of Natural Resources, 1986. Biological survey of the Red River of the North, Special Publication No. 142. Minnesota Department of Natural Resources, Division of Fish and Wildlife, p. 60. Moyle, J B 1940. A biological survey of the Upper Mississippi River System. Fisheries Research Investigational Report No. 10. Minnesota Department of Conservation, Division of Game and Fish. Pages imnumbered. Moyle, J. B. 1947. .A biological survey and fishery manage- ment plan for the streams ot the Saint Louis River Basin. Fisheries Research Investigational Report No. 69. Min- nesota Department of Conservation, Division of Game and Fish. 112 p. Nachtreib, H. F. unpublished. Report on the pearl mussel investigations of the Minnesota River and its tributaries, made in 1908. Report to the United States Bureau of Fish- eries. Over, W. H. 1913. Mollusca of South Dakota. The Nautilus 27:78-81, 90-95. Radke, S. 1992, Otter Tail River fishes and nnissels: an old connection. Plan B Master s Thesis, Lniveristy of Minne- sota. 31 pp. Sargent, H. E. 1895. Annotated list of the Mollusca found in the vicinity of Clearwater Lake, Wright County, Minne- sota: Part Second — Aquatic Species. The Nautilus 9:125- 128. Daniel L. Graf, 1997 Page 51 Simpson, C. T, 1896. The classification and geographical dis- tribution of the pearly fresh-water mussels. Proceedings of the United States National Museum 18:295-343 Smith, L. L., jr. and J. B. Movie. 1944. A biological survey and fishery management plan for the streams of the Lake Superior North Shore Watershed. Technological Bulletin No. 1, MN Department of Conservation. Di\ ision of Game and Fish. Southall, J B 1925 Mussel survey of Lake Pepin in 1924 with a discussion of the effects of the system of alternate closure of sections of the lake to the taking of mussels. United States Bureau of Fisheries Economic Circular 57: 1-3. Stern, E. M. 1983. Depth distribution and densit>' of fresh- water mussels (L'nionidae) collected with SCLiB-i^ from the Lower Wisconsin and St. Croi.x Risers. The Nautilus 97:36-42. Thiel, P. A. 1981. .-V survey of unionid mussels m the Upper Mississippi River (Pools 3 through 11). Technical Bulletin No. 124 Department of Natural Resources, Madison, WL 23 p. Turgeon, D. D., A E Bogan, E \' Coan, W K Emerson, W. G. Lyons, W. L Pratt, C. F. E. Roper, A Scheltema, F. G. Thompson and J. D Williams. 198S. Common and scientific names of aquatic in\ertebrates from the United States and Canada: moUusks. American Fisheries Society Special Publication, 16. 277 p. Underbill, J. 1989. The distribution of Minnesota fishes and Late Pleistocene glaciation. Journal of the Minnesota Academy of Science 55:32-37. van der Schalie, H. 1961. The naiad (fresh-water mussel) fauna of the Great Lakes. Great Lakes Research Division, Institute of Science and Technology, University of Mich- igan. Publication No. 7: 156-157. van der Schalie, H. and .A. van der Schalie. 1950. The mussels of the Mississippi River. American Midland Naturalist 44: 448-466. Walker, B. 1913. The unionid fauna of the Great Lakes. The Nautilus 27:18-23, 29-34, 40-47, 56-59. Williams, J.. M. Warren, K. Cummings, J. Harris and R. Neves. 1993. Conservation status of freshwater mussels of the United States and Canada. Fisheries: A Bulletin of the American Fisheries Societ\ 18(9):6-22. Wilson, C.B. and E. Danglade. 1912. Mussels of central and northern Minnesota. United States Bureau of Fisheries Economic Circular 3:1-6. Wilson, C. B. and E. Danglade. 1914. The mussel fauna of central and northern Minnesota. Appendi.x V to the Report of the U.S. Commissioner of Fisheries, 1913 (Bureau of Fisheries Document No. 803). 26 p. APPENDIX Figure Citations and Specimens E.xamined Provided below are figure references and the James Ford Bell Museum of Natural History (JFB) specimens used to construct Figure 2. For each lot, its catalog number and collection locality are provided. Whenever possible, specimens cited by previous authors have been selected, and these references are cited below. Also provided are corrections and re-identifications with regard to JFB specimens of historical significance. The following drainage system abbreviations are used: LMRS = Lower Mississippi River Subsystem LSS = Lake Superior System LWS = Lake of the Woods System MRS = Minnesota River Subsystem RRS = Red River System SCRS = St. Croi.x River Subsystem UMRS = Upper Mississippi River System Actinonaias ligamcntina (Lamarck, 1819). Figures: Cummings & Mayer (1992:105), Fuller (1985:36). LWS: Dawley's (1944, 1947) Sturgeon River record is JFB 2863 Lampsilis siliquoidea. RRS: Radke's (1992) Otter Tail River record is JFB 7569 Lampsilis siliquoidea. I'MRS: Dawley (1944, 1947) and Moyle's (1940) Crow Wing River record is JFB 2872 Lampsilis siliquoidea. LMRS: JFB 2847 Mississippi River, Dresbach, Winona Co.: Daw- ley (1944, 1947), Grant (1885). MRS: JFB 2859 Minnesota River, New Ulm, Brown Co.: Dawley (1944, 1947). SCRS: JFB 2878 Kettle River, Rutledge, Pine Co.: Dawlev (1944, 1947). Alasmidonta marginata Sa\, 1819. Figures: Cummings & Mayer (1992:85), Fuller '(1985:42). LMRS; JFB 2766 Mississippi River, Winona, Winona Co.: Dawley (1944, 1947), Holzinger (1887). MRS: JFB 2763 Minnesota Riv- er: Dawley (1944, 1947). SCRS: JFB 2769 St. Croi.x River at the mouth of Sunrise Ri\er, Chisago Co.; Dawley (1944, 1947). Amblema plicata (Saw 1817). Figures: Cummings & Mayer (1992:41), Fuller (1985:50). LSS; JFB 7567 Clo- quet River, Brimson, St. Louis Co. RRS: JFB 3105 Red River, 20 mi. N of Breckenridge, Wilkin Co.; Dawley (1944, 1947), Grant (1885). LMRS; JFB 3083 Mississippi River, Dresbach, Winona Co.; Dawlev (1944, 1947). Grant (1885). MRS; JFB 3093 Minnesota River. Ft. Snelling; Dawley (1944, 1947), Grant (1885). SCRS; JFB 3112 St. Croix River at the mouth of Sunrise River, Chisago Co.: Dawley (1944, 1947). Anodonta suhorhiculata Sav, 1831. Figures; Cummings & Mayer (1992:77), Fuller (1985:58). Anodontoidcs ferussacianus (Lea, 1834). Figures; Cum- mings k Maver (1992:81), Fuller (1985:57). LSS; JFB 2709 Cloquet River, St. Louis Co. RRS; JFB 6582 Cor- morant Lake, Becker Co.; Radke (1992). UMRS; JFB 3995 Rum River, Anoka, Anoka Co.; Dawley (1944, 1947), Grant (1885). LMRS; JFB 2717 Cedar River, Austin, Mower Co.; Dawley (1944, 1947). MRS: JFB 2742 Cot- tonwood River, Garvin, Lyon Co.; Dawlev (1944). SCRS: JFB 2753 Rush Creek; Dawley (1944, 1947). Arcidcns confragosus (Sav, 1819). Figures; Cummings & Mayer (1992:89), Fuller (1985:51). LMRS; JFB 2771 Mississippi River, Red Wing, Goodhue Co.; Dawlev (1944, 1947). MRS; JFB 2773 Miimesota River, Ft. Snelling: Dawley (1944, 1947), Grant (1885). Cumherlandia monodonta (Say, 1829). Figures; Cum- mings & Mayer (1992:23), Fuller (1985:16). SCRS; JFB 6690 Rush Creek, Rush City, Chisago Co. Page 52 THE NAUTILUS, Vol. 110, No. 2 Cyclonaias tuberculata (Rafinesque, 1820). Figures: Cummings & Mayer (1992:49), Fuller (1985:10). LMRS: JFB 3198 Mississippi River, Ft. Siielling: Dawley (1944, 1947), Grant (1885). SCRS: JFB 3199 St. Croix River at the mouth of Sunrise River, Chisago Co.: Dawley (1944, 1947). Ellipsaria lineolata (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:107), Fuller (1985:45). LMRS: JFB 2916 Mississippi River, Nininger, Dakota Co.: Dawley (1944, 1947). MRS: JFB 2915 Minnesota River, Ft. Snell- ing: Dawley (1944, 1947), Grant (1885). SCRS: JFB 4200 St. Croix River, Taylors Falls, Chisago Co. Elliptio complanaia (Lightfoot, 1786). Figures: Clarke (1973:56-57, plate 3). LSS: JFB 3258 Lake Superior, Du- luth, St. Louis Co.: Dawley (1944, 1947). Elliptio crassidens (Lamarck, 1819). Figures: Cummings & Mayer (1992:67), Fuller (1985:14). LMRS: JFB 3249 Lake IPepin, Goodhue Co.: Dawley (1944, 1947). MRS: JFB 3245 Minnesota River: Dawley (1944, 1947). Elliptio dilatata (Rafinesque, 1820): the spike. Figures: Cummings & Mayer (1992:69), Fuller (1985:15). LMRS: JFB 3225 Mississippi River, Winona, Winona Co.: Daw- ley (1944, 1947), Holzinger (1887). MRS: JFB 3242 Min- nesota River, Ft. Snelling: Dawlev (1944, 1947), Grant (1885). SCRS: JFB 3244 St. Croix River, Pine Co.: Dawley (1944, 1947). Epioblasma triquetra (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:163), Fuller (1985:44). SCRS: JFB 5701 St. Croix River, Sand Is., Chisago Co. Fiisconaia ebena (Lea, 1831). Figures: Cummings & Mayer (1992:43), Fuller (1985:12). LMRS: JFB 3057 Mis- sissippi River, Dresbach, Winona Co.: Dawley (1944, 1947), Grant (1885). MRS: JFB 6437 Minnesota River, Scott Co.: Bright et al. (1990). SCRS: JFB 4044 Lake St. Croix, Washington Co. Fitsconaia flava (Rafinesque, 1820). Figures: Cummings & Mayer (1992:47), Fuller (1985:11). RRS: JFB 3014 Red River, 20 mi. N of Breckenridge, Wilkin Co.: Dawley (1944, 1947), Grant (1885). LMRS: JFB 3045 Mississippi River, Winona, Winona Co.: Dawley (1944. 1947). MRS: JFB 3044 Minnesota River, Ft. Snelling: Dawley (1944, 1947), Grant (1885). SCRS: JFB .3047 St. Croix River, Marine on St. Croix, Washington Co.: Dawley (1944, 1947). Larnpsilis cardiuni (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:157), Fuller (1985:28). LSS: JFB 2448 Lake Superior, Minnesota Point, Duluth, St. Louis Co.: Dawley (1944, 1947). LWS: JFB 2411 Sturgeon Riv- er, Little Fork River watershed, St. Louis Co.: Dawley (1944, 19471 RRS: JFB 2396 Red River, 20 mi. N of Breckenridge, Wilkin Co.: Dawlev (1944, 1947), Grant (1885). UMHS: JFB 2400 Mississippi River, Brainerd, Crow Wing Co.: Dawley (1944, 1947), Grant (1885). LMRS: JF"B 2397 Vli,ssissippi River, Dresbach, Winona Co.: Dawley (1944, 1947), Grant (1885). MRS: JFB 2404 Minnesota River, Ft. Snelling: Dawlev (1944, 1947), Grant (1885). SCRS: JFB 2415 St. Croix River, Marine on St. Croix, Washington Co.: Dawley (1944, 1947). Larnpsilis higginsi (Lea, 1857). Figures: Cummings & Mayer (1992:153), Fuller (1985:33-34). LMRS: JFB 2454 Mississippi River, Dresbach, Winona Co.: Dawle\ (1944, 1947). MRS: JFB 6396 Minnesota River, Carver Co.: Bright pf a/., 1990 (cf. L. /ugginsi); Dawley's (1944, 1947) record is JFB 2455 Obovaria olivaria. SCRS: JFB 2453 Lake St. Croix, Washington Co.: Dawlev (1944, 1947). Larnpsilis siliquoidea (Barnes, 1823). Figures: Cum- mings & Mayer (1992:151), Fuller (1985:29). LSS: JFB 2291 Cloquet River, St. Louis Co.; Dawley (1944, 1947). LWS: JFB 2983 Sturgeon River, Little Fork River wa- tershed, St. Louis Co.: Dawley (1944, 1947). RRS: JFB 2305 Red River, 20 mi. N of Breckenridge, Wilkin Co.: Dawley (1944, 1947), Grant (1885). UMRS: JFB 2242 Mississippi River, Brainerd, Crow Wing Co.: Dawley (1944, 1947), Grant (1885). LMRS: JFB 2307 Rolling- stone Creek, Minnesota City, Winona Co.: Dawley (1944, 1947), Grant (1885). MRs! JFB 2253 Minnesota River, Ft. Snelling: Dawley (1944, 1947), Grant (1885). SCRS: JFB 2261 St. Croix River at the mouth of Sunrise River, Chisago Co.: Dawley (1944, 1947). Larnpsilis teres (Rafinesque, 1820). Figures: Cummings & Mayer (1992:149), Fuller (1985:31). UMRS: JFB 2232 Leech Lake, Cass Co. LMRS: JFB 2239 Mississippi River, Red Wing, Goodhue Co.: Dawley (1944, 1947). MRS: JFB 2236 Minnesota River, Ft. Snelling: Dawley (1944, 1947), Grant (1885). Lasmigona complanata (Barnes, 1823). Figures: Cum- mings & Mayer (1992:93), Fuller (1985:39). LSS: JFB 3308 Lake Superior, Minnesota Point, St. Louis Co.: Daw- ley (1944, 1947); Baker (1928): reporting Lea's type- locality. LWS: JFB 3305 Sturgeon River, Little Fork River watershed, St. Louis Co.: Dawley (1944, 1947). RRS: JFB 3302 Red Lake River, Crookston, Polk Co.: Dawley (1944, 1947). UMRS: JFB 7507 Mississippi River, Little Falls public access, Morrison Co. (collected by Bright et al.); reported by Dawley (1944) & Grier & Mueller (1922) that Wilson & Danglade (1914) found it in the I'MRS; however, the data can not be located in that publication. LMRS: JFB 3301 Mississippi River, Min- neiska, Wabasha: Dawley (1944, 1947). MRS: JFB 3299 Minnesota River, Ft. Snelling: Dawley (1944, 1947), Grant (1885). SCRS: JFB 3289 Lake St. Croix: Dawley (1944, 1947). Lasmigona compressa (Lea, 1829). Figures: Cummings & Maver (1992:97), Fuller (1985:41). LSS: JFB 3262 Clo- quet River, St. Louis Co.: Dawley (1944, 1947). LWS: JFB 3261 Fall Lake, Lake Co.: Dawley (1944). RRS: JFB 7196 Otter Tail River, Otter Tail Co. (collected by Bright etal.). UMRS: JFB 3265 Sauk River, Stearns Co.: Dawley (1944, 1947), Movie (1940). LMRS: JFB 5251 North Branch Middle Fork Zumbro River, Highway 57 Bridge, Dodge Co.: Bright ct al. (1989). MRS: JFB 6017 Min- Daniel L. Graf, 1997 Page 53 nesota River, Yellow Medicine Co.: Briglit ct al. (1990). SCRS. JFB 3266 St. Croix River at the mouth of Sunrise River, Chisago Co.: Dawley (1944, 1947). Lasmigona costata (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:95), Fuller (1985:4o"). RRS: JFB 3279 Red Lake River, Crookston, Polk Co.: Dawley (1944, 1947). LMRS: JFB 3280 Mississippi River, Winona, Wi- nona Co.: Dawle\ (1944, 1947). MRS: JFB 3274 Min- nesota River: Dawley (1944, 1947). SCRS: JFB 4003 Snake River at the mouth of the Groundhouse Ri\er, Kanabec Co. Leptodca fragilis (Rafinesque, 1820). Figures: Cum- mings & Maver (1992:121). Fuller (1985:20). LMRS: JFB 2919 Lake Pepin: Dawley (1944, 1947), MRS: JFB 2923 Minnesota River, Ft. Snelling: Da\\ lev (1944, 1947), Grant (1885). SCRS: JFB 4613 St. Croix River, Marine on St. Croix, Washington Co. Ligumia recta (Lamarck, 1819). Figures; Cummings & Maver (1992:137), Fuller (1985:18). LSS: JFB 2205 Lake Superior: Dawley (1944, 1947). LWS: JFB 2214 Lake of the Woods, Pine Is., Lake of the Woods Co.: Dawley (1944). RRS: JFB 2201 Red River, 20 mi, N of Breck- enridge, Wilkin Co.: Dawley (1944, 1947), Grant (1885). UMRS: JFB 2994 Mississippi' River, Brainerd, Crow Wing Co.: Dawley (1944, 1947), Grant (1885). LMRS: JFB 2989 Vlississippi River, Winona, Winona Co.: Dawlev (1944, 1947), Holzinger (1887). MRS: JFB 2998 Blue Earth Riv- er: Dawley (1944, 1947). SCRS: JFB 2213 Kettle River, Rutledge, Pine Co.: Dawley (1944, 1947). Mcgalonaias nervosa (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:25), Fuller (1985:52). LMRS: JFB 3071 Mississippi Ri\er, Red ^^ ing, Goodhue Co.: Dawley (1944, 1947). MRS: JFB 3065 Minnesota River, Cedar Ave., Hennepin Co.: Dawley (1944, 1947). SCRS; JFB 3063 St. Croix River, Hudson, St. Croix Co., Wisconsin: Dawley (1944, 1947). Lower Mississippi River below Minnesota: Baker (1903). Obliqiiaria reflexa Rafinesque, 1820. Figures: Cummings & Mayer (1992:101), Fuller (1985:48). LMRS: JFB 2808 Lake Pepin, Lake City, Wabasha Co.: Dawley (1944, 1947), Grant (1885). MRS: JFB 2489 Minnesota River, Ft. Snelling: Dawley (1944, 1947), Grant (1885). SCRS: JFB 2804 Lake St. Croix, Washington Co.: Dawlev (1944, 1947). Obovaria olivaria (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:109), Fuller (1985:35). LMRS: JFB 2829 Mississippi River, Dresbach, Winona Co.: Dawle\ (1944, 1947), Grant (1885). MRS: JFB 6520 Minnesota River, Pike Is., Hennepin Co : Bright et ai. 1990; Daw- ley's (1944, 1947) record is JFB 2832 Fusconaia ebena. SCRS: JFB 2826 St. Croix River at the mouth of Sunrise River, Chisago Co.; Dawle> (1944, 1947). Plethobasns cyphyus (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:51), Fuller (1985:49). LMRS; JFB 3203 Mississippi River, Dresbach, Winona Co.; Dawley (1944, 1947), Grant (1885). Pleurobema coccineum (Conrad, 1836). Figures: Cum- mings & Mayer (1992:59), Fuller (1985:13). LMRS; JFB 3212 Lake Pepin, Goodhue Co.; Dawley (1944, 1947). MRS; JFB 6068 Minnesota River, Chippewa Co.; Bright etal. (1990). SCRS; JFB 3219 Snake River: Dawley (1944, 1947). Potainilus alatiis (Say, 1817). Figures: Cummings & Mayer (1992:125), Fuller (1985:25). RRS; JFB 2946 Red River, 20 mi. N of Breckenridge, Wilkin Co.; Dawley (1944, 1947), Grant (1885). LMRS; JFB 2945 Mississippi River, Dresbach, Winona Co.; Dawley (1944, 1947), Grant (1885). MRS: JFB 2941 Minnesota River, Ft. Snelling; Dawley (1944, 1947), Grant (1885). SCRS; JFB 2957 Snake Ri\er, Kanabec Co.: Dawley (1944, 1947). Potamilus capax (Green, 1832). Figures; Cummings & Mayer (1992:129), Fuller (1985:27). LMRS; Johnson (1980) reporting Green's type locality at St. Anthony Falls, Minneapolis. Lov\er Mississippi Ri\er below Min- nesota; Grier & Mueller (1922:9); ". . .the consensus of opinion is that it ordinariK does not go much north of Davenport, Iowa. Potamilus ohiensis (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:123), Fuller (1985:22-23). LMRS JFB 2963 Mississippi River, Red Wing, Goodhue Co. Dawley (1944, 1947). MRS; JFB 2964 Minnesota River Ft. Snelling; Dawlev (1944, 1947); Grant's (1885) record is JFB 2920 Leptodea fragilis. SCRS; JFB 5543 St. Croix River, Interstate Park, Chisago Co., Minnesota; Dawley s (1944, 1947) record is JFB 2962 Leptodea fragilis. Pyganodon grandis (Sav, 1829). Figures: Cummings & Mayer (1992:79), Fuller'(1985;60). LSS; JFB 2471 Com- stock Lake, St. Louis Co.; Dawley (1944, 1947). LWS: JFB 2625 Sturgeon Lake, St. Louis Co.; Dawlev (1944, 1947). RRS; JFB 2568 Red Lake River. Crookston, Polk Co.; Dawley (1944, 1947). UMRS; JFB 2560 Rum River, Milaca, Mi'Ue Lacs Co.; Dawlev (1944, 1947), Movie (1940). LMRS; JFB 3327 Zumbro River, Wabasha, Wa- basha Co.; Dawley (1944, 1947), Grant (1885). MRS; JFB 2664 Minnesota River, Ft. Snelling; Dawley (1944, 1947), Grant (1885). SCRS; JFB 2660 O.xbow Lake, Marine on St. Croix, Washington Co.; Dawley (1944, 1947). Qnadrula fragosa (Conrad, 1836). Figures: Cummings & Mayer (1992:29). LMRS; JFB 3127 Mississippi River, Nininger, Dakota Co. [collected November, 1886 by Winchell; reported as Q. quadrula by Dawley (1944, 1947)]. MRS; JFB 6356 Minnesota River,'Sibley Co.; Bright et al. (1990). Quadrula metaixevra (Rafinesque, 1820). Figures; Cum- mings & Mayer (1992:35), Fuller (1985:6). LMRS; JFB 3152 Mississippi River, Dresbach, Winona Co.; Dawley (1944, 1947), Grant (1885). MRS; JFB 6357 Minnesota River, Sibley Co.; Bright et al. (1990). SCRS; JFB 4607 St Croix River, Marine on St. Croix, Washington Co. Quadrula nodulata (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:37), Fuller (1985:8). MRS; JFB 6424 Minnesota River, Scott Co.; Bright et al. (1990). Page 54 THE NAUTILUS, Vol. 110, No. 2 SCRS: JFB 4578 St. Croi.x River, Coppermine Dam, Douglas Co., Wisconsin. Quadriila pustiilosa (Lea, ISol). Figures: Cummings & Mayer (1992:39), Fuller (1985:9). LMRS: JFB 3136 Mis- sissippi Riser, Winona, Winona Co.: Dawley (1944, 1947), Holzinger (1887). MRS: JFB 3144 Minnesota River, Ft. Snelling: Dawley (1944, 1947). SCRS: JFB 3147 St. Croix River, Marine on St. Croix, Washington Co.: Dawley (1944, 1947). Quadrula quadrula (Rafinesque, 1820). Figures: Cum- mings & Mayer(1992:31), Fuller (1985:7). RRS:JFB3125 Red River, 20 mi. N of Breckenridge, Wilkin Co.: Daw- ley (1944, 1947), Grant (1885). LMRS: JFB 3121 Missis- sippi River, Red Wing, Goodhue Co.: Dawley (1944, 1947). MRS: JFB 3122 Minnesota River, Ft. Snelling: Dawley U944, 1947), Grant (1885). SCRS: JFB 4911 St. Croix River, Stillwater, Washington Co. Simpsonaias amhigua (Say, 1825). Figures: Cummings & Mayer (1992:91), Fuller (1985:56). MRS: JFB 6074 Minnesota River, Chippewa Co.: Bright et al., 1990. SCRS: JFB 5063 St. Croix River, Marine on St. Croix, Wash- ington Co. Strophitus iindulatiis (Say, 1817). Figures: Cummings & Mayer (1992:83), Fuller'(1985:61). RRS: JFB 2784 Red River, 20 mi. N of Breckenridge, Wilkin Co.: Dawley (1944, 1947), Grant (1885). UMRS: JFB 7415 Mississippi River, Crow Wing State Park, Crow Wing Co. (collected by Bright et al). LMRS: JFB 2793 Mississippi River, Red Wing, Goodhue Co.: Dawley (1944, 1947). MRS; JFB 6075 Minnesota River, Chippewa Co.: Bright et al. (1990). SCRS: JFB 2795 Grindstone River, Kettle River water- shed. Pine Co.: Dawley (1944, 1947). Toxolasma parvus (Barnes, 1823). Figures: Cummings & Mayer (1992:131), Fuller (1985:54-55). LMRS: JFB 2974 Vlississippi River, Wacouta, Goodhue Co.: Dawley (1944, 1947). MRS: JFB 2976 Minnesota River, Ft. Snell- ing: Dawley (1944, 1947), Grant (1885). SCRS: JFB 2973 St. Croix River, Marine on St. Croix, Washington Co.: Dawley (1944, 1947). Tritogonia verrucosa (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:27), Fuller (1985:53). UMRS: JFB 3190 Sauk River, St. Cloud, Stearns Co.: Bright (1988), Dawley (1944), Moyle (1940). LMRS: JFB 3189 Missis- sippi River, Dresbach, Winona Co.: Dav\ley (1944, 1947), Grant (1885). MRS: JFB 3181 Minnesota River, Ft. Snell- ing: Dawley (1944, 1947), Grant (1885). SCRS: JFB 3191 St. Croix River, Tavlors Falls, Chisago Co.: Dawley (1944, 1947). Truncilla donaciformis (Lea, 1828). Figures: Cummings & Mayer (1992:117), Fuller (1985:47). LMRS: JFB 4307 Mississippi River, Northern States Power Prairie Is. Nu- clear Generating Plant, 1.5 miles upstream of Lock & Dam #3, Goodhue Co. MRS: JFB 2933 Minnesota River, Ft. Snelling: Dawley (1944, 1947), Grant (1885). SCRS; JFB 4610 St. Croix River, Marine on St. Croix, Wash- ington Co. Trur}cilla truncate (Rafinesque, 1820). Figures: Cum- mings & Mayer (1992:115), Fuller (1985:46). LMRS; JFB 2896 Mississippi River, Winona, Winona Co.; Dawley (1944, 1947), Holzinger (1887), MRS: JFB 2897 Min- nesota River, Ft. Snelling: Dawley (1944, 1947), Grant (1885). SCRS: JFB 2900 St. Croix River, St. Croix Falls, Polk Co., Wisconsin: Dawley (1944, 1947). Utterbackia imbecillis (Sav, 1829). Figures: Cummings & Mayer (1992:75), Fuller (1985:59). UMRS: JFB 2683 Mississippi River, Fridlev, .\noka Co.: Dawley (1944, 1947), Moyle (1940). LMRS; JFB 2682 Lake Pepin, Wa- couta, Goodhue Co.; Dawley (1944, 1947). MRS; JFB 2679 Minnesota River, Ft. Snelling: Dawley (1944, 1947), Grant (1885). SCRS: JFB 2768 Oxbow Lake, Marine on St. Croix, Washington Co.: Dawley (1944, 1947). Vcnustaconcha ellipsiformis (Conrad, 1836). Figures: Cummings&Mayer(1992:141), Fuller (1985:38). LMRS; JFB 5280 Cascade Creek, Zumbro River watershed, Rochester, Olmsted Co.; Bright et al. (1989). SCRS; JFB 5736 St. Croix River, Polk Co., Wisconsin. THE NAUTILUS 110(2):55-75, 1997 Page 55 The Life and Malacological Contributions of R. Tucker Abbott (1919-1995) M. G. Harasewych Department of Invertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC 20560 U.S.A. Dr. Robert Tucker Abbott, one of the best known and most venerated figures in American Malacology, died November 3, 1995 after a protracted battle with pul- monary fibrosis. During his long and multi-faceted ca- reer, he was an eminent researcher, curator, editor, mu- seum director, educator, enterpreneur, and an ambas- sador of his science to a broad and appreciative audience of enthusiastic shell collectors and weekend naturalists. R. Tucker Abbott was undoubtedly the most widely known malacologist in the world. His unique career has been chronicled like that of no other malacologist. Dur- ing his life, articles and biographical sketches appeared in Reader's Digest [August 1961], Hawaiian Shell News \1(d)~ [1963] and again 24(3):3 [1976], Ttic Wall Street Journal [March IS, 1994] reprinted in Ihiwaiian Shell News 42(8):7-8 [1994] and Sniithsunian magazine [Au- gust, 1995] as well as in numerous newpaper and shell club articles. Following his death, countless tributes and obituaries have been published, among them those ap- pearing in American Conchologist . Journal of Shell- fisheries Research and The Sew York Times. Most have emphasized the latter portions of his career. Born on September 28, 1919 in Waterlown, Massa- chusetts, to Charles Matthew (a paint manufacturer) and Frances (a homemaker, nee Tucker) Abbott, Tucker was the third of four brothers. As a bo\. Tucker spent his summers in Bermuda, where an uncle. Col. Richard Tucker was Director of the Bermuda Biological Station. It was there that he met and was befriended by Dr. William Beebe, the ichthyologist and pioneer of deep- sea exploration. Dr. Beebe was to have a profound influ- ence on Tucker. Many aspects of Tucker's career as a scientist, popularist, author, and celebrit\ find parallels in the life of William Beebe. When Tucker was a young boy, his family moved to Canada. It was there that he and a friend built their first natural history museum in the basement of the Abbott home. Upon graduating from the Humberside Collegiate in Toronto in 1938, Tucker entered Harvard College. While at Harvard, Tucker met Dr. William J. Clench, who was to become his lifelong friend and mentor. Tucker's career as a malacologist began \\ hile he was still an undergraduate at Harvard. He was a Research Assistant in the Department of MoUusks, Museum of Comparative Zoology, Harvard University from 1938- 1942. During this time he helped establish the mono- graphic series Johnson ia. serving as its first Business Man- ager, and published his first mongraphs and taxa (co- authored with Clench) prior to graduation. He was given leave from his studies to serve as the malacologist on the Harvard-Archbold Expedition to Melanesia in 1940-41, collecting mollusks in Fiji for three months aboard the junk-yacht Cheng-Ho. After graduating from Harvard College with a B. S. degree in 1942, Tucker became a United States Naval Aviator, and served as a di\e-bomber pilot stationed in Guantanamo \a\al Station in Cuba until 1944, when he was transferred to the United States Naval Medical Re- search Unit 2 (NAMRU-2) based in Guam. As the unit's first medical malacologist. Lieutenant Abbott traveled throughout the Marianas, Philippines and the Yangtze Valley of China, examining liuiidreds of species of fresh- water mollusks to identify those capable of acting as intermediate hosts of treniatodes parasitic in man, to describe and study the life c\ cles of these parasites, and to develop eftective counternieasures to control them. In 1946, Tucker left the Navy, married Mary M. Sisler and was appointed Assistant Curator in the Division of Mollusks of the National Museum of Natural History, Smithsonian Institution. In the \ears that followed, the Abbott family grew with the births of Robert Tucker, Jr., Carolyn Tucker, and Cvnthia Douglas. Tucker en- rolled in the graduate program of George Washington University, and earned a M. S. degree in 1949, com- pleting his thesis "The Anatonn and Life History of the fresh-water snail, Thiara granifera (Gastropoda: Proso- branchiata)" under the direction of Ira B. Hansen. The same year, he was promoted to Associate Curator. De- spite maintaining an extensive publication record. Tuck- er found time in tlie succeeding years to publish the first edition of American Seashells (1954), one of his land- mark works, and to complete his Ph. D. at George Wash- Page 56 THE NAUTILUS, Vol. 110, No. 2 ington University. His dissertation, The Gastropod Ge- nus Assiminea in the Philippines ' was completed with Paul Simon Galtsoff serving as his faculty advisor. In late 1954, Tucker left the National Museum of Natural History to fill the new i\ established Pilsbr\' Chair of Malacology at the prestigious Acadenn of Natural Sciences of Philadelphia. Shortly after arriving in Phil- adelphia, Tucker founded and served as the first presi- dent of the Philadelphia Shell C^lub, as well as Editor of its Proceedings. Following Henr\ Pilsbry's death in 1957, he became co-editor of The Nautilus together with H. B. Baker. In 1959, Tucker established the monographic series Indo-Pacific Mollusca as a complement to John- sonia, w hich dealt with western Atlantic molluscan fau- nas, and wrote the first three monographs in rapid suc- cession. Under Tucker's leadership, and with support from the Natural Science Foundation, the Department of Malacology enjoyed unprecedented grow th, launching collecting expeditions to all areas of the Indo-Pacific ranging from Madagascar to the Cocos-Keeling Islands, the Bay of Bengal, tlie Philippines and the Cook Islands. It was in the middle of his stay at the Academy of Natural Sciences that I first met Tucker. He was 40, already at the top oi his field, and just returned from the du Pont — Academy of Natural Sciences expedition to the Philippines. My classmate and I were ten years old and very interested in man\ aspects of natural history, .^fter spending an afternoon with Tucker, we both be- Figure 2. Mr, R. Tucker Abbott in the moliusk collectiuiis of the National Museum of Natural History, Smith,sonian Insti- tution, 1950. came avid malacologists and volunteers, a conversion he effected regularly on enthusiasts of all ages. During the Philadelphia years. Tucker continued to publish prodigiously, yet found time to pursue his hobby, the collection and classification of paper clips. When he appeared as a guest on the television quiz shows "What's My Line'r*' and "To Tell the Truth the panel deduced that he w as a malacologist, but Tucker w as able to elude them when appearing in his capacity as a "clipologist!" Tucker was widowed in 1964 and married Sue Sweeney Darwin in 1966. Because of his prominence in the field and his large follow ing among collectors. Tucker was recruited by John du Pont to help build a new natural history museum in the suburbs oi Wilmington, Delaware. He could not resist the challenge and opportunit) to create a new Depart- ment of Malacology or a new natural history museum devoted in large part to moUusks. Tucker left the Acad- emy of Natural Sciences of Philadelphia in 1969 to take up the ilu Pont Chair ot Malacolog> and serve as Assistant Director ot the Delaware Museum of Natural History. Calling upon his professional colleagues and friends in the world of amateur concholog\ . Tucker assembled a collection and librar\ that toda\ is one of the ten largest in the western hemisphere. \\ bile at the Delaware Mu- M G. Harasewych, 1997 Page 57 Figure 3. Dr. R. Tucker Abbott marking proof pages of Sea- shells oj the World in his office at the Academy of Natural Sciences of Philadelphia, circa 1962. seum of Natural History, Tucker continued as editor of Indo-Pacific Mollusca, and assumed primary editorship of The Nautilus upon the death of H. B. Baker in 1970. He pubhshed numerous papers and books, among them Kingdom of the Seashel! (1972) and tiie second edition ot American Scashells (1974), and was appointed Ad- junct Professor at the College of Marine Studies, Uni- versity of Delaware in 1973. Despite a distance of some 100 miles between the Campus of the College of Marine Studies and the Delaw are Nhiseum, Tucker was an active faculty member, teaching graduate courses in Malacol- ogy (together with M. R. Carriker) and Evolutionary Biology, and serving on five graduate committees. His students, myself among them, remember his enthusiasm and skill in collecting moilusks in tlie field, as well as in transforming the "surplus' specimens into a variety of culinary delicacies, including his "'Seven Species Soup. Tucker was divorced in 1974. He married Cecelia White in the spring of 1977. With his monumental accomplishment of museum building behind him, Tucker "retired to Melbourne, Florida in 1978, to concentrate on writing and to build a successful publishing company, American Malacolo- gists. Inc. Since Indo-Pacific Mollusca remained at the Delaware Museum of Natural History, he started another monographic series, Monographs of Marine Mollusca, one not encumbered by zoogeographic boundaries. I re- member Tucker confiding that, as an undergraduate, he had been torn between pursuing a career in malacology Figure 4. .A \ iew of Tucker in the specimen preparation room of the Delaware \hiseum of .Natural History, circa 1972. (Photo courtesy of Delaware Museum of .Natural History). and one in writing. .At this point in Tucker's career, his passion for writing clearly blossomed with a seemingly endless profusion of books and articles for the amateur naturalist and collector. His primary mission now more than ever was to reach and educate an ever-expanding audience of avid readers. By the mid 1980 s. Tucker would occasionally talk of retirement, when he would devote his time to gardening and writing about gardening. But that was always "in another five years. Even at this phase of his career, he remained interested and active in various aspects of mu- seum work, consulting with the Florida Museum of Nat- ural History on several exhibits, and completely design- ing the Hall of Shells at the newly built Fernbank Mu- seum in .Atlanta. When he was in his seventies, which he used to say was middle-aged for a malacologist. Tucker accepted another challenge, to build another museum, this time one devoted entirely to moilusks and emphasizing edu- cation and outreach rather than research. Those who were aquainted with Tucker knew that he was never happier than when he had some new project on the drawing boards. He was grateful for having been given the opportunity to call upon the experiences of his long Page 58 THE NAUTILUS, Vol, 110, No. 2 Figure 5. R, Tucker Abbott ut the groundbreaking for the Bailev-Matthews Shell Museum, JuK 6, 1992. and varied career to design a museum that incorporated tlie best elements of all tlie museums that he had worked in or visited. First as Consultant to the Board, then as Founding Director, Tucker devoted himself to this new task with extraordinary energy, enthusiasm and inten- sity. The Bailey-Matthews Shell Museum, which had its formal opening No\ember 18th, barely two weeks after his death, embodies Tucker s concept of the quintessence of what a shell museum should be. He regarded it to be the "crowning accomplishment of his life. Many remember Tucker as "Mr, Seashell, ' a flam- boyant man of boundless wit and good humor, described in the Wall Street Journal as "partial to short sleeved Hawaiian shirts decorated with brightK colored shells" and wearing multiple pairs ol glasses tethered around his neck. He had an enthusiasm that was contagious and the uncanny abilit\ to transform even a casual acquain- tance into a co-conspirator in the most far-fetched of projects. Tucker was a man l)lessed with a love of his work and an unfailing corn iction in its importance. For him, the joy w as in conceiving and doing a project, rather than its completion. Tucker also had a very strong sense of duty and service to the malacological community. In addition to .serving as editor of The Nautilus { 29 \ears), Indo-Pacijic Mollusca (19 \ears) and Monographs of Marine Mollusca (7 years), he was President of the Amer- ican Malacological Union (1959), Founder and President of The Philadelphia and Wilmington Shell C^lubs, Hon- orarv Member of numerous shell clul)s in man\ countries. Figure 6. R, Tucker .Abbott autographic a cop\ of one of his books (May. 1992), Secretary of the Society of Svstematic Zoology (1956- 59), Publications Chairman and later Awards Chairman for the Conchologists of America, Councilman of the Harvard Graduate Society for Advanced Study and Re- search, Member of the Board of Trustees of the Bermuda Biological Society and the Paleontological Research In- stitution, and served numerous other organizations in many and varied capacities. Among his accomplishments. Tucker leaxes a legacy of many significant contributions to the infrastructure of malacology in .America. He enlarged collections, pio- neered curatorial s\stems, built new museums and De- partments of Mollusks, and is responsible for creating at least two positions in the field of malacology. He main- tained and created new venues for publication of schol- arly works. He trained malacologists b\ ser\ing on their graduate committees, and by organizing and adminis- tering award programs that continue to support students in malacology. Because of his interest in the historv of .American malacology, he was instrumental in establish- ing archives now administered by the .American Mala- cological Union and housed at the .\cadem> of Natural Sciences in Philadelphia. Perhaps his most obvious con- tribution was his strong commitment to popularize the science of malacolog\' and the hobb> oi shell collecting. He w as instrumental in building a large, supportive, w ell- educated, and well-organized constituency for mollusks, a constituency that few other invertebrate groups enjoy. In a "Letter to a Beginner." published in .American Conchologist [15(2): 17], Tucker wrote "You ma\ wonder why I've taken so much time to answer \ our letter, even though I am 300 years behind in my w ork and writings. It s because I wrote a letter similar to yours about fifty \ears ago to a Harvard shell professor. Dr. William J. M. G. Harasewych, 1997 Page 59 Clench. He patiently answered me in detail and started me on the right track. It's m\ turn to pass on his kindness. I hope \ou 11 have the chance to take your turn. " Tucker Abbott has done a great deal for malacology during his life. We would do \\ ell to remember his simple request. R. Tucker Abbott is survived by his wife, Cecilia White Abbott, his son, Robert Tucker Abbott. Jr., his daughters CaroKn Tucker Palmer, C>ntliia Douglas Sullivan, Cher\l Robin Nelson, Jonna Leigh liobson, and Erika von der Heyden, and b) his brothers John, Matthew and Peter. ZOOLOGICAL TAXA OF R. TUCKER ABBOTT Despite a prolific publication record, R. Tucker Abbott proposed relatively few new nomina, these being limited to one family-level taxon, ten genus-level ta.xa (seven as subgenera) and 71 species-le\el taxa (19 as subspecies). Of these, 60'"f of the genus-level taxa and 21 ^c of the species level-taxa were co-authored. Two taxa, the genus Laevistrombiis and the species Acanthochitona halesae. have been attributed to Abbott by subsequent authors because he associated a description and or figure v\ith a nude name of an earlier author. Family-Level Taxa PROPEAMUSSIIDAE Abbott, 1954, American Seashells, p. 369. For Propcainussiuin deGregorio, 1884. Genus-Level Taxa Clenchiella Abbott, 1948. The Nautilus 61(3):76. Type species: Clenchiella victoriac Abbott, 1948, by original designation. Amnicolidae. Echiniiui.'i Clench and .Abbott, 1942, Johnsonia 1(4):3. New name for \ina Gra\, 1850, not Sina Horsfield, 1829, nor Nina Gray, 1855. Littorinidae. Fukuia Abbott and Hunter, 1949. Proceedings of the Helminthological Society of Washington 16(2):79-80. Type species: Fukuia kurodai Abbott and Hunter, 1949, by original designation. Amnicolidae. Glohivasuiu .\bbott, 1950. As a subgenus of Vasum Rod- ing, 1798. Johnsonia 2(28):215-216. Type species: Tur- binelta nuttingi Henderson, 1919, by original designa- tion. Turbinellidae. Laevistrombus Abbott, 1960. As a subgenus of Strombus Linne, 1758. Indo-Pacific Mollusca l(7):47-48. Type spe- cies: Strombus canarium Linne, 1758, by original des- ignation. Abbott attributed this taxon to Kira, 1955, whose use was a nomen nudum. See Bieler & Petit, (1996) for a discussion of the authorship of this taxon. Strombidae. Namrutua Abbott, 1948. As a subgenus of Hua Chen, 1943. Bulletin of the Museum of Comparative Zoology Harvard 100(3):296-297. T\pe species: Melania ning- poensis Lea, 1856, by original designation. Thiaridae. Pitarenus Rehder and Abbott. 1951. As a subgenus of Pi tar Romer, 1857. Revista de la Sociedad Malacologica "Carlos de La Torre " 8(2):57. Type species: Pitaria cor- data Schwengel, 1951, by original designation. N'eneri- dae. Siphovasum Rehder and .Abbott 1951. As a subgenus of Vasum Rriding, 1798. Re\ ista de la Sociedad Malacolo- gica "Carlos de La Torre" 8(2):61. Type species: Vasum (Siphovasum) latiriforme Rehder and Abbott, 1951, by original designation. Turbinellidae. Tcctininus Clench and Abbott, 1942. As subgenus of Echininus Clench and Abbott, 1942. Johnsonia 1(4):4. Type species: "non umbilicated Echininus nodulosus (Pfeifter, 1839)", b\ original designation. Littorinidae. Texadina .\bbott and Ladd, 1951. As a subgenus of Lit- toridina E\doux and Soule\et, 1852. Journal of the Washington Academy of Science 41(10):335. Type spe- cies: T. sphinctostoma Abbott and Ladd, 1951, by orig- inal designation. Amnicolidae. Species-Level Taxa Abbreviations for repositories of type material: AM = Australian Museum, Sydney. AMNH = .'\merican Museum of Natural History, New York. ANSP = The .-Academy of Natural Sciences of Philade- phia. BM(NH) = British Museum (Natural History), London, [now The Natural History Museum, London] BMH = Bishop Museum, Honolulu. CM = Carnegie Museum, Pittsburg. DMNH = Delaware Museum of Natural History, Wil- mington. FSM = Florida State Museum, Gaines\ille. [now Florida Museum of Natural Histors ] MCZ = Museum of Comparative Zoology, Harvard Uni- versity, Cambridge. MZUM = Museum of Zoolog\, Universit)' of Michigan, Ann Arbor. NMV = National Museum of X'ictoria, Brisbane. PBS = Philippine Bureau of Science, Manila. PML'H = Poey Museum, University of Havana. RM = Raffles Museum, Singapore. UMML = Rosenstiel School of Marine and Atmospheric Science, L'niversitN of Miami. I'SNM = National Museum of Natural History, Smith- sonian Institution, Washington, DC. IT = Institute of Marine Science, L'niversity of Texas. WAM = Western .Australian Museum, Perth. afrobellatus. Strombus terebcllaius Abbott, 1960. Indo- Pacific Mollusca 1(2):88, plate 61, figure 2. Holotype ANSP 214295, paratypes ANSP 211423, USNM, ANSP, Coryndon Museum 1611. Pange Island, west side of Zan- zibar, Tanzania. Strombidae. alcaldei. Naniviirea Jaume and Abbott, 1948. Revista de la Sociedad Malacologica "Carlos de la Torre" Habana 6(l):5-7, figures la-c, 4. Holotype PMUH 12178, para- Page 60 THE NAUTILUS, Vol. 110, No. 2 types, Jaume collection 25202, USNM 488585. Cuba. Amnicolidae. alfredi, Turbonilla (Pyrgisciis) .\bbott, 1958. Mono- graphs of the Academy of Natural Sciences of Philadel- phia 11:104-105, text-figure 5, la-c. Holotype ANSP 198692, paratypes ANSP 198693, Ostheimer Collection. Grand Cayman Island. Pyramidellidae. andersoni. Phaliuin .Abbott, 1968. New name for Cassis (Phalium)dalli Anderson, 1929, not Phalium dalli (Dick- erson, 1917). Miocene, Caribbean coast of Colombia. Cassidae. areas, Terebra (Strioterebruni) Abbott, 1954. The Nau- tilus 68(2):40-41, pi. 2, figure 4. Holotvpe USNM 613882, paratvpes, USNM 613883, ANSP 191712, Steger collec- tion (now DMNH 104984, DMNH 118028). Areas Cays, Campeche Bank, Mexico. Terebridae. atlanticus, Pleuwbranchus (Susania) Abbott, 1949. The Nautilus 62(3):73-78, plate 5, figures 1-10. Holotype USNM 574352, paratvpes USNM 574342, ANSP 184350, MCZ 165951. Soldier Key, Bisca> ne Bay, Florida. Pleu- robranchidae. aiireofasciatus. Conns spuiius Rehder and Abbott. 1951. Revista de la Sociedad Malacologica "Carlos de La Tor- re" Habana 8(2):64-65, plate 9, figs. 3, 4. Holotype USNM 597521, USNM 597522, Sennott collection, Woolsey col- lection. Off Dry Tortugas, Florida, in 20 fms. Conidae. auricincta, Strombifortnis Abbott, 1958. Monographs of the Academy of Natural Sciences of Philadelphia 11: 106-107, text-figure 5, 4. Holotype ANSP 198840, para- types ANSP 199841, USNM 619550, ANSP 75855. Grand Cayman Island. Melanellidae. anstini. Conns Rehder and Abbott. 1951. Journal of the Washington Academv of Sciences 41(l):22-24, figure 7. Holotype USNM 603017, paratypes USNM 421721, USNM 603018. Dry Tortugas, Florida. Conidae. unstini, Stenoihijra Abbott, 1951. Journal ot the Wash- ington Academy of Sciences 41(1):14-16, figures 1, 3-7 Holotype USNM 603669, paratypes USNM 603670, USNM 603671, USNM 603672, MCZ. Leyte, Philippines. Stenothyridae. balcsae. Acanlliocliituna Abbott, 1954. American Sea- shells, p. 318. Holotype ANSP 349331, Bonefish Key, Florida. Abbott attributed this taxon to Pilsbry, 1940, whose use was a nomen nudum. See Lyons (1988):102 for a discussion of the authorship of this taxon. Acan- thochitonidae. bayeri, Olivella Abbott, 1951. The Nautilus 64(4):111- 1 12, plate 7, figures 2a-b Holotype USNM 595132, para- types USNM 412871, MCZ, ANSP, UMML. Off Ajax Reef, off Elliott's Key, Lower Florida Keys. Olividae. blandfurdi, ,\ssiniinea Abbott, 1958. I'roceedings of the Academv of Natural Sciences of Philadelphia 110:247- 249, plate 21. Holotype USNM 615212. paratypes USNM 513820, ANSP 193360. Samar Island. Philippines. Assi- mineidae. boettgeri, Assiminea Abbott, 1958. Proceedings of the Academy of Natural Sciences of Philadelphia 110:249- 251, plate 15, figure 3, plate 23. Holotype USNM 615214, paratypes USNM 613828, ANSP 193370. Leyte Island, Philippines. Assimineidae. bonnci, Indopijrgns Abbott, 1945. Occasional Papers on MoUusks l(l):3-4, plate 1, figures 1-6, plate 2, figures 1,2. Holotype, MCZ 147474, paratypes MCZ 147475, MCZ 147476. Lake Pose, Celebes. Hydrobiidae. boschi, Cijmatium (Ranularia) Abbott and Lewis, 1970. The Nautilus 83(3):86-88, figures 1-6. Holotype DMNH 22862, paratypes AMNH 139206. ANSP 188844, Bosch collection. Al Masirah Island, Oman. Ranellidae. branluunac. Fasciolaria distans Rehder and Abbott. 1951. Revista de la Sociedad Malacologica "Carlos de La Tor- re" 8(2):59-60, plate 8. figs. 4,5. Holotype USNM 597513, Branham collection, Raines collection. Off Puerto Alvaro Obregon, Tabasco, Mexico. Fasciolariidae. cardmca. Coralliophda Abbott, 1958. Monographs of the Academy of Natural Sciences of Philadelphia 11:166- 67, text-figure 3, plate 1, figures g-h. Holot>pe ANSP 197054, paratvpes ANSP 61054, (DMNH 24282 ex ANSP 61054), ANSP 194091, ANSP 29958, ANSP 36673, ANSP 196397, ANSP 29962, ANSP 29959, ANSP 190548, ANSP 197056, ANSP 197055, ANSP 195857, MCZ 133415, MCZ 38548. Vera Cruz, Mexico. Coralliophilidae. caribaea, Cosa .\bbott, 1958. Monographs of the Acad- emy of Natural Sciences of Philadelphia 11:112-113, text-figure 6. Holotype ANSP 200064, paratype ANSP 205419. Grand Cayman Island. Philobryidae. clarki. Stcnothyia ansiini Abbott, 1951. Journal of the Washington Academ> of Sciences 41(1):14-16, figure 2. Holot) pe USNM 603673, parat> pes USNM 303387. Min- doro Island, Philippines. Stenoth\ridae. clarki. Conns Rehder and Abbott. 1951. Journal of the Washington Academy of Sciences 41(1):22, figures 1-6. Holotype USNM 485740, parat\ pes USNM 488 465, MCZ. SSW of Marsh Island, Louisiana. Conidae. cookei, Omphalotropis Abbott. 1949. Occasional Papers of the Bishop Museum 19(I5):263-266. figure la-e. Ho- lotype USNM 591306, paratypes MCZ 182894, BMH 212195, numerous additional paratypes at USNM, MCZ, BMH. Saipan Island, Marianas. Synceridae. cosmani, Chicoreus .\bbott and Finlay, 1979. The Nau- tilus 94(4):159-162, figures 1-9. Holotype USNM 783323, paratypes AMNH, ANSP, DMNH. Cosman, Finla\. and Bijur C:ollections. Ocho Rios. Jamaica. Muricidae, dinagatcnsis. Synccra pscndoquadrasi .\bbott. 1948. Bulletin of the Museum of Ciomparative ZoologN Har- vard. 100i3):284-285. Holotspe MCZ 175323, paratypes MCZ 175324, USNM 485565. Panamoan, Dinagat Island, NE of Mindanao Island. Philippines. Synceridae. M. G. Harasewych, 1997 Page 61 cloiseae. Acteon Abbott. 197;3. The Nautilus 87(4):91- 92, figures 1-5. Holotype DMXH 27213, paratypes DMXH 67746, Bosch collection. Al Masirah Island, Oman. Acteonidae. emipowlusi. Ocenebra (Ocenebra) Abbott, 1954. The Nautilus 68(2):41-42, pi. 2, figure 3. Holotype USNM 613881, paratypes, ANSP 191711, Steger collection (now DMNH). Off Fort Myers, Florida. Muricidae. gcrrardi. Transeiiuella .Abbott, 1958. Monographs of the Academy of Natural Sciences of Philadelphia 11:130- 131, text-figure 7, plate 4, figures a-c. Holotvpe ANSP 199507, paratvpes ANSP 199508, ANSP 199509, (DMNH 21194 ex ANSP 199509), ANSP 199506, USNM 619551, MCZ, USNM, ANSP 199510. Grand Ca\ man Island. Ve- neridae. grandanus, Cantharus midtangulus Abbott, 1986. The Nautilus 100(4);120-121, figure 1. Holotype USNM 859096, paratypes ANSP, AMNH, MCZ, FSM. Black Isle, St. Joseph Ba\, Gulf Count\, Florida. Buccinidae. guamcnsis, Sijncera nitida Abbott, 1949. Occasional Pa- pers of the Bishop Museum 19(15):273-274, figures 8a- c, 9a-c. Holotype USNM 589997, paratypes USNM 589992, MCZ 183904, BMH 212193. Banks of the Pago River, Guam Island, Marianas. Synceridae. habci. Assiminea habei Abbott, 1958. Proceedings of the Academy of Natural Sciences of Philadelphia 110:251- 254, plates 16-18, 25. Holotype USNM 615213, para- types USNM 613801, ANSP 193367, USNM 613831, ANSP 193366, (DNMH 25311 ex ANSP 193366), ANSP 193368, ANSP 193369. Leyte Island. Philippines. Assi- mineidae. hectori, Phalium .\bbott, 1968. New name for Cassis muricata Hector, 1886, not Cassis muricata Menke, 1828. Miocene, New Zealand. Cassidae. hedgpethi, Pleurobranchaca Abbott, 1952. Florida State University Studies 7:1-2, plate 1, figures 1-8. Holotype IISNM 574607, paratype, Hedgpeth collection. Port Aransas, Texas. Pleurobranchidae. hoffmeyeri, Terebra (Strioicrcbntin) Abbott, 1952. The Nautilus 65(3):78-80, plate 5, figures 5-9. Holotype USNM 598077, paratvpes USNM 596972, MCZ, ANSP, (DMNH 24990 ex ANSP 234286), BMH, PBS, Hadley Collection. Basay Beach, Manila Ba\ , Luzon Island, Phil- ippines. Terebridae bummi, Pohjccra .\bbott, 1952. Florida State University Studies 7:3-4, plate 2, figures 1-8. Holotype, USNM 574647, paratypes USNM 574648, Humm collection. Al- ligator Harbor, Franklin Count\ , Florida. PoKceridae. indomaris, Lambis scorpius Abbott, 1961. Indo-Pacific Mollusca 1(3):165-166, plate, figure. Holotype ANSP 242210, paratypes ANSP 247564, ANSP 212384, ANSP 214263, ANSP 212968. Nossi-be, Madagascar. Strombi- dae. iredalci. Strombus vomer Abbott, 1960. New name for Strombiis australis Gra\, 1826, not Strombus austraUs Schroter, 1805. Strombidae. keelingensis, Siiccinea Abbott, 1950. Bulletin of the Raf- fles Museum 22:87-90, figures 1-7. Holotype USNM 488579, paratypes, USNM 488580, RM, BM(NH). North Keeling Island, Cocos-Keeling Islands, Succineidae. klineorum, Strombus Abbott, 1960. Indo-Pacific Mol- lusca 1(2):70, plate 20, figures 20-21. Holotype ANSP 247621, paratypes ANSP 211121, ANSP 211173, MCZ 224889. Powder Bay, Sri Lanka. Strombidae. kurodai, Fuhuia .Abbott and Hunter, 1949. Proceedings of the Helminthological Societ\ of Washington 16(2):80- 82, plate 1, fig. 6, plate 2, fig. 4.' Holotype USNM 593365, parat\pes USNM 488538, USNM 488517. Takeda River, near Maruoka City, Fukui Prefecture, Honshu, Japan. Amnicolidae. kurodai. PhaUum .Abbott, 1968. Indo-Pacific Mollusca 2(9):105-106, plate 87. Holot\pe NSMT, paratypes ANSP 241694. Tosa Bay, Shikoku, Japan. Cassidae. latiriforme, Vasum (Siphovasum) Rehder and Abbott. 1951. Revista de la Sociedad Malacologica "Carlos de La Torre" 8(2):61-62, plate 9, figs. 5,6. Holotype USNM 597517, paratypes, USNM 597518, Sennott collection, Raines collection. NW of Cape Ca- toche, Yucatan, Mexico, in 18-20 fms. Turbinellidae. luzonica, Assiminea habei Abbott, 1958. Proceedings of the Academy of Natural Sciences of Philadelphia 110; 254, plates 15, figure 9. Holotype ANSP 96035, paratypes ANSP 193356, ANSP 193355. USNM 201151. Luzon Is- land, Philippines. Assimineidae. margaritensis. Murex Abbott, 1958. Monographs of the Academy of Natural Sciences of Philadelphia 11:61, plate 1, figures n-o. New name for Murex imperialis Swainson, 1831, not Murex imperiahs G. Fischer, 1807. Muricidae. marshallensis. Assiminea nitida Abbott, 1958. Proceed- ings of the Academy of Natural Sciences of Philadelphia 110:256, plate 15, 'figure 8. Holotype USNM 615210, paratypes USNM 580909, ANSP 193364. Japtan Island, Enivvetok Atoll, Marshall Islands. Assimineidae. minirosea, Ocenebra (Ocenebra) Abbott, 1954. The Nau- tilus 68(2):43-44, pi. 2, figure 1. Holotvpe USNM 103380, paratvpes, USNM 161271, USNM 103413, USNM 416653, USNM 416655, USNM 416654. Jamaica. Muricidae. nworei. Olivella Abbott, 1951. The Nautilus 64(4):112- 114, plate 7, figure la-b. Holotvpe USNM 595133, para- types USNM 595134, MCZ, ANSP, UMML. Carysfort Reef, off Key Largo, Florida. Olividae. morrisoni, Assiminea Abbott, 1958. Proceedings of the Academy of Natural Sciences of Philadelphia 110:259- 260. Holotype USNM 615211, paratypes USNM 614191, ANSP 193357. Teteu Island, Raroia Atoll. Tuamotu Is- lands. Assimineidae. Page 62 THE NAUTILUS, Vol. 110, No. 2 multistriata, Fukuia Abbott aiul Hunter, 1949. Proceed- ings of the Helminthological Societ\ ot VV'ashiiigtoii 16(2): 82-83, Plate 1, fig. 5, plate 2, figs. 5 & 11. Holotype USNM 594185, paratypes USNM 488540, USNM 487383. Takefu in Kamiyama-mura, Nannjo-gun. Fiikui Prefec- ture, Honshu, Japan. Amnicolidae. nipponensis, Casmaria ponderosa .\bbott, 1968. Indo- Pacific Mollusca 2(9);200, plate 14, figures 13-14. Ho- lotype ANSP 313258, paratypes ANSP 241636, ANSP 227579, MCZ 233233. Tanabe, Wakayama Prefecture, Japan. Cassidae. ochroglottis, Stronibus inutabilis Abbott, 1960. Indo-Pa- cific Mollusca l(2):74-75, plate 20. figures 9-10. Holo- type ANSP 250187, paratypes ANSP 185466, USNM 2'6652a. USNM 465685, MCZ. Mauritius. Strombidae. orrae. Strombus urceits Abbott, 1960. Indo-Pacific Mol- lusca l(2):66-67, plate 20, figure 28; plate 41, figure 5. Holotvpe ANSP 247756, para^types MCZ, ANSP 232673, ANSP 233409, ANSP 240131, ANSP 240137. Augustus Island, Western Australia. Strombidae. ostheimerae. Emarginula .Abbott, 1958. Monographs of the Academy of Natural Sciences of Philadelphia 11:18, plate 1, figure c. Holotype ANSP 195983, paratypes ANSP 195982, Ostheimer collection. Grand Cayman Island. Fissurellidae. parkeri, Ithycythara Abbott, 1958. Monographs of the .\cademv of Natural Sciences of Philadelphia 11:96, text- figure 5. Holotype ANSP 198919, paratypes ANSP 209867 Grand Cayman Island. Turridae. patac. Conns Abbott, 1971. The Nautilus 85(2):49-51, figures 1-6. Holotvpe DMNH 44097, paratypes DMNH 44096, DMNH 40595, DMNH 44095, Finlay collection. Pompano Beach, Broward Count\ , Florida. Conidae. pihbryi, Lambis crocata Abbott, 1961, Indo-Pacific Mol- lusca 1(3);158-159, plate 128, figures 1-2. Holotype ANSP 156123, paratypes MCZ 49694, BMH 68268. Nuku Hiva, Marquesas. Strombidae, pseudoquadrasi, Syncera .Abbott, 1948. Bulletin ot the Museum of Comparative Zoology Harvard. 100(3):283- 284, plate 1, figs. 11-13, text iig. 12. Holotype MCZ 175325, paratypes MCZ 175326, USNM 485564. Bugo, Macajalar Bay, northern Mindanao Island, Philippines. Synceridae. rasilistoma, Tudicula (Tiuliculu) .Abbott, 1959. Indo-Pa- cific Mollusca 1(1):29, plate 4, figure 2, plate 10, figures A-C. Holotype NMV F18189, paratypes NMV, ANSP 227669. Tweed Heads, New South Wales, Australia. Tur- binellidae. saipanensis, Paludinella conica Abbott, 1949. Occasional Papers of the Bishop Museiun 19(15):271. Holotype ISNM 590891, paratypes MC;Z 182889, USNM, MCZ, BMH. North end of Saipan Island, Marianas. Synceridae. satlasi,Murex(Murexjrecurvirostris Hehderand Abbott. 1951. Revista de la Sociedad Malacologica "Carlos de La Torre" 8(2):58-59, plate 9, figs. 7, 8. Holotype USNM 597515, USNM 597516, Sennott collection. 60 miles NW of Cape Catoche, Quintana Roo, Mexico, in 18-20 fms. Muricidae. sennottoriim. Conns Rehder and .\bbott, 1951. Revista de la Sociedad Malacologica Carlos de la Torre 8(2):63, pi. 9, figures 1-2. Holotype USNM 597519, paratypes, USNM 597520, Sennott collection. SW of Campeche, Mexico. Conidae. sphinctostoma. Littoridina (Texadina) Abbott and Ladd, 1951, Journal of the Washington Academy of Science 41(10):335-338. Holotype USNM 596722, paratypes, USNM 596723, USNM 596724, MCZ, ANSP 187519, MZUM, UT. N of Webb Point, NW side of San Antonio Bay, 27 miles NE of Rockport, Texas. Amnicolidae. springeri, Anadara Rehder and .\bbott, 1951. Revista de la Sociedad Malacologica "Carlos de La Torre" 8(2):54. plate 8, figs. 8, 9. Holotype USNM 597375, paratypes USNM 538689, USNM 603848, USNM 603843. 12 miles off Dry Tortugas, Florida, in 66-60 fms. Arcidae. stegeri, Terebra (Myurella) floridana Abbott, 1954. The Nautilus 68(2):39-40, pi. 2, figures 5,6. Holotype USNM 613884, paratypes, USNM 613885, USNM 605184, ANSP 191709, Steger collection (now DMNH 105856). Off Campeche, Mexico. Terebridae. thielei, AssUninea Abbott, 1958. Proceedings of the Academy of Natural Sciences of Philadelphia 110:242- 244, plate 15, figure 5, plates 9, 22. Holotype USNM 609577, paratvpes USNM 613802, USNM 613813, USNM 613848, USNM 1018, ANSP 193358, ANSP 193359, MCZ. Leyte Island, Philippines. .Assimineidae. transversnnt, Microcardium Rehder and Abbott, 1951. Revista de la Sociedad Malacologica "Carlos de La Tor- re" 8(2):56, plate 9 figs. 9, 10. Holotype USNM 485734. 50 miles SSW of Marsh Island, Louisiana, in 29 fms (Pelican station 94-1). Cardiidae. vector. Phalinm (Seniicassis) Abbott, 1994, The Nautilus 107(3):94-96, figures 1,2. Holotype. USNM 860326, paratypes, Dan collection. Sa\ a de Malha Bank. Cassidae. victoriac. Clcnchiclla Abbott, 1948. The Nautilus 61(3); 76-80, plate 5, figures 1-7. Holotype USNM 488534, paratypes USNM 488535, MCZ. ANSP 18.3548. MZIM, CM. San Joaquin River, Le> te Island, Philippines. Am- nicolidae. virginensis. Latirns (Polygona) Abbott, 1958. Mono- graphs of the .\cadem\' of Natural Sciences of Philadel- phia 1 1 :76, text-figure 4, plate 2, figure r. Holot\ pe ANSP 196459, paratypes ANSP 34975, ANSP 34968, ANSP 34969. St. Thomas Island, Virgin Islands, Fasciolariidae. iLilsoni, SIrondins (Canarium) .\bbott, 1967. Indo-Pa- cific Mollusca l(7):455-456, plate 328, figures 1-3. Ho- lotvpe ANSP 253088. paratvpes ANSP 212695. WAM 847-66. WAM 840-66, WAM 838-66. WAM 851-66, ANSP, BMH. Zanzibar, Tanzania. Strombidae. M. G. Harasewych, 1997 Page 63 liinkuorthi. Fhalinrn Abbott, 196>S. Intlo-PaciBc Mol- lusca 2(9):179-181, plates 165-16b. Holot\pe AM, para- types AiNSP 272396, WAM 159-62. Abrolhos Islands, Western Australia. Cassidae. zanzibarica. Titdicula Abbott, 1958. \otulae Naturae 305:1-4, figures 1-7. Holotype AXSP 225261, paratypes ANSP 225262, 2 additional paratypes. Ras Nungwi, Zan- zibar, Tanzania. Turbinellidae. PUBLICATIONS OF R TUCKER ABBOTT R. Tucker Abbott was a prolific author, including among his works numerous newspaper articles, book reviews and sound recordings. Man\' of his popular works were pub- lished in multiple printings and editions, some were translated and published in several languages. The var- ious editions of his works are listed here, but individual printings are not. Man\' of Tucker's book reviews were published in se\eral different journals or shell club pub- lications. The following list of publications is arranged in chronological order. Dates of publication are provided when they could be determined. Dates of items published in publications of the United States National Museum are from Anonsmous (1971), those published in The Nautilus are from Coan and Harasewych (1993). Clench, \V. J. and R. T. Abbott. 19-11. The genus Strombus in the Western Atlantic. Johnsonia 1(1):1-15, [October 25] Clench, W. J. and R. T. Abbott. 1942. The genera Tectarius and Echininiis in the Western Atlantic, Johnsonia 1(4):1- 4. [May 1] Clench, W. J. and R, T. .Abbott 1943 Tiie genera Cypraea- cassis. Morum, Sconsia and Dalium in the Western \\.- lantic, Johnsonia 1(9):1-S. [July 7] Clench, W. J. and R. T. Abbott. 1943 The Cenera Gaza and Livona in the Western Atlantic. Johnsonia l(12l;l-9. [De- cember 31] Abbott, R. T. 1943. Guantanamo Bay. Cuba Johnsonia 1(12); 10-11. [December 31] .Abbott. R. Tucker, 1944. The genus Modulus in the Western Atlantic. Johnsonia l(14):l-6. [October 16] Abbott, R, T, 1945, A new Celebes Fresliwater Snail (Hy- drobiidae). Occasional Papers on Mollusks, Harvard, 1(1): 1-4, [February 27] Abbott, R, T. 1945, The Philippuie Intermediate Snail Host (Schistomophora quadrasi) of Schistosomiasis. Occasional Papers on Mollusks, Harvard 1(2):5-16. [April 30] Clench, William J, and R. Tucker Abbott. 1945. The Genus Strombus in the Western .Atlantic, Johnsonia 1(18):1, [No- vember 23] .Abbott, Lieutenant Tucker, 1946 The Egg and Breeding Habits of Oncomelania quadrasi Mlldff,. the Schistoso- miasis Snail of the Philippines Occasional Papers on Mol- lusks, Harvard 1(61:41-48, [February 14] Abbott, Tucker, 1947, Review ol "Morris, P A, 1947, A Field Guide to the Shells of our Atlantic and Gulf Coasts. Houghton Mifflin Co., Boston. 190 pp The Wood Thrush 3:21, [December] Abbott, R, Tucker, 1948. A New Genus and Species of Phil- ippine Amnicolidae. The Nautilus 61(3):75-80, plate 5, [March 2] Abbott, Lieutenant R, Tucker, 1948, Handbook of Medicallv Important Mollusks of the Orient and Western Pacific. Bulletin of the Museum of Comparati\e Zoology Harvard. 100(3):243-328, pis. 1-5. [April] Abbott, R, Tucker, 1948, Notes on the .Anatoni) of a giant Subspecies Mclongena corona from Florida Shell Notes by Frank Lyman 2(2&3):21-27, plates 1-5, [May] Jaume, M, L, and R, Tucker .Abbot [sic], 1948, A new Cuban species of the .Amnicolid genus i\^anivitrea. Revista de la Sociedad Malacologica "Carlos de la Torre" Habana 6(1): 5-8. [June] Abbott, R. Tucker. 1948. A potential snail host of oriental Schistosomiasis in North .America (Pomatiopsis lapidaria). Proceedings of the United States National Museum Smith- sonian Institution 98(:3222):57-68, pis. 3,4. [July 2] Abbott, R. Tucker. 1948. The Spread and Destructiveness of the Giant .African Land Snail Achatina julica. The Nau- tilus 62(1 ):31-34. [July 22] Abbott, R. Tucker. 1949. March of the Giant African Land Snail. Natural History 58(2):68-71. [February] .Abbott, R. Tucker. 1949. A .New Florida Species of the Tec- tibranch Genus Pleurobranchus. The Nautilus 62(3):73- 78, plate 5, figures 1-10. [March 18] Abbott, R. Tucker. 1949. Mollusks and Medicine in World War II. Report, Smithsonian Institution for 1947:325-;338, pis. 1-3. [presented as a paper at the 14th .American Mal- acological Union meeting, Pittsburgh in 1948. Presentation reviewed in American Malacological L'nion News Bulletin and Annual Report 1948:16-17 [March] [reprinted in Ster- kiana 9:28 (1963)] Abbott, R, Tucker. 1949. New S\ncerid Mollusks from the Marianas Islands (Gastropoda, Prosobranchiata, Synceri- dae). Occasional Papers ot the Bishop Museum 19(15):261- 274. [June 25] Abbott, R. Tueker and G. W, Hunter 111 1949, Studies on Potential Snail Hosts of Schistosonm japonicum I. Notes on the .Amnicolid snails of the genera Blanfordia. Tricula, and a new genus Fukuia from Japan. Proceedings of the Helminthological Societv of Washington 16(2):7;3-86, plates 1.2 [July] Hunter, III, G. W., R. Tucker Abbott, C Pan and E. Stray. 1949. Studies on Potential Snail Hosts of Schistosoma japonicum II. Infection Experiments on .Amnicolid snails of the genera Blanfordia. Tricula. & Fukuia. Proceedings of the Helminthological Societv of Washington 16(2):86- 89. [July] .Abbott, R. Tucker. 1949 Sexual dimorphism in Indo-Pacific Strombus. The Nautilus 63(2):58-61, [November 1] .Abbott, R. Tucker, 1949. An Indian Species of Clenchiella. The Nautilus 63(2):62. [November 1] Abbott, R. Tucker. 1949. Samoa, Shell-collector's paradise. The Scientific MonthU 69(5)319-327, [November 3] .Abbott, Robert Tucker. 1949 The .Anatonn and Life History of the fresh-water snail, "Thiara granifera" (Gastropoda: Prosobranchiata). Master of Science Thesis. George Wash- ington University, Washington, DC \i + 68, ill. Directed b>': Ira B. Hansen. [November] .Abbott, R. Tucker. 1950. The Genus Cyclostrcma in the West- ern Atlantic. Johnsonia 2(27):193-200. [January 6] Abbott, R. Tucker 1950 The Genera Xancus and Vasum in the Western Atlantic Johnsonia 2(28):201-219. [January 6]. Abbott, R, T. 1950, Voyages of the "Eolis", Johnsonia 2(28): 219-220, [January 6], Abbott, R, Tucker, 1950, Snail Invaders, Natural History .59(2): 80-85, [February] Page 64 THE NAUTILUS, Vol. 110, No. 2 Abbott, R. Tucker. 1950 Review of •C M Voiige. 1949. The Sea Shore. \\\ + 311 pp. Collins, LoikIoii" The Sci- entific Monthly. 70(2):135-136. [February] .Xbbott, R. Tucker. 1950. The Molluscan Fauna of the Cocos- Keeling Islands, Indian Ocean. Bulletin of the Raffles Mu- seum, Singapore 22:68-98. [.-Vpril]. Abbott, R. Tucker. 1950. The \'enomous Cone Shells. The Science Counselor 13(4)T25-126, 153. [December] Abbott, R. Tucker. 1951. New stenothyrid gastropods from the Philippines (Rissoidae). Journal of the Washington .i^cademy of Sciences 41(1):14-16. [January 15] Rehder, Harald A. and R. Tucker Abbott. 1951. Two new- Recent cone shells from the Western Atlantic (Conidae). Journal of the \\ ashington Academy of Sciences 41(1 ):22- 24. [January 15] .'\bbott, R. Tucker. 1951. The use of infra-subspecific names in Doctor de la Torre's essay on Polymita. The Nautilus 64(3):103-104. [February 15] .■\bbott, R. Tucker. 1951. Review of "de la Torre y Huerta, C. 1950. El Cencro Polymita. Memorias Soc. Cubana Hist. Nat. 20(11:1-20, pis. "l-ll." The Nautilus 64(3):107. [February 15] Abbott, R. Tucker. 1951. Review of "Moreno, A. 1950. Es- tudio Anatomico del genero Polymita Beck. Memorias Soc. Cubana Hist. Nat. 20(l):21-35, pis. 12-22." The Nau- tilus 64(31:108. [February 15] Abbott, R. Tucker. 1951. Review of "Colnian, J. S. 1950. The Sea and Its Mysteries. Norton, New York, xvi + 261 pp." The Scientific Monthly 72(2): 132. [February] Abbott, R. Tucker. 1951. In Search of the Golden Cowrie. Natural History 60(3):104-110, 144. [March] Abbott, R. Tucker. 1951. New Deep-Water Olivellas from Florida, with Notes on the O. jaspidea-nivea Complex. The Nautilus 64(4):110-1 16. plate 7, figures 1-5. [May 7] Abbott, R. Tucker. 1951. Operation Snail-Folk. Natural His- tory 60(6):280-285. [June] Abbott, R. Tucker. 1951. Review of "Pearse, A. S. 1950. The Emigration of Animals from the Sea. Sherwood Press, Drvden, NY xii + 210 pp." The Scientific Monthly 73(1): 68.' [July] Abbott, R. Tucker. 1951. Eastern Pacific Poromya and Ce- toconcha Names. The Nautilus 65(1 ):33. [August 27] Abbott, R. Tucker. 1951. Review of "Carson, R. L. 1950. The Sea Around Us. Oxford University Press, NY, vii -I- 230 pp." The Scientific Monthly 73(2):'l37-138. [August] Abbott, R. T. and H. S. Ladd. 1951. A New Brackish-water gastropod from Texas (Anmicolidae: Littoridina). Journal of the Washington Academs of Science 41(101:335-338. [October] Rehder, Harald A. and R. Tucker Abbott. 1951. Some New and Interesting Molhisks Irom the Deeper Waters of the Gulf of Mexico. Revista de la Sociedad Malacologica "Car- los de La Torre" 8(2):53-66, 2 plates. [November 12] Abbott, R. Tucker. 1951. Battle of the Snails. Science Digest, for November 1951: 15-19. [November] Abbott, R. Tucker. 1951. Genetic Analysis of Wild Popula- tions of the Genus Cerion. Colonial Inheritance of the multiple factor for shell length. [Abstract and Discussion of paper presented at 17tli Annual Meeting of the Amer- ican Malacological Union, August 22-24, 1951 ] American Malacological Union News Bulletin and .-Viinual Report 1951:10-11. [December) [Reprinted in Sterkiana 9:38 (1963)] Abbott, R Tucker. 1952. Re\ie« of "Klingel, G. 1951. The Bay. Dodd, Mead, New York \ -I- 287 pp " The Scientific Monthly 74(11:58. [January] Abbott, R. Tucker 1952. \ new Terebra (hoffmeyeri) from the Philippines The Nautilus 65(31:77-80, plate 5, figures 5-9. [Februarx 25] .■\bbott. R Tucker 1952 .\ Stud\ of an Intermediate Snail Host (Thiara granifera) of the Oriental Lung Fluke (Par- agonimus). Proceedings of the United States National Mu- seum 102(32921:71-116, pis. 8,9. [February 26] Rehder, Harald .A , R Tucker .Abbott, Joseph P. S. Morrison, David Nicol, Julia Gardner, W. P. Woodring, John B. Reeside, Jr., Ralph W. Imla\ , L. W. Stephenson, and W. A. Cobban. 1952 Objection to M. Gilbert Ranson's .Ap- plication in Regard to the Generic Name "Cryphaea" Lamarck, 1801 (Class Pelecypodal. Z. N. (S.)365. Bulletin of Zoological Nomenclature 6(61:188-189. [Ma> 22] Abbott, R. Tucker. 1952. Review of "Berrill, N. J. 1951. The Living Tide. Dodd, Mead, New York 256 pp " The Scientific Monthly 74(51:313. [May] .Abbott, R. Tucker 1952. Two New Opisthobranch MoUusks from the Gulf of Mexico belonging to the Genera Pleu- robranchaea and Polycera. Florida State I niversitv Stud- ies 7:1-7. [October 15] Abbott, R. Tucker. 1952. Review of "Morris, P. A. 1952. A Field Guide to Shells of the Pacific Coast and Hawaii. Houghton Mifflin, Boston." Atlantic Naturalist 8(2):I00. [November] Abbott, R, Tucker. 1952. Review of "Morris, P. A. 1952. A Field Guide to Shells of the Pacific Coast and Hawaii. Houghton Mifflin, Boston." The Scientific MontliK 75(6): 384. [December] Abbott, R. Tucker. 1952. Neu Federal Regulations on Im- porting Mollusks. [Abstract and Discussion of paper pre- sented at 18th .Annual Meeting of the .American Malaco- logical Union, August 20-23, 1952 ] American Malacolog- ical Union Annual Report 1952:9. [December] Abbott, R. Tucker. 1952. Opportunities in Medical Malacol- ogy. [.Abstract and Discussion of paper presented at 18th Annual Meeting of the .American Malacological L nion, August 20-23, 1952 ] .American Malacological I'nion .An- nual Report 1952:14-15. [December] Abbott, R. Tucker. 1953. Federal Regulations on Importing Living Mollusks. The Nautilus 66(31:104. [Februar\ 2 ] Abbott, R. Tucker. 1953. Review of "Hedgpeth, J. 1952. Between Pacific Tides. Ricketts and Calvin. 3rd Edition" Scientific Monthly 76(11:49. [February] Warinke, G. L. and R. Tucker .Abbott 1953. The Gross Anat- omv and Occurrence in Puerto Rico ol the Pelecypod Yoldia perprotructa. Journal of the \\ ashinglon .Academy of Science 43(8):260-261. [August] Abbott, R. Tucker. 1954. How to Collect Shells. Natural Hi.s- tory 63(I):32-37. [January] Abbott. R. Tucker. 1954. Section 10. MedicalK Important Mollusks. /)!. Hunter, G. W., W. W . Fr>e and J C Swa- trzw elder (eds.) .A Manual of Tropical Medicine. W B. Saunders C;o, Philadelphia, pp. 625-640. [Jaiiuar\ ] [Second Edition 1954, Third Edition 1960] Abbott, R. Tucker. 1954. The Habits and Occurrence of the Nudibranch, Armina tigrina. in southeast I nited States. The Nautilus 67(3):83-86. [Februar> 18] Abbott, R. Tucker. 1954. Review of the Atlantic Periwinkles, Sodilittorina. Echininus, and Tectarius. Proceedings of the Ignited States National Museum l()3(3328):449-464 [March 23] .Abbott, R Tucker [Illustrations bv Frederick M Ba\er] 1954 M. G. Harasewych, 1997 Page 65 American Seashells. D \'aii Ndstraiul t^o.. Int. New York xiv + 541 pp., 40 plates. [April 1.5] Abbott, R. Tucker. 1954. New Gulf of Mexico Gastropods (Tercbra anil Ocenebra). The Nautilus 68(2);37-44, pi. 2. [November 15] Abbott, R. Tucker [illustrations b\ Frederick M. Ba\er]. 1955. Introducing Seashells. D. \'an Nostrand Compan\, Inc. New York, vi + 64, 8 plates. [Februar\ 17] Abbott, R. Tucker. 1955. Secrets of the Pearl. Natural Histor\ 64(3):130-133. [March 7] Abbott. R, Tucker. 1955. The Titian R. Peale Shell Collection. The Nautilus 68(4): 123-127, plate 4, figures 1-8. [April 28] .\bbott, Robert Tucker. 1955. The Gastropod Genus As.si- minea in the Philippines Doctor of Philosophy Disserta- tion. George Washington L niversit\, Washington DC. i\ + 136 pp. Directed by Paul Simon Galtsoft. [June] .•\bbott, R. Tucker, George M. Moore, Jeanne S, Schwengel, and Margaret C, Tesks . 1955. Forward, p. i. In: Abbott, R. Tucker, M. K. Jacobson, and M. C. Teskey (Editors). How to Collect Shells. (A Symposium). American Mala- cological Union, Buffalo, NY [i\ ] + 75 + [vii] pp. [ p. i in second edition, 1961; p. i in third edition, 1966, p. ii,] .\bbott, R. Tucker 1955 Introduction p ii In: Abbott, R Tucker, M. K. Jacobson, and M. C. Teske\ (Editors). How to Collect Shells. (A S\ mposium). .\merican Malacological Union, Buffalo, NY [iv] + 75 + [vii] pp. [ p. ii in second edition, 1961; p. ii in third edition, 1966, p. ii, modified in fourth edition, 1974] Abbott, R. Tucker. 1955. The Shell collection, pp. 40-43. (reprinted in part from "American Seashells ), In: .Abbott, R. Tucker, M. K. Jacobson, and M. C. Teskey (Editors) How to Collect Shells. [\ Sv mposium). .\merican Mala- cological Union, Buffalo, NY [iv] + ~b + [vii] pp. [pp. 79- 83 in second edition, 1961; pp. 80-84 in third edition, 1966 and fourth edition, 1974] Abbott, R. Tucker. 1955. Oiitstandnig Shell collections, pp. 68-70. (reprinted in part from ".\merican Seashells"). In: .\bbott, R. Tucker, M. K Jacobson, and M. C. Teskey (Editors). How to Collect Shells. (.\ Symposium). American Malacological Union, Buffalo, NY [iv] -1-75-1- [vii] pp. [pp 85-87 in second edition, 1961; pp. 93-95 in third edition, 1966; pp. 94-95 in fourth edition, 1974] Abbott, R. Tucker. 1955, Review of "Abbott, R. T., M. K. Jacobson, and M. C Tesk\ . 1955, Hou to Collect Shells. American Malacological Union, Buffalo Museum of Sci- ence. New York" The Nautilus 69(1 );36. [August 1] Abbott, R. Tucker. 1955. Review of "Perry, L. M. and J. S. Schwengel. 1955. Marine Shells of the West Coast of Florida Paleontological Research Institution, Ithaca, NY". 198 pp, 55 pis." The Nautilus 69(1):.36. [August 1] Abbott, R, Tucker. 1955. Review ol "Bartsch, P. 1955. The Pyramidellid Mollusks of the Pliocene Deposits of Sorth St. Petersburg, Florida. Smithsonian Miscellaneous Col- lections 125(2):1-102, pis, 1-18." The Nautilus 69(I):iii. [.August 1] Abbott, R. Tucker. 19.55. Shells of Superstition. Frontiers 20(1); 12-13, inside back cover. [September 20] Abbott, R. Tucker. 1955. How to Catch Mice. Frontiers 20( 1 ); 32. [September 20] Abbott, R. Tucker. 1955. Anatomy of the Venezuelan Gas- tropod, Doryssa kappleri. The Nautilus 69(2):44-46, plate 4, figures a-j. [November 5] Abbott, R. Tucker. 1956. Comments on the Nomenclature of the Cypraeacea in Kaicher's "Indo-Pacific Sea Shells". Ha- waiian Shell News 4(7):68-69. [Mas 2] A[bbott], R. T. 1956. The Kimball \alentine Collection. The Nautilus 69(41:141. [May 10) .Abbott, R. Tucker. 1957. Conns niusicns nanus in Hawaii. Hawaiian Shell News 5(I0):101-105. [August 7] Abbott, R. Tucker. 1956. Notes on Mitrex scorpio and its allies. Hawaiian Shell News 4(1I):I13-114 [September 5] Abbott, R. Tucker 1956. Shell Clubs in America. Proceedings of the Philadelphia Shell Club l(l):l-5. [September] .Abbott, R. Tucker. 1957 The Tropical Western .Atlantic Prov- ince. Proceedings of the Philadelphia Shell Club 1(2):7- 11. [September] Abbott, R. Tucker. 1957. Prunnni ruscidum in New Jersey. The Nautilus 71(2):52-53, plate 4, Bgures 4, 4a. [Novem- ber 4] Abbott, R. Tucker. 1958. Western Atlantic Marine Mollusks [Title only]. Bulletin of the American Malacological Union 24:16. [.American Malacological Union Annual Report for 1957] [January 1] Abbott, R, Tucker, 1958, From the Pilsbr\ Chair of Mala- cology. The Nautilus 71(3):100-103. [March 4] .Abbott, R. Tucker. 1958. Redisco\er\ of a Rare Caribbean Conus. The Nautilus 71(4):117-118. [April 24] .Abbott, R. Tucker. 1958. A New Recent Species of Tudicula from Zanzibar (Gastropoda: X'asidae). Notulae Naturae 305:1-4, figures 1-7. [April 25] .Abbott, R, Tucker, 1958. Shells. National .Audubon Society Nature Program. Nelson Doubleday, Inc. Garden City, NY', 56 pp, 29 adhesive stamp plates [April; Second Edi- tion, 1966, Revised Edition, 1968] .Abbott, R. Tucker. 1958. The gastropod genus Assiminea in the Philippines. Proceedings of the .Academy of Natural Sciences of Philadelphia 11021.3-278, plates 15-25. [Au- gust 22] Abbott, R. Tucker. 1958. A Brief Histor> of Shell Auctions. Buttonwood Farm Shell Auction Catalogue pp. 11-12. [September 13] Abbott, R. Tucker. 1958. The Radula of Vuhitoconus grossi Iredale. Buttonwood Farm Shell .Auction Catalogue p. .35. [September 13] Abbott, R. Tucker. 1958. The Mollusk Diar> of Doc Bales. Proceedings of the Philadelphia Shell C;lub 1(3):6-12. [Oc- tober] .Abbott, R. Tucker. 1958. Notes on the .Anatomy of the .Aus- tralian Volutes, bednalli and grossi. Journal of the Mala- cological Society of Australia l(2):2-7. [November] Abbott, R. Tucker. 1958. Marine Mollusca of Rennell Island, Solomon Islands. The Natural History of Rennell Island, British Solomon Islands, Copenhagen, 2(28):203-206, plate 20 [December] .Abbott, R. Tucker. 19.58. The Marine Molluscs of Grand Cay- man Island, British West Indies. Monographs of the Acad- emy of Natural Sciences of Philadelphia ll:i-viii, 1-138, plates 1-5. [December 31] [.A second printing was pub- lished in October 1967, identical to the first, but with an inde.x to Common Names, consisting of 16 unnumbered pages, appended follow ing the plates]. Abbott, R. Tucker. 19.59. The FamiK \asidae in the Indo- Pacific. Indo-Pacific Mollusca l(l)T5-.32. [March 31] Abbott, R. Tucker. 1959. How to Collect Shells. /;!. Weyer, E. M. (ed.). The Natural Sciences Illustrated. J. J Little and Ives, Co., New Y'ork. pp. 2578-2589. Abbott, R. Tucker. 1960. Presidential Remarks. [Abstract of remarks at opening of presented at 26th .Annual Meeting Page 66 THE NAUTILUS, Vol. 110, No. 2 of the .\inerican Malacological I iiioii, June .jI-JuK 3, 1959] Bulletin of the .\merican Malacological I nion 26:2. [American .Malacological I nion .An- nual Report for 1959] [Januars 1] .•\bbott, R, Tucker. 1960 Expedition to the Philippine Islands. Proceedings of the Philadelphia Shell t:lub 1(4)1-6. [April] Abbott, R. Tucker. 1960. Basic Research — Is It Dying? . Proceedings of the Philadelphia Shell C:lub l{4):22-25. [April] .Abbott, R. Tucker 1960. Gift Irom the Sea of Japan. Frontiers 24(4):120-122. [March 22] A[tibott], R. T. 1960. Dr. Paul Bart.sch [Note]. The Nautilus 74(1 );33. [July 1] A[bbott], R. T. 1960. Norman T. Matto.x [Note], The Nautilus 74(1 ):33. [July 1] A[bbott], R. T. 1960. Department of Living Invertebrates re- established at the American Museum [Note]. The Nautilus 74(2):S1. [October 5] Abbott, R. Tucker. 1960. The Genus Strombus in the Indo- Pacific. Indo-Pacific Mollusca 1(2);33-144. [November 23] [Baker, H B., C. B. Wurtz and R. Tucker Abbott] Editors. 1961. Jeanne Sanderson Schuengel [notice ot death]. The Nautilus 74(4):165-166.[April 6] .Abbott, R. Tucker. 1961. Hov\ to Know American Marine Shells. Signet Key Book, The New American Library, Nev\- York. 222 pp. 12 plates. [May; Revised Edition, 1970] Warmke, G. L. and R. Tucker Abbott. 1961. Caribbean Sea- shells, A Guide to the Marine Mollusks of Puerto Rico and other West Indian Islands, Bermuda and the Lower Flor- ida Keys. Livingston Puhlisliing C^o,, Narbeth, Pennsyl- vania. i-.\ -I- 1-346 (44 plates). [May; Reprinted in 1975 by Dover Publications] [Baker, H. B., C. B Wurtz and R. Tucker Abbott] Editors. 1961. Jeanne Sanderson Schwengel, Sc. D., 1889 to 1961. The Nautilus 75(l);36-39, plate 5. [July 3] Abbott, R. Tucker. 1961. The Genus Lambis in the Indo- Pacific. Indo-Pacific Mollusca 1(3):147-174. [September 29] Abbott, R. Tucker 1961. Forward. In: Jacobson. M. K. and W. K. Emerson, Shells of the New York City Area. Ar- gonaut Books, Inc. New York. p. \ii. Abbott, R. Tucker. 1961. Comments on the proposal to place the generic name Gari Schumacher, 1817, on the official list unemended, Z, N. (S.) 1461. Bulletin of Zoological Nomenclature 1S(5):301. [November 10] Abbott, R. Tucker. 1961. Oldest Mollusk Research Center in America. Shells and Their Neighbors 8:1,3. [December] Abbott, R. Tucker. 1961. Review of ■Cameron, R. 1961. Shells. G. P. Putnam's Sons, New York. 128 pp. " Frontiers, December 1961:59. [December] .Abbott, R. Tucker. 1962. Historical Notes on the American Journal of Conchology. Sterkiana 6:1-4. [February] Abbott, R. Tucker. 1962. Review of "Jacobson, M. K. and W K. Emerson. 1961. Shells of the ^'eu: York City Area. .Argonaut Books, Larchmont. NY. 142 pp. Frontiers 26(4): 123. [March 22] Abbott. R. Tucker 1962 Recent I sesoi Non-binoniial Works. The Veliger 4(4);21.3. [April 1] Abbott, R. Tucker. 1962. U.se ol the Term "Hvpotypc". The Veliger 5(2);93-94. [October 1] Abbott, R. Tucker. 1962. America's oldest mollusk research center. P'rontiers 26(3):84-85. Abbott, R. Tucker 1962. Sea Shells of the World. A Guide to the Better-Known Species. .A Golden Nature Guide. Golden Press, New York, 160 pp, [Dutch Kdilion, 1963; French Edition, 1964; Norwegian Edition 1965; Italian Edition 1966; British Edition, 1967, Danish Edition 1968, Spanish Edition, 1973, German Edition, 1975, Revised .American Editions 1969. 1985] Abbott, Robert Tucker, 1962, Harvard Class ol 1942. Records of the Class, 1962, Harvard .Alumni .Association, Cam- bridge, MA. p. 1. Abbott, R. Tucker. 1963. Further comments on the name of the type species of Xenophora Fischer von Waldheim, 1807. Bulletin of Zoological Nomenclature 20(1):15. [March 18] Abbott, R. Tucker. 1963. .A Visit to Some European Shell Collections, Part I - Great Britain. Proceedings of the Philadelphia Shell Club 1( 7);25-31, [March] Abbott, R. Tucker, 1963, The Janthinid Genus Recluzia in the Western Atlantic, The Nautilus 76(4):151, [April 19] Abbott, R. Tucker. 1963. Expedition to the Bay of Bengal [Discussion of paper presented at 29th .Annual Meeting of the .American Malacological L nion, June 18-21, 1963] Bul- letin of the .American Malacological L'nioii 30:5, [Decem- ber 1] .Abbott, R, Tucker, 1963, Schelpen van der W ereldzeen, J, M. Meulenhoff, .Amsterdam. 160 pp [Dutch Edition of "Sea Shells of the World "] Abbott, R. Tucker 1963. Report on Collections made at Phu- ket Island, Thailand: International Indian Ocean Expe- dition, 1963. .Academy of Natural Sciences, Philadelphia, 33 pp, 1 map, [Mimeographed report] Abbott, R. Tucker, 1964, America's Shelling Mecca, Infor- mation for Visitors to Sanibel and Captiva, Special Shell Fair Edition:9, [March] Abbott, R, Tucker, 1964, Anolher Cypiaea gut tain. Hawaiian Shell News, 12(6):2. [April] .Abbott, R Tucker, 1964. Three Conus glorianiaris in Phil- adelphia. Hawaiian Shell News 12(8):6. [June] .Abbott, R. Tucker 1964. Littorina ziczac (Gmelin) and L. lineolata Orbignv. The Nautilus 7S(2):65-66. [October 15] Wagner, Robert J.\.'and R. Tucker Abbott. [Editors] 1964 Van Nostrand's Standard Catalogue of Shells. [First Edi- tion]. D. \an Nostrand Co. Inc., Princeton, N], 190 pp. [November 16, 1964; Errata, dated March 1965; Second printing with Errata (pages 192-195), May 1965] Abbott. R. Tucker. 1964. Indo-Pacific Mollusca. [Abstract and Discussion of paper presented at .30th .Annual Meeting of the .American Malacological L'nion. July 21-24, 1964] Bul- letin of the American Malacological l'nion 31:11. [.Amer- ican malacological L nion .\nnual Report for 1964] [De- cember 1] Abbott, R. Tucker. 1964. Report on the Marine Mollusks Collected in the Seychelles for the International Iiulian Ocean Expedition. .Academy of Natural Sciences, Phila- delphia. 6 pp. [Mimeographed report] Abbott, R. Tucker. 1964. Report on the Marine Mollusks Collected in the Bav of Bengal on Oiiise One ot the ".Anton Bruun". Department of Mollusks. The .Academv of Nat- ural Sciences of Philadelphia 34 pp. [Mimeographed re- port] .Abbott, R. Tucker. 1964. Cxjquillages. Especes du Monde Entier. Le Petit Guide, Hachette. 160 pp. [French Edition of "Sea Shells of the World"] Abbott, R. Tucker 1965. Review of "Kennelly, D. H. 1964. Marine Shells oj Southern Africa. Frontiers 29(3):94. [February] .Abbott. R Tucker 19()5, Miocene .S/ro)ii//!/.v {0(;/o;(i(7i(i 1 Iroiii India, Indo-Pacific Mollusca U6):401-402, [April .30] M. G. Harasewych, 1997 Page 67 Abbott, R- Tucker. 1965. Seashells from Cat Food. Hawaiian Shell News 13(12):6. [October] .\bbolt, R. Tucker. 1965. Cypiuca arenona C.ra\. 1824. Ha- waiian Shell News 14(2):8. [December] Abbott, R. Tucker. 1965. The wonderful world of shells Sci- ence Digest 5?>i6);66-73. [December] Abbott, R. Tucker. 1965. Giant Clams. Frontiers 30(2):46- 49. [December] Abbott, R. Tucker. 1965. Snegler og Skjell: Fra Hele Verden. Fredhois Forlag A/S, Olso. 160 pp. [Norwegian Edition of "Sea Shells of the World"] Abbott, R. Tucker. 1965. The Shell Desk t:alendar lor 1966. Shell Oil Co., New York. 120 pp. .-Vbbott, R. Tucker. 1966. Concholog): Queen ol the Natural Sciences. Frontiers 30(3):68-73. [Februar\ ] Abbott, R. Tucker. 1966. Curator's Choice: Glor\-of-India Cone. Frontiers 30(3):90. [Februar>] Abbott, R Tucker. 1966. Conchiglie, Specie di Tutto il Mon- do. Mondadori, Milan. 160 pp. [Italian Edition of "Sea Shells of the \\'orld"] [Februarv ] Abbott. R. Tucker. 1966. Quiz-Me: Seashells. .\ Junior Golden Guide. Golden Press, New \ork 4S pp [.\pril. Braille Edition, 1967] Abbott, R. Tucker. 1966. Comments on the proposed vali- dation of Valuta episcopalis Linnaeus, 1758. Z.N.{S.) 1728. Bulletin of Zoological Nomenclature 23(2/3 ):80. [July 29] Abbott, R. Tucker. 1966. Comment on A/i(ra per/a/a Roding, 1798, as a nomen oblitum. Z.N.{S.) 1726. Bulletin of Zoo- logical Nomenclature 23(2/3);90. [July 29] Abbott, R. Tucker. 1966. New England's Rarities, 1672. New York Shell Club Notes 1245. [September] Abbot [sic], R. Tucker. 1966. Shells. Nelson Doubledav , Inc. and Odham Books, Ltd. 63 pp.. 29 adhesive stamp plates. [Revised Edition, 1968] Abbott, R. Tucker. 1966. The Shell Desk Calendar for 1967, With Illustrations by Several Hands Showing Sea Shells in Aspects Practical, Scientific, and Ornamental Shell Oil Co., New York. 120 pp. Abbott, R. Tucker. 1967. Re\iew of "Dance, S. P. 1966. Shell Collecting: An Illustrated History." Frontiers 1(3): 66-67. [February 1] Abbott, R. Tucker, and Russell H Jensen. 1967. Molluscan Faunal Changes around Bermuda. Science 155(3763):687- 688. [Februar^ 10] Abbott, R. Tucker. 1967. Review of "Dance, S P. 1966. Shell Collecting: An Illustrated History." Natural Historv 76(3):66-67. [March] Abbott, R. Tucker. 1967. N'enom apparatus and Geographical Distribution of Conus glorumwn.'i. Notulae Naturae 400: 1-8. [May 3] Jung, Peter and R. Tucker .Abbott. 1967. The Genus Tere- bellum (Gastropoda: Strombidae). Indo-Pacific Mollusca l(7):445-454. [May 15] Abbott, R. Tucker. 1967. Strombus (Canarium) wilsoni. new- species from the Indo-Pacific Indo-Pacific Mollusca 1(7): 455-456. [May 15] Wagner, R. J. L. and R. Tucker Abbott. [Editors] 1967. \'an Nostrand's Standard Catalogue of Shells. [Second Edition]. Van Nostrand Co. Inc., Princeton, NJ, i-xi + 1-303. [Sep- tember 19] Abbott, R. Tucker. 1967. Review of 'Tebble, N. 1966. Brit- ish Bivalve Seashells. A Handbook for Identification. Trustees of the British Museum (Natural History) London. 212 pp + 12 pis." Quarterly Review of Biology 43(3):343- 344. [November 25] .Abbott, R. Tucker ami H, S. Zim. [co-authorship misattributed] 1967. Sea Shells of the World. A Guide to the Better- Known Species .\ Little Guide in Colour. Paul Hamlyn Ltd, London. 160 pp. [British Edition of "Seashells of the world "] Abbott, Robert Tucker. 1967. Harvard Class of 1942, Records of the Class, 1967. Harvard Alumni .Association, Cam- bridge, MA. p. 1. Abbott, R Tucker. 1967. Re\iew of "Dance, S. P. 1966. Shell Collecting: An Illustrated History." Quarterly Re- view of Biolog) 42:549-550. Abbott, R. Tucker. 'l967. Review of "Tebble, N. 1966. Brit- ish Bivalve Seashells. A Handbook for Identification. Trustees of the British Museum (Natural History) London. 212 pp -I- 12 pis." Frontiers 32(2):30. .Abbott, R. Tucker. 1967. Terebellum's Blue-E\ ed Periscope Frontiers December :8-9. [December] Abbott, R. Tucker. 1967. Quiz-Me; Seashells, .A Junior Golden Guide. Twin Vision, California. 37 pp. [Braille Edition] Abbott, R. Tucker. 1967. The Shell Desk Calendar for 1968, .A Journal Book Well .Adapted for the Purpose of .Noting Events and Expectations w itli the Greatest Readiness, Be- ing Calculated to Sa\e Much tune and Trouble. Shell Oil Co., New York. 120 pp. .Abbott, R. Tucker, and Russell H. Jensen. 1968. Portuguese Marine MoUusks in Bermuda. The Nautilus 8 1(3): 86-89. [Januar\' 25]. Abbott. R Tucker 19fj8. Giant Cowries. The Nautilus 82(1): 32. [July 24] Abbott, R.Tucker. 1968. The Helmet Shells of the World (Cassidae). Part 1. Indo-Pacific Mollusca 2(9):15-202. [Au- gust 30] .Abbott, R Tucker. 1968. [Editor] William Swainson's "Exotic ConchologN," .A Facsimile Edition of the 1841 Edition, with Essential Excerpts from the First and Second Issues U821-1835I. Delaware Museum of .Natural History, Re- production Series, Number 1. D. \'an Nostrand & Com- pan>, Princeton, NJ. i-xxi\ -(- 4-47 4- 48 plates. [.Abbott contributed the Preface (v) and the Modern Explanation of Plates (41-47)] [September 11] Abbott, R Tucker. 1968. Shells. Nelson Doubleday, Inc. and Odham Books, Ltd. 63 pp. 29 adhesive stamp plates [Re- vised Edition] [September] Abbott, R. Tucker 1968. Review ot "Knudseii, J 1967. The Deep-Sea Bivalvia. The John Murray Expedition, 1933- 34, Scientific Reports ll(3);235-343 -t- 3 pis." Quarterly Review of Biology 43(3):344. [September] Stix, Hugh, Marguerite Stix and R. Tucker .Abbott [photographs b> H. Landshoft ]. 1968. The Shell, Five Hundred Million \ ears of Inspired Design. Harr\ N. .Abrams, Inc. New York. 188 plates, plus unpaginated text. [October; French Edi- tion. 1969; Italian Edition 1969; .Abridged English Editions 1972. 1973, 1978, 1988] .Abbott, R. Tucker and Herbert S. Zim. [co-authorship misat- tributed] 1968. Fakta om Ivonkylier. Lademann, Copen- hagen 160 pp. [Danish Edition of "Sea Shells of the World '] [November 1] .Abbott, R. Tucker [recorded b\]. 1968. Pronouncing the Sci- entific .Names of Seashells of North .America. R. Tucker .Abbott and the Delaware Museum of Natural History, Greenville, Delaware. One, High Fidelit\, 33 13 RPM recording (36 minutes). .Abbott, R Tucker [Illustrations b\ George F. Sandstrom]. 1969. Seashells of North .America; .A Guide to Field Identifica- Page 68 THE NAUTILUS, Vol. 110, No. 2 tion. Golden Press, New York. 2.S0 pp [Revised edition, 1986] [January] Richards. Horace G , R. Tucker .\bl)ott. and Thomas Skymer. 1969. Marine Pleistocene Mollusks from Bermuda. The Geological Societ\' of .America, Abstracts with Programs for 1969 (4):69. [.\bstract] [April 9] Richards, Horace G., R. Tucker .Abbott, and Thomas Skymer. 1969. The Marine Pleistocene Mollusks of Bermuda. No- tulae Naturae 425:1-10. [October 10] Sti.\, Hugh, Marguerite Stix and R. Tucker Abbott [photographs by H. Landshoff]. 1969. Le Conchiglie. Garzanti, Italy. 188 plates, plus unpaginated text. [October 15] .Abbott, R. Tucker. 1969. Achatina fulica invades Florida. The Nautilus 83(2):75. [October 31] Stix, Hugh, Marguerite Stix and R. Tucker Abbot [sic] [pho- tographs by H. Landshoff]. 1969. Les Coquillages, les chefs-d oeuvre de la vie sous-marine. Seghers, Paris. 188 plates, plus unpaginated text. [November 25] Abbott. R Tucker. 1969. The Shell Desk Calendar for 1970 Shell Oil Co., New York. 116 pp. Abbott, R. Tucker and Hal Lewis. 1970. Cymatiitm boschi. New Species from the Arabian Sea. The Nautilus 83(3): 86-88. [January 23] Abbott, R. Tucker. 1970. .-Kmerican Malacological Union Symposium on the rare and endangered mollusks of North America, 7. Eastern marine mollusks. Malacologia 10:47- 49. [May] Abbott, R. Tucker. 1970. How to Know American Marine Shells [Revised Edition]. Signet Key Book, The New Amer- ican Library, New York. 222 pp. 12 plates. [May] Abbott, R. Tucker. 1970. Conchology: Queen of the Natural Sciences. La Conchiglia 11(1 1-12) [21-22]:14-15. [Decem- ber] Wagner, R. J. L. and R. Tucker Abbott. 1970. Van Nostrand's MonthK Supplement. Number 1. New subgenera of Con us proposed since 1966. Value changes and additions La Conchiglia II (ll-12)[21-22]:3, 22-23. [December] * Abbott, R. Tucker. 1971 The Shell Desk Calendar for 1971, with photographs suggesting the color, range and balance of life in the sea. The Shell companies. New York. 116 pp. Wagner, R J. L. and R. Tucker Abbott. 1971. Van Nostrand's Monthly Supplement. (Number 2, January, 1971). The Vo- lutes (Volutidae). La Conchiglia III (1)[23]:15. [January] * Wagner, R. J. L. and R. Tucker Abbott. 1971. Van Nostrand's Monthly Supplement. Continuation of Number 2. The Volutes (Volutidae), La Conchiglia III (2)[24];15. [Feb- ruary] * Wagner, R. J. L. and R. Tucker Abbott. 1971. Supplemento mensile al catalogo del Van Xostrand. La Conchiglia III (3)[25]: 15. [March]* Abbott, R. Tucker. 1971. .-^re marine mollusks endangeredr' La Conchiglia III (4)[26]:1 1-12. [April] Wagner, R J L. and R Tucker Abbott 1971 \an Nostrand's *.\ccording to an announceinent in La Coiwhiglia II( 1 1-12)[21- 22];3 (1970), supplements containing value changes in Van Soxtrand's Standard Catalog of Hhell.s (Second Edition) were originally published in issues of La Conchiglia. These were subse(|ueiitly reprinted (unpaginated) in a format suitable for insertion into the second edition ot Van Nostrand s Standard Catalog of Shells and sold separately. Monthly Supplement. (Number 5, .\pril, 1971 ) The Cow- ries (Cypraeidae). La Conchiglia III (4,l[26]:15. [.April] * Wagner, R. J. L. and R. Tucker Abbott 1971. Supplemento mensile al catalogo del \ an Nostrand, \ an Nostrand s Catalog Monthlv Supplement Familia Cvpraeidae. La Conchiglia III (5)[27]:15. [May]* Wagner, R. J. L. and R. Tucker Abbott. 1971. Supplemento mensile al catalogo del \'an Nostrand, Van Nostrand's Catalog MonthK Supplement. Familia Cvpraeidae La Conchiglia III (6)[28]:15. [June] * Abbott, R. Tucker and Charles B. Wurtz. 1971. Horace Bur- rington Baker, 1889-1971. The Nautilus 85(l):l-4. [July 20] Wagner, R. J. L. and R. Tucker Abbott. 1971 Family Cy- praeidae. Van Nostrand's Monthly Supplement. Number 3. La Conchiglia 30: insert of 8 unnumbered pages. [Sep- tember] * Abbott, R. Tucker. 1971. Conus patae, \ .New Caribbean Gastropod. The Nautilus 85(2):49-51. [October 4]. Wagner, R. J. L. and R Tucker Abbott 1971 Van Nostrand's Monthly Supplement. (Number 10, October 1971). Value Changes and Additions to Strombidae, .Additions to the Helmet Shells (Cassidae). La Conchiglia III (10)[32]:15. [October] * Abbott, R. Tucker. 1971. Seashell Safari to the Solomons. New York Shell Club Notes 176:3-5. [November] Abbott, R. Tucker. 1971. Bivalves, Magnificent Jewelry of the Sea. International Wildlife l(6):58-63. [November] Wagner, R. J. L. and R. Tucker Abbott. 1971. Van Nostrand's Monthly Supplement. (Number 11-12, Nov. -Dec. 1971). Additions to the Helmet Shells (Cassidae). La Conchiglia III (11-12)[33-34]:15. [November-December]* Abbott, R. Tucker. 1971. The Shell Desk Calendar for 1972, with ilustrations showing how the art of many ages has been enriched b\ the form and s> mmetrv of the shell. The Shell Companies, New York. 116 pp. Wagner, R. J. L. and R. Tucker Abbott. 1972. \aii Nostrand's Monthly Supplement. (Number 1, January 1972). .Addi- tions to the Helmet Shells (Cassidae). La Conchiglia I\' (1)[35]:15. [January]* Wagner, R. J L. and R Tucker Abbott. 1972. Van Nostrand's Monthly Supplement. (Number 2, Februarv 1972). .Ad- ditions to the Helmet Shells (C:assidae). La Conchiglia IV 2(36):15. [February] * Wagner, R. J. L. and R. Tucker Abbott. 1972. Van Nostrand's Monthly Supplement. (Number 2, February 1972). Ad- ditions to the Helmet Shells (Cassidae). La Conchiglia IV 3(37):15. [March]* Wagner, R. J. L. and R Tucker Abbott. 1972 \aii Nostrand's Monthly Supplement. (.Number 2, Februarv 1972). .Ad- ditions to the Helmet Shells (Cassidae). La Conchiglia I\' 4(38): 15. [April]* Abbott, R. Tucker. 1972. Review of "Keen, A. M. 1971. Sea Shells of Tropical West America." Quarterlv Reviev\ of Biology 47:240. [June] Abbott, R Tucker. 1972. C:onchologv— Hobbv ami Science. Jacksonville Shell Show Program:7, [JuK 29] Abbott, R. Tucker. 1972. New MoUusk Exhibits in America. La Conchiglia IV 7-8(41-42):220-21. [July-August] Wagner, R. J L and R. Tucker Abbott. 1972. Van Nostrand's Monthly Supplement. (Number 2, February 1972). .Ad- ditions to the Helmet Shells (Classidae). La Conchiglia I\' 7-8(41-42):23, [July-August] * Abbott, R, Tucker. 1972. Kingdom of the Seashell. Crouii M G. Harasewych, 1997 Page 69 Publishers, Inc., New York. 256 pp. [August] [French Edi- tion, 1974; Itahan Edition, 1974; Revised Edition, 1993] Keen, A, M. and R. Tucker Abbott. 1972. Problem of the type species of Lucina (Mollusca: Pelecypoda) Z.N.(S. ) 2001. Bulletin of Zoological Nomenclature 29{3):15S-161. [No\ ember 30] Abbott, R. Tucker. 1972. The Bermuda "VVilke." Newsletter, Bermuda Biological Station lor Research. 2(1)2. [Decem- ber] Salvini-Plawen, L. v. and R. Tucker Abbott. 1972. The Mol- lusks. pp. 19-28. In: Grzimek, B., O. Kraus, and R. Reidel (eds.) Grzimek's Animal Life Encyclopedia. Volume 3. Mollusks and Echinoderms. Van Nostrand Reinhold Com- pany, New York. 541 pp [paperback edition, 1984] Salvini-Plawen, L. v. and R. Tucker .\bbott. 1972. The So- lenogasters and Chitons, pp. 29-42. In: Grzimek, B., O. Kraus, and R. Reidel (eds.) Grzimek's .Animal Life En- cyclopedia. Volume 3. Mollusks and Echinoderms. Van Nostrand Reinhold Company, New York. 541 pp [paper- back edition, 1984] Salvini-Plawen, L. v. and R. Tucker Abbott. 1972. The Shelled Mollusks pp. 43-46. /().■ Grzimek, B , O. Kraus, and R Reidel (eds.) Grzimek's Animal Life Encyclopedia. \'ol- ume 3. Mollusks and Echinoderms. Van Nostrand Rein- hold Companv, New York. 541 pp. [paperback edition, 1984] Salvini-Plawen, L. \'. and R. Tucker .Abbott. 1972. The Gas- troverms. pp. 47-49. In: Grzimek, B., O. Kraus, and R Reidel (eds.) Grzimek's Animal Life Encyclopedia. Vol- ume 3. Mollusks and Echinoderms. \'an Nostrand Rein- hold Companv, New York, 541 pp. [paperback edition, 1984] Salvini-Plav\en, L. v. and R. Tucker ,\bbott 1972. The Gas- tropods, pp. 50-135. In: Grzimek, B , O Kraus, and R. Reidel (eds.) Grzimek's Animal Life Enc\clopedia. Vol- ume 3. Mollusks and Echinoderms. \an .Nostrand Rein- hold Companv, New York. 541 pp. [paperback edition, 1984] Salvini-Plawen, L. \ . and R. Tucker Abbott. 1972. Scaphopods and Bivalves, pp. 136-189. In: Grzimek, B., O. Kraus, and R. Reidel (eds.) Grzimek's .\nimal Life Encyclopedia. Vol- ume 3. Mollusks and Echmoderms. Van Nostrand Rein- hold Companv, New York. 541 pp. [paperback edition, 1984] Stix, Hugh, Marguerite Sti.v and R, Tucker .Alibott [photographs by H. Landshoff ], 1972. The Shell, Five Hundred Million years of Inspired Design. Ballaiitine Books, Inc. New York. [Abridged version of 1968 eilition] 135 plates, plus unpa- ginated text. Abbott, Robert Tucker, 1972. Har\ ard Class of 1942, Records of the Class, 1972. Har\ard .-Xlumni Association, Cam- bridge, \\.\ p 1. Abbott, R. Tucker. 1973. Review of "Halliday, P. 1972 Snailij Verse. \'olturna Press, Scotland. 48 pp. " The Nau- tilus 87(1 ):4. [January 24] Abbott, R Tucker. 1973. Review of 'Goodman, S. and L. Goodman. .1972. Art From Shells. Crown Publishing Co., New York. 210 pp." The Nautilus 87(1 ):23. [Januarv 24] Abbott, R. Tucker. 1973. Spread ot Melanuidcs tiibcrculata The Nautilus 87(1 ):29. [January 24] Abbott, R. Tucker. 1973. Review of "Boss, K. J. 1972. The Genus Semele in the ivestern Atlantic. Johnsonia 5(49): 1-32." The Nautilus 87(1 ):29. [Januar\ 24] Abbott, R. Tucker. 1973. Mollusks Dangerous to SCLBA di- vers. Of Sea and Shore 3(4):161-162. [January] Sandved, Kjell B. and R. Tucker Abbott. 1973 Shells in Color \'iking Press, New York. 112 pp. [June] Abbott, R. Tucker. 1973. Review of "Jenkins, M. M. 1972. The Curious Mollusks. Holiday House, New York. 224 pp." Quarterly Review of Biology 48:376. [June] Abbott, R. Tucker. 1973. Review of "Andrews, J. 1971. Sea Shells of the Texas Coast. Lhiiversity of Texas Press, Aus- tin, xvii + 298 pp." Quarterly Review of Biology 48:376. [June] Abbott, R. Tucker, 1973. In search of seashell treasures. Sci- ence Digest 74(1):16-21. [July] Abbott, R. Tucker. 1973. Mollusks Dangerous to SCUBA Di- vers. Delaware Medical Journal 45(6):161-164. [Jul\] Abbott, R. Tucker. 1973. Acleon etoiseae, a New Opistho- branch from Arabia. The Nautilus 87(4):91-92. [October 4] Abbott, R. Tucker. 1973 Review of "Furlong, M. and \' Pill 1972. Edible}' Incredible:' Ellis Robinson Publishing Co." The Nautilus 86(2-4):66. [December 1] Abbott, R. Tucker. 1973. Mollusks Dangerous to SCUBA Di- vers. The Shell Case [Naples Shell Club] 3(l);5-6. Abbott, R. Tucker. 1973. Review of "Bequart, J. C. and W. B. Miller. 1973. The Mollusks oj the Arid Southivest. The University of Arizona Press, Tucson, xvi -(-271 pp." QuarterK Review of Biolog\' 48:644-645. [December] Abbott, R. Tucker. 1973. What do Shells Tell':' Young Nat- uralist 13(12):25-31. [December] [In Russian — translated by V. Lesiyovsky] Abbott, R. Tucker [Editor]. 1973. American Malacologists, A National Register of Professional and Amateur Malacol- ogists and Pri\'ate Shell Collectors and Biographies of Early American Mollusk Workers Born Between 1618 and 1900. First Edition. American Malacologists, Falls Church, \'ir- ginia. i-iv + 1-494. Stix, Hugh, Marguerite Stix and R. Tucker Abbott [photographs by H. Landshoff]. 1973. The Shell. Harry N. Abrams, NY [Abridged version ot 1968 edition] 135 plates, plus unpaginated text. .•\bbott, R. Tucker. 1973. Conchas marinas, Ediciones Dia- mon, Barcelona, 1(50 pp, [Spanish Edition of "Sea Shells of the World"] .Abbott, R, Tucker. 1974. A tribute to a conchological bicen- tenary. New York Shell Club Notes 200:2. [March 18] .•\bbott, R. Tucker. 1974. American Malacologists — A Bio- graphical Survey (1618-1973). Bulletin of the American Malacological Union, Inc. for 1973:8-11. [May] Abbott, R. Tucker, 1974, .American Seashells, The Marine Mollusca of the Atlantic and Pacific Coasts of North .Amer- ica, Second Edition, \an Nostrand-Reinhold Publishing Company, New York. 663 pp. [October] .Abbott, R. Tucker. 1974. Au Royaume des Coquillages. Edi- tions des Deux Coqs d'Or, Paris. 255 pp [Kingdom of the Seashell, French Edition] Abbott, R. Tucker. 1974. II Meraviglioso Monde della Con- chiglie. .Arnoldo Mondadori Editore, Milan. 256 pp. [King- dom of the Seashell, Italian Edition] Abbott, R Tucker. 1975. Obituary, John Quincy Burch (1894- 1974). The Nautilus 89(1):12-13. [January 31] Abbott, R. Tucker. 1975. Beware the Asiatic Freshwater Clam. Tropical Fish Hobbyist 23(6): 15. [February] Abbott, R. Tucker. 1975. The Favored Shell. The Original Tide-ings 5(4/5):55. [April] Abbott, R. Tucker. 1975 Thomas Sav — America's First Ma- Page 70 THE NAUTILUS, Vol. 110, No. 2 laeologist. [.Abstract] Bullt-tiii o\ tht- .\merican Malacolog- ical Union, Inc. for 1974:66, [May) Abbott, R Tucker [Editor], 197.5, American Malacologists, A National Register of Professional and .\mateur Malacol- ogists and Pri\ ate Shell Collectors. Supplement. American Malacologists, Greenville, Delaware, pp. 495-609. [June or later] Abbott, R. Tucker. 1975. Shells and Our \ anishing World. Jacksonville Shell Show Program: 9. [JuK 26] .Abbott, R. Tucker, 1975, Note on Thais haemastoma. Te.xas Conchologist 12i2):30-31, [December] Abbott. R, Tucker, 1975. Muscheln und Schnecken des Meeres. Delphin Verlag, Stuttgart and Zurich. 160 pp. [German Edition of "Sea Shells of the World"] .Abbott, R. Tucker, 1976. Cittarium pica (Trochidae) in Flor- ida. The Nautilus 90(I):24. [January 30]. Abbott, R. Tucker. 1976. Review of "Kay, E. A. and W. J. Clench, 1975, A Bibliography of William Harper Pease. Malacologist oj Polynesia. Nemouria 16:1-50, Hawaiian Shell News 24( 1 ): 10 [January ] Abbott. R Tucker 1976 Seashells, Ridge Press, Bantam Books, New York. 160 pp. [subsequent printings, 1978, 1983] [February] Abbott, R. Tucker, 1976, Review of "Murray, S. B, 1975. Seashell Collector's Handbook 6 Identifier. Covering the Shells of the Coasts of the .Anxericas. Sterling Publishing Co., New York. 240 pp. (Quarterly Review of Biology 51(1):149, [March] Abbott, R, Tucker, 1976, Early Conchological Immigrants. Jacksonville Shell Show Program for 1976:16-17. [July] Abbott, R. Tucker [Editor]. 1976. The Best of T/ie iVauJi/us, \ Bicentennial Anthology of American Conchology. American Malacologists, Inc., Greenville, Delaware, i-viii, 1-280. Abbott, R. Tucker. 1977. In Memonum [W illard Lee Mo- horter and John Dyas Parker]. New sletter. .American Mal- acological Union 8(2):2. [Spring] .Abbott, R. Tucker. 1977, Shells — .An Emperor's Treasure. Palm Beach Life 70(7):17-19, 56-57. [July] Abbott, R. Tucker, 1977. Review of "Yonge, C. M. and T. E. Thompson. 1976, Living Marine Molluscs. William Collins & Son, London. 288 pp." Quarterly Review of Biolog) 52(4):438. [Decemljer] Abbott, R. Tucker and P. Alcolado M. 1977. Bivalves. FAO Species Identification Sheets. Fishing .Area 31. (W. Central Atlantic) 37 pp. Abbott, R. Tucker and P. Alcolado M. 1977. Gastropods. FAO Species Identification Sheets. P'ishing Area 31. (W. Central Atlantic) 18 pp. Abbott, R. Tucker. 1977. Chitons. F"AO Species Identification Sheets. Fishing .Area 31. (W, Central .Atlantic) 2 pp Abbott. Robert Tucker. 1977 Harvard C:lass of 1942, Records of the Class, 1977. Harvaril .\luinni Association. Cam- bridge, M.A, p 1, Abbott, R Tucker, 1977. EarK Shell ( iollet tors in Florida, The Shell Case [Naples, FLJ 5(1): 1-2 V\agner, Robert J. L. and R. Tucker Abbott [Editors], 1978. Standard t^atalogue of Shells. [Third Edition]. American Malacologists, liic , Greenville, Delaware i-v -I- 1-17 + misc. cosmically numliered pages. [February] Abbott, R, Tucker, 1978 Review of "Andrews, J. 1977. Shells and Shores of Texas " Science Books and Films 14(1):25. [May] Wagner, Robert J L and R Tucker Al)l)otl[EdU()rs]. 1978 Standard Catalogue of Shells, [Supplement 1] .\merican Malacologists, Inc, Greenville, Delaware, 28 pp, [Septem- ber] Sti,\, Hugh, Marguerite Stix and R Tucker .Abbott [photographs by H. Landshoff]. 1978, The Shell. Harry N, Abrams, Inc. New York. 163 pp. [a concise version of the 1968 edition; Japanese edition of this version, 1980] .Abbott, R. Tucker. 1979, The shell game. Sojourn, Summer 1979:31-33. [July] Richardson, L,, R, Tucker Abbott, and G. M. Davis, 1979. Early references to the figures in the Conchylien Cabinet of Martini and Chemnitz: Volumes 1-12. Tryonia 2(l):l-225; 2(2):226-427, [.August 1] Abbott, R, Tucker, 1979, The Shell Game. Diversion 7(8): 118-120. [August] .Abbott, R, Tucker, and C. John Finlay , 1979, Chicoreus cos- mani. .A new Muricid Gastropod from the West Indies. The Nautilus 94(4): 1.59-162. [October 30J .Abbott, R, Tucker, 1980. Further notes on Lambis. .New York Shell Club Notes 258:1-2. [January] Abbott, R Tucker, 1980. Are shells disappearing':' Shell Col- lector 2:47-49. [July] Abbott, R, Tucker, 1980, A misfit, tortuous tellin. Shell Col- lector 2:60. [July] Abbott, R. Tucker. 1980 Morris Karl Jacobson (1906-1980). An Obituary. The Nautilus 94(4):129. [October ,30]. Abbott, R, Tucker, 1980 The shell trade in Florida, status, trade, and legislation. Trade Records .Analysis of Flora and Fauna in Commerce (TR.AFFIC) USA Special Report 3:1- 85. [October] .Abbott, R. Tucker. 1980, Feed-back [letter], Conchologists of America Bulletin 22:2. [December] Stix, Hugh. Marguerite Stix and R. Tucker .Abbott [photographs by H, Landshoff ], 1980, Kai, sonobunkatobi, [Shells,Their culture and beauty] Harry N. .Abrams, Inc, New York, 163 pp, [Japanese edition of 1978 version] ■Abbott, R, Tucker 1981 Review of "Gordon, .A, and Lorna Strong Melvin, 1981, l.UUO World Sea Shells: Rare to Common, icith Values. Charles E, Tuttle Company , Rut- land VT " The Quarterly Review of Biology 56(2):182. [June] .Abbott, R. Tucker, 1981, Some comments on that mystery Hydatinid in color, Hawaiian Shell .News 29(1 1):6. [No- vember] .Abbott, R. Tucker and S Peter Dance. 1981. Compendium of Seashells, E, P. Dutton, Inc., New York, NY, x -I- 411 pp. [December; Copyrights. 1981, 1983. 1986, 1990; Sec- ond Edition 1983, Third Edition 1986, reprinted 1990; Japanese Edition, 1985; .Australian Edition, 1990] Abbott, R. Tucker 1982, Catalogue of the Recent Shells of Georgia— 1849, The Shell-O-Gram 23(3):3. [March] Abbott, R. Tucker, 1982. The C. O, A, Sanibel Sojourn of 1982, Conchologists of America Bulletin 10(3): 1,8-9, [Sep- tember] Abbott, R, Tucker 1982 Review of "Rehder. H A 1981 The .Kudubon Field Guide to Sorth American Seashells. Alfred .A. Knopf, New York. 894 pp " New York Shell club Notes 285:6. [December] Wagner, Robert J. L, and R. Tucker Abbott [Editors]. 1982. Standaril Catalogue of Shells. [Supplement 2], .American .Malacologists. Inc., Melbourne, Florida. 28 pp. Abbott, R, Tucker, 1983, Harrv S(tephen) Ladd, The Nautilus 97(1 );43, [January 31]. Abbott, R Tucker. 1983. C;harles B. Wurtz. The Nautilus 97(1):43, [January 31], M G. Harasewych, 1997 Page 71 Ahhdtt. R, Tucker. 1983. William Euwood) Old. Jr The Nautilus 97(l);4o-44. [Jaiuiarv 31] Abbott, R. Tucker. 1983. Reviews ol "Relider. H. A. 1981. The Audubori Society Field guide to Sorth American S/ie//s, Chanticleer Press.; Clarke, A. H. 1982. The Fresh- water Mollusks of Canada. The University of Chicago Press; and Smythe, K. 1982. Seashells of the Arabian Gulf" The Quarterly Review of Biolog> 58(1 ):70 [March] Abbott, R. Tucker and S. Peter Dance. 1983. Compendium of 8easliells. E. P. Dutton, Inc., New York, NY, x + 411 pp. [Second Edition] Deisler, Jane E. and R. Tucker .\bbott. 1984. Range Exten- sions of some Introduced Land Mollusks in the Bahama Islands, with First Reports for Four Species. The Nautilus 98a):12-17. [January 30J Abbott, R. Tucker and S Peter Dance. 1984. Compendium comment. Hawaiian Shell News 32(I):6. [January] .•Xbbott, R- Tucker. 1984. Giant Octopus comes Ashore. New York Shell Club Notes 290: 13. [March] Abbott, R. Tucker. 1984. A Farewell to Bill C:lench. The Nautilus 98(2):55-58. [April 27J Abbott, R. Tucker. 1984. Review of "Jokinen, E. H. 1983. The Freshuater Snads of Connecticut. State Geological and Natural History Survey of Connecticut, vii + 83 pp." The Nautilus 98{2):79. [reprinted New York Shell Club Notes 292:3-4] Abbott, R. Tucker. 1984. Collectible Florida Shells, American Malacologists, Inc. Melbourne, Forida, 1-64, [JuK] [Pub- lished in Regular and Waterproot Editions] [Re-issued the same year as "Collectible Shells ot Southeastern L .S., Ba- hamas & Caribbean"] Abbott, R. Tucker. 1984. "Collectible Shells ol Southeastern U.S., Bahamas & Caribbean [Regular and Waterproof Edition], .\merican Malacologists, Inc., Melbourne, Flor- ida. [Identical to above, except for title] Abbott, R T. 1984. Obituary [Zinaida A. Filatova], The Nau- tilus 98(4):166. [October 31] Abbott, R. Tucker. 1984. Seashells. pp. 103-109. In: Griffis, Nixon (ed. ) The Mariner's Guide to Oceanograpin . Hearst Marine Books, New York. 272 pp.. Ifci plates. [October] Abbott, R. Tucker. 1985. IRS Not Charitable to Some Con- chologists. Conchologists of .America Bulletin 13(2):22 [June] Abbott, R. Tucker. 1985. Review of "Anderson, R. 1985. Guide to Florida Seashells. Winner Enterprises. Florida. 56 pp." Shell-0-Gram [Jacksonville Shell Club] 26(4):6. [July-August][reprinted The Busycon [BrowartI Shell C'lub] 20(8);2-3; New York Shell Club Notes 29610-11, Ha- waiian Shell News 33(9):9.] .•\bbott, R. Tucker. 1985. Kirk .\nders — Shell Guide Supreme (1945-1985). Thatcheria 20(9):[2] [November] Abbott, R. Tucker. 1985. Kirk Anders, 1945-1985. Hawaiian Shell news 33(12):12. [December] Abbott, R. Tucker. 1985. Kirk Anders. New York Shell Club Notes No. 297:10-11. [December] Abbott, R. Tucker. 1985. Seashells ot the W orld, A Guide to the Better-Known Species. [Revised Etiition]. \ Golden Nature Guide. Golden Press, New York. 160 pp. Wagner. Robert J. L. and R. Tucker Abbott [Editors]. 1985. Standard Catalogue of Shells. [Supplement 3 World Size Records]. .American Malacologists, Inc., Melbourne, Flor- ida. 28 pp. Abbott, R. Tucker and S. Peter Dance. 1985. Compendium of Seashells. Japanese Edition [translated by T. Habe and T. Okutani], Heibonsha Ltd, Tokyo, pp. 1-445. [Abbott, R. Tucker]. 1986. The Nautilus, Its 100th Anniver- sarv and \oluine 100. The Nautilus 100(11:1-8. [January 31]' RTA. 1986. Tom Pullev and the Train. The Nautilus 100(1): 8. [January 31] Abbott, R. T 1986. Rev lew ol "Wells, F. R. 1986. Seashells of Western Australia. Western .Australian Museum, Perth. 207 pp. "The Nautilus 100(3): 113. [July 30, 1986] [reprint- ed New York Shell Club Notes 300:18] Abbott, R. T. 1986. Review of "Reid, D. G. 1986. The Littorinid Molluscs of mangrove Forests in the Indo- Pacific Region. British Museum (Naturl History), London. 228 pp." The Nautilus 100(31:113. [July 30, 1986] Abbott, R. T. 1986. Review of "Gate, J. M. and S. Raskin. 1986. It's Easy to Say Crepidula! Prettv Pretty Press, Santa Monica, 155 pp." The Nautilus 100(3); 113. [July 30, 1986] [reprinted New York Shell Club Notes 300:19] Abbott, R. T. 1986. Review of "Just, H. and M. Edmunds 1985. iVor(/i Atlantic Sudibranchs (Mollusca) Seen by Hcnning Lemche" The Nautilus 100(3);1 14. [July 30, 1986] .Abbott, R. Tucker 1986. Review of "Rios, E. C. 1986. Sea- shells of Brazil. Museu Oceanographico. Rio Grande. Bra- zil, 328 pp., 102 pis." Hawaiian Shell News 34(9):8. [Sep- tember] [reprinted New York Shell Club Notes 300:16] .Abbott, R. Tucker. 1986. Cantharus multangulus new sub- species grandanus from Northwest Florida (Buccinidae). The Nautilus 100(4);120-121. [October 31J Abbott, R Tucker. 1986. To Joe Rosewater— From R. Tucker Abbott. The Nautilus 100(4):152. [October 31] Abbott, R. Tucker[lllustrationsb> George F, Sandstrom]. 1986.A Guide to Field Identification, Seashells of North .America. Golden Press, New York. 280 pp. [Revised Edition] .Abbott, R. Tucker and S. Peter Dance. 1986. Compendium of Seashells. American Malacologists, Inc., Melbourne, Florida, x -I- 411 pp. [Third Edition] Abbott, R. Tucker. 1987. The Spell ot the Shell. Golden Years, Melbourne, FL 9(3):18-19. [January] Abbott. R Tucker. 1987. New hsln home for mollusks. The Capsule 19(4)1, 4. [Januarv] [Reprinted in COA Bulletin, March 1987] Abbott, R. Tucker. 1987. Concerning Olala lactea (Miiller, 1774). Festivus 19(2);12. [February 12] Abbott, R. Tucker. 1987. That Old Food Chain at Work. The Capsule 19(5):5. [March] Abbott, R. Tucker. 1987. Win moon-struck snails face east. Conchologists of America Bulletin 15(1 ):4. [March] ■Abbott, R. Tucker. 1987. New fish> home for mollusks. Con- chologists of .America Bulletin 15(1 ):8. [March] Abbott. R. Tucker. 1987. Letter to a Beginner— With Apol- ogies, [letter]. American Conchologist 15(2):17. [June] Houbrick, Richard S., Robert Robertson, and R. Tucker .Abbott 1987. .Anatomv and Systematic Position of Fastigiella carinata Reeve (Cerithiidae; Prosobranchia). The Nautilus 101(3);101-I10. [July 31] Abbott, R. Tucker. 1987. Marine Genus New to Atlantic Dis- covered. The Capsule 20(1):1. [Julv] .Abbott. R- Tucker. 1987. Letter [concerning editors of mala- cological journals]. Shells and Sea Life 19(7 ):4. [September] Abbott, R. Tucker. 1987. Review of "Petuch, E. J. 1987. Neic Caribbean Molluscan Faunas. CERF, Charlottes- ville, Virginia. 154 -I- 4 pages, 28 -I- 1 plates." The Festivus 19(10):98-99. [October 8] [Reprinted in New York Shell Club Notes 305:6-7] A[bbott], R. T. 1987. New LL S. Shell Stamp Laicovered. .American Conchologist 15(4 ):2. [December] Page 72 THE NAUTILUS, Vol. 110, No. 2 Abbott, R. T. 1987. Review of "Petucli, E. J. 1987. New Caribbean Molhiscan Faunas. CERF, C^liarlottesviile, VA. 154 + 4 pages, 28 + 1 plates." Conchologists of ,\merica Bulletin 15(4):19. [December] Abbott, R.T. 19S7 Review of 'Winner, B. E. 1987. A Field Guide to Molluscan Spawn, volume 1. E.B.M. North Palm Beach, PL. 139 pages. Conchologists of .\merica Bulletin 15i4);19, [December] Abbott, R. Tucker [Editor], 1987. Register of .American Ma- lacologists : .A National Register ot Professional and Am- ateur Malacologists and Private Shell Collectors. [Second Edition], American Malacologists, Inc., Melbourne, Flor- ida. i-.\iii, 1-168. Abbott, Robert Tucker. 1987. Harvard Class of 1942, Records of the Class, 1987. Harvard .\lunini Association, Cam- bridge, MX. p. 1, Abbott. R, Tucker. 1987, Pronouncing the Scientific Names of Scashelh of S'orlh America. (.Cassette), .American Ma- lacologists, Inc. Melbourne. FL Abbott, R. Tucker. 1987. E.xploring Collectible Shells. 90 minutes of commentary by the author. (Cassette). Amer- ican Malacologists, Inc. Melbourne, FL Abbott, R. Tucker. 1988. Dear COA Reader: [Editorial], American Conchologist 16(1 ):2. [March] Abbott, R. Tucker. 1988. Emerald Green Snails. American Conchologist 16(1 ):8. [March] Abbott, R. Tucker. 1988. Green Snails. American Conchol- ogist 16(1 ):9. [March] Abbott, R. Tucker. 1988. Teskey Collection Enhances Florida Research. New York Shell Club Notes 306:6. [March] [re- prmted, see below] Abbott, R. Tucker, 1988, Teske\ Collection Enhances Florida Research. Hawaiian Shell News 36(4 ):.5. [.April] Abbott, R. Tucker, 1988, 100Ct)nchological \ears Ago, Amer- ican Conchologist 16(2):10, [June] Wagner, R J L, and R, Tucker Abbott. 1988. World Size Records. American Conchologist 16(2): 17. [June] Court, I. and R Tucker Abbott. 1988. William Daniel Bledsoe (1920-1987). American Conchologist 16(3):16. [Septem- ber] Abbott, R, Tucker, 1988, What's New in Molluscan Research':' American Conchologist 16(3):17. [September] Abbott, R. Tucker, 1988. Ecphora Fever. American Con- chologist 16(3):20. [September] Abbott, R. Tucker. 1988. What's New in Molluscan Research? .American Conchologist 16(4):19. [December] Abbott, R. Tucker. 1988. Limerick Bargain. American Con- chologist 16(4):19. [December] Sti\, Hugh, Marguerite Sti\ and R. Tucker Abbott [photographs by H. Landshoff]. 1988. The shell: h\e hundred million years of inspired design. Harry .\ Abrams, Inc New York 188 plates -I- unpaginated text Abbott, R. Tucker. 1989. Review of "Long islanil Shell Club 1988. Seashclls of Long Island, Long Island Shell Club, Manhasset, NY. 209 pp." The Capsule 21(4):.\. [January] Abbott, R. Tucker, 1989, The Spectacular Scallop. .American Conchologist I7(l);4-5. [March] Abbott, R. Tucker. 1989. Renate Wittig Skinner, New York Shell Club Notes 31 1:4-.5 [June] Abbott, R Tucker. 1989. What's Neu in Molluscan Research'? .American Conchologist 17(2):9 [September] .Abbott, R Tucker. 1989. Museum with a Molluscan Mission. .American Conchologist 17(2): 17. [September] Abbott, R. Tucker. 1989. Renate Wittig Skiimer 1922-1989. American Conchologist 17(2):24. [September] Abbott, R. Tucker 1989, Shells Portland House, New York, 160 pp [December; German Edition, 1990; French Edi- tion, 1990] Abbott, R, Tucker. 1989. Compendium of Landshells, .A Color Guide to More than 2,000 of the World's Terrestrial Shells. American Malacologists, Inc , Melbourne, Florida, i-viii + 1-240. [December] .Abbott, R. Tucker. 1989. Snorkeling Sea Shells. .American Conchologist 17(3):17. [December] Abbott, R, Tucker, 1989. Corinne Elizabeth Edwards (1905- 1989). American Conchologist 17(3):19, [December] Wagner, Robert J. L, and R. Tucker Abbott [Editors], 1990. Standard Catalogue of Shells. [Supplement 4, World Size Records]. American Malacologists, Inc., Melbourne, Flor- ida. 80 pp. [April 27] Abbott, R. Tucker. 1990. Seashells Photo Postcards: 24 full- color ready-to-mail cards. Dover Publications, Inc, Min- eola, NY. 16 pp. [Ma\] Abbott, R. Tucker. 1990. Correcting the Book. "Shells". American Conchologist 18(2 ):25, [June] Abbott, R. Tucker. 1990. Yo\o Clams discovered m East Florida. New York Shell Club Notes No. 315: 10 [June] [Reprinted from Capsule 22(6) .Astronaut Trail Shell Club]. Abbott, R. Tucker. 1990. The Pocket Guide to Seashells of the Northern Hemisphere. Dragons' World Ltd., Limps- field and London, 176 pp. [.August] Abbott, R, Tucker, 1990. Comments on the proposed con- servation of Limax fimbnatus Martsn, 1784 and Merita hebraea Martyn, 1786 (currently Placostylus fimbriatus and Natica hebraea, MoUusca, Gastropoda). Bulletin of Zoological Nomenclature 47(3):202. [September 28] Abbott, R. Tucker. 1990. .American .Nature Guides. Seashells. Gallerv Books, New York. 1-176, [September] [.American Edition of Seashells of the Northern Hemisphere] Abbott, R, Tucker, 1990. Sowerb\ Reborn New York Shell Club Notes 317:8-9. [December] Abbott, R. Tucker. 1990. Zebra Mussel Watcli Invoked in Florida. New York Shell Club Notes 317: 11. [December] Abbott, R, Tucker and S. Peter Dance. 1990, Compendium of Seashells. Crawford House Press, Bathurst, Australia, 411 pp, [.Australian Printing] .Abbott, R, Tucker, 1990. Muscheln unil Meeresschnecken. Karl Muller, \'erlag. Erlangen, 160 pp, [German Edition of Shells, 1989] Abbott, R, Tucker. 1990. Coquillages. Editions Soline, Cour- bevoie. 160 pp. [French Edition of Shells, 1989], Abbott, R. Tucker. 1991. The Life Span of Mollusks. Amer- ican t;onchologist I8(4):25. [Januarv 23] Abbott, R, Tucker. 1991. Shellmg in Southeast Asia. Of Sea and Shore 13(4):148-154. [February 4] Abbott, R. Tucker. 1991. Seashells of the Northern Hemi- sphere. Gallery Books, New York. 1-191. [March] Abbott, R. Tucker. 1991 Review of "Sabelli, B.. R Gianuzzi Savelli and B. Bedulli [eds ]. 1990 .\nnotaled Check-list of Mediterranean Marine Mollusks, Vol. I. Soc. Italiana Malacologia, Bologna, ItaK. 348 pp." American Conchol- ogist 19(1):23. [March] Abbott, R. Tucker, 1991, Review of "Rios, Cedar Garcia 1990, Los (^)uintones de Puerto Rico. University de Hu- macao, Humacao, PR, 48 pp," .American Conchologist 19(1):23. [March] Abbott, R. Tucker, 1991, Seashells ol Southeast Asia, Tvnron Press, Thornhill, Dummfriesshire 1-145, plates 1-52 [Ma\ ] Lipe, Robert E, and R Tucker Abbott 1991 Living Shells M. G. Harasewych, 1997 Page of the Caribbean and the Florida Ke\s, American Mala- cologists. Inc. Melbourne, Florida. HO pp. [May 27] Abbott, R. Tucker. 1991. Mollusk Research ni Bermuda American Conchologist 19(3):7. [September] Abbott, R. Tucker. 1991. Review of "Dance, S. Peter and David Heppell. 1991. Shells — Classical Natural History Prints. Studio Editions, London, 128 pp.' American Con- chologist 19(3): 11. [September] Abbott. R. Tucker. 1991. Those Puzzling Olive Shells. Amer- ican Conchologist 19(3):23. [September] Abbott, R. Tucker. 1991. Review of 'Dance, S. P. and D. Heppell. 1991. Shells — Classical Natural History Prints. Studio Editions, London 128 pp." New York Shell Club Notes 323:8. [June] Abbott, R. Tucker. 1991. The Shell Collector Threat. Sea Frontiers 37(5): 14-19. [October] Abbott, R. Tucker. 1991. Was Christopher Columbus a shell collector'-' Shell Museum Musmgs. Island Reporter [Sani- bel-Captiva] 19(43):5B. [November 1] Abbott, R. Tucker. 1991. The first lady of Sanibel shelling. Shell Museum Musings. Island Reporter [Sanibel-Captua] 19(45);5B. [November 8] Abbott, R. Tucker. 1991. What's in a name'.-' Shell Museum Musings. Island Reporter [Sanibel-Capti\ a] 19(46):5B. [No- vember 15] Abbott, R. Tucker. 1991. Golden shells. Shell Museum Mus- ings. Island Reporter [Sanibel-Captiva] 19(47):4B, 5B. [No- vember 22] Abbott, R. Tucker, I99I. Your shell collection - dump it or save it':* Shell Museum Musings. Island Reporter [Sanibel- Captiva] 19(48):4B. 5B. [November 29] Abbott, R Tucker. 1991. Commensals, parasites, and pearls. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 19(49):4B. [December 6] Abbott, R. Tucker. 1991, Edgar Allan Poe and other Pizzle Editions. Shell Museum Musmgs. Island Reporter (Sanibel- Captiva] 19(50):5B, 14B. [December 13] Abbott, R. Tucker. 1991. Honey, I shrunk the shells! Shell Museum Musmgs Island Reporter [Sanibel-Captiva] 19(51): 4B, 5B. [December 20] Abbott, R. Tucker. 1991. With lose Irom Russia. Shell Mu- seum Musings. Island Reporter [Sanibel-Captiva] 19(52): 4B, 5B. [December 27] Abbott. R. Tucker. 1991. New C;0.\ Grants. American Con- chologist 19(4):17. [December] Abbott, R. Tucker. 1991. Let's go shellingl Or shall we'r" New York Shell Club Notes 321.14-17. [December] [Reprinted from "Sea F'rontiers " October 1991 under the title "The Shell Collector Threat"] Abbott, R. Tucker. 1992. Showers of Shells. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 20(1):4B. [Jan- uary 3] Abbott, R. Tucker. 1992, Dangerous mollusks and zebras. Shell Museum Musings. Island Reporter [Sanibel-C^aptisa] 20(2):4B. [January 10] Abbott, R. Tucker 1992. Museum acquisitions. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 20(3):8B-9B. [January 17] Abbott, R. tucker. 1992. The Sanibel Loosa Beads. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 20(4): 13B. [January 24] Abbott, R. Tucker. 1992. Bhie-Ribbon conchologists. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 20(5): 6B-7B. [Januarx 31] Abbott, R. Tucker. 1992. Shells kill birds Shell Museum Mus- ings. Island Reporter [Sanibel-Captiva] 20(6):11B. [Feb- ruary 7] Abbott, R. Tucker. 1992. The Spectacular Scallop Shell Mu- seum Musings. Island Reporter [Sanibel-Captiva] 20(7): 6B-7B. [February 14] Abbott, R. Tucker. 1992. The day an oil spill hits Sanibel. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 20(8): 17B. [February 21] Abbott, R. Tucker. 1992. Dogs and shell collectors. Shell Mu- seum Musings. Island Reporter [Sanibel-Captiva] 20(9): 17B. [February 28] Abbott, R. Tucker. 1992. It's show and shell time! Shell Mu- seum Musings. Island Reporter [Sanibel-Captiva] 20(10) Shell Fair Guide: 8-10. [March 6] Abbott, R. Tucker. 1992. Fallen Angels, part one. Shell Mu- seum Musings. Island Reporter [Sanibel-Captiva] 20(11): lOB-llB [March 13] Abbott, R. Tucker. 1992. Fallen Angels, part two. Shell Mu- seum Musings. Island Reporter [Sanibel-Captiva] 20(12): 9B. [March 20] Abbott, R. Tucker, 1992. Earl\ Florida shell collectors. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 20(13): 16B-17B. [March 27] Abbott, R, Tucker. 1992. Review of "Leal, Jose H 1991. Marine Prosobranch Gastropods from Oceanic Islands off Brazil. V. B. S. Publishers, Holland 418 pp." New Y'ork Shell club Notes 322:5 [March]. The identical review appeared in Hawaiian Shell News 40(3):8 [March] and American Conchologist 20(2):20. [June] Abbott, R. Tucker. 1992. The miracle of Granular Ark Mus- sels. Shell Museum Musings. Island Reporter [Sanibel-Cap- tiva] 20(14):5B. [April 3] Abbott, R. Tucker. 1992. [Untitled]. Shell Museum Musings. Island Reporter[Sanibel-Captiva]20(15):4B-5B [April 10] .Abbott, R. Tucker. 1992. Museum mission — inspiring kids. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 20(16):5B. [April 17] .Abbott, R. Tucker. 1992. Giant African snails — again! Shell Museum Musings. Island Reporter [Sanibel-Captiva] 20(17): 5B-6B. [April 24] .Abbott, R. Tucker. 1992. Thomas Sa\ — America s first con- chologist. Shell Museum Musings. Island Reporter [Sanibel- Captiva] 20(18):6B. [May 1] Abbott, R, Tucker. 1992. The shell people - the Calusa In- dians. Shell Museum Musings. Island Reporter [Sanibel- Captiva] 20(19):5B, 9B. [May 8] Abbott, R. Tucker. 1992. [Untitled]. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 20(20):4B. [May 15] .Abbott, R, Tucker 1992. Pearl buttons from river mussels. Shell Museum Musings. Islaiul Reporter [Sanibel-Captiva] 20(21):10B. [May 22] Abbott, R. Tucker, 1992. Pioneer in nature's garden. Shell Museum Musings, Island Reporter [Sanibel-Captiva] 20(22): 2B. [May 29] Abbott, R. Tucker. 1992. Review of the new journal RL TH- ENICA. Hawaiian Shell News 40(5):7 [Ma\ ] Abbott, R. Tucker. 1992. Review of "Rice, T. 1992. Catalog of Dealers' Prices, including Freshwater and Terrestrial Shells 1 1th Edition. Of Sea and Shore, Port Gamble, WA. 112 -H 44 pp." Hawaiian Shell News 40(5):7 [May] Abbott, R. Tucker. 1992. Shelling laws and your license. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 20(23): 2B, 7B. [June 5] Abbott, R. Tucker. 1992. Museums aren't attics Shell Mu- Page 74 THE NAUTILUS, Vol. 110, No. 2 seum Musings, Island UepDrtiT [banibei-tJaptiva] 20(24): 3B, 5B. [June 12] .■\l)l)ott, R. Tucker. 1992, Biodiversity — buzzword or boon- doggle? Shell Museum Musings. Island Keporter [Sanibel- Captiva] 20(25):2B-3B. [June 19] Abbott, R. Tucker, 1992, VMien the eyes have it. Shell Mu- seum Musings, Island Reporter [Sanibel-Captiva] 20(26): 2B-.3B, [June 26] Abbott, R, Tucker, 1992. COA Grant Helps Scallop Study. American Conchologist 20(2): 19. [June] Abbott, R. Tucker. 1992, Shell \Uiseum Musings. American Conchologist 20(2 ):22. [June] Abbott, R. Tucker. 1992. RLTHENICA — a new journal. New York Shell Club Notes 323:8, [June] Abbott, R, Tucker. 1992. Scallop Aquaculture. Shell Mu- seum Musings, Island Reporter [Sanibel-Claptiva] 20(27): lOB, [JuK 3] Abbott, R, Tucker, 1992, Supreme Court and shells. Island Reporter [Sanibel-Captiva] 20(28):2B, 5B, [July 10] Abbott, R. Tucker. 1992. Dreams do come true. Shell Museum Musings. Island Reporter [Sanibel-Captiva] 21(28):2B-3B. [July 17] Abbott, R, Tucker. 1992. Endangered Florida species. Shell Museum Musings, Island Reporter [Sanibel-Captiva] 21(29): 5B. 9B, [July 24] Abbott, R. Tucker, 1992, Calilornia I, O, Us and Pismo Clams. Island Reporter [Sanibel-Captiva] 21(30):4B-.5B. [July 31] Abbott, R. Tucker, 1992. Those sex-crazed squids. Island Re- porter [Sanibel-Captiva] 21(32):2B, 4B. [August 7] Abbott, R. Tucker. 1992, Honey, I blew up the shells! Island Reporter [Sanibel-Captiva] 21(33):3B. [August 14] Abbott. R. Tucker, 1992, The making ol a Florida shell book. Island Reporter [Sanibel-Captiva] 21(34):2B, .5B, [August 21] Abbott, R, Tucker, 1992, Ghosts ot Clornell's conchological past. Island Reporter [Sanibel-Captiva] 21 (35):.5B, 6B. [.-Vu- gust 28] Abbott, R. Tucker. 1992. Subtropical red tides; Sanibel shells. Island Reporter [Sanibel-Captiva] 21(.36):2B, [September 4] Abbott, R, Tucker. 1992. Psychiatr> , shells and Sanibel Island Reporter [Sanibel-Captiva] 21 (.37 ):3B-4B, [.September 11] Abbott, R, Tucker, 1992. The making ol a museum exhibit. Island Reporter [Sanibel-Captiva] 2) (38):2B-3B. [Septem- ber 18] Abbott, R. Tucker, 1992. The Sacred Chank. Island Reporter [Sanibel-Captiva] 21(39):2B-3B. [September 25] Abbott, R. Tucker. 1992 Review ol "Winner, Beatrice B. 1992. A Field Guide to MoUuacun Spawn. Volume II., E. B. M. Publishing, North Palm Beach, FL94 pp." Amer- ican Conchologist 20(3):20, [September] Abbott, R, Tucker, 1992, Is the Shell Museum relevant '' Island Reporter [Sanibel-Captiva] 21(40):2B-3B, [October 2] Abbott, R. Tucker. 1992. Museum traveling exhibits. Island Reporter [Sanibel-Captiva] 21(41):2B-3B, [October 9] Abbott, R Tucker 1992. Atlanta's new hall of shells. Island Reporter [Sanibel-Captiva] 21(43);2B-3B. [October 23] Abbott, R, Tucker, 1992. Spondvlus Medicine— good and bad American Conchologist 20(4): 1 8 [December] Abbott, R. Tucker, 1992, Dreams do come true. The Junonia December 19927-9 Abbott, R, T, 1993, Robert J, L, Wagner (190.5-1992). Amer- ican Conchologist 21(I):18, [March] Abbott, R. Tucker. 1993. The makings ol a shell show. The 56th Annual Shell Fair (iuide [Supplement to the Island Reporter, Sanibel] pp. 1.3-15, 22, 24. [March] .■\bbott, K, Tucker, 1993, "\\ orld Size Records Continues. American Conchologist 21(3):5. [September] Abbott, R. Tucker 1993, Review of "Winner, Beatrice B. 1993, Life Styles of the Seashells. E, B. M, Publishing, North Palm Beach, EL 61 pp, " .-\mericaii Conchologist 21(4):18, [December] Abbott, R. Tucker. 1993. Seashells ol Great Britain and Eu- rope. Junior Nature Guide. Dragon's World, Ltd., London, 80 pp, [Edited b\' T, Rowland-Entwistle] Abbott, R. Tucker, 1993. Kingdom of theSeashell, the color! ul story of shell collecting and living mollusks. [revised edi- tion]. American Malacologists, Inc. Melbourne, Florida 256 PP Abbott, R, Tucker, 1993, Robert J, L, Wagner (190.5-1992). New York Shell Club Notes .326:16, [March] Abbott, R, Tucker, 1994, Phalium (Semica.ssis) vector, a New Deep- Water Species from the Central Indian Ocean, The Nautilus 107(3):94-96. [February 2] .Abbott, R. Tucker. 1994. You can fool some of the people. . . Island Reporter [Sanibel-Captiva] 23(16):10A. [April 22] Abbott, R. Tucker, 1994. Review of ■'Abbot(t), R.Tucker, 1993 Seashells of North America. Dragon's World, Lon- don , 81 pp. American Conchologist 22(1 ):18, [June] Abbott, R, Tucker. 1994. Review of "Ingoglia, G, 1994. Sea.ihells. Science Close-Up Series. Western Publishing CO., Racine, WI, 24 pp.. .American Conchologist 22(1): 19, [June] Abbott, R. Tucker. 1994, Fareu ell to a Shelling Friend. [Obit- uary— Mary Kline]. Hawaiian Shell News 42(9):9. [Sep- tember] Abbott, R. Tucker. 1994. The best beaches for shell collectors. Travel & Leisure 24(12):76, 80, [December] Hess, D F,, R T, Abbott, J, Hamann. K Meyer, S Millen. T Gosliner, N. Sefton and Roger T, Hanlon, 1994 9, Ma- rine Molluscs of the Cayman Islands. In: Brunt, M, .\. and J. E. Davies (eds. ) The Cayman Islands: Natural History and Biogeography Kluwer .Academic Publishers, Nether- lands, pp, 139-189, Abbott, R, Tucker and Perc> A. Morris. 1995. A Field Guide to Shells, .Atlantic and Gulf Coasts and the West Indies. Fourth Edition, Houghton Mifflin Compain. New York 350 pp., 74 pis, [.April] Abbott, R. Tucker, 1995, My Favorite Seashells, Boys Life, June 1995:24-27, [June] Abbott, R, Tucker. 1995. A Guide to Shelling, Siiiuu Day Guide 3(1 ):28-30,32, 34. Abbott, R. T, 1995. Bailey-Matthews Shell Museum, Cultural Jewel of Sanibel, Sunny Day Guide 3(1):.33, Abbott, R. Tucker, 1996, Some Children's Shell Books— A Review. Of Sea and Shore 19(3):150, [Fall— published posthumously] ,At the time of hi.s death, R. Tucker .Abliott was working on a number of books. Among these was our collaboration on the third edition of American Seashells, on wiiicli he w ill be a co-author. BOOKS EDITED BY R. TUCKER ABBOTT .Abbott, R, Tucker, M K, Jacobsciri. and M G Teskc\ (Editors). 1955. How to Collect Shells (A Symposium). .American Malacological Union, Bulfalo, New York, [iv] -t- 75 + [vii] pp M. G. Harasewych, 1997 Page 75 Abbott, R. Tucker, M. K^ Jacobson, and M. C. Teske\ (Editors)- 1961. How to Collect Shells. (A Symposium). Second Edi- tion. American Malacological I'ninn, Marinette, Wisconsin iv + 92 + [i\ ] pp. .Abbott, R. Tucker, M. K, Jacobson, and M. C:. Teske\ (Editors). 1966. How to Collect Shells. {.\ S\niposiuni). Third Edi- tion. American Malacological L nion, Marinette, Wisconsin iv -I- 101 -1- [v] pp. Abbott, R. Tucker, M. K. Jacobson, and M. C. Teskey (Editors). 1974. How to Collect Shells. (A Symposium), Fourth Edi- tion. American Malacological Union, Wrightsville Beach, North Carolina [vi] + 107 pp. Hasting, Louise Burrell and Margaret Crockett Teske\ . 1979, [Edited b\ R. Tucker Abbott], Indexes to The Nautilus: Geographical {\'ols. 1-90) and Scientific Names (X'ols, 61- 90). .-Kmerican Malacologists, Inc. Melbourne, Florida. i\ -I- 238 pp. Sutty, Lesley. 1986. [Edited b\ R. Tucker Abbott]. Seashell Treasures of the Caribbean. Dutton, New York. 128 pp. Bratcher, T. and W. Cernohorsk). 1987. [Edited b> R. Tucker Abbott]. Living Terebras ot the World : .\ Monograph of the Living Terebridae of the \\ orld .American Malacol- ogists, Melbourne, Florida. 240 pp. Naught, K, C, 1989, [.\bbott, R. Tucker and Kenneth J. Boss, Editors], .\ Classification of Living Mollusca, .American Malacologists, Inc, Melbourne, Florida, i-,\ii + 1-195, ACKNOWLEDGMENTS I am most grateful to Cecelia W, .Abbott for allowing access to Tucker's personal files and scrapbooks and for supplying many of the photographs as well as biograph- ical information, Jose H. Leal and Edith Chippeau.x brought to m\ attention some of Tucker's notes and card files that were found at the Bailev -Matthews Shell Mu- seum. Ellen Strong kindly pro\ ided detailed information on his Master s Thesis and Doctoral Dissertation. I thank Richard E. Petit for comments on tiie manuscript, ad- dition of citations, and information on editions and sup- plements. Alan l\. Kabat brought to my attention several Abbott ta.\a. Melbourne R. Carriker kindly made avail- able a draft of his "In Memoriuni manuscript. Paula M. Mikkelsen and Robert Robertson generously provided photographs of Tucker. Special thanks are due Cecelia Abbott, Kenneth J. Boss, Melbourne R. Carriker, Russell H. Jensen, Jose H. Leal, Paula M. Mikkelsen, and f^obert Robertson for their comments on a draft of this manu- script. LITERATURE CITED .Anonymous, 1971, Publications of the I'nited States National Museum (1947-1970). Bulletin 298 (final volume of series). Smithsonian Institution Press, Washington, DC. 77pp. Bieler, R. and R. E. Petit. 1996. Additional notes on nomina first introduced by Tetsuaki Kira in Coloured Illustrations of the Shells of Japan. Malacologia 38(l-2);33-34. Coan, E. V. and M. G. Harasewvch, 1993. Publication Dates of The Nautilus. The Nautilus 106(4):174-180. Lyons, W. G. 1988. A Review of Caribbean Acanthochiton- idae (Mollusca: Polyplacophora) with descriptions of si.x new species of Acanthochitona Gray, 1821. American Malacological Bulletin 6(1):79-114. THE NAUTILUS 110(2): 76, 1997 Page 76 110(2) PetitiUa, New Name for Petitella Wise, 1996, a Preoccupied Name (Mollusca: Gastropoda: Pyramidellidae) John B. Wise Houston Museum of Natural History Oiif Hermann Circle Drive Houston TX 77030-1799 USA In a recent study of pli\logeneti(.' relationships within the famiK PyramideUidae, I proposed the generic name Petitella \\'ise, 1996, witiiiii the new subfamily Sayel- hnae. The editors of Zoological Record have since brought to my attention that Petitella is preoccupied, having previously been apphed to a genus of fish (Gery & Bou- tiere, 1964). A replacement name is therefore proposed. PetitiUa. new name for Petitella Wise, 1995:495, not Petitella Gery & Boutiere, 1964:474. Type species, by original designation of Petitella Wise, 1996, Sayella crosseana Dall, 1885, Recent, Western At- lantic. The new name honors Richard E. Petit for his many invaluable contributions to malacology, as did the name it replaces. 1 thank the eilitors of Zoological Record lor bringing this to m\ attention. LITERATURE CITED Dall, W. H. 1885. Notes on some Floridian land and fresh- water shells with a revision of the .'\uriculacea of the east- ern United States. Proceedings of the L'nited States Na- tional Museum 8:225-289, pis. 17-18. Gery, J., and A. Boutiere. 1964. Petitella georgiae gen. et. sp. nov. (Pisces CypriniformesCharacoidei). X'ieet Milieu, Supplement 17:47:3-484. 6 figs. Wise, J. B. 1996. Morpholog) and pin logenetic relationships of certain pyramidellid ta\a (Heterobranchia). Malaco- logia 37(2):443-511. INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes papers on all aspects of the biology' and svstematics of mollusks. Manuscripts de- scribing original, unpublished research as well as review articles will be considered. Brief articles, not exceeding 1000 words, will be published as notes and do not re- (juire an abstract. Notices of meetings and other items of interest to malacologists will appear in a news and notices section. 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Harascwych Division of Mollusks National Museum of Natural Histoiy Smithsonian Institution Washington, DC 20560 MANAGING EDITOR Dr. Jose H. Leal Bailev-Matthews Shell Museum 3075 Sanibel-Captiva Road Sanibel, FL 33957 CONSULTING EDITORS Dr. Riidigcr Bieler DepartnuMit of Invertebrates Field Museum of Natural History Chicago, IL 60605 Dr. Arthur E. Bogan Freshwater Molluscan Research 36 Venus Way Sewell NJ 0S080 Dr. Robert T. Dillon, Jr. Department of Biologv' College of Charleston Charleston, SC 29424 Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natmal History New York, NY 10024 Dr. Eileen H. Jokinen Institute of Water Resources University oi Connecticut Storrs, Ct 06269-4018 Mr. Richard I. Johnson Department of Mollusks Museum oi Comparativi' 7.i » ili ii_r\ Harvard University Cambridge, MA 02138 Dr. Aurele La Rocque Department of Geology Tlie Ohio State University Columbus. OH 43210 Dr James H. McLean Department of Malacology Los Angeles County Museum of Natural Historv' 900 Exposition Boulevard Los Angeles, CA 90007 Dr. Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoology Harvard Universits' Cambridge, MA ()2L38 Dr. Paula M. Mikkelsen Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Dr. Doniild R. Moore Division of Marine Geology and G(^ophysics Rosenstiel School of Marine and Atmospheric Science University of Miami 4600 Rickenbacker Causeway Miami, FL 33149 Dr, Ciustav Paulay Marine Laboratoiy University of Guam Mangilao, Guam 96923 Mr. Richard E. Petit PO. Box 30 North Myrtle Beach, SCJ 29582 Dr. Edward J. Petuch Department of Geologv' Florida Atlantic University Boca Raton, FL 33431 Dr (Jai-y Rosenberg Department of Mollusks The .Academy oi Natural Sciences 1900 Benjamin Franklin Park-way Philadelphia, PA 19103 Dr. David H. Stansbeiy Museum of Zoolog)' The Ohio State Universit)' CJol.unbus. OH 43210 Dr. Ruth D. Turner Department o( Mollusks Museum oi Comparative Zoology Harvard University Cambridge, MA 02138 Dr. Geerat J. Vermeij Department of Geologv' University of California at Davis Davis, CA 95616 Dr. G. Thomas Watters Aquatic Ecologv' Laboratorv' 1314 Kinnear Road Columbus, OH 43212-1194 SUBSCRIPTION INFORMATION The subscription rate per volume is US $28.00 for individuals, US $45.00 for institutions. Postage outside the United States is an additional US $5.00 for surface and US $15.00 for air mail. All orders should be accompanied by payment and sent to: THE NAUTILUS,' PO. Box 7279, Silver Spring, MD 20907-7279, USA. Chanii^c of address: Please inform the publisluM' of your new address at least 6 weeks in advance. All communications should include both old and new addresses (with zip codes) and state the effective date. THE NAUTILUS (ISSN 0028-1344) is publislied (luarterly by Trophon Corporation, 8911 Alton Parkway, Silver Spring, MD 20910. Periodicals postage paid at Silver Spring, MD and additional mailing offices. POSTMASTER: Send address changes to: THE NAUTILUS PO. Box 7279 Silver Spring, MD 20907-7279 TH EfcNAUTI LUS CONTENTS \\>litntc HI). Xitmhcr 3 juhj IS, 1997 ISSN '002S-1344 Claudia Julia del Ri'o (>\'n()/.()if Bioijeotiraphic Histon' ot the Eiintlit'iinal (hmius Rclroldpcs. New Cieiius (SuhtainiK" Tapctiiiae) Iniiii Sdutlii'in Soutli America and Antarctica 77 Iphittts roheiisi (Epitoniidae: Nvsticllinae). A New Species of Deep-Sea Gastropod from the Gnlf oi Mexico 94 N'erification of the Specific Status of tlie Eutlangered Anthony's River Snail. Atlicdniin (iiilliniii/i. Using Alloz\ine Electroplioresis 97 Inilnced Metamorphosis of Fresliwater Mussel Cilochidia on Nonliost Fisli 102 Bruno Sabelli Marco Ta\ iani Robert T. Dillon, Jr. Steven A. Alilstedt Sheila G. Kirk James B. Lavzer THE NAUTILUS 110(3):77-93, 1997 Page 77 Cenozoic Biogeographic History of the Eurythermal Genus Retrotapes, New Genus (Subfamily Tapetinae) from Southern South America and Antarctica Claudia Julia del Rio Centre de Investigaciones en Recursos Geologicos Ramirez de Velazco 847 (1414) Buenos Aires ARGENTINA and Facultad de Ciencias Exactas y Naturales L'niversidad de Buenos Aires Ciudad Universitaria Pabellon 2 Nunez (1428) ARGENTINA ABSTRACT Retrotapes. new genus, comprises a group of Neoaustral bi- valves that appeared in the southern circumpolar regions b\ the Eocene and ha\e since been confined to the high latitudes of the southern hemisphere. Its presence in the Tertiar\ strata of Antarctica and southern South America reflects an active fauna! interchange between both regions during the Eocene. Retrotapes is here proposed to include to those Recent and Tertiarv representatives of the Subfamily Tapetinae (Family Veneridae) from southern South .America and .Antarctica that had been previous!) placed in Venus Linne. 1758, Marcia H. and .•\. .Adams, 1857, Eurhomalea Cossmann,1920, Samarangia Da!!, 1902, and Kalelijsia Romer, 1857. The Argentine Tertiar> species Retrotapes r^infasiensis, new species, R. fuegoensis. new species, R. striatolaniellata (Ihering, 1897) and fi. scutata (Ihering, 1907) are here described and illustrated. The Recent Argentine Venus exalbida Chemnitz, 1795 and Venus lenlic- ularis Sowerby, 1835, the Antarctic V. antarctica Sharman and Newton, 1894 and \', newtoni Wilckens, 1911 (Eocene-early Oligocene', La Meseta Formation), and the Neogene Chilean species V. navidadis Philippi, 1887 and \'. colchaguensis Phi- lippi, 1887 are also included in this new genus. Key words. Tapetinae, Retrotapes, new genus, biogeograpin, Neoaustral, Tertiary, .Argentina, Chile, .Antarctica. INTRODUCTION The Subfamii\ Tapetinae (FamiK Veneridae) shows a moderatel) high degree of endeniism in Recent as well as in Tertiary faunas. The known geographic distribution of most living tapetines is mainly restricted to the south- ern hemisphere. This subfamiK is particular!)' abundant in the southern Indo-Pacific region, where it is repre- sented by more than sevent) species. Most of the Indo- Pacific genera, among them Katelysia Romer, 1857, Marcia H. & A. Adams, 1857, Granicorium Hedley, 1906, Hemitapes Romer, 1864, Notirus Finlay, 1928, Notopaphia Oli\er, 1923, Eumarcia Iredale, 1924, Pa- phirus Finia), 1927, and Gomphinella Marwick, 1927, are restricted to New Zealand and Australia. Others, including Gonjp/nna Morch, 1853, Venerupsis Lamarck, 1818, Ruditapes Chiamenti, 1900, Tapes Mergele von Miihlfeld, 1811, and Paphia Roding, 1798, are also found beyond those regions. In contrast. Recent tapetines are poorly represented in the littoral zones along both coasts of the American continents. The subfamily is known from only twelve living North American species distributed among the genera Riulitapca, Liocyma Dall, 1870, Psephidia Dall, 1902 and Inn Schmidt, 1818, as well as five South Amer- ican taxa assigned to the genera Eurhomalea Cossmann, 1920 and Retrotapes. new genus. The fossil record in the .Americas reveals that Tape- tinae were more abundant in the Tertiary than in the Recent fauna. Two diverse and different assemblages appeared during the Tertiary, one restricted to North America, the another to austral latitudes. The northern assemblage comprises the endemic genera Liocyma, Cy- clorisma Dall, 1902, Psephidia and Sii}onia Stephenson, 1952, as well as taxa with european affinities such as Mercimonia Dall, 1902, Flaventia Jukes-Browne, 1908, Paraesa Case) , 1952 , Legumen Conrad, 1858 and Tex- tiveniis Cossmann, 1886. The less diverse austral assem- blage, characterized by Eumarcia, Katelysia, Atamarcia Marwick, 1927 and Retrotapes. appeared during the ear- ly Tertiary in the southernmost region of South America and Antarctica. Page 78 THE XALTILLS. \ ol 110, No. 3 v'^ 1 \P E « U 70°) 60* C 50° ^^—d 20°. ( / BOLIVIA j ^ .20" \ Vj ~V ^•. "'="- \ S /■ ^■■v *> u < ■"•^ 7 \ '■ •/ .^ r^ o 1 ,' '—>■ >' f o / / y 9- J '^ COQuimboJO; y * y 30°. ..0°. 1 1^/ ' ^\ /^^ 2 ValporoisoV 'v ^ OHUGUArf y^to Grond« O SANTIAGO] •! • K'' ( / o / ^\ ^k Malvinos Klonds ^7^^J 50°. '41 V ■^r^ L__b»1' 1 ~^-' .»• ■ ■- 60° 70° 60° ,1/5'i Stymour • EOCENE LATE OLIGOCENE- ^/ Ulond ■ EARLY MIOCENE / c y A MIDDLE MIOCENE z/ del Rio (1995). Only one fairK- well- preserved valve of Retrotapes is known from the up- permost, highly-fossiliferous, orange coquinoid sand- stones that alternate with green or yellowish gra\ silstones and sandstones exposed at Cafiadon El Lobo. Retrotapes is poorly represented in the El Chacay Formation, exposed at Cerro Puntudo (Lago Cardiel). Chiesa and Camacho (1994) placed this unit in the late Eocene based on biostratigraphic correlations with the San Julian Formation. Specimens described in this paper are housed in the Museo Argentino de Ciencias Naturales "Bernardino Ri- vadavia (MACK), Buenos Aires, Argentina, in the Cen- tro de Investigaciones en Recursos Geologicos (CIRGEO- PI), Buenos Aires, Argentina, Facultad de Ciencias Ex- actas y Naturales of the Universidad de Buenos Aires (CPBA), Buenos Aires, .Argentina, Direccion Nacional Servicio Geologico (DNSG), Buenos Aires, .Argentina and in the Department of Earth and .\tmospheric Science, Purdue University (PU), Indiana, USA. SYSTEMATIC PALEONTOLOGY Phylum MOLLUSC A Linne, 1758 Class BIVALVIA Linne, 1758 Subclass HETERODONTA Neumavr, 1884 Order VENEROIDA H. & A. Adams, 1856 Superfamilv N'ENEROIDEA Rafinesque, 1815 Familv \'ENERIDAE Rafinesque, 1815 Subfamilv TAPETIXAE H. & A. Adams, 1857 Diagnosis: Members of this subfamily are characterized b\ having smooth inner ventral margins and three car- dinal teeth in each valve, with at least two of them grooved or bifid. Most Tapetinae have exterior surfaces with commarginal lines and/or grooves, while others have polished surfaces or very fine radial threads. Though no attention has been paid to the presence of a lunule or the arrangement of cardinal teeth, these characters, along with the general outline of the cardinal platform, are of taxonomic importance at the generic level. When present, and viewed dorsally, the lunule may be concave or convex, and is limited b> a groove and/or ridge that is distinguished from the remaining surface of the shell b>' its different ornamentation, or b> the presence of a groo\e. Hinge characters ha\e pro\en to be the most useful morphological feature for distinguishing among genera. Cardinal teeth ma\' be placed on a thin, slender, cardinal platform with a straight ventral margin, or situated on a short, broad, strongly arcuate platform. While some genera are characterized b\ basing teeth that diverge from a point situated below the beaks, others ha\e non- divergent teeth (Figure 3). Divergent teeth are charac- terized by a strongly forward-inclined anterior tooth, a vertical median tooth, and a backward-inclined posterior tooth. Genera with non-divergent teeth may have three cardinal teeth sloping backwards (posterior tooth nearly Page 80 THE NAUTILUS, Vol 110, No. 3 pas divergent teeth non-divergent teeth Figure 3. Hinge teeth t\ pes in Tapetinae. aas, anterior atlductor muscle scar; lu, lunule; pas, posterior adductor muscle scar; l,3a,3b, cardinal teeth. horizontal) or have the median and posterior teeth in- clining backwards, with the anterior tooth being vertical. Genus Retrotapes, new genus Patagornalea del i^io, 1991 ;93, nomen nudum. Diagnosis: Shell large to medium sized, highly variable in outline. Escutcheon facing opposite valve, usually wid- er, smoother on left valve than on right in adults. Lunule large to medium sized, deeply impressed, strongly dif- ferentiated from remaining shell surface, highK to mod- erately concave, nearly vertical, with longitudinal me- dian sulcus, bounded b\ deep lunular groove. Hinge heavy, short, arcuate behind teeth, with three long car- dinal teeth. Right and left posterior teeth nearly hori- zontal in some species, left and right anterior teeth nearly vertical to strongly sloping backw ards, nearK paralell to posterior teeth; left median tooth, right median and pos- terior teeth bifid or grooved. Type species: Retrotapes ninfasiensis , new species. Type locality: Valdes Peninsula, Chubut Province, Ar- gentina, Puerto Madryn Formation (Middle Miocene). Stratigraphic range: Eocene to Holocene, southern South America and .Antarctica. Etymology: retro- L. backwards. Referring to the incli- nation of non-divergent cardinal teeth towards the pos- terior region of the shell. Remarks: Recent as well as fossil species belonging to Retrotapes ha\e pre\ iousK- been placed in Venus Linne, 1758, (Philippi, 1887; Ihering, 1897), in Marcia (Dall, 1902; Ihering. 1907; Riveros & Gonzalez, 1950), in Sa- marangia Dall, 1902 (Jukes-Browne, 1908; C^arcelles, 1944; Carcelles & Williamson, 1951; Castellanos, 1970 Rios, 1975), in Eurhonialea (Keen, 1954; Soot-Ryen, 1959 Raniorino, 1968; Fisher-Piette & Vukadinovic, 1977 Malumian el al.. 1978; Zinsmeister, 1984; Stilwell & Zins- meister, 1992) and in Katelifsia (Dall, 1902; Jukes-Browne, 1908; Riveros & Gonzalez', 1950). The taxonomic placement ot the Holocene South American species Venus exalbida (Figures 22-23, 41) and Venus lenticularis (Figures 19-21), here assigned to Retrotapes. has long been debated, and the phylogenet- ical relationships of these species within the family Ve- neridae were misunderstood. Dall (1902) incorrectly in- LEFT VALVES 46 2b 2o RIGHT VALVES UM 3o 1 3b / ropes //ferafus {Lmnoeui] 2- Katelysia scalarina (Lomorck) /vAA fvAJX 3- Eufhomalea rufo (Lamarck) tyJXh aiu\ 4 - Retrotapes ninfosiensis n. sp. /u/nn KS\J\ 5- Retrotopes striotolomelloto (Ihering) fvnsK /uui 6- Retrotapes fuegoensis n. sp. 7- Retrotapes lenticularis (Sower by) /UHA AAn 8- Retrotopes exalbida (Chemnitz) Figure 4-. Schematic cross-sections of hinge teeth. Tapes lit- eratiis (Linne). M.-\CN 266-1, Indian Ocean, Recent 1,2.4 x); Kalclysia scalarina (Lamarck). PU 863-5, N'ictoria, Australia. Recent (2.4 x); Eurhumalea rufa (Lamarck). PL 165, Chile, Recent. (1.2 x); Retrotapes ninfasier^sis. new species. CPBA 15.110, Fondeadero Ninfas, middle Miocene, Puerto Madr\n Formation, Argentina (1.2 x); Retrotapes striatolamellata (Ihering). MACN 2639, Yegua Quemada, late Oligocene, Mon- te Leon Formation, Argentina (1.2x); Retrotapes fuegoensis, new species. PU 355-12, Cerro Castillo, late Oligocene-early Miocene, Carmen Silva Formation, Argentina (1.2 x); Retro- tapes lenticularis (Sowerby), PL' 165, Chile. Recent (1.2 x); Retrotapes exalbida (Chemnitz) MACN 21 172, .Argentina. Re- cent (1.2 X). terpreted Fischer's (1887) statement about V. exalbida being the type species of the genus Marcia, considereil Sainarangia (t\ pe species Venus quadrangularis Adams & Reeve. 1850) to be a section of Marcia. antl placed V. lenticularis in Sauiarangia. He thought V. exalbida, V. lenticularis and V. quadrangularis to be part of a closeK related group of species related to Marcia Jukes-Browne Claudia Julia del Rio. 1997 Page 81 Figures 5-9. Tapes litcratus (Linne). MACX 266-1. Indian Ocean, Recent. 5. Exterior view of left valve (IX). 6. Interior view of right valve (IX). Enlargements of 7. right and 8. left hinges (2X). 9. Dorsal view of an articulated specimen (IX). Figures 10- 14. Eiirhomalea nifa (Lamarck). PU 165, Chile, Recent. 10. Exterior and 11. interior views of right valve (IX) 12. Dorsal view of an articulated specimen (IX). Enlargement of 13. left and 14. right hinges (2X). Page 82 THE NAUTILUS, Vol. 110, No. 3 (1908) designated Venus pinguis Chemnitz, 1782 (= V. opima Gmelin, 1791 ) as the t\ pe species of Marcia, and excluded V. exalbida from this genus. Venus quadrangularis (see Fischer-Piette & Vukadi- novic, 1977:22, figs.207-211, for illustrations of the spe- cies) cannot be related to either \'. exalbida or \'. Icn- ticiilaris because of the presence in V. quadrangularis of a smooth exterior surface, a pustular left anterior lat- eral tooth and an entire pallial line, characters that al- lowed Keen (1969) to segregate Samarangia in the Sub- family Samarangiinae. Other diagnostic features, in- cluding the hinge and lunular characteristics of V, ex- albida, V. lenticularis, as well as those of their Tertiary ancestors, require the erection of the new genus Retro- tapes. It is placed in the SubfamiK Tapetinae on the basis of the presence of shells with a smooth inner ventral margin, moderate to very deep pallial sinus, three car- dinal teeth in each valve with two of them deeply grooved or bifid, and the absense of lateral teeth. Morphological features that distinguish Retrotapes from the remaining genera of the subfamiK are the presence of non-diver- gent cardinal teeth and a well defined, concave lunule, bounded by a deep lunular groove and ridge. Eurhomalea, represented by its t\pe species E. rufa (Lamarck, 1818) (Figures 4, 10-14) and by £. salinensis Ramorino, 1968 (see Ramorino. 1968:218, pl.3, fig.2, pi. 9, figs.2-3 for illustrations of the type species), is distributed along the Peruvian and Chilean coasts and has recentlv been placed in the Subfamily Chioninae (Fischer-Piette & Vukadinovic, 1977). However, the presence of a smooth inner ventral margin in Eurhomalea indicates that it is more accurately assigned to the Subfamily Tapetinae. It clearly diff^ers from Retrotapes in lacking both lunule and escutcheon, in having a straight dorsal margin, a more antero-posteriorly elongate shell, and a narrower hinge plate than Retrotapes. Moreover, the cardinal teeth are markedly smaller than those of Retrotapes and quite divergent from a point situated below the beaks. The anterior cardinal teeth of both valves in Eurhomalea slope forward. The left posterior and right anterior car- dinal teeth are much lower than in Retrotapes. and the left middle, and right posterior and middle teeth are only shallowly grooved. Katelysia (type species: Venus scalarina Lamarck, 1818) (Figures 4, 25-28), a middle Miocene-Holocene genus confined to New Zealand and Australia, differs in having a diff^erent outline, a narrower lunule and es- cutcheon, a shallower pallial sinus, and a shorter cardinal teeth than Retrotapes. Marcia opima. the Indo-Pacific type species of Mar- cia, was illu.slrated by Abbott and Dance (1986:363). The genus differs from Retrotapes in having a smooth exterior shell surface, a high umbonal area, a weakly defined escutcheon, and a nearly smooth and lightK impressed lunule that lacks a lunular groove. These genera may also be separated becau.se Marcia has lower, narrower and more widely divergent teeth than Retrotapes with the anterior cardinal teeth inclining forward. Eurnarcia (t\pe species Venus fumigata Sowerby, 1853, illustrated by Lamprell & Whitehead, 1992, pl.74, fig.589; Abbott & Dance, 1986:363) a common genus in New Zealand and Australia, is easih separated from Re- trotapes on the basis of its oval and smooth shells, a lunule that is not impressed and that is bounded by a weak line, divergent teeth with left anterior and middle cardinal teeth deepK grooved and equal in size, and in having a left posterior tooth that is fused to nymph. The Tertiary New Zealand Atamarcia (type species Eurnarcia sulcifera Marwick, 1927, figs. 200, 203, 205, earl\ Oligocene-late Pliocene) differs from Retrotapes in being sculptured with commarginal grooves, in having a fairly impressed lunule, with a shallo\\- lunular groove, divergent teeth with curved posterior teeth, and with the left anterior and middle teeth being of equal width. The monot\pic, lower Pliocene Opimarcia Marwick, 1948, (type species: O. healyi Marwick, 1948, pi. 5, figs. 1, 2, 4) is distinguished from Retrotapes in having oval, inflated shells, very prominent beaks, a poorly defined lunule not bounded by a lunular groove, a fairly well- developed escutcheon, fine irregular commarginal ridges on the posterior and anterior ends of valves, a shorter pallial sinus than Retrotapes, and divergent teeth with a strongly curved right posterior tooth and a triangular, deeply grooved left anterior tooth. Tapes (type species: Venus literatus Linne, 1758) (Figures 4-9) has shells that are more antero-posteriorK elongated than those of Retrotapes, and have a straight, horizontal dorsal margin, acuminate anterior margin, a rounded pallial sinus, straight or slightK arcuate hinge margin and a grooved left anterior cardinal tooth. The oldest presentK known record of Retrotapes dates to the late Eocene. This genus occurs in the San Julian Formation and the El Chaca> Formation (.\rgentina), where is represented b\ R. scutata and R. striatolamel- lata, and in the lower and middle sections of the La Meseta Formation (late Eocene-Oligocene":*, ,\ntarctica) where R. aniarctica and R. newtoni have been found. Retrotapes ninjasiensis, new species Figures 15-18, 40 Marcia strialolamellata Frengueiii, 1926, not M striatola- mellata Iliering, 1897. Diagnosis: Shell thick, ovate to subrectangular. Lunule deeply concave, nearly vertical, strongly inclined toward opposite vaKe. Cardinal teeth straight, high, long, thick. Median and anterior teeth sloping backwards, posterior cardinal teeth horizontal. Pallial sinus triangular, mod- erateK short. Description: Shell thick, large, o\ate to subrectangular, weakly convex, ranging from longer than high to nearly equidimensional. Umbones small, at anterior quarter of length. Dorsal margin moderately to strongly convex, posterior margin truncated to weakK convex, ventral margin nearly straight to w eakly convex, anterior margin convex. Lunule concave, with longitudinal median lu- nular groove, nearK' vertical, strongK inclined toward opposite val\ e, bounded b\ deep groove and w ell marked Claudia Julia del Rio, 1997 Page 83 .<.:/ ^^ - >iJv \ -.V # 15 J S:^ "' - \^ ;^ 1 7 18 X i /A**^ 19 20 Figures 15-18. Rctrotapes ninfasiensis. new species. Puerto Madryn Formation, middle Miocene. 15. Interior and 18. exterior views of right vaKe of holotype, CPBA 13.573, Cerro Prismatico. 16. Interior view of left valve of parat>pe, CPBA 15.110, Fondeadero Ninfas. 17. Interior view of left valve of paratype, CPBA 15 090, Punta Norte Figure§ 19-21. Retrotapes lenticularis (Sowerby), PU 165, Cfiile, Recent. 19. Exterior, 20. interior and 21. dorsal views of left valve. .\11 figures 1 x. Page 84 THE NAUTILUS, Vol. 110, No. 3 22 23 /y 24 26 30 25 8W3'^ 27 31 Figures 22-24. Retrotapes exallnda (Chemnitz) MACN 21.172, San Matias Gulf. Argentina, Recent Ail figures 1 x. 22. Interior and 2.1. exterior views of left valve. 24. View of right hinge. Figures 2.5-28. Kateltjxia scalarina (Lamarck), PU 865-5. Victoria, .Xustralia, Recent. All figures 2X. 25. Interior, 26. exterior, and 2K. dorsal views of left valve, 27. Interior view of right valve. Claudia Julia del Rio, 1997 Page 85 ridge. Escutcheon on left \alve wide, long, inclining to- ward right valve and only slightly sculptured with fine commarginal ridges; escutcheon on right valve poorly differentiated and ornamentated as remaining shell sur- face. Hinge short, narrow , stronglv arcuate behind teeth, with three high, thick cardinal teeth per vaKe (.Hgure 4). Anterior and median teeth strongly inclined back- wards, posterior teeth nearK' horizontal. Right valve with deepK groo\ed posterior tooth separated from nymph h\ socket, followed b\ ridge, median cardinal bifid with lamellar posterior section, anterior tooth thick, equal in height to median tooth; socket for left median tooth w ith radial lamella. Left hinge with strong, smooth, anterior tooth, median tooth deepK' grooved, with both parts equal in size, lower than anterior tooth. Posterior tooth lamellar, slightl) arched, with superimposed ridge on posterior face, separated from nymph by shallow groove. Anterior adductor muscle scar o\ al, more deepK impressed, larger than posterior muscle scar; anterior pedal retractor mus- cle scar small, deeply excavated, placed above adductor. Pallial sinus relativeK short, triangular, with horizontal dorsal margin, straight, ascending ventral margin; ape.\ pointed. Exterior ornamented with wideK-spaced, thin, lamellar commarginal ridges, much more numerous near ventral margin; interspaces sculptured with ver> fine radial threads. Material examined: Holotype, CPBA 13.573, right valve, Cerro Primatico, heiglit 66.6 mm, length 76.2 mm. Para- tvpes, CPBA 15.090, left valve, Punta Norte, height 77.1 mm, length 81. 9 mm, CPBA 15.110, left valve, Fon- deadero Ninfas height: 60,0 mm, length: 66,0 mm; 19 left valves, 16 right valves, one articulated specimen, Punta Norte (PN 2): CPBA 15.087-15.089, CPBA 13.287- 13.288, Fondeadero Ninfas (F 9): CPBA 12.345, CPBA 15.1 1 0, Cerro Prismatico (N 2): CPBA 12.501, CPBA 11.646-11.650, CPBA 12.343. CPBA 12.502, CPBA 13.572, 13.574-13.575 (del Rio collection) Stratigraphic and geographic distribution: Puerto Mad- ryn Formation, middle Miocene. Valdes Peninsula, Chu- but Province. Remarks: Specimens of Retroiapes ninfasiensis have erroneously been attributed to Mania striatolamellata (Frenguelli. 1926). a closely related, late Oligocene spe- cies that occurs in the Monte Leon Formation of Santa Cruz Province. Retroiapes striatolamellata (Figures 33, 37-39, 42-47) ma>' be differentiated b\ its longer, more convex shells, straighter dorsal margin, a lunule that is more concave and not inclined toward the opposite valve, less prominent teeth, a left anterior cardinal tooth that is less inclined backw ards, being slightK arched forwards in most specimens and higher or equal in height than the median cardinal (Figure 4), and a pallial sinus that is tongue-like and longer than in R. ninjasiensis. Retro- tapes antarctica (Sharman & Newton. 1894. fig. 3). from the La Meseta Formation (Antarctica. Eocene- early Oli- gocene?), differs in having a trigonalK suboval outline with the dorsal margin strongK sloping backwards and rounding to the posterior margin, by the presence of a shallow ly conca\e, narrow er lunule not inclined toward the opposite valve, and a narrower, shorter pallial sinus than R. ninjasiensis. Moreover, R. antarctica has nar- rower teeth than R. ninfasiensis, the right anterior tooth inclines forwards, the left anterior tooth is grooved and higher than the median one, and the left median tooth is deepK grooved, with both sections equal in width. Retrotapes ninfasiensis is easily separated from R. na- vidadis (Philippi. 1887. pi. 14. fig. 4. Navidad Formation) because the Chilean Miocene species has a thinner and smaller shell with an acuminate anterior margin, straighter dorsal margin, more prominent beaks and a narrower lunule that is not inclined to the opposite valve. Cardinal teeth of R. navidadis are narrower and much shorter than in R. ninfasiensis. the right anterior cardinal tooth is vertical, but its anterior face inclines forward, while the left anterior cardinal tooth is lightly curved and also slopes forward. Retrotapes lenticularis (Sow- erby) (Riveros & Gonzalez, 1950:fig.30) (Figure 19-21), a species distributed along the Chilean literal from Co- quimbo to N'alparaiso, and also present in the Pleistocene outcrops in Central Chile (Herm, 1969:pl.l3, figs. 1-4). has subcircular shells with straight to lightly convex dor- sal margins, smaller lunules not inclined toward the op- posite valve, and much narrower and shallow er grooved cardinal teeth than R. ninjasiensis. The right posterior cardinal tooth inclines backwards and is separated from the nymph by a groove, the right anterior tooth varies from being vertical to slightK inclined forward, and the left anterior and median teeth are curved. The exterior surface is covered by fine commarginal ridges, widely spaced, deep commarginal grooves, and very fine radial threads. The Recent R. exalbida (Castellanos, 1970:250, pi. 22, figs. 4-5) (Figures 22-24, 41) is characterized by subrec- tangular and more inflated shells, with a shallower lunule, lower and shorter cardinal teeth with a deeper, grooved, left median tooth than in fi. ninfasiensis (Figure 4). Retrotapes striatolamellata (Ihering, 1897) Figures 33, 37-39. 42-47 Marcia itriatolamcUata Ihering, 1897:253, pi. 7, fig. 44; Ihering, 1907:305, Mania navidadis Philippi, Ortmann, 1902:141, pl.27, fig.l2. Marcia ortmanni Ihering, 1907:;504 Figures 29-31. Retrotapes scutata (Ihering). .\\\ figures 2X, Holot\pe M.^CN 429. Canadon El Lobo, San Julian Formation, late Eocene. 29. Left hinge. 30. exterior and 31. dorsal views of left valve. Figure 32. Retrotapes fuegoensis, new species. 1 x. Exterior view of left valve of holotype. PL' 356-12. Cerro Castillo. Carmen Silva Formation, late Oligocene-early Miocene. Page 86 THE NAUTILUS, Vol 110, No. 3 Figure 33. Retrolapes striutolanwllata (Ihering) Interior view of left valve, CPBA 9359, Monte Entrada, Monte Leon Formation, late Oligocene ( 1 X ) Figures 31— 36. Retrotapcs fiu'goensis. new species. Carmen Silva Formation, late Oligocene-early Miocene. 34. Kniargement of right hinge, DNSG 16.500. f:stancia La Federica (2X), 35. Interior view of left valve of holotype PL' 356-12, Cerro Castillo (IX). 36. Dorsal view of an articulated paratype DNSG 16.501, Estancia La Federica (IX). Figures 37-39. Rctrulapes Claudia Julia del Rio, 1997 Page 87 Diagnosis: Shell thick, o\ate to subrectangular in outline, longer, more convex than R. ninjasiensis. Lunule strong- ly concave, not inclined toward opposite valve. Left an- terior tooth arched (in most specimens), equal in height to median cardinal. Pallial sinus tongue-shaped, longer than in R. ninjasiensis. Description: Shell thick, large, ovate to subrectangular, moderatelv convex, longer than high. L'mbones small, at anterior quarter of length. Dorsal and anterior margins convex, ventral margin slightly convex, posterior margin slightly convex to subtruncated. Lunule broad, deeply concave, nearly vertical, not inclined toward opposite valve, marked b\ deep groove and distinctive ridge; lon- gitudinal, median, lunular groove impressed. Escutcheon on left valve better defined than on right, consisting of wide, flat surface facing opposite valve, sculptured with very faint commarginal ridges. Escutcheon on right valve narrower than on left, heavily ornamented like remain- ing valve surface. Hinge with three narrow cardinal teeth; right valve with cardinal sloping backwards, lamellar anterior tooth, median cardinal narrow K and shallowly grooved, as high as or higher than anterior tooth. Pos- terior tooth broadly grooved, separated from nymph by ridge; anterior tooth of left hinge lamellar, arched, ver- tically or slightly inclined backwards, median tooth deep- ly grooved, broad, as high as, or higher than anterior tooth, narrow, straight or arched posterior cardinal tooth. Muscle adductor scars equal in size, shallowK- impressed. Pallial sinus deep, tongue-shaped. Commarginal sculp- ture of thin lamellar ridges, much more closeK spaced near ventral margin. Interspaces smooth, except for growth lines. Material examined: Holotvpe, MACX 437, right valve, Yegua Quemada, height 67.5 mm, length 79.3 mm; Para- tvpes, two left valves, Yegua Quemada, MACN 2639, height 65.0 mm, length (38.0 mm. and MACN 2640, height 65.0 mm, length 68.0 mm; two left and two right valves from Monte Entrada, CPBA 9.359, PU429, Las Cuevas, CPBA 9.39L and from Cerro Puntudo (Lago Cardiel) CIRGEO-PI 2.513. (Ihering, Ortmann, Medina and Camacho collections) Stratigraphic and geographic distribution: Monte Leon Formation (late Oligocene), from the mouth of the Santa Cruz River to Yegua Quemada, and El Chacav For- mation (late Eocene), Cerro Puntudo (Lago Cardiel). Remarks: Retrotapes striatolamellata comes from out- crops of the Monte Leon Formation at Monte Entrada, Las Cuevas and Yegua Quemada (Figure 2). This species has also been recorded from exposures of the El Chacay Formation at Cerro Puntudo (Lago Cardiel) (Santa Cruz Province). The type material of R. striatolamellata had been collected b\ Carlos Ameghino at Yegua Quemada, but this fossiliferous localitv has never been recognized by subsequent authors. Ortmann (1902) placed fi. stria- tolamellata in synonymv with R. navidadis (Philippi, 1887). Ihering (1907) considered Ortmann's specimens to represent a new taxon that he named Marcia ortmanni Ihering, 1907. However, a re-analysis of Ortmann's ma- terial (1902, plate 27, figure 12) (Figure 47), reveals them to be voung specimens of R. striatolamellata. Material coming from Cerro Puntudo (Lago Cardiel) (Figures 37-39) is limited to a poorly preserved right valve with an eroded hinge. It is placed in R. striato- lamellata because of the arrangement of teeth, shape and size of pallial sinus, characteristics of lunule, es- cutcheon and exterior commarginal ornamentation. The onl) difference between this specimen and those coming from the Monte Leon Formation is in the outline of the shell. While the Cerro Puntudo specimen has a narrowly convex posterior margin w ith an ovate-subtriangular out- line, most representatives of R. striatolamellata from eastern Patagonia have shells with outlines ranging from subrectangular to ovate, with a broadly convex or trun- cated posterior margin (Figures 33, 42-46). Retrotapes navidadis (Philippi, 1887) is distinguished from R. striatolamellata in having a thinner and smaller shell with a straight dorsal margin, narrower and less concave lunule, shorter and narrower teeth and more widely spaced and homogeneousK' distributed commar- ginal ridges than in R. striatolamellata. Retrotapes len- ticularis has a subcircular shell w ith a narrower and less concave lunule, lower and more shallowly grooved teeth than in R, striatulaniellata. a vertical right anterior tooth, right and left posterior teeth that slope backwards, and a pallial sinus with an acute apex and concave ventral margin. Retrotapes striatolamellata may be easily sep- arated from R. antarctica (Sharman 6i Newton, 1894), which has a trigonally suboval shell, slightly concave lunule, narrower teeth, a right anterior tooth that is in- clined forward, and a shorter and triangular pallial sinus than R. striatolamellata. Retrotapes scutata (Ihering, 1907) Figures 29-31 Marcia scutata Ihering, 1907:303, pi. 11, fig.76. Diagnosis: Shell, elongate-oval in outline, anterior mar- gin convex. Lunule narrow, slightK' concave. Description: Shell ot medium size, elongate-oval in out- line, weakly convex, longer than high. Dorsal margin striatolamellata (Ihering), CIRGEO-PI 2513 Cerro Puntudo, El Chacay Formation, late Eocene, All figures IX. 37. Exterior, 38. interior and 39. dorsal views of right valve. Figure 40. Retrotapes ninjasiensis, new species, CPBA 15.087, Punta Norte, Puerto Madryn Formation, middle Miocene. Dorsal view of articulated specimen (1 x ). Figure i^X. Retrotapes exalbida (Chemnitz), MACN 21 172, San Matias Gulf. .Argentina, Recent, Dorsal view of an articulated specimen (1 x ). Page 88 THE NAUTILUS, Vol. 110, No. 3 ^^Sa^ % 44 47 45 46 Figures 42-47. Retrotapes strialolamellata (Ihering), Monte Leon l'\irmation, late Oligocene. 42. Interior view ot riglil valve of paratype, MACN 2639, Yegua Quemada (1 x ). 43. E.xterior, 44. interior, and 46. dorsal views of the holotype, MACN 4.37, Yegua Quemada (1 x ). 45. Interior view of left valve of parat\ pe, MACN 2640, Yegua Quemada (1 x ). 47. P-429, Monte Entrada. interior view of left valve of juvenile specimen illustrated by Orlniann (1902:27, fig.l2) (2x), Claudia Julia del Rio, 1997 Page 89 steeply sloping donwards, rounding to posterior margin; anterior margin narrowly convex. Lunule narrow, shal- lowly conca\e, not inclined toward opposite valve. Es- cutcheon on left \aKe strongly sloping toward right valve. Left hinge narrow, with lamellar anterior cardinal tooth sloping backwards, median tooth deeply grooved, pos- terior tooth lamellar, horizontal, slightly curved. Exterior v\ith lamellar commarginal ridges. Interior unknown. Material examined: Holotype, NIACIN 429, left valve, Canadon El Lobo, height 32,0 mm, length 3i>,0 mm. (Ihering Collection). Straligraphie and geographic distribution: San Julian Formation (late Eocene) from Canadon El Lobo (Santa Cruz Province). Remarks: Rctrotapcs scutata is known only- from the holot\ pe, a left valve whose interior is unknown and the hinge partially covered with marl. The elongate-oval outline, narrow, slightly impressed lunule and thin car- dinal teeth, separate this species from fi. striatolamellata and R. ninfasieuf^is. Retrofapes scutata and R. newtoni (Wilckens, 1911: plate 1, figure 16; Zinsmeister, 1984: figure 9 I, H, K, Eocene-early Oligocene? La Meseta Formation, Antarctica), share similar features including outline, ornamentation, characters of the lunule, and ar- rangement of cardinal teeth. These similarities suggest that these species are very closely related. A more thor- ough comparision between R. scutata and R. newtoni is not possible until more material of R. scutata is collected. Retrotapes fuegoensis new species Figure 32. 34-36 Eurhomalca '' cl juenzalidai . Malumian et al., 1978:278. pi. 3, tig. 4, not Venus fuenzalidae Pliilippi. 1887 Diagnosis: Shell elongate-oval in outline, anterior margin subtruncated, posterior margin convex. Cardinal teeth lower, narrower than in both R. striatolamellata and R. ninfasiensis, right anterior tooth straight, vertical, right median tooth shallowly grooved. Lunule very narrow, shallowly concave. Pallial sinus tongue-shaped, much deeper than in R. striatolamellata. Description: Shell large to medium sized, weakly convex, elongate-oval, longer than high. Umbones at 1 /9 of dorsal length. Dorsal margin slightly convex, rounding to con- vex posterior margin; anterior margin subtruncated. Lu- nule very narrow, shallowly depressed to nearly flat, not inclined toward opposite valve, broader in left valve than in right valve, with slighth marked median radial sulcus, bounded b\ moderately deep lunular groove. Escutcheon narrow, slightlv broader in left valve than in right valve, with strongly sculptured commarginal ridges on both valves. Hinge with anterior tooth vertical, median and posterior teeth inclined backward. Left hinge with strong, high anterior tooth, median cardinal rectangular, bifid, with both parts equal in size, as high as anterior tooth. posterior cardinal lamellar, curved. Right valve with thin anterior tooth, narrow grooved median tooth, thin, deep- ly grooved posterior tooth. Exterior with strong, rounded, commarginal ridges, more crowded toward ventral mar- gin. Material examined: Holotype, PU 355-12, left valve, Cerro Castillo height 62.0 mm, length 73.0 mm (Zins- meister Collection); Paratvpes, articulated specimen, DNSG 16.501, height 48.3 mm, length 58.8 mm, right valve DNSG 16.502, height 32.8 mm, length 27.0 mm; six articulated specimens, one left valve, two right (Mal- umian Collection), and one fragment with left hinge from Estancia La Federica and Cerro Castillo PU 357- 16, PU 355-13, DNSG 16.500, 16.503-16.509 (Zins- meister and Malumian Collections). Stratigraphic and geographic range: Carmen Silva For- mation (late Oligocene-early Miocene), Cerro Castillo and Estancia La Federica, Isia Grande de Tierra del Fuego. Remarks: Retrotapes fuegoensis comes from the up- permost conglomerated beds of the Carmen Silva For- mation's exposures at the quarrel of Estancia La Federica and at Cerro Castillo, where it is associated with a highly diverse and well preserved molluscan fauna previously studied by Malumian, Camacho and Gorrofio (1978). The outline of R. fuegoensis is the most distinctive character that readily distinguishes this species from its congeners. Based on examined material, the Chilean Neogene spe- cies Venus colchaguensis Philippi, 1887 ( = V .fuenzalidai Philippi, 1887) ma\' be also included in the genus Re- trotapes. It differs from R. fuegoensis in having an acu- minate anterior margin, a poorly defined escutcheon, and commarginal sculpture that consists of wideK and uni- forniK spaced lamellae. Retrotapes ninfasiensis may be distinguished from R. fuegoensis by its ovate to subrec- tangular, posteriorly truncated shell, a more concave and broader lunule facing the opposite valve, much larger cardinal teeth, median and anterior teeth that strongly slope backward, and a triangular and shorter pallial sinus than in R. fuegoensis. Retrotapes striatolamellata differs from R. fuegoensis in having a shell that is ovate, more convex, and more acuminate anteriorly, with a lunule that is more concave and broader, and teeth that are larger than in R.fuegoensis. Retrotapes scutata has a more elongate and ovate shell, with a more concave lunule than R. fuegoensis. Retrotapes antarctica is char- acterized by a trigonally suboval shell with umbones placed more posteriorly, an anterior margin that is not truncated, and a smaller triangular pallial sinus than that of R. fuegoensis. as well as a right anterior tooth that inclines slightly forward. Retrotapes newtoni has an ovate, smaller shell that is not subtruncated anteriorly. It also has a more concave lunule, thinner teeth and a shorter pallial sinus than R. fuegoensis. The Recent species R. exalbida and R. lenticularis have subrectangular and subcircular outlines respectively, deeper, concave lunules and shorter pallial sinuses than R. fuegoensis. Page 90 THE NAUTILUS. Vol. 110. No. 3 DISCUSSION Fleming (1963) proposed that Neoaustral faunal ele- ments originated in the low latitudes of the Pacific mar- gins during the late Tertiary. However. Zinsmeister (1982.1984) pointed out that at least some components of this fauna would have originated during the early Tertiar\ in high latitudes of circumpolar regions, and subsequently radiated northward. Antarctic Eocene re- cords of Aulacomya Morch, 1853, Gaimardia Gould, 1852, Gomphina Morch, 1853 and Eurhomalea Coss- mann.1920. led Zinsmeister to consider these to be Neoaustral taxa that originated in the higher austral lat- itudes much earlier than Fleming suspected. The transfer to Refrotapes of Antarctic species pre- viously assigned to Eurhomalea (Zinsmeister. 1984; Stil- well & Zinsmeister. 1992) limits Eurhomalea to strata younger than the Pleistocene. Eurhomalea most likely originated in Chile during the Pleistocene, later reached the Peruvian littoral fauna, and is now restricted to the eastern Pacific coast. Ret rot apes, however, is a Neoaus- tral genus that appeared in Patagonian and Antarctic regions during the early Tertiary. According to present data. Retrotapes first appeared in the southwestern Atlantic Ocean by the Eocene. It occurs in the late Eocene San Julian Formation and the El Chacay Formation of Patagonia, as well as in the Antarctic La Meseta Formation of late Eocene-Oligo- cene? age. By the early Miocene, Retrotapes expanded northward into the middle Chilean region, reached northern Patagonia b\ the middle Miocene, and attained its northermost distribution (R. lenticularis and R. ex- albida) during the Holocene (Figure 1). Fleming (1963) and Zinsmeister (1982. 1984) agreed that cool seawater temperatures were related to the presence of Neoaustral taxa. Zinsmeister (1982) stated that the development of Neoaustral faunal elements in the early Tertiary of Ant- arctica reflects the cool water temperatures that existed in that region during the late Eocene. He also considered the absence of Neoaustral genera in earlier Tertiary de- posits of South America to indicate the lack of suitable climatic conditions for the development of this fauna. The present study records a new. Tertiary and Ho- locene. Neoaustral genus in southern South America, and suggests that it has adapted to a wide range of water temperatures that fluctuated from cool-temperate to warm. Estimates of Antarctic late Eocene paleotemper- atures range wideK among authors. Kennett (1977) cal- culated Eocene sea-surface temperatures around Ant- arctica to have been 9-12°C higher than today, while Feldmann and Zinsmeister (1984) indicated that cool temperate conditions occurred in the area. More re- cently, Stilwell and Zinsmeister (1992) indicated that, although cool to warm conditions may be inferred from marine invertebrate taunas, the molluscan fossils indicate warm-temperate conditions during the deposition of the La Meseta P'ormation. Despite discrepant inlercnccs ol paleotemperatures. mollusks reveal that Antarctic conditions during the late Eocene were cooler than those in southern Patagonia during both the Eocene and Oligocene, as well as those recorded during the middle Miocene of northern Pata- gonia. Del Rio (1990, 1994 a,b), inferred warm temper- atures for northern Patagonia during the middle Miocene based on the presence of tropical and subtropical genera as Amusium, Flahellipecten, Chionopsis, Antinioche, Hexacortnila. Miltha. Egeta, Area. Dosinia s.str. and Lucinisca. According to del Rio (1990. 1994 a.b), middle Miocene temperatures in northern Patagonia would have been similar to those of the tropical Panamic Molluscan Province and the warmest regions of the Gulf and Ca- ribbean Molluscan Provinces. Along the Patagonian li- toral, temperatures decreased steadily, with cooler con- ditions similar to those occurring today having been es- tablished by the end of middle Miocene times. Almost 70% of the middle Miocene genera became extinct, in- cluding all of the above mentioned tropical and sub- tropical genera. Retrotapes. howe\er. continued to live in this region adapting to the new climatic conditions. Its present distribution is restricted to the warm tem- perate Argentinian Province and to the cool-temperate Magellanic Province. Recent advances in our knowledge of Antarctic and Patagonian molluskan assemblages (Zinsmeister & Ca- macho, 1980; Zinsmeister. 1976.1981.1984; Camacho & Zinmeister, 1986; Griffin. 1991; Stilwell & Zinmeister. 1992) show that the ocurrence of Retrotapes in Patagonia and Antarctica, along with several other genera and sub- genera, including Periploma (Ae/ga ) Slodkewitsch. 1935. Pteromijrtea Finla\. 1927. Lahillia Cossmann. 1899. Crassatella Lamarck. 1799, Au/aconij/a Morch. 1853, the gastropods Struthiolarella Steinmann and Wilckens. 1908 and Eoscaphella Stilwell and Zinsmeister. 1992. as well as nuculoids. arcoids (del Rio & Camacho. 1997) and veneroids that are presently being studied, reinforces the similarities in paleoclimates of southern South America and the Antarctic continent during the Paleogene. even after the final break-up of the Weddellian Province. ACKNOWLEDGMENTS 1 wish to express my gratitude to G. Parma (Facultad de Ciencias Exactas y Naturales of the liniversity of Buenos Aires) for her assistance in the accurate stratigraphic determination of fossil specimens from the San Jorge Gulf area and to M. G. Harasewych (National Museum of Natural Histor\. Washington. DC) and A. Oleinik (Purdue Lhiiversity. Indiana) who improved the English text. I am also indebted to W. Zinsmeister (Purdue Lhii- versity). F. Medina (Centre de Investigaciones en Re- cursos Geologicos. Buenos .Aires. Argentina) and N. Mal- umian (Ser\icio (ieologico Nacional. Buenos .Aires. .Ar- gentina) for allowing me to study their collections. Thanks also to the curators of the Ortmann and Ihering Colle- citons housed in Purdue L'niversity and in the Museo Argentino Argentino de (Ciencias Naturales "Bernardino Rivadavia respectiveK'. for permitting me access to the material described in this paper. This research has been Claudia Julia del Rio, 1997 Page 91 partialK' carried out through a Research Fellowship of the CONICET (Consejo Nacional de Investigaciones Cientificas y Tecnologicas) carried out in the Department of Earth and Atmospheric Sciences of Purdue University. C. Gambedotti took the photographs and Amalia Gon- zalez (Centro de Investigaciones en Recursos Geologicos, CONICET) drafted the figures. LITERATURE CITED Abbott, R. T. and S. P. Dance. 1986, Compendium of Sea- shells. American Malacologists, Inc. Florida, 411 p. Adams, H. and \. .•\dams. 185.3-1858, The Genera of Recent Mollusks arranged according to their organization. vol, 2 (1854-18.58), John Van Voorst, London, 661 p, Adams, A. and L, Reeve, 1850, Mollusca. In: A. Adams, (ed.) The Zoology of the vosage of H, M.S. Samarang, under the command of Captain Sir Edward Belcher during the years 1843-1846, London, Reeve and Benham, Each sec- tion paginated separately; Mollusca, X, 87 p. 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Journal of Paleontology 54:1- 14. THE NAUTILUS 110(3):94-96, 1997 Page 94 Iphitus robertsi (Epitoniidae: Nystiellinae), A New Species of Deep-Sea Gastropod from the Gulf of Mexico Bruno Sabelli Dipartimento di Biologia Evoluzionistica e Sperimentale Universita d Bologna Via S. Giacomo 9 1-40126 Bologna ITALY Marco Taviani Istituto di Geologia Marina, C.N.R. Via Gobetti 101 1-40129 Bologna ITALY e-mail; taviani boigm2.igm.bo.cnr.it ABSTRACT Iphitus robertsi. new species, is described on the basis of nine specimens collected in the northern Gulf of Mexico. This new species, which differs from its six congeners in having a bicar- inate shell with weakly canceliate sculpture, represents the first record of this genus in the Gulf of Mexico. The co-occurance of this species with deep-sea scleractinian corals suggests that Iphitus robertsi may, like its New Zealand congener, may be parasitic on corals. Key words: Gastropoda, Epitoniidae, new species. Gulf of Mexico, deep-sea. INTRODUCTION A recent survey of deep-sea cold seeps of the continental slope of the Gulf of Mexico using the research submers- ible Johnson-Sea-Link I has led to the discovery of a new species of Epitoniidae from off Louisiana and Texas. The specimens were collected in a vent-free area at a site known as Green Canyon (Roberts et al., 1990), Lease Block 189, during Johnson-Sea-Link I dive 3306. The sea bottom at a depth of 184 m was composed of mixed rocky and biogenic (silty-sandy shell debris) substrate. The description of this new species is based on nine empty shells in different stages of growth that were re- covered while sorting three cubic decimeters of biogenic sand under a dissecting microscope. The species diversity of the molluscan fraction of this sample was very high (n=197). SYSTEMATICS Family Epitoniidae Berry, 1910 Subfamily Nystiellinae Clench and Turner, 1952 (Jenus Iphitus Jeffreys, 1883 Iphitus robertsi nev\ species Figures 1-4 Description: Shell small, relatively tragile, trochilorm, hoiostomatous, umbilicate, with carinate, angular whorls rapidiv increasing in diameter. Suture subcanalicuiate. Protoconch I of 1 '/i smoothly rounded w horls. Protoconch II (larval shell) of 3V2 whorls, strongly sculptured by axial ribs and weaker spiral cords. Protoconch brownish, axis inclined by about 10° to teleoconch axis. Teleoconch of 3'/2 whorls, with strong, spiral sculpture of widel\- spaced carinae and strong spiral cords. Color w hitish, w ith few, irregular, pale brownish blotches. First half-whorl some- what rounded, subsequent whorls with two carinae, one in upper third of w horl, the second suprasutural. Strong spiral cords, one between suture and shoulder, one or two between carinae, appear beginning with second whorl. Finer spiral threads visible on body whorl of larger specimens. Shell base with 5-6 spiral threads, innermost bordering deep, wide, infundibuliform umbilicus. Sur- face of umbilicus sculptured with thin, spiral threads. Axial sculpture of densel\- spaced threads, producing rectangular, canceliate pattern at intersections with spi- ral cords, threads, and carinae. .\perture attached to pre- ceding whorl along one face of thin, roughly pentagonal peristome. Only one of nine specimens retained a com- plete protoconch, the remaining specimens lacked em- bryonic portion of the protoconch (protoconch I). Type locality: Green C^anvon, Block 189, Gulf of Mexico (27°46.48'N, 93°17.74'\V)'in 184 m. Type Material: Holot\ pe. National Museum of Natural History, Smithsonian Institution, Washington, DC, USNM 880185, Paratype USNM 888186; 3 paratypes, Museo di Zoologia, Bologna lWii\ersit\-, MZB 11621 a,b, MZB 11622; 1 paratype Museum National Histoire Naturelle, Paris; 1 paratype, Houston Museum of Natural Sciences, HMNH 42544, all from the type localit\ . Additional Material Kxamined: 1 shell (Figure 2, acci- dentalK broken), from the t\pe localit>. Etymology: This species is named after Dr. Harr> Rob- erts (Louisiana State University), the Chiel Scientist of the research crui.se and Scientist-in-Charge of scientific operations during dive JSL-I-3306. B. Sabelli and M. Taviani, 1997 Page 95 Figures 1-4. Iphitus robertsi, new species. Green Canyon, Block Holotype, USNM 888185, Scale bar = 3 mm 2. Adult shell, scale bar of holot\'pe. Scale bar = 3 mm. 189, Gulf f Mexico (27°46,48'N, 93°17.74'W) in 184 m. 1. 3 mm. 3. .Apical and 4. Lateral views of the protoconch ['age 96 THE NAUTILUS, Vol. 110, No. 3 Remarks: Bertolaso and Palazzi (1994) synonymized Iphitus Jeffreys, 1883 with Stylotrochus G. Seguenza, 1876, which is unavaliable because it is preoccupied by Stylotrochus Haeclcel. 1862. To date, oni\ six (four extant and two extinct) species of the genus Iphitus have been described. The Recent Iphitus tuberatus Jeffreys, 1883, /. cancellatus Dautzenberg and Fischer, 1896, and /. tuarshalli (S\ kes, 1925) inhabit the eastern North Atlan- tic, while /. neozelanicus (Dell, 1956) occurs off New Zealand. The fossil species Iphitus asperatus (G. Seguenza, 1886) and /. papillosocinctus (G. Seguenza, 1886) are both from upper Pliocene-lower Pleistocene bathyal deposits of Sicily (Palazzi & \illari, 1996). All of these species have been well illustrated (Beu, 1978; Taviani & Sabelli, 1982; Bouchet & Waren, 1986, Bertolaso & Palazzi, 1994, Palazzi & V'illari, 1996), and show little similarity to the new species described here. Iphitus rohertsi superficially resembles 7. marshalli. \\ hich also has strong carinae and spiral cords, but these differ in number and arrangement in the two species. Iphitus marshalli also differs in lack- ing cancellate sculpture. Iphitus cancellatus. the only species to be reported from the western Atlantic, differs from /. rohertsi in having a much coarser reticulate sculp- ture and a rounded aperture lacking carinae. The fossil legacy of Iphitus is quite scant. The oldest record (as Iphitus sp.) is from an upper Miocene coral-assemblage in southeastern Spain (Barrier et ai, 1991). Iphitus as- peratus (G. Seguenza, 1886) and /. papillosocinctus (G. Seguenza, 1886) are known from Plio-Pleistocene de- posits in the Messina region of SiciK' ( Bertolaso & Palazzi, 1994. Palazzi & Villari, 1996), while Iphitus tuberatus occurs in glacial-Pleistocene deposits of southern Italy (Rindone & N'azzana, 1989) and submerged last glacial (Pontinian) tanathocoenoses of the Sicilan Channel (Tav- iani & Sabelli, 1982). Flabitat: .\vailable ecological data indicates that species of Iphitus are parasitic on deep-sea scleractinian corals. Beu (1978) discovered /. neozelanicus living embedded in the calices of Goniocorella dunwsa (Alcock, 1902). While there are no direct observations on any of the Atlantic species, a relationship with deep-sea scleracti- nian corals has been suggested by Taviani and Sabelli (1982). Bouchet and Waren (1986) noted that I. tuber- atus occurs with Lophelia and that an association is prob- able. Bertolaso and Palazzi (1994) recorded one shell of /. asperatus still associated with a piece of Lophelia from the glacial Pleistocene of Calabria (southern Italy). A similar habitat is inferred for /. robertsi. As supported by visual observations (H. Roberts, personal communi- cation, September 1992) and sample content, dead and living scleractinian corals (e.g. Madracis sp., Caryophyl- lia sp., Coenosmilia arbuscula Pourtales, 1874, and Bal- anophyllia sp.: H. Zibrowius, in litt. 29 October 1993) were abundant at the sampling site. Among the possible hosts, the branching scleractinian Madracis. the com- monest coral, is the most likeK candidate, although no traces attributable to a secondary ectoparasite (Beu, 1978) were visible in our material. ACKNOWLEDGMENTS We are very grateful to Dr. Harr\ Roberts for inviting one of us (M.T.) on the research cruise, and for obtaining the samples on site. The Captain, crew and scientists aboard the R/V Edwin Link are gratefully acknowl- edged for their help and cooperation. We thank P. Bouchet , A. Waren and H. Zibrowius for criticalK read- ing the manuscript. Scanning Electron Micrographs are by A. Bonfitto. Thanks are due to H. Zibrowius for the scleractinian determinations. This is Istituto di Geologia Marina scientific contribution number 952. LITERATURE CITED Barrier, P., Zibrowius, H , Lozouet, P.. Montenat, C, Ott d Es- tevou. P., Serrano, F and Soudet, H.J. 1991. Une faune de fond diir du bathyal superieur dans le Miocene terminal des Cordillers Etiques (Carboneras, SE Espagne). Mesogee 51:3-13. Bertolaso, L. and S. Palazzi. 1994. /p/iifus Jeffreys, 1883, un sinonimo di Stylotrochus G. Seguenza, 1876 (appunti di malacologia neogenica:!). Bollettino Malacologico 29(9- 12):286-290. Beu, A. G. 1978. Habitat and relationships of Iphitctia neo- zetanica (Dell) (Gastropoda; Epitoniidae). New Zealand Journal of Marine and Freshwater Research 12(4):391- 396. Bouchet, P. and A. Waren. 1986. Revision of the Northeast Atlantic bathyal and abyssal Aclididae, Eulimidae, Epi- toniidae (Mollusca. Gastropoda). Bollettino Malacologico, Supplemento 2:299-576, Palazzi, S. and .\. X'lllari. 1996. Malacofaune batiali plio- pleistoceniche del Messinese.2: Capo Milazzo. Naturalista siciliano, ser.4, 20 (3-4):237-279. Rindone, V. and A. N'azzana. 1989. Alcune specie di niolluschi delle argille batiali del piano siciliano (Pleistocene inf.) della cava di Archi (Reggio Calabria). Bollettino Mala- cologico 25 (5-8): 233-240. Roberts. H. H., P. Aharon, R. Carney, J. Larkin and R. Sassen. 1990. Sea floor responses to hydrothermal seeps, Louisiana continental slope. Geo-Marine Letters 10:232-243. Taviani, M. and B, Sabelli. 1982. Iphitus (Mollusca, Gastrop- oda) a deep-water genus new to the Mediterranean sea. Lavori della Societa Malacologiea Ilaliana, .\tti del \'°Con- vegno della Societa Malacologiea Italiaiia 191-131. Vazzana, A. 1995. Malacofauna batiale del Pleistocene in- feriore del Vallone Catrica (Reggio Calabria, Italia). Bol- lettino Malacologico 31 (5-8); 143-162. THE NAUTILUS 110(3):97-101, 1997 Page 9" Verification of the Specific Status of the Endangered Anthony's River Snail, Athearnia anthonyi, Using Allozyme Electrophoresis Robert T. Dillon. Jr. Department of Biolog\, College of Charleston Charleston, SC 29-424 USA Steven A. Ahlstedt' Tennessee \'alley Authority Clean Water Initiative Norris, TN 37828 USA ABSTRACT Although nominally the single surviving representative of a unique pleurocerid taxon, Athearnia anthonyi (Budd, in Red- field, 1854) is so rarely collected that even its specific status has been uncertain We used allozy me electrophoresis to compare a population of A. anthonyi to the similar pleurocerid snail, Leptoxis praerosa (Sa\. 1821), co-occurring with it in the Se- quatchie River of Tennessee, and to a second population of L. praerosa collected approximately 500 km distant. Observed levels of heterozygosity offered no evidence of inbreeding or unusually severe population bottlenecking in any of these pop- ulations. Strikingly different allele frequencies at five of the eleven enzyme loci examined, together v\ ith differences in shell morphology especially apparent in young individuals, con- firmed that A. anthonyi and L. praerosa are distinct species. Their similarity at the six loci remaining supports previous suggestions that Athearnia may be a subgenus of Leptoxis. Key words: Athearnia. Leptoxis. Pleuroceridae, freshwater gastropods, proteins, enz\ mes. Tennessee. INTRODUCTION The pleurocerid genus Atliearnia was proposed by Mor- rison (1971) to include the two species, Anculosa an- thontji (Redfield, 1854) and Anculosa crassa (Haldeman, 1842), previously but incorrectly included in the genus Eunjcaelon (Goodrich, 1931). Some authors (Davis, 1974; Burch, 1982) have subsequently considered Athearnia to be a subgenus of the widespread genus Leptoxis, but we follow Bogan and Parmalee (1983), Garner (1992), and the Federal Register in retaining its generic rank. The distinction between the two nominal Athearnia species, A. anthonyi and A. crassa, has been confused for some years. In any case, since A. crassa is generally considered to be e.xtinct (Bogan & Parmalee, 1983), A. anthomji ' Present Address: U.S. Geological Survey, 1820 Midpark Drive, Knoxville TN 37921 USA appears to be the sole existing representative of this dis- tinctive pleurocerid taxon. Historicallv , populations of Athearnia have been re- corded from the Tennessee River and its larger tribu- taries upstream from Muscle Shoals, Alabama, including the clinch, Powell, Nolichucky, Little Tennessee, French Broad, Sequatchie, and Elk Rivers (Goodrich, 1940). However, the impoundment of the Tennessee drainage that began in the 1930's, improvement of the stream bed for navigation, and general habitat degradation due to farming, mining, and industry eliminated Athearnia from the great majority of its former habitat. Bogan and Par- malee (1983) feared that Athearnia "may be extinct throughout its range except for a possible relic popula- tion. More recently , it has become clear that small but ap- parently viable populations of A. anthonyi still inhabit lower regions of the Sequatchie River in Marion County. Tennessee, and Limestone Creek in Limestone County, Alabama (Garner, 1992). These snails also survive in at least one channel of the main Tennessee River down- stream from the mouth of the Sequatchie (Jenkinson, 1994; Garner, 1994). In early 1994, A. anthonyi became the first pleurocerid gastropod formally listed as "en- dangered" by the United States government (Federal Register 59:17994-17998). In addition to protecting ex- tant populations, the A. anthonyi recovery plan (U.S. Fish & Wildlife Service, 1996) calls for the future re- establishment of at least a few populations into portions of the snail's historic range. .\lthough great interest has focused on the distribution and abundance of A. anthonyi, questions remain re- garding its ta.xonomic status. Adult A. anthonyi may be confused with large, senescent Leptoxis praerosa (Say, 1821). a much more common pleurocerid with which Athearnia often occurs. Juveniles of A. anthonyi have not often been described. Given that trematode infection may induce extreme and anomalous growth in gastropod hosts (Sturrock, 1966; Hodasi, 1972), it has seemed pos- sible that Athearnia may be parasitically castrated, gi- gantic, L. praerosa. Allozyme electrophoresis has become established as an Page 98 THE NAUTILUS. Vol. 110. No. 3 important tool for measuring genetic di\ersit\ both with- in and among populations of pleurocerid snails (Cham- bers. 1978; 1980; Stiven & Kreiser, 1994). Published work to date has. howe\er. concentrated e.\clusi\el\ on the diverse genus Goniobasis, primarily an inhabitant of smaller streams. Genetic variation seems to be unusually low within Goniobasis populations, but unusually high between them (Dillon & Davis, 1980). The levels of both intra- and interpopulation gene flow also seem to be unusually low (Dillon. 1988a). Significant gene frequen- cy differences have been reported between samples of Goniobasis taken at distances as short as 500 m (Dillon, 1988b). GeographicalK' distant Goniobasis proxima (Say, 1825) populations may share no alleles at as many as si.\ enzyme loci of seven studied, yet show no evidence of reproductive isolation (Dillon, 1984; 1986; 1988a). Dis- tinct species of Goniobasis generalK' share alleles at very few enzyme loci (Chambers, 1980; Dillon & Davis, 1980). So although no data have been published on divergences in other pleurocerid genera to this date, there is reason to e.xpect that if A. anthonyi is indeed different from L. praerosa, differences in alloz\ me frequencies will be ap- parent. In the present work, we use allozyme electrophoresis to compare a population of A. anthonyi from the Se- quatchie River to a co-occurring population of L. pra- erosa and to a population oi L. praerosa from the Duck River 500 river kilometers distant (appro.ximately 350 km downstream, then 150 km upstream). In addition to verifying the specific status oi A. antlionyi, our data also bear on two ancillary questions: the systematic placement of the genus Athearnia and the degree to which an im- portant population of A. anthonyi may be inbred or bottlenecked. MATERIALS & METHODS Athearnia anthonyi and Lcptoxis praerosa were col- lected from a site on the Sequatchie River in Marion County, Tennessee. Leptoxis praerosa were most com- mon in riffle areas, while A. anthonyi were more com- mon on larger submerged objects (rocks, snags, etc.) in the transition areas between rifiles and pools. A second sample of L. praerosa was collected from the Duck River at Lillard Mill, Marshall County, Tennessee. Voucher specimens of A. anthonyi have been deposited in the Ohio State University Museum of Biological Diversit\ (catalog number 19820). Individuals were transported alive to Charleston, S.C, where the shells were cracked and the digestive glands examined for parasites. Tissues were then frozen in a tris-phosphate bufter at — 70°C and examined electrophoreticalK within one week. Horizontal starch gel protein electrophoresis was per- formed on whole animal homogenates as has been pre- viously described (Dillon, 1985; 1992). We initially screened 15 A. antlionyi and 14 Secjuatchie River L. praerosa for variation in 10 enzyme systems: aspartate aminotransferase (AAT), esterases (EST a-napthyl ace- tate as substrate), glucose phosphate isomerase (GPI), alcohol dehydrogenase (ADH, hexanol as substrate), mannose phosphate isomerase (MPI), octopine dehydro- genase (ODH), phosphogluconate deh\ drogenase (6PGD), phosphoglucomutase (PGM), sorbitol dehydro- genase (SDH), and superoxide dismutase (SOD). Simple Mendelian inheritance of codominant alleles has been demonstrated at the 6PGD and PGM loci in Goniobasis floridensis (Reeve, 1860) by Chambers (1980) and at the GPI, ODH, and ESTl loci in G. proxima by Dillon (1986). It should be noted that a great many esterase allozymes are generally detectable in pleurocerids. but that to date only the strong, slowly-migrating products ot the "ESTl" locus are genetically interpretable. An effort was made to examine alloz>me phenot>pes in as wide a range of buffers as possible. The following gel buffers were emploved: Tris-Cit 6.0 (for PGM, GPI, ODH, and MPI), AP 6.0 (for 6PGD), TEB S.O (for EST and 6PGD), Poulik (for SDH, ODH, and GPI), and TEB 9.0 (for SOD, ADH, AAT, PGM, and SDH). Recipes for most of these buffers and stains were obtained from Shaw and Prasad (1970) or Harris and Hopkinson (1976), mod- ified for agar overlay in many cases. Also included in the initial survey for allozyme vari- ation were C. proxima standards (population SL'GR of Dillon, 1984). Leptoxis and Athearnia bands were la- beled by their mobilities in millimeters relati\e to this population in a standard buffer. As was the case in 1984, some "hidden variation was detected at the ODH locus - bands not resolved b\ the standard Tris-Cit 6.0 buffer were detected with a Poulik buffer. These isozyme classes were labeled "S". "F", and "VF" for their migration in Poulik gels. After the initial screening, we concentrated our in- vestigations on the five putative enzyme loci apparently varying in our comparison of A. anthonyi and L. pra- erosa. Ultimately we examined 37 A. anthonyi, 29 L. praerosa from the Sequatchie River, and 25 indixidual L. praerosa from the Duck Ri\er. Gene frequencies and observed heterozygosities were calculated using BIOSYS version 1.7 (Swoftord & Selander, 1981). RESULTS Figure 1 compares representati\ e shells from Sequatchie populations of A. anthonyi and L. praerosa. Athearnia anthonyi is distinguishable b\ the higher shoulder of its w horls, beginning as a pronounced keel (or "carina ) but becoming less distinct with age. The oldest A. anthonyi are quite smooth, and come to resemble large L. pra- erosa. \er\ little evidence of parasitism was detected in ei- ther Sequatchie pleurocerid population. OnK 2 oi 29 L. praerosa showed obvious trematode infections, while none of the 37 .4. anthonyi appeared to be infected. Thus the large sizes attained by individual A. anthonyi do not appear to be due to parasitic gigantism. R. T. Dillon, Jr. and S. A. Alilstedt, 1997 Page 99 .#^ a I 10 mm Figure 1. Growth series for Athi-anna nntlionyi (abose— live specimens) and Lepioxis praerosa (below) from tfie Sequatchie River, TN. No variation was detected between populations of A. anthomji and L. praerosa at AAT, ADH, 6PGD, SDH, SOD, or at a faster-migrating PGM locus. Divergence was striking, however, at the five loci listed in Table 1. The two svmpatric populations share no alleles at ESTT GPI, MPI,' or at the slower PGM locus, 'PGMS". And at the ODH locus, A. anthomji appears to be fixed for an allele present at a frequenc\ of onl\ 0.345 in Seciuatchie /,. praerosa. Although spatially separated by over 500 km, the two L. praerosa populations seem to have remained rather similar genetically. Table 1 shows that the differences in alloz) me frequenc\ at the GPI, MPI, and ODH loci are significant, but not fixed. The Sequatchie population ap- pears substantially more polymorphic. Rather low levels of genetic variation were detected within these three populations of pleurocerids, as has been commonly reported in the past, jutlging b\' the 95% criterion, onl\ four of the 15 loci show n in Table 1 could be considered polymorphic. The fit to Hardy- Weinberg expectation at these four loci was verv good, however. DISCUSSION It is clear that previous misgivings about the specific status of A. anthomji were unfounded. Although not as divergent as typical species of Goniobasis, the data pre- sented in Table 1 show clearly that A. anthomji and L. praerosa are reproductiveK isolated in the Sequatchie River. Their shell morphologies are distinct. The greater size attained by A. anthomji appears to be a natural Table 1. .Allele frequencies and observed heterozygosities (H) at variable loci in populations of Athearnia anthomji from the Sequatchie River. Leptoxis praerosa from the Sequatchie River, and L. praerosa from the Duck River, Sample sizes 37, 29, and 25, respectively. .\llele Sequatchi ieR. Duck R Locus A anthomji L . praerosa L, praerosa ESTl 105 0.0 1.00 1.00 102 1.00 0.0 0.0 H 0.0 0.0 0.0 GPI 102 0.392 0.0 00 97 0.0 0.5S6 1.0 95 0.608 0.0 0,0 90 0.0 0.414 0 0 H 0.622 0.552 0,0 MPI 100 0.0 0.034 0,0 95 0.0 0.966 10 90 1.0 0.0 0,0 H 0,0 0 069 0 0 ODH 115 OO 0.086 0.460 113\F DO 0.310 0,0 113F 1,0 0.345 0.520 113S 0,0 0.034 0.0 107 0,0 0,224 0.020 H 0,0 0.670 0.440 PGMS 103 0,0 1-0 1.0 90 1,0 0 0 0.0 H 0,0 0,0 0.0 Page 100 THE NAUTILUS, Vol. 110, No. 3 feature of development, not a consequence of parasitic gigantism. The level of intrapopulation variation in A. anihoniji is low compared to that of most other organisms, but comparable to that seen in L. praerosa and in other pleurocerids. We did discover one highly polymorphic locus in A. arithonyi, GPl, showing genotype frequencies not significantK different from expectation under Har- dy-VVeinberg equilibrium (Yates-corrected x' = 2.53). Thus we are unable to detect evidence of inbreeding or an unusually severe population bottleneck. Previous electrophoretic investigations of Goniobasis populations have generally detected higher levels of di- vergence than we report here between Sequatchie River and Duck River populations of L. praerosa. Multiple fixed differences are often observed between G. proxima populations isolated in small creeks at distances as great as 500 km. The less dramatic differences between con- specific Leptoxis populations reflected in Table 1 may be a consequence of their adaptation to larger rivers. Populations of Leptoxis ma\ have inhabited occasional rocky shoals down the length of the Tennessee River before its impoundment, connecting such tributary pop- ulations as those of the Sequatchie and Duck Rivers in stepping-stone fashion. Previous studies have shown greater divergence be- tween pleurocerid species than uncovered here. Dillon and Davis (1980) and Chambers (1980) reported that typical species of Goniobasis rarely share any similarity at any allozyme locus. However, L. praerosa and A. anthonyi were indistinguishable at six of the eleven loci initially screened, and seem to share one fairly common allele at a seventh locus (ODH). This constitutes some support for the nomenclature ot Burch (1982) and Dillon (1989) in which Athearnia is placed as a subgenus of Leptoxis. Regardless of whether they represent a unique genus or a unique subgenus, the populations of A. an- thonyi now restricted to just a tew rivers of central Ten- ne.ssee are a valuable resource for evolutionary study, and warrant protection at the highest levels. ACKNOWLEDGMENTS Charles Saylor assisted in collecting the snails, Vivian Johnson delivered them to Charleston, S.C., and Amy Wethington was a great help running the gels. Technical reports were provided by John Fridell. LITERATURE CITED Bogan, A., and P. Parmalee. 1983. Tennessee's rare wildlife, volume II: The mollusks. Tennessee Wildlife Hesnurces .Agency, Nashville, TN. pages 81-84. Burch, J. 1982. North Auierican fre.shwater snails; identifi- cation keys, generic synon\ my, supplenicntai notes, glos- sary, references, index. Walkerana -}:l-'36.5. Chambers, S. M. 197S. An elfitidpiiorelically delected siljimg speeiesof " Coniolmsis floi ulcnsi.s" (Mesogastropoda: Pieu- roceridae). Vlalacologia 171.57-162. Chambers, S. M. 1980. Genetic divergence between popu- lations of Goniobasis occupying different drainage sys- tems. Malacologia 20:11.3-120. Davis, G. 1974. Report on the rare and endangered status of a selected number of freshwater gastropoda from south- eastern U.S.A. U.S. Department oi Interior, Fish and W'ild- life Service, Washington, DC. 51 p. Dillon, R. T., Jr. 1984. Geographic distance, environmental difference, and divergence between isolated populations. Systematic Zoology 33:69-82. Dillon, R, T,, Jr 1985, Correspondence between the buffer systems suitable for electrophoretic resolution of bivalve and gastropod isozymes. Comparative Biochemistry and Physiologv 828:643-645. Dillon, R. T , Jr. 1986. Inheritance of isozyme phenotype at three loci in the freshwater snail Goniobasis proxima: Mother-offspring analysis and an artificial introduction. Biochemical Genetics 24:281-290. Dillon, R. T., Jr. 1988a. 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Survey of mollusks, Tennessee River mile 412.1, .Aquatic Resources Center, Franklin, TN, 2 p. Goodrich, C. 1931, The pleurocerid genus Eurycaelon. Oc- casional Papers of the Museum of Zoology, University of Michigan 223:1-9, Goodrich. C, 1940, The Pleuroceridae of the Ohio River system. Occasional Papers ol tlie Museum of Zoology, Uni- versity of Vlichigan 417:1-21, Harris, H., and D Hopkinson. 1976. Handbook of enzyme electrophoresis in human genetics. Elsevier, New York. Hodasi, J. K. M. 1972. The effects of Fasciola hepatica on Lymnaea Iruncatula Parasitology 65: 359-369. Jenkinson, J. 1994 PYeshwater mollusk survev at CSX railroad bridge, near Bridgeport, .Alabama, Tennessee River Mile 414.5. Tennessee \'alley Authority, Chattanooga, TN. 15 p. Morrison, J. 1971. Athearnia a new name for a genus of pleurocerid snails. The Nautilus 84:110-111. Shaw, C. R., and R. Prasad. 1970. Starch gel electrophoresis ot enzymes — a compilation ot recipes. Biochemical Ge- netics 4:297-320, Stiven, A , and B, Kreiser. 1994. Ecological and genetic dif- ferentiation among populations of the gastropod Gonio- basis proxima (Say) in streams separated by a reservoir in the Piedmont of North Carolina, Journal ol the Elisha Mitchell Scientific Society I10:.53-67, Sturrock, IV M, 1966, The intluence ol inlection with Schis- tosoma mansoni on the growth rate and reproduction of R. T. Dillon, Jr. and S. A. Ahlstedt. 1997 Page 101 Bioniphalana pjeiffch. Annals ot Tropical Medicine and United States Fish & Wildlite Service. 1996. Technicai/Agen- Parasitology 60: 187-197. cy Draft Anthony's Riversnail Recovery Plan. Atlanta GA. Swofford, D. L.. and R B Selander. 1981 BI08YS-1; a FOR- 31pp. TRAN program for the comprehensive analysis of elec- trophoretic data in population genetics and svstematics. Journal of Heredity 72:281-283. TliE NAUTILUS 110(3);102-106, 1997 Page 102 Induced Metamorphosis of Freshwater Mussel Glochidia on Nonhost Fish Sheila C. Kirk James B. Layzer National Biological Service Tennessee Cooperati\e Fishery Research Unit Tennessee Technological University Cookeville, TN 38505 USA ABSTRACT Intraperitoneal implants of Cortisol, an immunosuppressant, suspended in liquid cocoa butter were administered to nonhost fish species. Fish were then infested with glochidia of fresh- water mussels to determine if transformation \\ ould occur after immune system manipulation. Glochidia ot Venustacoitcha sima (Lea, 1838) {Villosa iris complex) transformed on orangethroat darters, Etheostoma spectabile (Agassiz, 1854), after injection of Cortisol at concentrations ranging from 0.005 to 0.040 mg per g of fish weight. Banded sculpins, Cottus carolinae (Gill, 1861), transformed glochidia of Villoaa taeniata {Conrad, 1834) following Cortisol injections ranging from 0.005 to 0.020 mg- g. Creek chubs, Scinolilus atromaculatus ^Mitchill, 1818), failed to transform glochidia of either mussel species after Cortisol administration. No juvenile mussels were collected from non- injected or sham injected fish during any experiment. Cortisol- induced immunosuppression facilitates metamorphosis of glo- chidia on some nonhost fish species. Refinement of this tech- nique could provide an alternative means for propagating freshwater mussels, especial!) for those endangered species that utilize unknown hosts. Key Words. Cortisol, inmiunosuppression, transformation, glo- chidia, implants, Linionidae. INTRODUCTION The obligate parasitic stage in the life cvcie of freshwater mussels on specific host fish ma\ limit reproduction of many populations. Because adequate biological inven- tories were not conducted before impoundment of most river systems, many fish species were extirpated before identification of glochidia! hosts could occur (Schindler, 1989). Hosts are known for onl\ one-quarter of the Nortli .American species of mussels (Hoggarth, 1992; Walters, 1994). Although glochidia of a few mussel species have laeen transformed in a sterile culture medium (Isom & Hud.son, 1982), metamorpliosis of other species has been less successful. Consequently, artificial propagation of many freshwater mussel species that utilize unknown hosts has not been possible. Other than the use of known hosts and artificial media, few advancements have occurred in methods of artificial propagation of freshwater mussels. With the relatively recent threat of the zebra mussel, Dreissena pohjmorpha Pallas, 1771, concern for the fate of our native fauna has increased efforts to sustain diversity and protect existing populations, especially endangered or threatened species without identified hosts. Efforts include assessment of relocation into areas with little risk of zebra mussel in- vasion and the use of mussel refugia such as hatcheries (Shannon ct ai. 1993). Host specificity of freshwater mussels is believed to have an immunological basis (Reuling, 1919; .Arey, 1932; Meyers et at.. 198U). Isom and Hudson (1982; 1984) demonstrated that components in fish blood necessary for initiation of glochidial transformation are not species specific and can be found in the blood of all fish. More- over, horse and neonatal calf serum are also suitable media for glochidial transformation (Keller & Zam, 1990). The iiuplications of these studies suggest that the inuiume response of the host is the determining factor in mussel- host specificity. Immune responses of both host and nonhost fish to glochidia of freshwater mussels ha\e been documented (Reuling, 1919; Arey, 1932; Meyers et at., 1980; Bauer & Vogel, 1987). Responses of host fish consist of host tissue proliferation which encysts the glochidium within a few hours after attachment (d'Eliscu, 1972; Waller & Mitchell, 1989). In host and nonhost species, humoral and cell mediated responses of the immune sy stem occur. These immune responses include the presence of specific antibodies to the parasites (Reuling, 1919; Meyers et ai. 1980; Bauer & \'ogel, 1987), an increase in eosinophils and other leukocv tes around glochidial c>sts (Are) , 1932), hyperplastic sloughing of host epithelial tissue (Meyers et ai. 1980), and in some cases, cytolitic destruction of the glochidium (Reuling, 1919; Arey, 1932). C^ortisol, a major corticosteroid in teleosts (Idler & Truscott, 1972) is released during stress (Bennett & Wolke, 1987; Schreck, 1990) and produces immunosuppressive S. C; Kirk and J. B. Layzer, 1997 Page 103 effects. Cortisol can be administered to produce a dose- dependent elevation of plasma Cortisol and immuno- suppression (Pickering & Duston. 1983). Cortisol reduces K mphocNte concentrations circulating in the blood, re- duces KmphocNte participation in inflammation and re- duces other inflammator\ cells (Pearson et al.. 1978). We h\ pothesized that the effects of Cortisol might inhibit the abilit> of nonhost fish to respond to glochidia b> de- creasing the number of antibodies and reducing inflam- mation around glochidial cysts, thus allowing glochidia to attach, enc\st, and metamorphose into juveniles. In this paper, we e\aluate the effects of Cortisol, adminis- tered in a vehicle of liquefied cocoa butter, on glochidial metamorphosis. MATERIALS & METHODS Collection of fish and mussels: For our experiments, we selected tw o brad\ ticitc species of mussels based on their narrow host specificity and availability. Gravid Veniistaconcha sima (Lea. 1838)', were collected from the Collins River in Grundy County, Tennessee. Gra\ id Villosa taeniata (Conrad, 1834) were collected from the Roaring River in Overton County, Tennessee. Rockbass, Ambloplites rupestris (Rafinesque, 1817), is the only known host of V. taeniata (Gordon et al., 1994), and banded sculpin, Cottus carolinae (Gill, 1861), is the only known host of V. sima (unpublished data). To avoid premature expulsion of glochidia, mussels were kept at approximately 4°C in pans of shallow water until needed. Laboratory established nonhost fish species were used as experimental fish, with the exception of creek chub, Sf- motilus atromaculatus (Mitchill. 1818). The suitabilit\ of creek chub as hosts for ^'. sima has not been previousK- tested in the laborator\-; howe\er, examination of wild fish did not re\eal an\ infestations (unpublished data) A backpack electrofishing unit was used to collect fish from the East Blackburn Fork, Putnam County, Ten- nessee. Populations of banded sculpins, creek chubs, rockbass, and orangethroat darters, Etheostoma spect- ahile (Agassiz, 1854), are abundant in this stream. No mussels occur in the East Blackburn Fork, and an im- passable waterfall downstream prevents any possible up- stream movement of fish that ma\' have been exposed to glochidia. Fish were transported back to the laborator\', acclimated to laborator\ conditions, and held in 38 liter aquaria Implantation and infestation: Pure cocoa butter was melted at a temperature of 40°C. A predetermined amount of Cortisol was dissolved in ethanol. Ethanol was used at a rate of 10^7 the volume of cocoa butter. The 'This species is generally considered a member of the Villosa iris complex but was elevated by Gordon (1995). In this paper, we use Venustaconcha sima to clearly identify which of the two members of the Villosa iris complex occurring sympatar- icaii\ in the Collins River were used in our experiments dissolved Cortisol was suspended in the liquid cocoa but- ter and allowed to mix in a water bath. Average fish weight was used to calculate the amount of Cortisol per gram of fish weight needed to obtain the desired con- centration within the fish. Orangethroat darters received 50 m1 implants and creek chub and banded sculpins re- ceived 100 /ul implants. Infective glochidia were obtained from gravid females and exposed to salt to determine maturity (Zale & Neves, 1982). After fish were anesthetized with tricaine meth- anesulfonate (MS 222), they were placed on a wet paper towel to reduce the removal of mucus during the injec- tion process. The liquid cocoa butter implant was injected into the peritoneal cavit\ of the fish with a 1 ml tuber- culin hypodermic syringe. The liquid cocoa butter solid- ified within the cavity of the fish and acted as a solid implant during the experiment. PresumabK, Cortisol leaked slowK from the implant, but the kinetics of this transfer were not addressed, Pickering and Pottinger (1985) observed maximum mean plasma Cortisol levels of 9 and 15 ng/ml for brown trout, Salmo triitta (Lin- naeus, 1758), that received 10 and 20 mg Cortisol re- spectively. ImmediateK after Cortisol injection, glochidia were pipetted onto the left gills of each fish. Fish were then transferred to fresh water and revi\ed. Each ex- perimental group was kept in a separate aquarium. Experimental design Treatments varied among exper- iments, but generalK' included a known host control group to measure glochidial viabilit\. The no-injection (NI) group \erified that the test species did not normally serve as a host. Sham injected fish received injections of pure cocoa butter only, and were used to determine if the injection process or cocoa butter had an effect on glo- chidial transformation. Cortisol injected fish received concentrations that ranged from 0.005 to 0.040 mg/g. Preliminary experiments determined the feasibility of working with these concentrations, which we initially based on those used b\ Pickering and Duston (1983) and Pickering and Pottinger (1985). Aquaria were siphoned daily and the siphonate was examined under a dissecting microscope at lOX to 20X magnification. The number of glochidia and juveniles, water temperature and fish mortality were recorded. Criteria for recognizing juvenile mussels included the presence of tw o adductor muscle scars, closed valves, and movement w ithin 24 hours of collection. If these criteria w ere not met, organisms w ere considered untransformed or partially transformed glochidia. Experiments were terminated when no juveniles w ere collected from host fish for one week after the last juvenile was collected, or examination of the gills revealed no enc\sted glochidia. .\11 fish were used for onl\ a single infection experiment, RESULTS Metamorphosis of I enustaconcha sima on nonhost fish: .\ total of 58 juvenile Venustachona sima were trans- formed on Cortisol injected orangethroat darters. Trans- formation occurred only on orangethroat darters injected Page 104 THE NAUTILUS, Vol. 110, No. 3 Table I. NumlKTi ot ju\eiiiles collected troni Cortisol injected orangethroat darters infested with glochidia of Venustaconcha sirna in each experiment. Dates indicate v\ hen fish were injected and infested. Numbers in parentheses are the average number of juveniles transformed per fish (NI = No-injection) Table 2. Nuinljers ol ju\ eniles collected from Cortisol injected banded sculpins (October and May) and orangethroat darters (December) infested with \ illosa taeniata in each experiment. Dates indicate when fish were injected and infested Numbers in parentheses are the average number of juveniles transformed per fish (N! = No-injeclioii). Total Cortisol img/g) Date Cortisol June 6 Au- gust 5 March 31 May 17 img/g) October 27 December 28 May Nl 0 0 0 0 0 0 0 0 0 Total 0.000 Nl (1 0 0 0 0.00.5 17 (1,89) 0 5 (0.36) 3 (0.25) 25 0.00 0 0 0 0 0 010 ~ 0 4 (0.29) — 4 0,005 1 (0,13) — 0 1 0020 — — 7 (1,00) 5 (0.50) 12 0,010 0 0 1 (0,10) 1 0030 — — 0* — 0 0.020 — 0 4 (1,001 4 0.040 — — 17 il 70) 0* 17 0,030 — 0 — 0 * Fish mortalitv was 2:88' in March, May, and June (Table 1). Glochidia from V. sima transformed on fish injected with 0,005 to 0.040 mg, g Cortisol. Metamorphosis never occurred in sham injected and no-injection (NI) treatment groups during any experiment. Glochidia did not transform on either orangethroat darters or creek chubs injected in August. Juvenile transformation varied among experiments. For instance, the average number of juveniles trans- formed per orangethroat darter injected with 0.005 mg/ g Cortisol ranged from 0.00 to 1,89 among experiments (Table 1 ). There w as a positive correlation between Cor- tisol concentration and metamorphosis within experi- ments; however, this relationship was confounded by high mortality in some treatment groups. Mortality var- ied within and among experiments; most deaths occurred soon after injection, and were likely the result of physical injury. Metamorphosis of \ illosa taeniata on nonhost fish: A total of six juvenile X illosa taeniata were collected from banded sculpins injected with Cortisol (Table 2). Trans- formation of V. taeniata glochidia did not occur on orangethroat darters or creek chubs injected with Cortisol. In one experiment, nonhost species sloughed glochidia quickly after the infestation. After examination of their gills revealed no remaining glochidia, the fish were rein- fested 1 1 days after the initial infestation. Although creek chubs and banded sculpins retained glochidia longer after the second infestation, metamorphosis did not occur. Metamorphosis of glochidia on host fish: Transfor- mation of glochidia of Venustachoncha sinia on the host, banded sculpins, varied between experiments Banded sculpins transformed 35 juvenile mussels in March and six juvenile mussels transformed on banded sculpins in May. When used as a host control, all banded sculpins were held in one aquarium during each exi)eriment, and individual variation in juvenile transformation could not be asse.ssed. Because of the aggressive nature of rockbass, the host of Villosa taeniata, individuals were kept in separate aquaria. Transformation of glochidia of V. taeniata on rockbass varied greatly among individuals and among experiments (Table 3). Ju\enile transformation on in- dividual fish ranged from 0 to 13 juveniles in one ex- periment, to a range of 7 to 262 ju\eniles in another experiment. There was no clear relationship between the number of glochidia used to infest host fish or water temperature and the number of juveniles recovered from individual rockbass. DISCUSSION Cortisol affects leukocyte circulation, influences immune effector mechanisms in lymphocytes, modulates acti\i- ties of inflammatory mediators, and modifies protein, carbohydrate, and fat metabolism (Tizard, 1988). In fish, this results in immunosuppression and an increased sus- ceptibility to infectious diseases such as furunculosis and bacterial fin-rot (Pickering & Duston, 1983; Pickering & Pottinger, 1985). The present study has shown that cortisol-induced im- munosuppression can facilitate glochidial metamorpho- sis on nonhost fish species. Presumably, Cortisol sup- pressed the humoral and cell-mediated responses ob- served in glochidial infestations by Arey (1932) and Mey- ers ct al. (1980). .Although the strength of fish immune responses after Cortisol administration was not measured, transformation occurred on experimental nonhost species only when the immune system was compromised. These results are consistent with the hypothesis that host spec- ificity of freshwater mussels is immunologically con- trolled. Glochidia of Venustachoncha sima transformed on orangethroat darters in three of four experiments at the lowest concentration (0.005 mg, g) used. Transformation of glochidia of Vtllusa taeniata occurred on banded scul- pins during two experiments. For this species, fewer ju- veniles were collected from fish injected with the same concentration used for orangethroat darters. Creek chubs failed to transform glochidia from either mussel species S. G. Kirk and J. B. Layzer, 1997 Page 105 Table 3. Total It'iigth. luunber ot glochulia reco\erecl, niinilier ot ju\eniles recovered, metamorphosis period, and water temperature (±S.D.) for iinlix iiliial rockl>ass infested with glochidia ot \'illosa tacniata. Dates indicate when fish were infested. Xumlifr III Number ot Total Length Temperature Glochidia Juveniles Metamorphosis Date (mm) (±S.D.I Recovered Recovered Period (Days) Oct 27, 1993 165 23.75 ± 1.54 9S6 (J — 165 798 0 — 155 693 13 29-31 169 659 3 29-32 115 741 1 32 Dec. 2S, )993 51 22.04 ± 1.80 230 7 43-52 77 124 12 39-52 63 121 2 34-52 Mav 27, 1994 160 22.32 ± 1.33 453 249 21-31 118 421 14 22-28 98 441 7 23-26 177 1,221 262 21-39 when injected with Cortisol concentrations of 0.005 and 0.010 mg/g Cortisol, suggesting that responses to (or up- take of) Cortisol varies with species. Threshold levels of Cortisol in the blood may exist that allow glochidial trans- formation, and these levels ma\' not have been reached in creek chubs. Fish experience periods of natural elevation ot plasma Cortisol (Idler & Truscott, 1972). Spring and siunmer elevations of plasma Cortisol and other circidating cor- ticosteroids coincide with migration, smoltification of ju- venile anadromous salmonids, se.xual maturation and spawning of salmonids (Thorpe el al.. 1987; Pickering & Pottinger, 1983; Pickering & Christie, 1981; Anderson, 1990). Cortisol concentrations used in this stud\' should have elevated Cortisol le\els to the ph\siological range of fish undergoing stress or periods of natural Cortisol elevation (Pickering & Duston, 1983; Pickering & Pot- tinger, 1985; Thorpe et al., 1987), Periods during the life cycle of fish where Cortisol elevation has been observed may correspond with the presence of some species of host fish over mussel beds during the discharge of glo- chidia (Farzaad, 1991). Perhaps mussels evolved to take advantage of weakened immune responses of host species during spawning or other periods of natural immuno- suppression. Host and nonhost fishes exhibit humoral responses of similar strengths, but host and nonhost species reject glo- chidia at different rates (O Connell, 1991 ). Although Cor- tisol injected fish transformed relati\el\' few juveniles during these experiments, transformation per fish was comparable to hosts in some cases. For instance, 0,020 mg/g Cortisol injected orangethroat darters transformed an average of 0.50 juvenile \'. sirua per fish in Ma\ . Host fish (banded sculpins) onl\' transformed 1.00 juvenile per fish in the same experiment. Individual variations within and among experiments in glochidial transformation of \'. tacniata on host fish may also reflect a seasonal aspect in the susceptibility of the host (e.g., host spawning sea- son) or the ability of glochidia to be infective at times of the year other than the normal discharge period. We have observed similar variation among trials and indi- viduals in the numbers of juveniles transformed per host for other mussel species. Variation of an order of mag- nitude or more in the mean number of juveniles trans- formed per fish has been reported in other studies as well (e.g., Zaie & Neves, 1982). Although the number of glo- chidia attaching to fish is difficult to control and may contribute to some of the variation in the numbers of juveniles transformed, there was no relationship in the apparent numbers (glochidia + juveniles recovered) of Villosa tacniata glochidia attaching to individual rock bass and the numbers of juveniles produced, .Additional development of the cortisol-induced im- munosuppression of nonhost fish is needed before the technique can be applied on a large scale as a culture method for freshwater mussels. Nonetheless, refinement of this technique could have widespread management implications, especially as a means for propagation of endangered species. This method could also be used to reestablish populations of nonendangered mussel species into new and reclaimed habitat. Application of Cortisol immunosuppression techniques to host species could per- haps increase juvenile transformation for reintroduction studies. Also, application of this technique to non-host fish would provide an alternative means of propagating mussel species that have rare or endangered hosts. The need for further research on the ecological re- quirements, including identification of fish hosts, habitat, and ph\sicochemical information of mussel species can- not be over emphasized. These ecological requirements hold the answers to developing self-sustaining popula- tions of freshv\ater mussels. Because survival and fitness of juvenile mussels produced b\' means of artificial prop- agation has not been determined (O Connell, 1991), ar- tificial propagation methods should be used with caution. Any long-term artificial propagation could alter the ge- netic integritv of mussel species, and possibh' prove more harmful than beneficial (Kennedy, 1975; O Connell, Page 106 THE NAITILUS. Vol. 110, No. 3 1991 ). As a short-term solution to the immediate problem of endangered species, the technique developed in this stud> offers hope for propagating many species for which hosts have not been identified. ACKNOWLEDGEMENTS Funding for this project was provided by the Upper Mississippi Science Center of the National Biological Ser- vice, and the Center for the Management, Litilization, and Protection of Water Resources at Tennessee Tech- nological University. We thank Michael J. Redding for his suggestions and comments on the study, and the stu- dents and staff of the Tennessee Cooperative Fishery Research Unit for their assistance in the field. LITERATURE CITED Anderson, D. P. 1990, Immunological indicators: effects of environmental stress on immune protection and disease outbreaks. American Fisheries Society Symposium 8:38- 50. .\rey, L. B. 1932. A microscopical study of glochidial im- munity. Journal of Morphology 53:367-379. Bauer, G, and C. V'ogel. 1987. The parasitic stage of the freshwater pearl mussel (Margaritifera margaritifera L.) 1. Host response to glochidiosis. . . Archiv fur Hydrobiol- ogie Supplement 76 4:393-402. Bennett, R. O and R. E. W olke. 1987. The effect of sublethal endrin exposure on rainbow trout Salmo gairdneri Rich- ardson. 1. Evaluation of serum Cortisol concentrations and immune responsiveness. Journal of Fish Biology 31:375- 385. d'Eliscu, P. N. 1972. Obseryations of the glochidium, meta- morphosis, and juveniles of Anodonta californiensis Lea, 1857. The Veliger 15:57-58. Farzaad, M. 1991. Microhabitat utilization of Veniistaconcha sima (Biyalyia:L'nionidae) and its host, Cottus carolinae, in the Collins River, Grundy t^ounty, Tennessee. Master of Science Thesis. Tennessee Technological University, Cookeville. Gordon, M. E. 1995. Venuslacuncha sima (Lea), an over- looked freshwater mussel (Bivalvia: L'nionoidea) from the Cumberland River Basin of Central Tennessee. The Nau- tilus 108:55-60. Gordon, M. E., J. B. Layzer, and L, M Madison, 1994. Glo- chidial host of Villosa laeniata (Mollusca:Llnionidea). Mal- acological Review 27:113-114. Hoggarth, M. A. 1992. An examination of the glochidia-host relationships reported in the literature for North American species of linionidae (Mollusca:Bivalvia). Malacology Data Net 3:1-30. Idler, D. R. and B. Truscott. 1972 Corticosteroids in fish In: D. R Idler (ed). Steroids in Nomnannnalian N'ertebrates. .\cademic Press, New York. pp. 126-252 Isoni, B. G. and R. G. Hudson. 1982. In vitro culture of parasitic freshwater mussel glochidia. The Nautilus 96: 147-151. Isom, B. G and R. C, Hudson. 1984. Freshwater mussels and their fish host; physiological aspects. Journal of Parasitol- ogy 70:318-319. Keller, .-V. E. and S. C; Zam. 1990. Simplification of i;i vitro culture techniciucs for freshwater mussels. Enyironmeiital Toxicology and Chemistry 9:1291-1296. Kennedy, C. R. 1975. Ecological animal parasitology. John Wiley and Sons, New York. Meyers, T. R., R. E. Millemann, and C. A. Fustish. 1980. Glochidiosis of salmonid fishes. IV. Humoral and tissue responses of coho and chinook salmon to experimental infections with Margaritijcra margaritifera (L.) (Pele- cypoda: Margaritanidae). Journal of Parasitology 66:274- 281. O'Connell, M. T. 1991 Immunological responses of fishes to glochidia of freshwater mussels. Master of Science Thesis. Virginia Polytechnic Institute and State University, Vicks- burg. Pearson, C. M., P. J. Clements, and D. T. Y. Yu. 1978. The effects of corticosteroids on 1\ mphoc\ te functions. Euro- pean Journal of Rheumatolog\ and Inflammation 1:216- 225. Pickering, A. D. and P. Christie. 1981. Changes in the con- centrations of plasma Cortisol and thyroxine during sexual maturation of the hatchery-reared brown trout, Salmo trutta L. General and Comparative Endocrinology 44; 487-496. Pickering, A.D. and J Duston. 1983 .\dministration of Cor- tisol to brown trout, Salmo trutta L., and its effect on the susceptibilit) to Saprolegnia infection and furunculosis. Journal of Fish Biology 23:163-175. Pickering, .\. D. and T. G, Pottinger. 1983. Seasonal and diet changes in plasma Cortisol levels of the brown trout, Salmo trutta L. General and Comparative Endocrinology 49: 232-239. Pickering, A. D. and T. G. Pottinger. 1985. Cortisol can in- crease the susceptibility of brown trout, Salmo trutta L., to disease without reducing the white blood cell count. Journal of Fish Biology 27:611-619. Reuling, F. H. 1919. Acquired immunity to an animal par- asite. Journal of Infectious Diseases 24:337-346. Shannon, L., R C. Biggins, and R. E. Hylton. 1993. Fresh- water mussels in peril: perspective of the U. S. Fish and Wildlife Service. In: K. S. CAimmings, A. C. Buchanan, and L. M. Koch (eds. ). Conservation and Management of Freshwater Mussels. Upper River Conser\ation Commit- tee. Rock Island, Illinois pp. 66-68. Schindler, D, W. 1989. Biotic impoverishment at home and abroad. Bioscience 39:426. Schreck, C. B. 1990. Ph\siological, behavioral, and perfor- mance indicators of stress. American Fisheries Society Symposium 8:29-37. Thorpe, J. E., M. G. McConway, M. S. Miles, and J. S. Muir 1987. Diet and seasonal changes in resting plasma Cortisol le\els in juvenile .-Xtlaiitic salmon, Salmo salar L. General and Comparati\'e Endocrinolog) 64:19-22. Tizard, I. R. 1988 Inimunolog\: an introduction Saunders College Publishing, Philadelphia. Waller, D. L. and L. G. Mitchell. 1989. Gill tissue reactions in walleye Stizostedion vitreum vitreum and common carp Cyprinus carpio to glochidia of the freshwater mussel Lampulis radiata siUquoulca. Diseases of .Aquatic Organ- isms 6:81-87. Watters, G. T. 1994. .\iinolaled bibliograplu of the repro- duction and proi.iagatioii ol the Uiiioiioidea. Ohio Biolog- ical Sur\e\ Miscellaneous C Contributions 1:1-158. Zale, A. \ . and R. J. Neves. 1982. Fish hosts of four species of lampsiline mus.sels (Mollusca:Unionidae) in Big Moc- casin Creek, Virginia. Canadian Journal of Zoology 60: 2535-2542. INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes papers on all aspects of the biology- and systeniatics of mollusks. Manuscripts de- scribing original, unpublished research as well as revdew articles will be considered. Brief articles, not exceeding 1000 words, wall be published as notes and do not re- cjuire an abstract. Notices of meetings and other items of interest to malacologists will appear in a news and notices section. Manuscripts: Each original manuscript and accompa- nyang illustrations should be submitted in triplicate. Text nni.st be typed on one side of 81/2 X 11 inch white paper, double spaced throughout (including literature cited, ta- bles and figure captions), with at least 1 inch of margin on all sides. 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Leal Bailey-Matthews Shell Museum .3075 Sanibel-C^aptiva Road Sanibel, FL 33957 CONSULTING EDITORS Dr. Riidiger Bieler Department of Invertebrates Field Museum of Natural Histor\' Chicago, IL 60605 Dr. Arthur E. Bogau Freshwater Molluscan Research 36 Venus Way Sewell, NJ 08080 Dr. Robert T. Dillon, Jr. Department of Biolog\' (College o( Charleston Charleston, SC 29424 Dr. William K. Emerson Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Dr. Eileen H. Jokinen Institute of Water Resources Uni\('rsity of Connecticut Storrs, CT 06269-40 IS Mr. Richard I. Johnson Department of Mollusks Mnseiun of Comparative Zoology- Harvard University Cambridge, MA 02138 Dr. Aurele La Roctjue Department of Geology The Ohio State University' Columbus, OH 4.3210 Dr. James H. McLean Department of Malacology Los Angeles County Museum of Natural History 900 E.xposition Boule\ard Los Angeles, CA 90007 Dr. Arthur S. Merrill % Department of Mollusks Museum of Comparative Zoology Harvard University Cambridge, MA 02138 Dr. Paula M. Mikkclsen Department of Living Invertebrates The American Museum of Natural History New York, NY 10024 Dr. Donald R. Moore Dixasion of Marine Geology and Geophysics Rosenstiel School of Marine and Atmospheric Science University of Miami 4600 Rickenbacker Causeway Miami. FL 33149 Dr. Gusta\' Paulay Marine Laborator\' University of Guam Mangilao, Guam 96923 Mr. Richard E. Petit RO. Box 30 North Myrtle Beach, SC 29582 Dr. Edward J. Petuch Department of Geolog)' Florida Atlantic University Boca Raton, FL 33431 Dr. Garv Rosenberg Department of Mollusks The .\cademy of Natural Sciences 1900 Benjamin Franklin Park^xax Philadelphia, PA 19103 Dr. David H. Stansber)' Museum of Zoology The Ohio State Universit)' Columbus, OH 43210 Dr. Ruth D. Turner Department of Mollusks Museum of C]omparati\e Zoology Haivard Univi'rsitv Cambridge, MA 02138 Dr. Geerat J. 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Box 7279 Silver Spring, MD 20907-7279 TH Et7NAUTI LUS CONTENTS Volume 110, Ntiinbcr 4 November 6, 1997 ISSN 0028-1344 Arnold G. Eversole Cianu'tom'iu'sis oi Mcrcciunia nicrcciiarid. M. r(iiiii)ccliicn\i\ and Their Ihlirkls 107 James L. Theler The Modern Terrestrial Gastropod (Land Snail) P\iiina ol \\'('stern Wisennsiii's Hill Prairies Ill Edward J. Petuch A New (Gastropod Fauna From An Oliiiocene Back-Reef Lagoonal Environment In West Central Florida 122 -i rt.jifc ■•.Jo.,c:r'OQrapr,,- Ir NOV 1 7 1997 Notice This is the final number ot THE NAUTILLLS to lit- pulilislifd In Troplinn (.'oiporation ot Sil\er Spring, Manlanil. Beginning with the next issue (Volume 111, Number 1) THE NAUTILUS will be published by The Bailey-Matthews Shell Museum. .\1I correspondente pertaining to subscriptions, purchase of back issues, and missing issue claims should be addressed to: Dr. Jose H. Leal THE NAUTILUS P. O. Bo.x 1580 Sanibel, FL 33957 LISA Voice: (941) 395-2233 FAX: (941) 395-6706 e-mail: LeaI@aolc(>in Beginning with \olume 112. I^r. Leal will become Editor-in-Chief of THE NAUTILUS. .\\\ new manuscript submis- sions should be addressed to Dr. Leal at the abo\e address. THE NAUTILUS 110(4):107-11(), 1997 Page 107 Gametogenesis of Mcrcenaha mercenaria, M. ciinipechiensis and Their Hybrids' Arnold G. Eversole Department of Aquaciilture, F'isiieries and U'ilcllife Clenison Uni\"ersit\ Clenison. SC: 29634-0362 USA ABSTRACT N(in-nati\t' stocks ot A/crcc/K/n't/ nwrtciiunn, M (V/i/i/iic/dc/isiv and their reciprocal Inbrids were enltured in Sontli (Carolina. Histolog;ic-al examination of the gonads revealed a himodal ga- nietogenic pattern with a major spring spawiiing peak that con- tinued into smnmer, followed by a smaller spawning peak in fall. The spring spawning peak of A/, iiiciniinriii and its ma- ternal Inhnd ocenrred one month earlier than that ot M. cam- pccliicnsis and its maternal hybrid. The length of the spring spawaiing peaks of the hybrids were hvo months shorter and less intense than those obsen.ed for M. lueiTctuiria or M. aiiu- prcliii'iisis. Kfi/ aorils: M. iiu'rcriinriti, M. crimpeclufusis. gametogenesis. h\brids. (juahogs. reprodncti\e C'\cle INTRODUCTION Lod.saiioll (1954) was the first to suceesslulK Inhridize Mcnriuihii iiwiwitaiia {Linnueiis, 175S) and A/, ciiin- pcchicnsis (C-inelin. 1791). Progeny of intra- and inter- specific niatings were cultured in Virginia (Ha\"en & An- drews, 1957), North Carolina (Chestnut el al . 1957) and Florida (Menzel, 1964). In general, M ((unpci-lnciisis grew the fastest, hvbrids were internietliate and A/, incr- cctKiria was slowest, hut M. cainpccliieiisis and its hy- brids had lower sunival. These early studies confirmed that there were some aquacultnre lienefits ot M nur- ccitiiria and M. cauipcchiensis Inhridization. The distributions ol M iiwrcciunid and A/, canipt- chii'H.sis mcrlap along the southeastern United States (Abbott, 1974). Dillon and Manzi (1989) found that hy- !)rids composed as nuich as 88*^ oi 179 clams collected from Indian Ri\er, Florida. Soniewiiat lower hybridiza- tion rate (31%) was obsened Irom a larger sample (n = 1,141) of clams collected from four sites in tlie Indian River by Bert et ill (1993). In Hamlin Creek, South Car- olina, pure A/, (-(uitpvchiensis were present at a irequen- c\' of O.l'yf in large lieds of A/. iniTirnari/i with negligible 'Technical Contribution No. 4291 of the Siiutli tlarolina Ex- periment .Station Inliridization (Dillon, 1992). This suggested that repro- ductixe isolation lietx\een the hvo species may be more pronounced in Soutli Carolina than in Florida. Dalton and Menzel (1983) docinnented the gameto- genic cycle ol' soung male progeny from laboraton crosses of A/, lucircii/iria and A/, raiupeclucniis cultm-ed in .alligator Harbor, Florida. The bimodal spermatogenic cvcle of the female A/, campecluensis X male M. mcr- ccnaria progeny was similar to the A/, caiupcchiensis pa- rental group, whereas, the spermatogenic pattern of the reciprocal hxbrid was similar to M ineiveuaria parents. Hesselnian li id. ( 1989) reported a bimodal gametogenic cvcle for adult quahogs collected from Indian River, Florida; howexcr, no differentiation of taxa was attempt- ed in that stucK. Information on the ganietogeuic cycle ol Mercriiiiriii species is limited and there are no studies on adult (jua- hogs of known pedigree. Also, nothing is known ol the gametogenic cvcle of A/, ciinipcciiicnsis as far north as South Carolina. The objectixe of this study xvas to com- pare the gametogenic cvcle of adult clams of the species M. lucrcciuiria. M. ciiinpn-liicnsis. and reciprocal cross- es. MATERIALS AND METHODS Mcncniirid iiurciii/niii troui Millcn-d, Connecticut, and M (Y/»i/)(y7»V/i.s(s IroTu Port St. joe. Florida, were spawnecl in November 1985. Januan' 1986 and Februarx 1986 at Waddell Mariculture Center, South Carolina. Crosses included: A/ incrrrniiiiii lemale X A/, inercc- naria male (MM); A/, ciiiupcrhiensis lemale X A/, cam- pcclticitsi.s male (CO; M iiurccnaiiii lemale x A/, cinn- l>cclui'n.sis male (MC); and A/, cuntpechivmis female x A/, nwrceiwria male (CM). Indixiduals from spawns and crosses xvere held separately in recirculating sxstems im- til the tjuahogs xx'ere mo\ed to a nurserx' at Folly Rixer, South Carolina. Quahogs from the three spawnis were combined for each cross on 5 May 1986 and maintained in recirculating systems mitil planting (10 October 1986). Quahogs of each cross were planted in the Folly River at about 960/m2 in separate protected trass. Ax- Paw 108 THE NAUTILUS, \ol. 1 10, No. 4 eragc shell icii^tli.s (SD com- puted loi- e:ich pedigree b\ sumiiiing :ili the dexelop- inental stage values lor male and lem:iie i|iuihogs, and dividing this figure bv llie number ol i|uahogs in the sam]ile. This procedure has been snccesslnllv uscmI In others to sludv the gametogenic cvele ol M. lucrcouniri in New York (Kassner & Malouf, 1982) and in (ieorgia (Hel'fernan rl al.. 1989: Walker & Hefferuim, 1994). These montliK gonadal index v:ilnes were then fitted i)V spline inter])olations to follow liie g:mietogenie cvele over aTi annual cvele. KKSUITS CJonadal index eunes lor progenv Irom each cross are given in I'"ignre 1. These results indic:ite a bimodal I'a- metogenie I'vele lor the progenv ol all lour crosses: how- ever, the timing ol g:unetogenesis dillers among the loui' progcTiv. .\ rapid decline in gouiulal index v:ilues indi- cated :i spring S[)awning tluit contimied into summer lor M cainpci-h'wnsis ((KJ) and its materiuil hvbrid (('M). Average gonadal index values lor (X," and (^M (juahogs were 3.62 and 2.28 in April, and 0.50 in August and 0.22 in [nne, respectivelv. A second smaller spawning peak Wiis obsened in the fall when average gonadal index val- ues lor (X: and CM decreased li'om November (3.50 and 1.30) to |;iuuan/Februan- (2.50 and 0.fS7). The pattern ol gonadal index values exhibited bv A/. 1 1 writ' Italia (MM) and its maternal hvbrid (MC) also in- dicated a decrease in spring, but roughlv one month ear- lier than CC, and CM (|ualiogs. .Average g()n;idal index values for MM and MC decri'ased Irom M;n'cli (2.94 and 3.25) through August (0.64) and |nne t 1.30), respective- lv. The decrease in gonadal index values obsened Irom September through November also indiciited a l:iil spawning. Although the trend of gametogenesis in the fall for MM and MC was not as clear cut as that of (X,' and CM, it appears that spawning occurred 1 to 2 months earlier in MM and MC. The patterns in gonadal index values loi' tlie hvbrids :ip[)e:n' to tr:ick tluit ol the m:iteiiKil parent, but at re- dncetl levels and for shorter periods. For ex:nnple, the decreases in gonackil index values lor spring and tail sp:ivvning periods were 3.12 and 1.00 lor C(>' compared to 2.06 :ni(l 0.63 for CM (|uahogs and 2.30 and 1.57 lor MM comp;n'e(l to 1.95 and 0.50 lor the M(,' hvbrid, re- spectively. The lengths ol spring spawning periods (i.(>., tim(> of declining gouatlal index values) were approxi- mately 5 mouths lor (X" ;uk1 MM comp:ned to .'■) montlis lor CM and MC (juahogs. DISCUSSION The sjiring domin;int bimodal ganietogenii- cvele ob- seiAed in this study has been reported loi- Wciri-iuniii t:L\a in Noith ( iiU'olina (Porter, 1964). South (Carolina lEver.solee/ f//., 1980: M;mzi <>/ r;/., 1985), (;eorgia ( llef- fernan .■/ ai. 1989: Walker & Heffern:m. 1994) and Flor- ida (Diilton & Menzel, 1983: Hes,selm:ui rl al . 19891. l,, contrast to Diiltou :uul Menzel's (1983) findings, tlie tim- ing of the sp;iwning pe;iks for M iiicrcciiaria. M. cam- jiciliiciisis and the recipicnal hvbrids Wiis not the s:ime; the spring sp:iwiiiiig pe.ik ol .\/ inirccnaria :ind its nui- teru:il hvbrid oecniied one month earlier than IIkiI ol M. lainpccliirii.sis :ind its maternal hvbrid. n:ilt()n :nid McMizel's (1983) lailnre to detect a dillerenee betxveen the two Mrrrciiaria species :iud e.ich ol the spe( ies ma- lernal hvbiid m.iv have resulted in [lart li'om using siiuill, voimg. and mostlv 111. ile qiKihogs lor histi liogK'.il evalu- ation. Mcrciiiaiia undergo a juvenile male sexual ph:ise (I,oos:molf. 1937) before :icliieving sexiuil maturitv (I'a- ersole ,7 al . 19801. Sex nitio (1.0 : 1.1, M : F) :uid av- erage SL (40.5 + 6.4 nun. SD) indicate that tlu' sanijiled qiiahogs had com|)lete(l the juvenile nuile phase (F\er- si>\v (I al. 1980; Knaiib \- Fversole 1988). A. G. Ex 1997 Fa; 109 X 0) ■D _C ■o CO c. o O 3.5 2.5 1.5 0.5 3.5 2.5 - 2 - 1.5 - 0.5 10 12 2 4 Month 8 10 Figure 1. Me ctiinpcclticnsis livbiid (CM). an gdiiLulal inik'\ with oiu' staiidaixl crnir liar fur crosses of M iitci'iriiiirid friiialc ■ ,\/ inircnunid male (MM), A/. female X A/, camprchiiiisis male (CC). A/, inercciitnia female ■' A/ rdinpcrhiiiisis male (MCJ) and the reciprocal Line fitted b\' spline inteq«)lation. Hcsselmaii ct a]. (1989) encoiiiitc^n'd eoiisidcralilc variations in tin- f^anieto<4eiiic cxiAv ainont; samples of (jiiali(iij;s eolli'ctecl li'diii Iiuliaii Rixcr. Florida. Clixfii that liotli species of Mercenaria occih' in Indiaii Ri\cr, Hfs- selnian et nl. (19S9) speculated one source ol the \ari- alion coiikl he related to genetic ditlei'ences in repiii- dncti\c acti\it\- among the taxa. Since then, i5eit el nl (1993) \'erifii'd tiiat Indiaii River contained hotli Mcr- ceiKiria species and their hvhrids. Ditficiilts associated with defining a ganietcjgenic cvcle wonld he expected il tlie reproduction cvcle is imder some genetic coiitiol and if closely-related species coexist and liNhridization occurs. Currently, there is exidenct' that dillereiit stocks of A/, nwrccnarid and Cnissostrca lin^inica (Cimelin. 1791) exhibit geiieticallv distinct cxries o( gametogenesis aiid spawning when cultured in a common enxironment (Knauh & ENersole. 198S; Barber c/ ,il . 1991). Bai-ber and his colleagues (1991) suggested th.it osster stocks had geneticalK' different emii-oinnental re(|nirenients necessarx' lor initiating gametogenesis and excntualK' lor cuing spawning. nillcin and Manzi (19S9) sugg<'sted that spawning time and habitat separation could he possible mecha- nisms inaintaininsi the genetic inteirritN' ol the two Mer- cenaria species. In the Indian Ri\ei', Floi'ida, where Mer- crnaria coexist, S.S7f and llWc of 179 and 1,141 collected (juahogs were natnralK occurring hxbrids, respecti\t'l\' (Dillon & Manzi, 19S9'; Bert et at.. 1993), Farther north in Hamlin CJreek, South Carolina, oulv about 0,1% of tfiousands oi tjuahogs examined were M. antipecluensis. and Inbrids wei'e i-arer still (Dillon, 1992). Although the timing ol the gametogi'uic cvcles ol introduced stocks of A/, mercenaria and M. cantpecltiensis differ in the spi'ing, sufficient o\erlap inav exist for hvbi'idization of native Mercenaria in South C'arolina. The fact that Mer- cenaria hybrids were encountered less frequently in South Carolina than in Florida indicates that ,spatial sep- aration or somi- other fact may he luore important for I'epioductixe isolation than the tenipoi'al diffei'ences in spawning. It is interesting that the M. campechiew>is and h\bi-ids identified bv Dillon (1992) were collected from a latlier uniijue South Cai'oiina habitat, one character- ized by ocean-dei'i\ed high-saliiie water, firm substrate covei'ed by deep water and a comnimiitx' of stenohaline biota (e,g,, ('.liinne cancellala Fimiaeus, 1767). A suncx' of M. mercenaria habitat in South Carolina rexealed that most ol the areas with clams were mud and sand covered In sh.illow mixohaline waters (Anderson et iiL. 197S). It Page 110 THE NAUTILUS, \ol. 110, No. 4 is .uiticipatfd tliat Miriiiitiriii li\ lnidizatioii will hccome a rarer exent in higlici' l.ititudcs ol tlic Atlantic coa.st as contignous habitats nicctinii; the sp( 'tics-specific require- ments ol both species become less available. Earlv stndies indicated a growth advantage ol M nwr- cenaria and M. campccliiciisis hxbridization (Haven & Andrews, 1957; Chestnut c/ tiL. 1957; Menzel, 1964) possiblv because tlie hybrids Ivdvc a shorter and less in- tense spawning period than tlie hvo species. However, Eversole and Heffernan (1995) also obsened that go- nadal neoplasia occurred more frequently and in a more advanced stage in tlie hvbrids tlian in M. mercenaria and A/, nnnpcchiensi.s. ,\(juacultnrists need to be aware of both the favorable and unlavorable attributes associated with liybridization [irograms when trxing to improve (juahog performance. ACKNOWLEDGMENTS The author thanks Nancv II. Hadlev and her fellow workers lor the clam husl)andn. Special thanks go to John Hudson who assisti'tl with the dissection, Yvonne Bobo for help with histological preparation and Chris Kempton helped with the illustration. Connnents by Hob Dillon and Randv' Walker on an earlier dralt greatly improved this manuscript. The research was supported by S.C. Sea Grant Consortium and S.C^ Agricultural Ex- periment Station. LITERATURE CITED Abbott, R. T. 1997. Aiiiericaii Scasliclls, 2ik1 cd Win Nostrand Rlieinliold, New York. Anderson, \V. D., W. ]. Keith, F. H. Mills, M. E. Bailey & J I,. Steinuiever. 1978. A snnev of South Carolina hard clam resources. South Carolina Marine Resources Center. Technical Report Number .'32, Charleston, South Carolina. Barber, B. J., S. E. Ford & K. N. Wargo. I99I. Genetic vari- ation in the timing ol gonadal maturation and spavvaiing ol the eastern oyster, ('mssostrcti r/rg/ii/rr; (Cmelin). Bi- ological Bnllctiii' 1SI:2I(>-22I. Bert, T. M.. D. M. llcssclnian, VV. S. .\rnold, W. S. Moore, II. Cruz-Lopez & D. (;. Marelli. 199.3. High Irequencv ol gonadal neoplasia in a hard clam (Mcnciuiiia spp.) hybrid zone. Marine Biologs 117:97-104. Chestmit, .\. F, W^. E. Fahv ^■ 11 ] I'orter. 1957. Growth ol vonng Wntis mcnrniirm \iiiiis r{iiiii>crhicii'i;;\ ami Arcliaeolog\' llni\ersit\' of Wisconsin-La Crosse La C^rosse. Wisconsin 54601 ABSTRACT This pajx-r describes the first snncv and (|uantified unaKsis ol the terrestrial gastropod fauna associated with \eric "hill prai- rie" and related dr\ habitats in western W'isconsin's Driftless Area. A region of 35,000 km- in the Midwestern U,S.A.. the Driftless Area was surrounded on three sides, but ne\er co\- ered b)' late Pleistocene glacial ice. In all, 44 separate \egeta- tion detritus samples, each with a xohune of 0.5 to 2.0 liters, were collected in nine western Wisconsin counties. These .sam- ples contained 10,900 gastropods, with 29 ta\a represented. The sampled hill prairies were found to ha\c localized, high- densit\' gastropod populations characterized bv a small mmiber of abimdant t;L\a. One species, Qaatrocoptii proccra (Gould. 1840) is currently listed as Threatened in Wisconsin. This spe- cies occurs at disjunct prairie habitats along the dissected \alle\ of the Mississippi Ri\er that mav represent a relict distribution for this species Ai'i/ words: Laiifl snails. biot;eograpli\, Midwestern L!.,S..\. INTRODUCTION This report describes tile results oi a liascliue sunev to assess the living terrestrial gastropod lauiia assnciatt'd with the xeric "iiil!" prairies (Curtis, 1959:275-276) ol Wisconsin's Driftless Area. The studv provides the first .systematic assessment of gastropod abuudaui-e and spe- cies diversitv' at lull prairie and associated dn loi-ations in nine western Wisconsin counties. From the stand- point of terrestrial gastropod distril)ution, this area is one of the least known regions in eastern North America (Hubricht, I9S5:2), although some researeli lias been undertaken in the past (I. P. E. Morrison, 1929; Levi & Levi, 19.50). The phvsical setting of western Wisconsin is varied and complex. The heavily dissected hill coimtn- of this region has been assigned to the Western Upland phys- iographic province iiv Martin ( 19fi5:42-43). Southwest- ern Wisconsin is within the Driftless Area (Martin, 1965: 82-8.3), located primarily within the Western Upland, with smaller portions extending into the adjacent states of Iowa, Illinois, and Minnesota. The Driltless .Area, as its name implies, lacks evidence ol glaciation during the later Plei.stocene (MickeLson it r;/. 1982:155-169), and covers an area of 35, ()()() km-, about 75% of which is in southwestern Wisconsin (Roosa, 1984:43). The Driftless Area's topography is characterized by steep-sided, stony valleys dissecting the uplands with dendritic patterns of small stream development. The narrow upland ridges and escaqinients of the larger valleys e.xhibit up to L50 ni of relief. The two prominent rivers crossing the region are the Mississippi, which lornis the western boundan' of Wisconsin, and the Wisconsin River, which drains much of central and southwestern Wisconsin. Western Wisconsin is situated between the tall-grass prairies to the west and the deciduous forest to the east. The southw-estern portion of the state has been assigned to the "prairie-forest province" King to the south of the Horistic tension zone of Curtis (1959:15) and is charac- terized by its mosaic of vegetation communities. The structure and position of tliese connnunities have re- sulted from a variety of factors, including topography, climate, peiiodic biu-ning of prairies, and underlying soils and bedrock. At the time oi Enro-.'Vnierican settle- ment, four distinct vegetation connnunities were pre- dominant: prairie, oak savanna, southern oak forest, and southern mesic forest. Tlie most extensive prairie com- munitv was the mesic prairie tvpe, dominated by grasses such as big bluesteni {Aiulropo^on gcrardi). Indian grass {Soriihristnini iiut/ms). and forbs. Prairies bv definition support fevvi'i- tli;m one tree per 0.4 ha. In western Wis- consin, mesic prairii'S iormed large tracts on gentiv roll- ing uplanils, particularlv south ol the W'isconsin River in Iowa, CTrant, and Lafayette counties. Nineteenth-centun,- travelers who journeyed from forested regions in eastern North America often described the extensive tall-grass prairies they first encountered in western Wisconsin. One of the best-known mesic prairies was the Militarv' Ridge Prairie, which ran Irom the town of N'erona in Dane County, to the northwestern corner of Cwant Countv- (Curtis, f 959:262-264). Xeric prairie coTimumities in the Driftless Area were locally conspicuous but spatially circumscribed (figures I, 2). These xeric hill or "goat" prairies occupied steep south- f)r southwest-facing slopes and made up only a Pa2e 112 THE NAUTILUS, Vol. 110. No. 4 J^ I \ Figure 1. Brack's Bliitt liill prairie in Trempealeau Couuh', Wisconsin. Arrow points to the saniplini^ location (l'\\.\-3()2(ii shown in figure 2. \'iew is toward the north, with the Mississippi River in the foreground. .small traction ot tlie Diiltlcs.s Areas vc^ctatioiial laiul- scapc. Hill prairies dexelopi'tl (jii the tliiii soils o\erKing (loloiiiite outcrops with exce.ssive ruuoH. These locales lia\'e a nniforiiilv high calcium couteut and a pH of S.O or more (CJurtis, 1959:275-276). The four most common plant species on these hill prairies, in order of abun- dance, are little bluestem {:\iitln)j). Gtistnu-oplii procira is a species associated with prairie/savanna vegetation connmmities in regions with a frost-free season of 160 ckivs or moie (Reigk', 1963; Baerreis, 1980:108-109). In 1985 two addition. il hill prairie vegetation detritus sam- ples were collected in La Oosse Countv. Wisi-onsin. One of these sam|)les .ilso contained G proccrii. Per- sonnel at the Wisconsin Chapter of The Nature Cou- senancv' (TNC) and the Wisconsin Department ol Nat- ural Resources (WDNR) showed interest in de\elo|)iug aTi iu\entor\ of hill prairie gastro[)od connnmnlies, lead- ing the autlioi to (olleit and analvze 10 .uldilioii.ii s.im- ples from western Wisconsin hill pi.iiii<'s and rel.ited habitats during 19.S6 and 1987. Hill pniiries sampled hn gastropods nicliided .i \,uiet\ J. L. Theler, 1997 Paw 113 Figure 2. BraiK's BIiitT hill prairir in rreiiipeiileaii Coimtv, Wisconsin. Sampling location l!\\A-o()2H is at tlic iipsiope side ot the ilftachcd lK]nkler. Photo looks west, with the Mississippi Ri\er on the lett. of TNC, WDNR, antl pri\atel\ (iwiicd \cric scttiiiy;s in nine western Wisconsin counties, eight of uliicli l)order the Mississippi Ri\er; Ironi soutli to nortli, thi'\' are Grant. Lafayette, CJrawfonl. N'crnon, La (^ros.se, Trem- pealeau, Buffalo, Pepin, and Fierce (table 1). Whenex'er possible, TNC anil WDNR propertii'S were emphasized, under written agreement with those agencies. In all, 35 samples were taken and anaKzed at 19 separate hill prai- rie settings (figure 3). Specific sampling locations were selected 1)\ searching each hill prairie to identii\' areas with living gastropods or fresh shells. The most prodiicti\-e locations occurred where relativeh dense stands or clinnps of prairie grass- es aiiutted isolated bedrock exposures, forming "pock- ets" that would hold accunuilations of \egetation detri- tus, .Althougli some of the sampled hill prairies (e.g.. Hush (Ireek South in X'ernon Countv) coxcr st-M-ral acres, the gastropods were primariK limited to discrete concentrations within the larger areas. Tlii'se localizt'd gastropod populations, often restricted to less than a sfjuare meter, seem to represent small, isolated com- mmiitii's within the hill prairie setting. Clixen this distri- bution pattern, random sampling would liaxc resulted in the recoveiT of fewer species and lower individual snails, in addition to the hill prairie settings, nine sampU-s were taken in four counties (Cirant, Lafavette. CJrawford. and N'ernon) from what were consitlered to be "iiigh- stress" habitats for gastropods (table 2). The habitats tiiat are referred to as high-stress appeared to represent the driest locations that could be expected to support lixiiig gastropod commmiities. These locations geueralK lacked xeric prairie vegetation and consisted of a thin layer (<2 cm) of organic detritus on dolomite bedrock exposm^es. The\' were olteu adjacent to some canop\ co\er from wood\- \egetation (birch, cedar, or oak), but were otli- en\ise exposed to the extremes of Wisconsin's climate. One to four samples, eatli with a xolume of 0.5 to 2.0 liters, were taken at each hill prairie or high-stress hal)- itat. Each sample consisted of vegetation detritus and approximateK' 1.0 cm of underKing soil taken among prairie plants adjacent to or against rock exposures. Sam- pled locations were marked on IISGS topographic maps and usnalK photograpiied. Each sample was measured In solnme in the laboratorx' and then water-screened, with the gastropods aiul other material retained in geo- logic siex'es (T\'ler #40. mesh size 0.425 mm). The screen-captured residue was then dried and .systemati- cally se-arched for gastropods under a low-power (lOX) binocular microscope. .^11 gastropod shells and poten- tialK identifiable shell hai'inents were remo\ed from the Page 114 THE NAUTILU.S, Vol. 110. No. 4 t2 a, o S o = L L L J 2 3 5 t/3 C« C/5 ■*- u u u o !\j Oj .^ ." ;=; z: ;: i^ -^ ;rr ^ ^ = ^ ^ -a ?i c^ c5 c^ "B ^ ii o O .J ^ sJ = = "^ >. O o ~> "^ ^ _o -O _0 -Q Ul -e-e o -0 ~ ^ ^ Wl _0 _g to -^ "Z :: '^. *T^ 5 1 ^--2^ '-£ ^ ^ . :? ^ C:^ t-X .= C/l W C/i r/> r/l t/i t/i c/1 (/^ r/l :r r/^ r/1 f/i f/i r/1 r/1 r^ ir C/1 ly; ly; t/^ O O O O C^ Oj Oj 0; Cj lb lb (U (U 0; 0; a> 0; lb yj ■«b th y/ ■ ^ ^ C/) i/5 t/2 V2 I 'Xi ,^ '^. .^ 'y^ >^ u^. \r. j^. ir. j^. j^. .n .r. ^ > > > :> CAl CO . . . '^ «^ > -o tLltLlc022;S:SZ::;coy; Z Z >■ Z Z >• ^ >- >- UUZ^EoUWZt^ZZ ;/}C/:i C/!jC/1 _^ ., ,, oi o] 1^ -t -t lO lO CO CO :r cc :^rz, y. 7, yy.^^ ^. y^ ^, y. . — ^ , — ^ ~f r"^ '"^ ""^ ""^ ~f ~f lO lO ol CI r-1 r-'l CI cl c'l cl cl c'l ci Hf— HHHHHE— f— t— f— I- ^ I- 35 3^ I- t- e; ^ lO I- 1~ IC IC ^ ^ lO ^ ^ ^ I~ I- f ^ tC CC (c CC CC CC CC t- I^ 1^ 1~ ^ccxi-i~cc;^a^>:cccccccooacjcocjccccococooc^cz~a'c^occccca?arjcjccccc NE. NE, SW NE. NE 5W. SW SW. NE. NE SW, NE, NE W NE, NE W NE, NE SE, SW SE, SW NW, SW W SW, NE W, SW, NE S; SW NE, NE S; SW. NE, NE : SW, NE, NE '); SE. NE : S. NE, NE ); S, NE, NE 4; E, SE, NW 4; E, SE, NW 4; E, SE, NW 4; E, SW, NW S; NW, NW, SE ); N, NE. SW CO^H O^a^COCOCtOCO'^-^fMO-lKcoScOcOcOcOCOfMfM _H ^ ■ ' ^ ^ CO CO '>] . ^ 1) ^ r o o o 03>0' CO CO CO cl C] CO CO CO CO CO CO CO C] cl CO C] CO o^ocii'-tcc-rioco^^^^cco;!- ClCO^lJO^CICICICICO't-t't't CO CO CO CO ci CO CO CO CO CO CO CO CO CO — ~ ^ _ s: -'. O - :c uu U. a. CO EC -J CO lo — : 'C - O J o f- 22 a. ■1/ h <■■ — R ST. "5 1 '" tt: r^ s C T ^. c; — r^ Zj ^r. t- ^ 1^ c II II II rC m So -* J. L. Theler. 1997 Page 115 t2 = ; c tt '•^ '■^- O 1/ _:r "^ c - ■J; ^ w X^ O 1^ Ij ^"^ -- *- a; ^ li o 0/ > -t: :- '^ l/l V. ^ • _ y v: v; t/; -r— y: v^ c/5 ;> i- ;/: ;/5 c/2 y. ^ y^ z z z ^- >• :^- a M w z z > c/; j^ c/2 Z Z Z z z ^ >■ >' is-' ti a w > z z z =^ ^ ^ c^ c^ ^ „• _• _• o, c, c, o o -■ Q^ O 1> 1/ iJ 1> Qj 1/ 3j C« C/2 C/2 C/2 c/5 ;/2 '^ > > > > > > c/; c/2 ;^ ^ u r^ ft r^ g eS g lO IP — ; a: a: cs 2: cc K cd K 2: ^, z z z z z z Z Z Z a^ cp o^ -t -t -1- ^ r- r- H H ^ 1 - 1- J- tc iC iC cc tc CC ^ .'■ -/■ l- J- J- J^ ^' ;/- vT' -f ^ 1- -? -? •? d5 ^ — 01 r CI r'-^ -r 1- .7- ^ ~ in \^ ci •^1 CO '-^.' CO C.^ CO s u Page 116 THE NAUTILUS, \\>\. 1 10. No. 4 Table 3. (;;istn)|)iKl Li.s.s('inl)l;ii.ii'S troni wcstcni Wisconsin liill ])rairies. Lula- Crant vette C'lawtorcl X'enion Counh : J Dew- M- Lcitner Battle ev Ziin- Mo- (Jran (Jrac Hollow Rush Oeek BluH X'icton Sariiplf location; Hts. nier untl UVVA:3051 3014 3040 2547 2576 3055305fS 303530363058 30433053 296229633037 Taxon: Sample volume (1.): 1.1 0.75 1 NR NR O.S 0.7 0.8 0.8 1 2 1.8 1 1 0.75 Stcnotrema harlwtiiin (G.H. Clapp, 1904) 2 — — — — — — — — — — — — — — Mesodon claiisns (Say, 1821) 1 — — — — — — — — — — — — — — Euconiilws /uIdus (Mullen 1774) — — — — — — — — — — — — — — — Ghipluinlinia hulcntnta (Say, 1823) 2 2 6 _ 3 — — 7 14 ^ 60 7 1—2 Scscnitrca ricciriiia (Gould, 1841) — — — — — — — — — — — — — — — Hawaua minnsada (Binney 1840) — 7 — 2 4 2 14 1 1 1 7 12 3 6 9 3 'Ainitoidi-s nrhorciis {Sa\, 1816) — — — — 13 — 2 2 S — 5 — — 1 1 Sirifiturii iiiiliuiii (E,S. Morse, 1859) — — — — — — — — — — — — — — — Striiiturii sp. — — — — — — — — — — — — — — — Deroccras laeve (Miiller, 1774) — — — — — — — — — — — 1 — — — Anugimpira (dtenmta (Say, 1816) — — IS — — — — — — — — — — — — Hrl tod isms pamlMus (Sav, 1817) 9 — — 4 35 1 29 — 16 8 79 26 — 3 20 Hclicodisrus sin^lciinnus (Pil.shn. 1890) 2 — — 1 — — 5 — 23 4 4 36 3 — 12 Piiiictuiii iiiintifissiiiiiiiii (I. Lea, 1841) 4 6 — — 2 — — — 6 1 8 1 10 4 5 Snccineidae 3 — 2 — 1 — 2 1 1 — — — — — — Strohdops lahtp-iiithira (Sa\-, 1817) 9 6 — 9 14 4 2 13 45 2S — — 3 3 2 Sirohdops (iffiitis Pil.shiT, 1893 6 8 — ____________ Strohdops sp. ■ _ 17 _ _ _ _ :3 1 42 5 — — 2 1 — Gastrocopta amiifcra (Say, 1821) 11 32 IS .39 27 12 98 2 3 59 34 11 4—2 Gastrocopta contrmia (Sav, 1822) 5 2 6 1 — — 5 2 16 — — 1 2 11 Gastroroptti Iwlzinocri (Sterki, 1889) 39 9 5 101 14—2 13 27 34 30 33 69 37 53 Gdstrocopia pcntodon (Say, 1821) 7 8 — 7 29 — — — 9 13 23 12 16 21 3 Gdstrocoptii corticaria (Say, 1816) 3 — 1 2 1 — 3 — 1 — — 1 — — — Gastrocopta proccra (Gcnild, 1840) 10 13 — 2 5 14 4 3 7 1 21 12 8 — 10 I'lipoidrs albdahris (C.B. Adams, 1841) 4 10 2 — — 13 22 — 6 4 51 9 — — — ('oliimella cndcittida iDrapamaud, 1805) — 1 — — — — — — — — — — — — — Valloiiia pidchclla (Muller, 1774) — — — — — — — — — — — — — — — Valloiiia costata (Muller, 1774) — — — — — — — — — — — — — — — Vallouia panula Sterk-i, 1893 _ _ _ 75 14 2 29 2 5 27 51 33 1 6 10 Valloiiia pcrspcctiia Sterki, 1892 1 — — 1 — — — — — — — — — — — Vallouia sp, 20 — 7 108 15 7 53 1 6 54 68 64 5 3 13 ('ochlicopa Inhrica f Muller. 1774) — — — — — — — — — — — — — — — Juwnilcs ,30 15 4 NR NR 20 28 5 24 35 58 33 56 46 61 NR = Not recordeil. (Ictritus and held lor identification. Spceinicn.s were later identified, fatalo])t(i armifcra (Say, IS21), Gastrocopta hohJuocri (Sterki, 1889), and Gns- tfocoptfi })ciito(loii (Sav, 1821). Each of these species contributed more than 11% of the total number of snails and was present in 30 or more ol the 35 samples: iwn ol the three were represented in all nine counties sam- pleil (table 4). Each ol the next 12 most .ibnndant t,L\a represented 1.3'^ to S.T'/r ol the total number ol snails, occurred in 14 to 2S ol the 35 samples, and was lound in at least six ol the counties suncved. Together, these 15 most abundant taxa Tuack' up oxer 96*^ ol the hill prairie gastrojiods. The niajoritx of the snails rejiresent- ed are taxa that UpicalK occur in prairie nv sav.uina hab- itats. The rem, lining 14 species made up less than 4'"'f of the gastiopods, and most occmri'd in rel,iti\el\ lew sam|)ies. The nine sam|)les taken Irom high-sticss h.ibitats con- tained 2.236 gastropods of 20 taxa it.ible 5). All of lliese J. L. Theler. 1997 Page 117 Tabli- 3. Kxtciulrd. Iniii- pe- La Crosse aleaii Pepin Pieree 7. ■ i n Buffalo 77. 777, , , , hA]ieniiieiital Bra- hi\e-.\lile Maid- Fanii Hixtiiii Forest Bin! (l\-'s Landfill Rd. Stock- l^rairie Fk Pepin en Rusli Hajj;er(:it\ Bliitf Bluff liolm Rock Riv. ^ ~ 29S7 3005 :50()fi 3029 3030 3042 3057 2V}W :)020 3024 3025 3023 3031 3()4f 3()4S 3049 3047 3046 3044 3045 1 1,5 1 1 f.2 f,f O.S OS I 1 1 1.5 1,1 f 1.5 ().(i5 1 1.7 1.1 0.6 _______ __ !________ — _ ___ 2— 4— — 1 4 1 — 1—— !____ — 5 2 11 22 3 1 4 5 12 — 3 5 15 11 11 3 20 — — _______ ______ !______ _ _ 1 27 1 3 — — 1 — 2 5 — 4 89 36 27 42 6 43 13 1 5 2 1 — — 2 20 1 — — — 5 3 1 — — 2 _______ ____________ 4 _______ !____________ _______ _____!_______ — S 12 61 5 6 — — 19 41 5 4 41 9 40 4 10 5 1 1 9 — 2 22 20 23 23 — 12 36 — — 31 20 13 24 23 14 4 — _ _ _ 36 1 14 ] — 3 4 — 1 — — — 2 3—12 1—1 2 2 — — — — 1 1 4 10 23 24 14 5 2 — 12 6 — — 15 4 — — — — 5 — 1 59 31 55 101 — 1 3 13 _ _ 29 5.S 4 141 11 5 15 11 16 42 1 4 13 7 13 80 56 97 1 _ 1 3 3 7 3 _ _ IS — 1 17 .33 6 7 4 16 1 1 ;54 2 63 — 24 3 11 33 5 — 10 31 — — 19 45 23 53 27 4 23 5 13 79 25 9 12 1 U 4S — 13 51 23 76 53 11 41 22 22 _____ 7- ___2— — — — — 8 — — — 1 _ 10 14 — 1 5 _ 14 _ 1 _______ 6 3 _ _ 22 28 — 9 — 1 6 2 12 13 3 — 9 12 3 45 15 10 _______ fi____________ — — ——— — 1 71 3 — 19 32 — — 9 19 8 60 12 — _______ _________ i__4 _ _ _ _ _ 54 — 480 6 — 42 43 — — 10 19 — .54 41 21 _______ ______ 5______ 41 4 53 83 10 102 44 78 15 55 21 48 31 16 31 77 21 79 45 44 tiLxa were also represented at hill prairie sites; however, tiie rank order of the species was different in tlie liigli- stress samples. The first- and second-ranked species at hill prairies, Gastrocopta amiifern and Gastwcopta liol- zinaeri, ranked second and fifth in the hiifji-stress sam- ples and contrihnted 3S.5% of all snails (talile 6). Three other species, Gastrocopta roiiicaria (Sax; ISlfi), Valloii- la perspectiva Sterki, 1S92, and Vallonia cii.\tala (Miiller. 1774) contributed 52.5% of the gastropotls in the lii'j;h- stress samples, ranking first, thiril, and fourth respec- tively, wliilc ranking sixteenth, nineteenth, and twentx- first in tile hill prairie samples. Together, these fixe most ah\mdant forms represented 91.19?- of the species re- coxered in the high-stress habitats. The difference in species composition between the assemblages probablx reflects the differences between the sampled habitats. DISCUSSION The hill prairies of xx'estern Wisconsin host a fairly di- xerse connnuuitx' of terrestrial gastropods, with some 15 species being relatixelx' abundant and xxddespread. A comparison of these sunex' results xxith Hubricht's (1985) countx' listings for eastern North America re- xealed that the hill prairie surxey contributed 1.30 new countx^ records, and eiglit species that had not been re- corded prexiously in Wisconsin (table 4). One taxon xx'orthy of special comment, Gastrocopta proccra. is currently listed in Wisconsin as a state Threatened species. Gastrocopta procera finds its pre- ferred habitat in prairie and sax'anna settings. This spe- cies does not lixe in the closed-canopy moist xx'oodlands of xx-estern Wisconsin (Theler, 1992). The nortliernmost Page 118 THE NAUTILUS. \ol. 110, No. 4 LANDFORMS OF WISCONSIN Gaological and Natural History Survay Q*or9a F Hsnkon. DirvcWf and Sut* Oaoloaist uwiviaairf cxtcnsion. uKiviasmr or Wisconsin 1971 Hager Cit Five-Mile Prair Rush Riv Maiden Ro Lk. Pepi jtoc'kho Landfill R Brady's Blu Bird Blu Hixon Fore Experimental Far Cedar Valle Vicfo Battle Blu Rush Cre Leifner Hollo Gran Gra Zimme' Dewey Hts. -*' M-Mound KiHiirc 3. Saiiipliiiii liK.itioii in wi-sUiii \\ iscoiisiii. I Biisi- map n\i(l ii illi jicriiiissioii aj llir W isroiisin Crolo^ical iiiid Xahiriil llisliin/ Siiirfijl. iiiodcni range ol Cdsli'niojiln pnitcni in North .Vnierica is represented hv tlie small popnlations lonnd at hill prairie locations in western Wisconsin described in this paper. These hill prairies are ii innanls ol more extensive njiland prairies and sa\'annas that c<)\ered ridgetops and well-drained sonth- and west-lacing exposures in much ol southern and western Wisconsin al the time ol Euro- American .settlement (Anderson. 1954; Curtis, 1959: 262-264). These prairies became established during die mid-1 loloccTie warm/dr\' Atlantic climatic episode, al)out S.OOO to 5.()()() years ago (Wendland, 197S:27S: Wend- land (S: iiiAson, 1971). The Atlantic episode witnessed the eastward expansion ol the wedge ol prairie .iiid sa- vanna often called the Prairie Peninsula (Wright. I96S). The ilnnalic episodes that lollowed the Atlantic were cooler and/or moistcr. and the prairie/savanna vegetation conmuuiities were reduced (Wendland. I97S) to stands that were either fire maintained or situated on topo- graphicalK controlled droughtx exposures, such as hill prairies. The cessation ol prairie fires with the onsel ol Enro-Ameiican seltleineut dining lln' ninitiintii n Rank iiecnired specimens <7c reeorded records records Ciislrocoptii iinnift'ra 1 32 955 l(i.5 9 8 CdstriHOjiid litilziii^t'h 2 30 862 14.9 9 9 Gdstn'coptd pcutdilou 3 30 686 11.8 8 7 Hfliciuhscus piirtillclus 4 28 502 8.7 8 7 Hawaiia iiiiiiuscnla 5 27 368 6.4 S 8 Gii.strocojitd conlrdcid 6 27 164 2.8 S 7 Strohihips Idhi/riiitliird 7 26 240 4.1 7 5 Ghjpliijdlinid intlciddtd 8 26 237 4.1 9 9 * Ptinctmn ininiitissiiniiiii 9 25 323 5.6 S 8 * Ptipoiilo. iiUnldl>ris 10 24 311 5.4 9 9 * Vdllonid pdinila 11 22 489 S.4 ( 7 * Gdstrocoptd proccrd 12 22 165 2.8 1 7 * Tj-mitoidcs drJum-ns 13 20 79 1.4 1 6 Suctineidae 14 17 78 1.3 ~ ? Hflicinlisciis siitilht/dnus 15 14 115 2.0 6 6 * Cast roccptd ct>i-ticdiid 16 10 29 (»,5 7 7 * Eiuo)iulus fiiliiis 17 7 14 0,2 4 Adi^iiispird ditrnidtd 18 6 56 1,0 2 \'dllt>iiid pcrspfdiva 19 4 7 0.1 3 Willtniid (■(i\tdliliips dffiiiis 21 2 14 0.2 1 Sfniotreiiid Ixirlnitum 22 2 3 <(1.1 2 Dcroccrds Idcvr 23 2 2 <0.1 2 * Vdllonid pulchclld 24 1 6 <().l 1 CochliiDpii Inhricd 25 1 5 <0.1 1 Stridltii'd inilhiin 26 1 4 <().! 1 Mcsixliiit ildtisiis 27 1 1 <().] 0 Scsot itrcd I'li'iirifid 28 1 1 <0.1 1 C'Ohnnilld nlrutuhi 29 1 1 <0.1 0 Ta\a not inilndt-d: Vdllonid sp. ||u\t'niles] 24 1,194 Stmhilops sp. [Juveniles] 18 365 Stridtiird sp. [|n\eniles] 1 1 [n\eniles [undefined t;L\a[ 33 1.313 Irdst-tice period ikji tliward aloiii; tlic xallcx eoiridoi". Hill prairies along this eorritlor ari' the oiiK kuowii umd- ern locations in Wisconsin snpportiuS .30.52 .50.5:1 .5( ).54 3027 .502S .50.39 \R 0.5 0.1 0.4 2.0 0.(S \R Eticoiuilus ftlllU.S Glijpliii\ sp. Qastrocoptii annifcni Gastrocoptd contractu Gastrocoptii holzingeri Gastrocopta pcntodun Gastrocopta coiiicaria Pnpoich's alhiliihhs Viilltiititi citstiilii Vdllonid pdnidu Xdlloniii pcrspi'cthd \'dll<>nid s]). Iinc'iiilcs 2 — — — 96 98 37 15 4 3 47 13 74 — 24 1 6 2 — 2 — 5 1 40 38 1.58 — 83 86 265 9 .1- 70 .5S U) 13 — 6 1 3 5 — 1 2 13 39 5 1 1 — — 1 11 1 2 _ 3 1 26 2 120 .3 120 8 51 13 _ 5 1 2 4 lis .3.3 15 12 51 1 1 15 NR = Not recoriled. diger Bieler at tlie Field Miisciiiii ol Natural llistiii-\ lor ollcring coiiiinents on this paper and accepting \'oiicher specimens. I would also like to thank Jerenn' Nienow for help on the tables and figures. An earlier xcrsion ot this paper was presented at ati annual meeting oi the Wis- consin .\cadeni\' ol Sciences, Arts, and Letters, held in Platte\ille, Wisconsin, on ,\[)ril 21. 1090. litp:h,ature cited .'\ndersoii, O. 1954. The pliNtosociologN ol dn liiiie prairies of Wisconsin. Pli.D. Dissertation, Universilx ol W Iseonsin at Madison. Bartlein, I'. ]., and 1'. Wcl.h 111 I0S2. Ilolortne eliniatic change estimated Irom pollen -18. J. L. Tlieler, 1997 Page 121 Table (i. Hank order ami asseinl)lai;e irequencs ol suiiiined terrestrial t^astropcids iroiii nine liii;li-stress luiliitats in western Wisconsin. Nnniher of assem- blages in Xnmher wiiicli ol species speci- •Species Rank occurretl mens % Gastroioptii ciii-ticiirm 1 9 218 17,4 Gastrocoptii (innifcid 2 8 380 30.3 Vtillonia pcrsprrtiia 3 6 244 19.5 Vdllonia ctifitiita 4 5 197 15,7 Cast rocop tii holziuj^c ri 5 4 10:5 8.2 Succineidae 6 3 1(1 0.8 Ga.strocopta pviitodon 7 3 8 0.6 Zmiitoidis (irhorcn.s S 3 ( 0.6 Helir(xli\ais parallclii^ 9 3 1 0.6 Valloiiiti ptiixnla 10 3 3 0.2 Pupoich\ alliildhiis 11 0 US 1.3 Gtislrocopta contnicta 12 0 12 1.0 Strohilops Jahijtinthicn 13 0 6 0.5 An^iiispirii altcrnata 14 1 27 2 2 Euconuhis fiilviis 15 1 9 0.7 Glyplii/dlinia iiidcntata 16 1 3 0.2 Hnwiiiia iniiiitscnhi 17 1 1 0.1 Hclicodisai\ s7(ii;/ci/r;;ii(.v 18 1 1 0 1 Piinrtiiiii iiii)iutissimiii>i 19 1 1 0 1 Strnl>iltip\ (iffiiiis 20 1 1 0 1 Tiixa not inclnded: Vdlliiiiiii sp. [Ju\eniles] 9 614 Strohilops sp. [|u\eniles] 2 4 |ii\eniles [nndefined taxa] 9 364 Roosa, D. M. 1984. Xalnral heritage protection in the Dritt- less Area. Proceedings ol tlu' Iowa Acadenn ot Science 91(11:42-46. Theler, J. L. 1992. The status of three Vei-fioo species in Wis- consin. Report to the Wisconsin Department of" N'atnral Resources, Bureau of Endangered Species, Mathson, Wis- consin. 42 pp. Turgeon, D. D.. A. E. Bogan, E.. V Coan, W. K. Emerson, W. G. Lvons, W. I,. Pratt, C:. F. E. Roper, A. Scheltema, F, G. Thompson, and J. D. Williams. 1988. Common and scientific names of acjuatic in\ertel)rates from the United States and Canada; mollusks. American Fisheries Societv, Special Publication 16. Bethesda, Mar\land. 277 p. VVendland, W. M. 1978. Holocene man in North America: the ecological setting and climatic backgromid. Plains Anthro- pologist 23:273-278. Wendland, W. M., and R. A. Br\son. 1974. Dating climatic episodes of the Holocene. Quatenian Research 4:9-24. Wright, H. E., Jr. 1968. Histon of the Prairie Peninsula. In: R. E. Bergstorm (ed.). The Quateniar\' of Illinois, Special Report 14, College of Agriculture, Universit\- of Illinoi,s, Urbana. p. 78-88. THE NAUTILUS 1 1()(4):122-13S, 199'] A New Gastropod Fauna From An Oligocene Back-Reef Laeoonal En\dronnient In West Central Florida o Edward J. Petuch DfjKutineiit of Gt'olos^' Florida Atlantic Uiii\f'rsitv Bota Raton, Florida 33431 utkI Departnjent ol Pakontoloin,- Graves Miisfinn ol Arcliacoloi^x ,uul Natural History 481 SoutI] Federal Iliis ki'iuhvici n. sp. (Cvpraeidae). Spiniftilf^^iir i^c'inmulatiiin n. sp. (Bus\conidae), Soh'no.steira sit- iciiiiiifcnsis n. sp. (Buccinidae), Vasuin siiwaiincensi.s n. sp. (Turl)inellidae), Ftilsih/ria kciidrcui n. sp. (\'olutidae), Con- oiiiitra ki-iiilnwi n. sp. (N'olntomitridac), D<'nliiiuira- loosahatehian Mollusean Provinces (Petuch, 1982; 19S8). As presently understood, the richest and best-preserved Oligocene assemblages are found onlv in southwestern Mississippi State (MacNeil & Docken', I9S4), and this area is considered the arehetvpe tor the earlv Oligocene. To date, three hmidred and ninet\-se\en species and subspecies of gastropods are known from the \icksburg Group (MacNeil & Docken-, 1984). In the earlv part of tlie Twentieth CyenttuA, N'icksburg- aged gastropod faunas were discovered along the Flint Ri\er and Blue Springs areas near Bainbridge, Georgia (Dall, 1916). These were found to be associated with the rich and extensive Oligocene coral reefs that were de- scribed from the same area bv Vaughan (1900). Later, Mansfield (1937) described an e\en richer Floridian Vicksburg-aged gastropod fauna, primarilv from (|uarries in Pasco and Hernando Counties (west central Florida^ and showed that several Flint Ri\er species occurred that far south. Although eontaining some elements ol the classic X'ieksbnrg lanna ol the .Mississippi Embasment. these two so\ithern loral reel-associated lamias also housed numerous characteristic endemic elements, in- chidiiig the stidinbid Chiliilttlax, the \olutid Fiilsilijrui. and a distinctixe assemblage ol o\er filtc'en species ol cerithiids and fi\e potamidids (combiiu'd Flint Ri\eraiid Suwannee Foriii.ilions). Since Mansfields work in 19.37, liowe\er, no new eompreliensi\e stiuK ol the s\steni.ities ol the Suwannee gastropods has bei'ii published. l^eeeiilK. an e.vposme ol a pre\ ioiisK -miknow ii, higlllv lossililercnis hu'ies ol the Suwannee Fcnination has been louiid in a limcslone (|uaii\ in extiemc western Polk CoiiuIn, west central i'lorida (The Terramai' Pit on l''lor- ida Highwav 54. approximatcK' 5 miles east ol '/eph\r- hills. i'a.sco CoimtN) (E. X'okes, 1992). This localit\ rep- E. J. Pftucli. 1997 Paw 123 resenti'il a liatk-rci-i laij;()(inal t'n\ iinniiifiit that liad lonned lii'liiml a icct tract to the west and cciiitaincd the largest miinlier oi gastropod species vet loiiiid at aii\ single Suwannee exposure. All specimens ot corals and niollnsks ironi the Terraniar site are silicified (siliceous pseudonioqihs), much like the Flint Hi\-er niaterial, and are generalK" well-presened. Throu<4li the iiitensi\t' col- lecting of Mr. Eric Kendrew ol X'alrico, Florida, and througli additional collecting hv niyseli, se\'ent\■-t^\•o spe- cies of gastropods (listed in Appendix 1 ) are now knowTi from the Terraniar Pit, more than douhling Mansfield's total of tlnrt\-one species. Nine of the Suwannee species reporteil In' Mansfield (1937) and one h\- E. N'okes (1992) were not collected at the Terrainar site, and these are listetl in Appendix 2. When Mansfield's and N'okes' additional species are combined with the list of those from the back-reef lagoonal enxironnicnt. the total num- ber of known Suwannee gastropods increases to eightv- two, less than one third of the numlier of X'icksbvu'g spe- cies. Of the se\c'nt\-tx\'o species collected at the Terraniar Pit, txvent\-one are new to science. Three new genera, all of which appear to be endemic to the coralline en- \ironments of the Flint Rixer and Suwannee Forma- tions, are also new. Following a discussion of the paleo- ecologv of the Terramar Oligocene site, tliese new spe- cies and genera are described. PALEOECOLOGY OF THE RAf:K-REEF LAGOOxNAL ENVIRONMENT OF THE SUWANNEE FORMATION The stnd\ material from the Teiramar Pit was collected either as perfectlv-present-d spt'cimeus looseK embed- ded in a residual unconsolidated calcilntite or as speci- mens emiiedded in blocks of a dense, partialK-silicified limestone. These specimens were often remoxed in a fragmentary state. Judging from the biotic assemblage collected in both lithofacies. the entire area represented a shallow (less than 5m deptli), quiet-water carbonate lagoon. This lagoon formed behind a still-mistudied cor- al reef tract that had developi'il farther to the west. E\- idence for this unmapped Oligocene ri'cf tract, including the presence of massixe coral heads and reef niollnsks such as the bixaKes Lima luilcitsis Dall. 191fi and Clianui sp. and the stromboidean gastropod Teirlielliim luniaii- docnsis Mansfield. 1937. has been nnco\ered in quarries in central Pasco and Hernando (bounties (Mansfield, 19.37). The Terramar lagoonal emironment, existing be- txveen this reef tract and the Oligocene western coast of Florida, contained four main substrate txpes: 1. bio- herms composed of Sti/lnpliorn corals: 2. deeper lagoon- al, open-bottom areas; 3. sea grass beds, and: 4. \er\ shallow-water o\ster beds and intertidal mud flats. Of the four biotopes that existed within the Terramar back-reef lagoon, that of the Sti/lopliora coral bioherms xvas apparentK the most extensive. Composed almost entirelv ol the small branching coral Sti/Iophora iiiiiiu- tissiiiui N'autilian, 1900 these bioherms formed denscK iiitermeshed beds that sheltered a wide \arietx' of gas- tropods. Manv blocks of limestone examined contained literallv hundreds of small pieces of this branching coral antl loose fragments were the most aiiundant component of the residuum facies. As exidenced bv haxing been collected in silii in Stt/h>phora-r\L-\i limestone blocks, a mmiber of Suwannee endemic niollnsks appear to ha\e been associated with these biolierms. Some ot the most characteristic gastropods inclndi- Calliastoma siliciitiiiti Mansfield, 1937 (figure 19) Cypriicarhis kcmlrcwi n. sp. (figures 22, 23), Salmosteira .siiwannccnsis n. sp. (figures 33, 34), \'ri\uiii Mtuiiiinccnsis n. sp. (figures 48, 49), Fal- silipia kviidrcwi n. sp. (figures 37, 38), and Conus (As- prcU(i) kciiilrcwi n. sp. (figure 72). Sexeral of the large Suwannee cerithiids. including Ccsliniircritliiitin liroohs- lillciisis (.Mansfield. 1937)(figures 3. 10) Ccsfiintn-ri- lliiiiiii pdscot'iisi^ (Mansfield, 1937)(figure 2), and Pii\- macerithuun prisma n. sp. (figures r>, fi, 11) were also associated with the Sti/lophora bioherms. Iiiterestinglv, sexeral specimens of Paclujcrommiiuit mansficldi n. sp. (figure 30) x\cre also collected in Sti/lophora-rich lime- stone, possibly indicating that this naticid may haxe lixed on, or in close proximitx to, the coral bioherms. The rare muricid Foiricria (Diillimiircx) nifinipicolus (Dall. I91R) (figure 31), wliii'h was prexiousK' known onlx' as an in- ternal mold from the Flint River Formation, was also found to be associated xvith this biotope. Small heads of the star corals Montastrea haiiihridf^cnsis (\'auglian. 1919) and Antiffiastrea siliccusis X'anghan. 1919 were also present, although uncommoiiK. throughout the Stt/- Icphora bioherms. Interspi'ised betxxT^en the coral bioherms in deepi-r water (approximatelx' 4-5 m) areas were large patches of open, fine-grained carbonate sand bottom. This biotope supported a rich gastropod fauna of xx'hich the Sux\aunee endemics Oliva (Omot^i/mna) hrotih.siillcnsis (Mansfield, 1937) (figure fi5) and A/.)/r(//c/ Iniwciiae (Mansfield. 1937) (figure 13) were the most abundant. A large \'icksbnrg- aged marginellid fauna also was associated xsith this bi- otope, and included species such as Persicida dackcn/i n. sp. (figures 52,53), Pninitm cricac n. sp. (figure fi4). Pniuiim jcssicac n. sp. (figures 56. 57), and Pniniun saii- dnic n.sp. (figures 58, 59). The strombid OiihauUix lirr- iiandofii'iis- Mansfield, 1937 (figures 20, 21) which is also present in the coral reef enxironments of the Flint Rixcr Formation, was abundant on these open bottom areas, fixing in large colonies much like Strnmlms s.s. species do in the Recent. Sex'eral of the new Suwannee species described here, such as Pachi/rnimittiiim dalli n. sp. (fig- ure 26). Sj)inifnl polhnisi.s n. sp. (figure 18). Ccrithitim insulatuin Dall, 1916, (fig- ure 12) and Modulus liveoakensis Mansfield. 1937 (fig- ure 16), and a large fauna of small, TlialassiaAovin^ mar- ginellitls such as Dcutiinan^o dalli n, sp. (figures 62,63), Hi/aliiKi silicifiuvia (Dall. 1916)(figure 51), Pcrsiculn uiacncili n, sp. (figures 54,55), and Fcrsicula suuainiccii- sis n. sp. (figures 60,61). Living within the grass beds were larger gastropods such as Cerithioclava culcxtilc (Dall. 1916)(figure 9), the Suwannee endemic Tiirhiiirlla suutninciisis (Mansfield, 1937)(figure 42, 43), and a host of small bivalves, priinarilv Cdtione haiul>rid<^cii.sis Dall, 1916, Pliacoidcs hcniandoensis Mansfield, 1937, and Ghjci/iiicris suwauucusis Mansfield, 1937. .\ living ana- logue to the Suwaimee grass Ix'd connnunitv is seen in the Recent along Roatan Island, Honduras, where spe- cies of Ccritliioclaia. Turhhulla. Astraca. Modulus. Pii- sirula. Hi/aliiia. and Phacoidvs all co-oi'cur in 'Phalassia beds. The unusual Suwannee endeniii.- bubble shell, .S'/(- uauiu:scaj)l>a liudac n. sp. (figures 70,71 ). max also have been associated with the sea grass beds. Faimal evidence also points to the existence of shallow intertidal mud Hat enviromnents within the Terramar la- goon system. This is seen in the presence of a tlistinctive potamidid fauna composed of tA\'o species of TelcscDpi- uui, [T. hlrukwatereusis (Mansfii'Id. 1937) figure 15. and T Itcniandiicusis (Mansfield. 1937) figure' S]. and Pi/ra- zisiniis kciidrcwi n. sp. (figmv 14). bi the Recent bido- Mahusian region, the amphibious genus I'lLscopiuui lives on tin' highest mud Hat areas near mangroves and is ex^50secl to air lor long periods of time. Often living along with Telescopiuui on these exposed flats is the ge- nus Tcrchralia. which is closelv related to the Floridian endemic genus Pipazisiuus and probabK occupies the same ecological nic-he as did its extinct New World coun- teqjart. The ovster Ostrca jjaroxis (Lesueur) Dockeiy, 1984 is conmionlv encountered in large masses in some blocks of limestone, indicating that small ovster bars also grew on these intertidal nuid flats. The melongenid Mipislira cn/ssicdinula (Conrad. 1848)(figures 35, 36) and the muricid Plcii/uctus propeposti (Mansfield, 1937)( figure 64) were associated with these ovster bars and were probably the major predators on Ostrca pa- roxis. A Recent analogue to this Oligocene mud flat and ovster bar enviromnent is seen in northwestern Australia along the .Aralura Sea. where Ptcn/uotus iPhcdualli). Tcrchralia. and Telescopiuui species occur togetiier on nmd flats and oyster bars near mangrove forests. systematic; section The holotvpes ol the new species described here are deposited in tlie lossil invfrtebrate collection of the Florida Museum of Natural Historv. University of Flor- ida, Gainesville, Florida, and bear UF numbers. Para- tvpes are depositeil in the research collection of the au- thor, as is a voucher collection of each species collected at the Terramar site. All specimens are siliceous pseu- domoiphs. .\ vouciier collection is also deposited at the Cu'aves Museum of Archaeology and Natural Iliston. Dania. Florida. Class Gastropoda Superfamilv Trochoidea Familv Turbinidae Suiifamily Astraeinae Genus Astraca Rikling. 1798 Subgenus Astraliuiu Link. 1807 Astraca (A.straliuut) jxilki'itsis ni'w species (figure 18) Description: Shell small for gemis and suligenns. con- ical, witli elevated spire whorls: suture indistinct; pe- riphen of body whorl flattened, bladelike, romided. with faint serrations; peripheries ol earlv wiiorls and spire whorls edged with 15-20 small, roimded knobs: whorls strongly ornaniented with 5 strong, beaded spiral eords; appro.xiniatelv one-halt of spire vvliorls). length 45 nun. Itvpc iit the new genus Pri.\iii(ici htliimiil 7. Sciiiiiciidtiiis iiicnfliafi'iitis MacNeil. 1984, length 15 mm. 8. Tricscitpiiiiit luriuniiliuiisis (Mansfield. 1937). length 4VJ nun. 9. d ritliiochica cutcxlih- (Dall, 1916), leni^tli 57 nnn. 10. Ccstuiiiiiiitluiiin l>niiik\i illnisis i Mansfield. 19.37). heavilv-sculptnrt'd variant, length 29 nun. H. Pris- iiKicirithiuin prisma Petnch, n.sp.. paratvpe (spire vvliorls. missing part of body wliorl ), length 19 mm. 12. Ccrithium iTlicriciuin) insulainrn Dall. 1916. length 22 nnn. 13. Apiciila hmccnac (Mansfield, 1937), length 39 mm. 14. Pt/razisiiuis kciulrcici Petnch, n.sp., Iiolotvpe (UF 75977). length 62 nnn. 15. Telescopiuui hhiekieatcrensis (Mansfield. 1937), length 43 nnn. 16. Modulus live- Odkensis Mansfield, 1937. height 15 nnn. 17. S<'iiiii(iiaiJ,us silieiion (Dall. 1916). length 15 nnn. 18. Asfrrien lAstmlium) polkensis Petncli, n.sp.. Iiolotvpe (UF 7.5976). lieii^lit I I iniii 19. ('ulliosUniiii silientiou Mansfield, 1937. lirit;lit 14 luiii. Page 126 THE NAUTILUS. N'ol. 110. No. 4 I to 3 c'.xtrc'tiieK fiiic tlircadlikc sccoiulan spiral cords present between 5 main coids; hasi' oi shell Hattened, ornanienteil witii 10 strong spiral cords: nnihilical region sliglitK' depressed. Material examined: 1 1( )L()TVFE— Height 11 mm. diameter (incomplete) 13 mm. in hack-reef lagoonal fa- des of the Sii\\'annee Formation. Tcrramar Pit, western- most Polk Count\, Florida, UF 75976. Etv'mologj': Named for Polk Countx. Florida, localit) ol the Tcrramar Pit. Discussion: This new species is the first A.striica known from the Eastern North American Oligocene. Of the kncmii Cenozoic American A.stralinin species, A. polkcnsis is closest to the late Eocene (Jackson Stage) Astraea (Astralium) withlacoochcnsis Palmer, 1953 from the Inglis Formation of the Floridian Oeala Group. The new species differs from this prol)aiile Eocene ancestor in ha\ing a smaller shi'll with a Hatter, less trochitorm shape, stronger spiral scnlptnri' on the whorls, and a re- duced, nonprominent peripheral keel. Snperiamilv Cerithioidea Faniilv Potamididae Genus Pt/razisinii.s Heilprin, 1SS6 Pt/razisimis kciulrcivi new species (figure 14) Description: Shell of average size for genus, elongat- ed, cerithiform; whorls ornamented with large, promi- nent. e\enl\ -spaced knobs; knobs distinctly angled at shoulder. a\eraging 10 per whorl; whorls scnlptureil with 5 large, prominent, raised spiral cords; base ot shell flat- tened; suture impressed, slightly canaliculate; apertural region of unique holotvpe broken, so exact natine o( adult lip is not known. Material examined: HOLOTYFE — Length 62 nun, width 28 nun, in nnid flat and ovster bar (acies oi the Snwamiee Formation, Terramar Pit, westernmost Polk County. Florida, UF 75977. Etvinologv': Named for Mr. Eric Kendrew ol N'alrico. Florida, who coll(>c'ted the holotvpe in the Terrainiu' l^it. Discussion: Pi/riizi'-iiniis koitlnu i is the olilcst-known member of its endemic Floridian gemrs. The ni'w spe- cies is most similar to the A(|nitaniau Miocene P. cor- imttis (Ilcilprin. 1.SS6) from the Tampa Nh'mberof the Arcadia Formation, but diliers in being ;i narrower more elongated shell with more numerous knobs per whorl. in having an angled shoulder area and angled knobs. ;uid in having a much more scnlptuicd shell, with (i\c dis- tiTict spiral cords. Familv Cerithiidae Gemis ('/■•ihniiccrilliiiiin new genus Diagn<)sis: ('eritli shells ol average-to-large size (25- 45 nun), slender and elongated: sutures indented, pio- dncing tnrreteil spire whorls: shell stiilptnre character- istically composed oi hvo distinct sections, with early whorls ha\ing a reticulate sculpturi' composed oi strong, oiten beaded, spiral cords intersected In longitudinal ribs, and with tlie later whorls and bod\' whorl being smootli, with fi\'e or six \ei-v large, ver\' prominent belt- like spiral cords: on adults, belt-like cords e.xtend beyond edge of lip, producing strong dentate, serrated appear- ance (shown here in figure 2); siphonal canal short, stnb- In-. Ty|Je species: Ccnth'uiiii pascoerms Mansfield, 1937, Suwannee Formation. earl\ Oligoceue of Florida (figure 2). Other species in Cestumcerithitim: Ccritliiiiin brooksiillctisis Nhiusfield. 1937 (figures 3.10), Cerithiiini liveoakciisi.s Mansfield, 1937 (figure 4). and Ccritliiiiin siiwannccnsis Mansfield, 1937. all from the Suwannee Formation. I'arK Oligoceue oi Florida, and Cfiithiinn vd'^iiKilinti IDall, 1916, Flint Ri\er Formation. earK Oli- goceue of southern Georgia and .Alabama, and the Su- wannee Fornuition of Florida. Etymologv: A combination oi the Latinized Greek "ccstii.s" (belt) and "Ccritliiiiin" (little horn), pnuhiciug "belted little horn", in reference to the [ii-omiuent and characteristic cords on the bodv whorl. Discussion: With the exception of the Flint Ri\er and Suwannee (\:stiiiiiceiitliiiiiii vaginatiinr all other knowni species oi this new genus ha\e been iound only in the Floridian Oligoceue. Here, tlie\' appear to luiM' been closelv associated v\ith the extensive coral reels and coral bioherms that dominated Latdorfian western Florida. Four oi'the fi\e known species are restricteil to tlie Su- wann(X' Formation and can be considered classic index fossils lor the Floridian earK Oligoceue. The genus aji- pears to ha\e been restricted to the Oligoceue. as no known members ha\i' been collected in the o\'erl\ing Tampa Member of the Arcadia Formation (Aquitaniau Miocene) or irom the underKing Eocene formations. The amount ;md degree of the characteristic l)elt-like corded sculpture appears to be \ariable. sometimes co\- eriTig onK the hist two whorls and at other times co\- ering most ol the kiter whorls ol members oi the same species. This is particnhirK noticeable oti the t\pe spe- cies, Ccstiiincciitliiiiin pascocn.sis. which can be hea\il\- belted (like the holohpe. Mansfield. 1937. plate 7, figure 4) or relativcK smooth, with onk' the bod\ whorl being belted (like the specimen illustnited here in figure 2, or like Mansfield's "Ccrilliiiiin s]i. afi'. iii}uiiiiniii iiiiiii\fiildi new species (figure 30) Description: Shell of ;i\crage size for genus, inflated, globose, with lounded bod\ whorl: spire protracted, sca- lariform; spire whorls relatively straight-sided; sutiu'es higliK' indented, producing narrow, \en' tleepK' canalic- ulate subsutnr:il border; edge of canaliculate bolder Page 128 THE NAUTILUS, Vol. 1 10. NO. 4 Figures 2()-34. Strombidae, Cvpraeidae, Ficidae, Ca.ssidae, Naticidae, Muricidac, and Hnccinidai- fniin llic .Siiwaiiiirc liack-rcei lagoons. 20,21. Ortluiuinx Iwrnamlornsis Mansfield, 1937. lenf^lh 70 nnn. 22,23. Cijprdcorlns kcndnui rclucli. n.sp., liolotApe (UF 75981), length 26 mm. 24. Fiats mississippicnsi.s Cdniad, IS4S. lengtli 17 mm dragmentan ). 2.5. Amptillinopsis fiintcnsis (Mansfield, 1937), lengtli 27 mm. 26. Pticliiicroniiiiinm iliilli Petucli, n.sp., Iiolotype (UF 75979), Icngtli 1^ imti. 27. Phnlium awlatura (Conrad, 1848), lengtli 29 mm. 28. ricnjnolns piopeposti (Mansfield, 1937), length 35 mm. 29. Tdlitijpliis iiii^sissippicnsis E. J. Petiicli, 1997 Faec 129 liiij;lil\ raised, proiluciiig sliarp, tlistiiict (.■aiiiia: iipcitiirc (i\al; Tio uinhilicMis prest-nt. Material examined: HOLOTYPE — Length 27 imii. widtli 21 iiini, ill hack-rt'el lagoonal iacics (in association with Sti/lophora liioherins?) of the Suwannee Forma- tion. Terraniar Pit. we.sternniost Polk (Jonnt\'. Florida. UF759S(): P.\R,ATYPE— length 24 nnn. same locality as liolot\])(', in the research collection ol the author. EtMiiologv: Named in honor oi Wendell (-'. Mans- field, the pioneer paleonialacologist who first descrihed the Suwannee gastropod ianna. Discussion: This new Pachifcroiiiiiiiiiiii closeK resein- hles, and is probabK' ancestral to. /' jiiuUland (Dall. 1892) from the .-Xquitanian Miocene Tampa .Member of the Arcadia Formation. The new species differs from P. jioriddiui in ha\ing a less-inflated shell with straighter sides to the whorls, a more protracted spire, and a much wider and deeper subsntmal channel. ,\t ])resent, P luaiisfichli is known onl\' from tlu- coral leef lagoonal facies ol the Suwannee Formation. SuperfamiK' C\praeoidea Family Cyjjraeidae Subfamily Cypraeorbinae Genus Cypmeorbis Conrad. 18fi.5 Cyprncorbis kcmlreui new species (figures 22. 23) Description: Shell of a\erage size for genus, ovate, slightly subcviindrical: base flatteni'd: aperture narrow, widening slighth' toward anterior end, almost straight, recuning slightK at posterior end; labial teeth propor- tionalK' large, e.xtending onto base ol lip. 23 in munber: columellar teeth smaller than labial teeth. 21 in number: extremities reduced, projecting onlv slightK hom main shell bod\': spire and posterior extremits covered with broad, low callus: fossula poorK'-de\eloped. supported In single large tootli along anterior edge. Material examined: HOLOTYPE— Length 2rs mm. width IS nun, in back-reef lagoonal facies {Sti/lnpliurd bioherm) oi the Suwannee Formation, Terraniar Pit, westernmost Polk Countv; Florida, UF759S1; Parat\pe- lengtli 26 mm, same localit\' as holotspe, in the research collection of the author. Etymology: Named for Mr. Eric Kendrew of X'ahico, F"lorida, who collected the holotvpe in the Terramar Pit. Discussion: Ct/praeorbis kciidnwi. one ol the last species of the Cypraeorbis Hneage, is most similar to C. vcnfiipotcns (Cossmann. 1903) from the [ackson Eocene .Moodvs Branch I'^ormation ol .Mississippi. The new s[)e- cies differs honi this older cowrie in ha\ing a iimch more c\iindrical, less iTiilati'd shell witii straighter sides, more rountled. less projecting e.xtremities. a straighter. less recin"\ed aperture, and more munerous and finer apertural tei'th. Tlie \'icksburg Oligocene cowrie listed as "Cypmeorbis all. i'. icutripotens (Cossni.in. 1903)" by MacNeil and Docken (19S4, plate 16, figs. 1().12.]5) is veiv similar to C kciuln-ui, !)ut iliffers in being a more inflated, rounded shell with a slightlv more recuned ap- ertiue. F'urther stniK' ma\ show that the Mississippi shell (from the Mint Springs Foiination) belongs to a north- ern iioimlation ol (' kciKbiui. SuperfamiK Hneciuoidea FaiiiiK' Busxcouidae SubfamiK Busx'couiuae Genus Spinifiili^iir Petnch. 1994 Sjiiiiifiil^iir ^cimiiuliilniii new species (figure 471 Description: Shell ol average size for genus, with dis- tinctly sloping shoulders and high pagodiform spire; edge of shoulder shaipiv angled, bordered bv promi- nent, thick, undulating carina; undulations on carina pro- duce small, rounded, evenlv-spaced knobs, numbering between 14 antl IS per whorl; suture highly indented, causing carina on previous whorl to overhang subse- quent whorl; sloping spire whorls heavilv ornamented with 8 strong spiral threads between which are interca- lated verv fine secondan' threads; bodv whorl (portion exposed in matrix, just anterior of shoulder carina of ho- lotvpe) ornamented with 5 (extrapolated from holohpe) ven' large, thick, beaded spiral cords: aperture and si- phonal canal unknown. Material examined: 11()L()TYPE — Length (frag- mentan'. missing portion oi body whorl and siphonal ca- nal) 24 mm. width 16 nun, in back-reef lagoonal facies of the Suwannee I'ormation, Terramar Pit, westernmost Polk Colmt^■, Florida, UF75928: PARATY'PE-spire only, width 16 nun, same localit\ as holotvpe, in research col- lection of the author. Etymology: "Gennned". in reference to tlie charac- teiistic gemlike knobs on tlie shoulder c;nina. Discussion: Sj)iiiijiil^iir ij^cininulatnin. although known onK honi Iragnieutan specimens, is so distinc- tive that its lormal description seems warranted at this time. The new species is similar to slope-shouldered var- iants ol the N'icksburg Oligocene Spinifulffir spini^cr (Conrad, 1848) (such as the tmv illustrated bv MacNeil (Gertiiiaiiii. 1969), lerigtli 24 inni. 30. Pnrlnirniiiiiiiiuiii iiumsficUli Petnch n.sp,. holotvpe (UF 75980), l<-iigtli 27 iiiiii. 31. Pcuriiria (Didlii)uuTx) nifinipicolus (Dall. 1916), tragniciitaiy spccinieii, length 21 mm (missing portion of bodv whorl and siplional canal). 32. Clui-«r(u\ stctopus (de Gregorio, 1890), length 32 mm. 33,34. Solenosteira suutnutecnsis Petucli, n.sp.. holotvpe (UF 75983). leiiirtli 27 mm. Page 130 THE NAUTILUS. \ol. 110. No. 4 m^i Figure 35,36. IciiHtli 42,43. Iciiiltli s 3.5-49. Mcloiigeriidae, BiLsycoiiidue, X'olutichie, Volutoniitridac. uiid Turhim-llidae from the Siiwiuiiiff back-reef lagoons. Mijrislica cra\sironuita (Conrad. 1848). length 49 nnn. 37,38. luilsili/ria kciidmvi Petuch, n.sp.. hoIot>pe {VF T59S5), 37 nnn. 39,40. FuUihjriu munsfiddi (l)all, 1916), length 32 mm. 41. Coiwiiiitm cirnulatn Dockeiy. 1984, length 21 nnn. Tiirhinclla suwtiniicnsis (Mansfield, 1937), length 107 nnn. 44,4.5. Conoiiiitm kviulivui Petneh. n.sp.. holot\7x- (UF 75986), 14 inni. 46. Clriiiolitlics vickshuixen.sis (Conrad, 1848). length S9 nnn. 47. Spinif'uU^ur Hfiniiiiildtiiin I'etneh. n.sp.. holot\pe E. J. Petuch, 1997 Paec 131 .nul Doeken, 19^4. plate 54. fifj;.H'. Iiut dirirrs in lae-king distinct spines, having, instead, low ronnded knobs ar- ranged in an undulating pattern. Spinijidour ^riiiiuiila- tint) also differs from S. spinii^cr. and an\- other known Sphiifulfiiir spi'cies. in being liea\ily sculptured with wide, thick spiral cords on the boiK whorl. At present, this highlv ornate species is known onK lidin the Su- wannee Formation. FamiK l-^uccinidae SublamiK Buccininae Genus SolcniKtcira Dall. 1890 Solcnostririi sinLdiiiiccn.'iis new species ( figures 33. 34) Description: Shell of average size for genus, liroadiv fusiform, inflated, with liigh. protracted spire; shoulder angled, spire whorls sloping; bodv whorl with 7-S \er\ large, rounded, e\enl\-spaced axial folds; folds become largest at shoulder producing large, rounded knobs at shoulder periphen'; sutme prominent uutlulating as it overlaps .slioulder knobs of spire whorls; bodv whorl or- namented with 12 large, raised spiral cords; spire whorls ornamented with 5 spiral cords; siphonal canal well-tle- veloped, broad, ornamented with 3 large spiral cords; pseudonmbilicus present, partially covered by parietal shield of siphonal canal: aperture wide, oval in shape. Material examined: HOLOTYPE — Length 27 nmi. width f7 mm, in back-reef lagoonal facies (Sti/lo))lu>r(i bioherm) of the Suwannee Formation, Terramar Pit, westernmost Polk Count), Florida. UF75984; PARA- T\TE — length 25 mm. same localitv' as holotv|ie. in the research collection of th ithor Discussion: Solowsteiiyi fnnvdiiiiciiisis is the oldest- known member (jf its genus to be found in Eastern North America. The new species is most similar to S. iiiiiniiilii Dall. 1890 from the Aquitanian Miocene Tam- pa Member of the Arcadia Formation, but differs in iiav- ing a lower, less protracted spire, in ha\ing a nnich coarser shell sculpture composed of strong spiral cords, and in having more numerous and smaller axial folds and shoulder knobs. Superfamilv Volutoidea Family Tnrbinellidae Subfamily \'asinae Genus \'asiiin Roding. 1798 Vasiiin siiurinnceusis new species (figures 48. 49) Description: Shell small for genus, with broad, shaip- Iv-angled shoulder; spire stepped, proportionally low. with spire whorls slightK sloping; suture indented, bor- dered by broad, nndnlatiug subsutnral collars; slioulder ornamented with 10 large, evenly-spaced, open spines: body whorl sculptiued with 8 large primarx' spiral i-ords: smaller secondan spiral cords present betxveeu some priman,- cords: spire whorls sculptured with 3 largi' spiral cords; siphonal canal well tlexcloped. ornamented with single row of veix' large, opi-n spines tliat correspond to the spacing of the shoulder spines: siphonal canal sculp- tured with 5 large spiral cords, 2 posterior of row of spines and 3 anterioi of spines; aperture open, flaring; columella with 3 \cr\ huge, promini'iit plications; on lio- lotx'jje, a louitli Ncix laint plication is present between anterior ;md media! plic;itious. Material examined: HOLOTYPE— Length 39 nun, wiiltli (tragmiMitaix, missing last third ol body whorl) 23 nun. in back-reef lagoonal facies {Sti/Iopliora bioherm) of the Suwannee Formation, Terramar Pit. westernmost Polk Countx-. Florida UF75984. Etymology: Florida. Named for the Suwannee Formation of Discussion: \'asiiiu siiwnnncensis is the first known \ase shell to be collected in the Eastern American Oli- gocene. The new species is most similar to \' humcro- smu Vaughan. 189fS from the Jackson Eocene Moodys Branch Formation, but differs in having a lower, flatter spire, in having more luunerous primary spiral cords on the body whorl, in having three veiy large and promi- nent spiral cords on tlii' spire whorls, and in having a larger, much more prominent, and more elevated snb- sutural collar. The new species is also similar to V. sith- capitclhnii Heilprin. 188fS from the ,\rti(in nl n.sp.. holotxpe (UF 7549S4), length oVJ mm. l)()(l\ wliiui anil siphonal canal), length 24 nmi. 48,49. Viisiiin Miuciniwi'nsis Petuch, Pajic 132 THE NAUTILUS, Vol. 110, No. 4 Fij;uri's 50-72. Mitridac, Mariiincllidac. Olividac. C.'Diiidac, Tnriidac, and (Aliclniidac Iroiii the Suwannee liaek-reef lat;oons. 50. i-'nsimilra coiujuislu iCJonrad, 1S4S), lenpe (UK 75988), leni;th 14 nnii. 54,55. Pcrsindti inariuili I'etneh. n.s|)., holotNpe lUF 75989), lent;th 11 nmi. 5G,57. Pnmum jcssicac Petucli, n.sp., h()l()t\pe (UK 75992), lenijtli Hi i 58,.59. I'ninnm sandnic relneh, n.sp.. iiolotxpe (V¥ 75993), length IS mm. 00,61. I'l-rsiruhi simnnncrn.si.t Petneh, n.sp., holotvpe {UK 75990) length 9 nun. ()2,()3. Dcntiiniin^o diilli E. J. Petucli. 1997 Paec 1:33 3 lai"m' s[)iral cords aromul aiitciiiii' ciul; spire wlinrls ornaiiu'iitfd (particularK' ovi last wliorji with siiii^lc, lariat' spiral cord: aperture wide, Haiiiiu;: colmnclla with S large, e\enlv-spaced, headed plications: protoconch pro- poitioualK' ven' sinall, composed ol 2 narrow, smooth whorls. Material examined: HOLOTVFE — Length 37 mm. witlth 21 nun. in hack-reef lagoonal facies (Stt/lophora hioherni) oi tlie Suwannee Formation, Terramar Pit. westernmost Polk Countw Florida, UF759.S5: PARA- TYPE — length 39 mm, same locality as holot\pe, in the research collection oi the author. Et\Tnolog\': Named for Mr. Eric Kendrew of \'alrico, Florida, who collected the holot\]ie at the Terramar Pit. Discussion: FdlsiUjiia koiihvwi occurs together in the Ti-rramar fauna with F iiuin.^fiilili (l^all. 1916) (figures 39, 40). hut can he readiK distinguished h'om its hetter- kuown congener in having a consisteutK broaden more inflated, and less elongated shell, with prominent spine- like shoulder knobs, and in biiug a smoother shell that lacks the spiral cord sculpture on tlu' bod\ whorl. Like- wise, two Fahihjria species occm' togt'ther in the ri'efal eu\ironments of the Burdigalian Miocene C^hipola For- mation. Of these t^\•o, F. uuinsfuhli appears to be an- cestral to the smooth-shouldered F. anoptos Hoerle and X'okes, 1978 while F. kcndrewi appears to be ancestral to tlie knobbed-shouldered F. pijcnopleura (Gardner, 1937). The smooth-shouldered and lu'a\il\-corded F. inusiciiKi iHeilprin. ISSfi) from the .\(|uitanian Miocene Tampa Member of the .\rcadia Formation is \erv similar to F iiuinsfn'hli and is the most probable descendant species. \o Falsilijria with strong shouldi'r knobs has \et been tounti in the Tampa fauna. At present, Ftilsili/hii kciulrcwi is known onl\- from the Terramar Stylophora biohernis. FamiK N'olntomitridae Genus Couoiuitra Conrad. lSrS5 Couoinitra keiuliTwi new species (figures 44, 45) with spire; Description: Shell small tor gi'uus. biconic shaqjiv-angled shoulder and proportiouallv low- anterior t^^■o-tllirds of bod\ wiiorl smooth, unoruameut- ed; spire whorls and posterior one-third ol bod\ uhoii iiea\il\' sculptureil with large. e\euK spacetl rib-like pli- cations, averaging 20 per whorl; aperture narrow; colu- mell.i with 5 large plications, with the largest at posterior i']]d. becoming smaller toward anterior end: anterior- most tip of siphonal iuca ornaTuented with 4-5 im- pressed spiral thrciids. Material examined: HOLOT^'PE — Length 14 nun. width 8 mm, in b;ick-rei'f lagoonal facies of the Suwan- nee Formation, Teiramar Pit, westernmost Polk (>)unt\, Florida, UF759Sfi; PAH.AT>TES— lengths 14 nun and 17 nun. same loc'alit\ as holot\'pe, in the research col- k-ction ol the .iiithoi'. Et%'niologv: Named lor Mr. Eric Kendrew of \alrico, Floiitla. who collected the holot\pe at the Terramar Pit. Discussion: This distinctive new Suwannee Conoiiii- tni species is most similar to the svmpatrie and wide- spread \icksburgian C cniudata Docken'. 1984. but dif- fers in having a nuicli lowfr spire vvitli a sliaqilv -angled shoulder and cone-shapetl appearance. Couomitra ken- (Irewi also differs from C. cniudata in having shoulder and spire plications that arc proportionally larger and fewer in number. In these characters, the nevy species is also similar to C. inodesia Docken, 1984 from the \icksburg-ageil Forest Hill Formation of Mississippi, but differs in having a much lower spire and much more shaiplv-angled slionldei'. Family Margint'llidae Subfamily Cdabellinae Genus Dentimargo Cossmann. 1899 Dcntimargo dalli nevy species (figures 62, 63) Description: Shell ol .i\er;ige length lor genus, slen- der, witli high, elevated spire; suture indented, protluc- ing slightlv stepped spire wiiorls; shoulder of bodv whorl only slightly angled; aperture narrow at posterior end. rapidly expanding and lu'coming flaring and opi'u at an- terior end; columella with 4 large plications; labnun of adult tliickened ;nid (.iillused. with smooth inner edge. Material examined: IIOLOTYPE— Length 15 nun, width 8 mm, in b;ick-reel lagoonal facies (sea grass bed) of the Suwannei' Formation, Terramar Pit, vyesterumost Polk Count\, Florida, UF 75987: PARATYPES— 2 spec- imens, both 14 mm, in length, same localitv as holotvpe. in the research collection o( the auti lor EtMnoIogv: Named in honor ol W illiam Heaiev Dall, the foremost pioneer Floridi;ui paleonialacologist. Discussion: Dciitiinargn dalli is tlie oldest known member of its genus to be found in the Floridian Ce- nozoic. The new siiecii'S is most similar to D. vlenantida Petiicli. 11, s[).. liiilotvpe (UF 75987), leiigtli 15 nun: tsis I^ockeiy, !9S4, length 10 iiini. 67. Piciintfii.sia hrooksvillensis (Mansfield, 1937), length 26 iniii. 68. Piciirofnsia doiiliiif:,! Petucli, n.sp., holotvpe (UF 75995), length 26 mm, 69. Conorbis porcellanus (Conrad, 1S4S), length 36 nini. 70. Siiudiiitcscdphii lindac Petucli, n,sp., holotvpe (UF 75996), lengtli 17 nun (t\pe of the new genus Simaniifscaphfi). 71. Snwaniic^cdplia lindiic Petuch, n.sp., view of spire of iioloKpe; 72. Conns {A.sjircll(i) kcndrewi Petucli, n.sp,, lioldtvpe (UF 75994). length 35 iniii. Paw 134 THE NAUTILUS. \ol. 110. No. 4 (Dall, 1890) iioni llic .\(|uitaiiiaii Miocene Tampa Meiii- l)cr of the Arcadia Foniiation, hut (litU'is in being a larg- er more slender and elongated .shell with a more sloping shonlder and in haxing a nuich more piotracted. dis- tiiK'tK scalariform spire. At present. Dcidiinan^o dalli is known onK' irom the Snwannee Formation. Genus Pcrsiculu Scinnnaeher 1S17 Persiaila dockcn/i new species (figures 52,53) Description: Shell large for genus, oxoid, inflated, slightK' Hatteu(xl dorso-ventralK'; posterior extremities well-developed, greatly projecting, recun-ed strongly to- ward right side of shell; aperture narrow, widening slightlv toward anterior end; posterior end of aperture recuned sharjilv toward right side of shell; spire whorls not \isible. buried in thick callus; columella with 5 large plications: lahrum thickened, smooth along inner edge; shaip-edged callus preseTit along posterior of columella, forming open canal in sharpK -recuned posterior end of apertme. Material examined: HOLOTYPE — Length 14 mm, width 9 mm, in back-reef lagoonal facies (open sand bot- tom) of the Suwannee Formation. Terramar Pit, west- ermuost Polk C'ount\-, Florida, iJV 75988. Etymology: Named in honor of Dr. David T. Dockery ill, of the Mississippi State Bureau of Ck-olog)', in rec- ognition of his extensive and important contributions to the systematics of the southeastern U.S. Paleogene mol- lusks. Discussion: Of the known Paleogene anil earlv Neo- gene Prrsirula species, P. dochcn/i is Tuost similar to P. proamvida (Gardner, 1937) from the Buriligalian Mio- cene Oak Grove Formation of southern Georgia and Tiorthwestern Florida. The new Oligocenc" species differs Irom this possible Miocene descendant in biing a larger, more elongated shell, in ha\ ing a fai' more recun-ed pos- terior end of the apeitnre, and in having luuch better developed, more conspiiiions, and more-projecting pos- terior extremities. At present. I'lisicula ditckcriji is know^n only from the Suwannee P'ormation. PcrsicuUi iiifintcili new species (figures 54. 55) Description: Shell ol a\er;ige si/e for genus, elonga- tely o\ate. with lelaliveU str;iight sides; extremities rouniled. onl\' laintK' produced, not projecting be\ond spire; aperture narrow throughout. slighlK recuived to right side of shell at posterior end; columella with 5 pli- cations; labrnm thickened, with smooth inner edge. Niil. kite of the U.S. Geological Sune\. in recognition ol his important eoTitributions to the s\stematics of the Vicksburg mollusks. Discussion: Pcrsiculu iiuiciicili is similar in appear- ance to P. Iwlensis (Dall, 1916) from the contempora- neous Flint Rixer Formation, but differs in Iwing a larg- er, more elongated shell with distinctk straighter sides. The new species is also similar to the \icksburgian P. virkshtir<^cnsis Docken-, 1984 from the Mint Springs Formation of Mississippi, but differs in being a umch larger, more elongated shell with a lower and more rounded spire. Persicnlfi .'iiiwanuccnsis new species (figures 60. 61) Description: Shell of a\erage size for genus, extreme- 1\ t'longated and cxlindrical. with straight sides; aperture narrow throughout, recuned to right side of shell at ex- treme posterior end; spire area flattened: posterior ex- tremitx on labial side better developed than posterior extremit)' of columellar side, projecting be\ond plane of spire; extreme posteriormost edge of columellar area with thin, sharp-edged blade, producing open posterior canal with labial projection; columella with 5 plications; labrum thickened, with \en fine crenulations along in- ner edge. Material examined: HOLOTYPE— Length 9 mm, width 5 miu. in back-reef lagoonal facies (sea grass beds?) of the Suwaimee Formation, Terramar Pit, west- ernmost Polk Gounty, Florida, UF75990; PARATYPE— length 10 nun, same locality as holotype, in the research collection of the ;iuthor Etvmolog>': Named for the Suwannee Formation of Florida. Discussion: Pcrsicula \iiu(initcciisis is the most slender ;iiid elongated of all known Oligocene and Miocene Pcr- .\icul(i species. This new Suwannee marginella is simiku' onl\ to the sxnipatric /' iiuioicili, but diffei's in being even narrower and more elongated, in liasing the pos- terior end of the ;iperture being more recuned, and in having the dislmrlK piiijectiug posterior labial extremit\'. Material examined: HOLOTYPE— Length llm. w idth 7 mm, in back-reel lagoonal facies (sea grass bed? ol the Snwannee Formation, Ternnnar Pit. westennnost Polk County Florida, UF 75989. Etymology: N;uned in honor of l)i'. I''. Stearns Mac- Genus Pniiniiii I leriiuannsen, 1852 Pill III I III criciii new sp(>cies (figure 641 Descripti«>n: Shell small for genus. eloTig;iteK o\ate, with high, pvramidal spire; shonlder slightK angled: ap- erture nairow ;il posterior end. expanding rapidly toward anterior end. becoming wide, flaring, and opi'ii; colu- mella distinctk arcuate, with 4 large plications; labrum ) thickeiieil. smontli aioii',; inner edge. Material examined: IK )LOTVPE— Length If mm, w idth 6 mm, in b.uk-ieii lagoonal facies (open sand bot- tom) of the Suwannee l'"oriiuitioii. Terramar Pit. west- Petuch, 1997 Page 135 tTiiiiiost Pt)Ik Counts. Florida. UF7.5yyi: PAR.Ji- TYPES — Lengtlis S nun and 9 mm. .same localit\' a.s lio- lot\pe, in till- resfatc'li LoIk'ctioTi ol tlic autlioi-. Etymolog^•: Named tor Ms. Erica Kendrew. danghter ol Mi-. Erie Kendrew. ol N'alrico, Florida. Discussion: Ol the three Fniiiiiiit species now known from tlie Suwannee Formation, P. cricac is tlie smallest and most slender in outline. In general size and shape. P. ericae is most similar to F iiifccta iDall, 1915) from the A([uitanian Miocene Tampa Meniher ol the Arcadia Formation, hut differs in heing a more slender shell with a w'i(k-r aperture. The new species mav he ancestral to till' radiation ol small, high-spired Pnuuim species lonnd in the' Tampa Member and including such distinctixc tiixa as P. infccta. P. impagina (Dall, 1915), P posti (Dall. 1915), andF inepta (Dall, 1890). The small un-named. high-spired PniniiDi illustrated bv MacNeil and Docken- (1984, plate 35, figures 4,5. as "Margiitclla sp.?'") Irom the Bvram Formation ol the Vicksburg C^roup is similar to P criaic and ma\' be conspecific. Pniituiii jcfisicdc new species (figures 5fi, 57) Description: Shell of avei'age size tor genus, cN'lindri- cal in form, with distinctly angled shoulder: spire high, protracted, slightlv scalaritorni; spire whorls and shoul- der of b()d\' whorl sloping; aperture narrow, wiilening onlv slightU" toward anterior end: columella straight, with 4 large plications: labrum (missing on holotspe, but pres- ent on parat\pe) slightK thickened, smooth along inner edge. Material examined: HOLOT^'PE — Length 1(S nnii. width 9 mm, in back-reef lagoonal iacies (open sand bot- tom) of the Suwannee Formation, Terramar Pit. west- ernmost Polk County. Florida. UF75992: PARA- TYPES — lengths 14 nnn and 11 nnn (fragmentan). same locality as holotvpe. in the research collection ol the author. Etvniologv: Named t(jr Ms. Jessica Kendrew, ilaugh- ter ot Mr. Eric Kendrew. of Vahico. Florida. Discussion: With its large size and high scalaritorm spire. P. jcssicac is the most easilv-recognizable ol the Suwannee Pninmn species. The new specii's is most similar to, and is possibly the ancestor of the ,\<|nitaniaii P silicata (Mansfield. 1937) from the Tampa member ol the .'\rcadia Fonriation. Pniniimjessicac dilfers Irom the early Miocene species in having a more inllated shell with a more-angled shoulder, and a highei-. more pro- tracted spire. Pniniim mndrnc new species (figures 58, 59) Description: Shell ol average size for genus, inllated. ovately subcvlindricak slionlder sli.upK -angled: spire proportionalK low : spire whorls slightK sloping: aperture narrow, widening slightly toward anterior end: columella straight, with 4 large plications: labnnn thickened, smooth along inner edge. Material examined: HOLOTYPE— Length IS nnn. width II iimi. in back-reel lagoonal Iacies (open sand bottom) ol tin' Suwannee Formation. Terramar Pit, westernmost Polk Countv, Florida. UF75993: PARA- TYPES — three specimens, lengths 13 mm, 15 nun. and 19 nnn, same locality as holotvpe. in the research col- lection ol the author. Etymology: Named lor Mrs. Sandra Kendrew. wile ol Mr. Eric Kendrew. ol X'ahico, Florida. Discussion: Pniiuiiii sandrav is very similar to, and is midoubtedlv the ancestor ol, the A(|nitanian Miocene P. gregaria (Dall. 1915) from the Tampa Member ol the Arcadia Formation. Both sliells liave tfie same large size, lovy spire, and narrow aperture. The nevy species dillers from its descendant, however, in being consistently a wider and more inllated shell. Pniniiin sanclrac appears to be the commonest marginellid in the Suwannee For- maticjn at Terramar. Superiamily Conoidea Family Conidae Genus C'())i(/s Linnaeus. 1758 Subgenus Aspnlla Schantuss, 1869 Conus (A.sprclla) kciidrcwi new species (figure 72) Description: Shell of average size for subgenus, elon- gated, tapering, with straight sides; spire flattened, with faintly stepped, slightlv projecting whorls; shoulder sharply-angled, boundeil bv thick, lidgelike carina; ca- rina present on spire whorls, producing depressed sca- lariform appearance; bodv wlmii heavily sculptured with (on unique holotvpe) 24 large, evenly-spaced, highly- raised spiral cords: spire whorls ornamented witli 5 fine, evenlv-spaced spiral thri'ads: aperture straight, narrow. Material examined: HOLOTYPE — Length 35 mm, width 21 mm. in b;ick-reef lagoonal facies (Stt/Iopliora bioherm) ol the Suwannee Formation, Terramar Pit. westernmost Polk ( loinitv. Florida. UF 75994. Etymology: Named lor Mr. Eric Kendrew ol X'alrico, Florida, who collected the holotvpe at the Terramar Pit. Discussion: ('.(iiiiis kciiiln'ui, with its e.xtremelv heavy corded bodv sculpture, wide shoulder carina, and de- pressed scalarilorm spire, is the most distinctive cone shell known from the Eastern American Oligocene. The only contemporaneous cone that even remotely resem- bles this new species is the svnipatric Suwannee and Flint River C. (Lcptoronus?) cookci Dall. 1916. Conns kiiiilirui differs from this more wide-ranging species in having a much coarser and stronger spiral cord sculp- ture, in having a distinctiv stepped spire, and in having Pajie 136 THE NAUTILUS. \<.l. 110, No. 4 a nroiiiincnt shoulder faiiua tliat fxteuds Ijcyoiui the IkkK' wliorl outline. Famil\ Tiirridae Suhlainil\- Turriculinae CJenu.s ricurofiisid de (Jrcsjorio. 1890 Ph'tirofiisia (hncliiif^i new .specie.s (figure 6S> Description: Shell of average size for genus, elonga- telv rusiloriii. with high, protracted, .scalarifonii spirc", roiiiKled hod\, and tliiek. elongated siplional canal; shoulder slightK' angled: suture indented; hotK' whorl or- namented with 9 large, thick spiral cords that lieconie thinner toward anterior; siplional canal oriiainenteil with 10-12 fine, thin spiral cords; suture houniled with single, large spiral cord; sntural cord separated from shoulder spiral cord hv wide, depressed gap, pioducing slightly canaliculate appearance; spire whorls ornamented with 3 large spiral cords; aperture pniportionalK small. o\al in shape. Material examined: HOLOTYPE — Length 26 mm width, in liac'k-reet lacies of the Suwannee Formation, Terramar Pit, westernmost Polk Countw Floritla. UF75995; PARATYPES-lengths 19 mm and 23 nun. same localitx' as liolot\'pe. in the icseareh collection ol the authoi'. Et^■nlologv: Named for .Mr. William Dowiing of Lake Woith. Morida. who assisted me at the Terramar Pit and who collected the holot\|)e. Discussion: Flctirofiisiii (louliuiii occurs together with the congeneric P. hnxiksiilli'iisis (Mansfield. 1937){fig- ure 67) at the Terranuu' Pit. hut is nex'cr as connnon. The new species dilTers from the better-known P. hrooksvilli'iisis in being a i-onsistentlv more heavilv- scul[)tured shell with coarser and thicker spiral cords, in lacking \arical undulations on the spire and bod\' whorl, and in ha\ing a nmch narrower ga[) beh\'een the sutnral cord and shoulder cmd, ( )rder ( iephaiaspidea Snperiamilv ( Aliclmoidea I'^amiK (!\lichnidae (ieuus Siiufiiincsc(ijili(i new genus Diaj^nosis: (Aclichnid Imlible shells ol greater-thau- axcrage si/.e. wideK inlhited. o\ate. \er\ thin and Iragile; spire llattened. planar in profile; siitinc extremek de- nre.s.sed, [)roducing \('i"\ deep, wide, open c h.iunel (see figure 71); edge o( shoulder along channel de\eloped into \ei"\ thin, shaq), erect blade; bod\ \\li(irl smooth, laintK sculptured with extremcK' numerous, \('n fine s|)iral threads; aperture ven- wide and llaring, o\al in sha])e; apical perforation absent, re])laced l)\ raised pro- tocouch; anal notch well-developed, corres|ionding to raised shoulder blade along spire clianiu>l. Type species: Siiii iiiiiii sciijiliii I'uhIiic Petuch, new species, Suwannee I*"oiuiatiou, earl\ Oligoceue ol Flor- ida (figures 70,71 ), At present, known ouK Irom the Ter- ramar Pit. Polk Oountx. Florida. Et>'m<)logy: A combination ol 'Suwannee' and ".s(v/- phd" (barrel). Discussion: This unusual cxiichnid most closeK resem- bles members of tlu' genus Scapliandcr Montlort. 1810. paiticularK' in shape and size, but is inuuediatcK' sepa- rable bv possessing the characteristic w idel\ canaliculate spire. In luuing a flat, channeled spire. Siiutniiicscnphd somewh;it resembles a giant version of the diminutive c\iiclniid geims Aclnirinti Gra\. 1847. but differs in be- ing a nuich more inflated, rotund shell and in having a deeper ;iud much wider spire channi'l. Morphologically, Suwiniiicsraplia combines the large size and inflated shell form of Scaphdndcr and the flat, channeled spire ol ArtcdfiiKi. SinL(iiiius(/ij)liii liiuldi new species (figures 70, 71 ) Description: As for diagnosis of genus. Material examined: HOLOTYPE— Length 17 nun, width I 1 nun, in back-reef lagoonal tacies (sea grass beds?) ol the Suw;muee Formation, Terramar Pit, west- ernmost Polk Countv, Florida, UF75996; PARA- TYPES — lengths 19 mm, ami 21 nnn. same localitv as the holotvpe. in the lesearch collection of the author. Etvnn>logv: Named for mv wife, Linda |ovce Petuch. who stoitaliv watched over our familv while I was off e\[iloring the Suwannee Formation. ACKNOWLEDCMFNTS 1 wish to thank Mr. Eric Kendrew of X'alrico, Florida, for his generous donation ol research material collected ;it the Terramar Pit and for his valuable assistance in the field. Thanks also go to Mr. William Dowiing of Lake Worth. Floiida, lor Ins assistance in collecting in the Ter- ramar Pit, Special thanks go to Mrs. |ude Turner. Pho- tographv Lai) ;it Florida .Atlantic I'niversitv. for devel- oping and [)rinting the photographs used here and to Mrs. CAiithia Mischler. I')epartmeut of (n-olog)-, Florida Atlantic Universitv. for tvping the mamist'ri[)t. LITFHATUHE CITEH Ca.sev, '1", !,, 1903. Notes on tlic {.iorn-.iil rollrctioii of \ icks- liurg fossils. v\illi (Icscriptioiis ol new species. Proceedings of the Acadeiiiv ol N.ilur.il Scii'iucs of Pliihnliipliia 55: 261-2S3. (liiniail, '1'. .\, ISIS. Olisrnations ciii the Koci'ue lonnatioii, .111(1 (l('srriptii)iis of iiiie liiindri'd and five new fossils ol (ImI pcndif troni llic viciiiilv ol X'icksliurg, Mississippi. Willi .III .qipiiiiliv |(iiiiii.il 111 llic \ca(lriiiv of Niiliinil Sei- i-iirrs (il iinkiilclpliia. Seidiid Series lii!!4 I I l.vl, I), ill. \\ II n)lli \ ( eiiliiliiiticiii to ihi' liivrrleliiate l''aiiiia ol llic Oli'Mirciii Beds 111 I'linl Kivrr < leort^i;!. Proceed- E. J. Pctiicli, 1997 I'ai 137 iiii^s 1)1 tile L'liited Stat<-s Xaliiiiial Mumuiii 5ll2lfS2l: 487-524. MacNeil. F. S. ami D, T, Dockt-n, 1984. Limci ( )lit^i)cene GastropDila. ScapliopocLi. and ( 'cplialopoila ol the N'icks- biirg (Iroiip in Missis.'iippi. .Mississijipi Departnu'iit of Natural Rcsiinrces (Bnreau ol CH-()li)t:;\ '. Bnllctin 124, 415pp Man.sfiflil. W. C^. 1937. .\Iollnsks oi llic- T.nnpa and .Suwannee Liniestone.s ol Florida. State ol Florida Department ol Con.senation. Geologieal Bulletin 15. 334 pji. Petueli, K. |. 19S2. Geographical Heteroeln'ons: (Contempo- raneous (>)e\i.stence of Neogene and Reient Mollnsean Fanna.s in the Americas. Palaeogeograpli\, Falaeoclima- tolog\, and Paleoecolog)' 37:277-312. Petncli, E. |. 1988. Neogene Historv of Tropical .\nieriean Mollusks; Biogeographx' and Flvohitionan Patterns ol Tropical Western Atlantic Mollusca. The < 'oas(al Educa- tion anil Research Foimdation. ( lharlottes\ illc. \irginia. 217 pp. \anglni. T. W. 1900. A Tertiar\ Coral Reel' Near Bauihridge. Cleorgia. Science (New Series) 12; 873-875. X'okes, E. H. 1992. Cenozoic Muricidae of the W'esteni .\tlan- tic Region, Part IX. Ptcn/iiotiis, Ptnricrui. AspcUa. I)< r- DWiniin'X, Calotrophon. Artintliolahin. mitl Alliliosa: .\d- ditions and Corrections, Tulane Studies in (;eolog\ anil Paleontoli)g\- 25(1-31:81-82. APPENi:)l.\ I CJa.stropoil.s From The Bai'k-HccI Lagooua! Facies Ol The Suwannee Foniiatinn in The Terraniar Pit, Polk Countv', Florida. Y = Widespread X'ick.slinrt^ian Species; F = Species Also Known From The Flint Hi\i-r For- mation: S = Species Known ()nl\ Frcuu Tin' Suwannee Formation. Trochidae Calliostoiiiii siliciitiiiii M.uisfield. 1937 S Turhinidae AstriDii lAsli'dliiiiiil i>(>lkiii\i.\ I'etueh, n.S[). S Potamididae Piinizisimis kciiilrcwi Petuch, n.sp. S Tvhscojihnn hlnckaiitcn-ims (Mansfield, 1937) S Tch'sciijiiuiii liciiuniildciisis (Mansfield. 1937) S Cerithiidae Bittiiiiii ciisci/i MacNeil, 1984 V Ccritliuirlinii nitixtilc (Dall, UJlfi) F ('rrilliiiiin iTiiiriiiiiiii} iitsitliitiiiit Dall. UJlfi F ('chlltiiiiii I new genus?) cooA'c; Dall. UJlfi I*" ('■(■^tiiiiictrilliiinu hnxiksvillciisis (Mansfield, 1937 1 S Ccstiiiiictrilliiiiiii lirmakfHsi.s (Mansfield, 1937) S Ccstuiun'rilliiinu pascociisis (Mansfield, 1937) S Cisltiiiucritliiuin la^inatitiu (Dall, 1916) F PrisDKiiTrithiuiti prisma Petuch, n.sp. S Scniiiciiardc<>rliis kciiihrui Petuch, n,sp. (Y?) S .Muricidae Cliinirciis sMopits (de Gregorio, 1890) \" Poiriciid (Ddlliiitiircx) nifintpicohts (Dall, 1916) F PtiTipuifiis pnipcposti (Mansfield, 1937) S TdJUijphis iiiississipp'uiisis (Certmann, 1969) \' Bu.s\coniilae Spimftdffiir •^iinmiildliun l\-tueh, n.sp. S Bnccinidae PdUdcera caseiji (MacNeil, 1984) Y P(dlacerd i-ickshiir'^cnsis (Com'ad, 1848) \' Solcnostcini siiuduiu'ciisis Petuch, n.sp. S Melongenidae Mi/hsticd crdssicotiuttd (Comad. 1848) \' Turlnnellidae Chnolitlics I irkshiii'ff'itsis (CJonrad, 1848) Y Tiirl)iiiillii suwd}iii('iisis (Mansfield. 1937) S Vi7.s7/;/( snudiinrciisis l\'tuch. n.sp. S \'()lllti(Lie ludsilipid kciidrcui I'etuili. n.sp. S F(dsilip-i(i iiidiisficlili I Dall. 1916) F N'olutomitridae Coitdinili'd crrinildld Docket, 1984 \' Cononiitrd kciidrcui Petuch, n.sp. S Coiioiiiitrti stdiiiincd ((Com'ad, 1848) Y Mitridae (Pleioptxgmatidae?) Fusiiiiitra ciuujuistd (faini'ad, 1848) \' Oli\idae OliidiOiitoi^i/iiiiid I hniiiksiillcnsis (Mansfield, 1937) S Oliiclld liicDdkciisis Mansfield. 1937 S Olivclld iickshur;^ciisis iOoeken 1984 \' Marginellidae DcntiiiKir^Ht dnlli Petuch. n.sp. S Hijdiiiid silicijliivia {DaW. 1916) F Pcrsicidd dtickcn/i Petuch. n.sp. S Pcrsiciild ludciicili Petuih. n.sp. S Pcrsiculd suudiiiiccnsis Petuch, n.sp. S Pniuuiii cricdc Petucli, n.sp. S Pnniiiiu jcssicfie Petuch, n.sp. S Pniiiiiiii sdiidrdc Petuch, n.sp. (\') S Conidae Coiius {Lcjiliicniius) cddkci ]~)all, 1916 F Page 13S THE NAUTILUS, \<)1. 110. No. 4 C.oniis (Asprclla) kcnilrcni I'ctiicli, ii.sp Tcrebriclae Tcrehra (TcrcbirltiiKi) diii-'Hini (Joiirad, 1 TurricUif Coiiorlii.s porccllaniis (Conrad, 1S4S) V Crassisjiiirlla lyoplciira (MacNcil, 1984 S >S48 \' y .7 S 1.37) S Euclatluurlla Uvcoakctisis Maiisfii'kl, Plcurofusia hrooksiillcusis ( Mansfield Ph'urofiisia (lowliiif^i Petncli. n..sp. .S Plcurofusia phttonica (Ca.sew 1903) \' Pli'unifusiii sciTdta (C^onrad, 184S) V Poh/stini snhsiiHilis (Ca.sev. 1937) V ( A'lic'hnidae SuiLdinwsraplui liiuluc Pctucii, n..sp. S APPENDIX 2 .\dditional Suwannee Formation Ga.stropod.s Tliat Were Reported B\- Mansfield (1937) and E. V'okes (1992) But Were Not Collected (In Tliis Study) At The Terramar Pit. V = Widespread N'icksburgian Species: F = Species Also Known From The Flint River Formation: S = Spe- cies Ktiowu Onl\' From The Suwannee Formation. Fissurcllidae "Dioihini cliipolanii (Dall)" (prohahK = D nnssissip- picHsis (Conrad, 1848)) X Cerithiidae Cciithiuiii iiforiiumnni Lvell and Sowerhw 1S45 F "i'lTitliiuiii suaaiiurcnsis" Mansfield, 1937 Turritcllidae "Turriltlld cl. Iiiilriisis Dall" '= Apiculii ci. iKilciisis) V Xenophoridae "XenopJiora roncliiilinjiliiini Horn" l= .V. Iiuiuilis (Conrad. 1848)) ' Terei)ellidae Tcrcbclluiu lienuiuclocn.'iis Mansfield, 1937 S Cassidae "Cassis sp." (= C. flinfcnsis Mansfield. 1940i F Muricidae Takia poiirlli (E. \'okes, 1992) S Buccinidae (?) "Latinis sp." (appears to be a Caulhanis sp.) S V'olutidae Liiria iiiississippicnsis Conrad, 1848 V 2 3 5 3 i' ; / 0' INSTRUCTIONS TO AUTHORS THE NAUTILUS publishes papers on all aspects of tiie bioiog)- and systematics of niollusks. 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