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Mussels from the ++ 1.04 m sampling horizon have heavier shells than those from the 2 lower sampling horizons. The RMAs of the height-width, width-length, shell weight-length, and soft part weight-shell weight plots differ more in slope than in Y-intercept (as evident in the appearance of the plotted data). The RMAs of the shell-height plots differ neither in slope nor Y-intercept. In animals frozen while immersed with open valves, the edge of the outer (secretory) mantle fold was found to be even with, or slightly outside the shell margin (Fig- ure 3). In animals frozen while emersed with closed valves, the edge of the outer fold was withdrawn within the shell margin. Figure 3 Schematic representation of mid-abaxial sections of mussel ventrum. (A) mussel with shell closed (animal emersed) (B) mussel with shell open (animal immersed). m — mantle s — shell P — periostracum A brief, general look at the data yields the following cor- relations. Abiotic parameters, as a whole increasing with the tidal height of the samples, can explain the similar general mussel shape and weight trend. Emersion per- centage or duration, or both, alone could also explain the shape and weight data trend, as these 2 sets of data vary together. However, wave stress alone cannot cause the shape and weight trends, because it is discontinuous. Fur- ther, a discontinuity resembling that of the wave stress data is not found in the shape and weight data. The results also establish the possibility of some causative role for population density. Mantle retraction in response. to stress (SEED, 1968; PANNELLA & MacC.intock, 1968; present study) likely effects shell growth more toward the midline and less in height and length (SEED, 1968). Thus the relative in- creased width in this study’s high-shore animals can be related to the relatively greater amount of their life- cycles spent emersed. This also concurs with the conten- tion that “normal” outer shell deposition can occur only with open valves and subsequent mantle extension well along the margins (PANNELLA & MacCuinTock, op. cit.). Study of short-term accretion in stressed and unstressed Mytilus californianus, as was done with Mercenaria mer- cenaria (Linnaeus) (ibid.) would help to understand this process, although these studies would be more difficult with the reflected margin of M. californianus. Support for a direct, causal connection between emer- sion time and shell growth patterns is found in studies of Vol. 22; No. 1 species differing in their emersion responses. Shell shape changes with tidal height (Mytilus californianus: Fox & CoE, 1943; present study; Mytilus edulis Linnaeus, 1758: SEED, 1968) have been found in species where valve closure always or almost always accompanies emersion (M. californianus: personal observation; M. edulis: COLEMAN, 1973) whereas no such shell shape changes were found (LENT, 1967) in Modiolus demissus Dillwyn, 1817, which air-gapes excessively (COLEMAN, op. cit.). Evidence that shell growth during closure is greatly re- tarded but continues (PANNELLA & MacC.intock, 1968) or is net positive in spite of interim dissolution (Bamp & DrINNAN, 1957) seems to leave open the possibility of modest width accretion during closure, especially when growth in height and length are likely even slower during closure. Wide mussels high on the shore are equally explicable by having growth in height and length attenuated either ontogenetically (SEED, 1968), or by a longer time spent emersed (this report). Data from old mussels found low on the shore would seem to offer arbitration between these two explanations. The rarity of such animals has been attributed to the effect of the feeding patterns of desicca- tion-intolerant predators (Mytilus edulis: SEED, 1968; M. californianus: PAINE, 1976). Even in the absence of pre- dation, however, the survival of M. edulis in the Menai Straits, North Wales, has been found to be markedly poorer at the 2% (spring extreme low water) exposure level than at the 6- 20% exposure levels (P J. Dare « G. Davies, unpublished data). Hence a search for arbi- tration in the form of old mussels from low on the shore may be fruitless, and therefore contraindicated. In either case, the result of allometric growth would be enhanced by a high shore shift toward longer-lived animals (M. edulis: SEED, op. cit.; M. californianus: PAINE, op. cit.), as the difference would accumulate with time. If population density (Fox « Cor, 1943; LENT, 1967; SEED, 1968) or overall growth rate (CoE & Fox, 1942; CoE, 1945) were each the sole or primary cause of shell shape changes with tidal height, we may well predict that mussels either show a simultaneous diminution of height and width relative to length with increasing popu- lation density, or changes in shell size alone with no shell shape changes, respectively. The former prediction is based on the assumption by animals of an orientation with long axes normal to the substrate in high population den- sities (LENT, 1967; SEED, op. cit.; present study). The trend of height relative to width, in addition to the in- dependent comparisons of height relative to length, and width relative to length, deny strict realization of these expectations. However, mussel girth (height and width THE VELIGER Page 55 taken together) does in fact decrease with an increase in population density, thereby leaving open the possibility of some causal link between these parameters. Wave impact may cause thickened shells (CoE & Fox, 1942), but the wave data reported here eliminate wave stress as the sole cause of the trend of shell weight relative to length. However, shell width (zbid.; present study) cor- relates well with shell weight relative to length, and could be the cause of the weight trend, when it is considered that a wider mussel of a given length and height is likely to be heavier. Whether wave stress and emersion act in concert or separately on shell thickness and width is unknown. Study of mantle position under wave stress would certainly help arbitrate putative causal links between these para- meters. Growth in height and length are equally affected by, and hence exhibit no relative variance along the emersion gradient. This results naturally from the relationship be- tween the emersion-affected mantle position and shell growth advanced in this study, because accretion to both height and length occurs when the mantle is protracted, and little or no accretion occurs to either when the mantle is retracted. Efforts to measure the tissue/shell weight relationship, acrucial gross energetic parameter, are plagued with prob- lems. The trend revealed in this study (also in Bamp & DRINNAN, 1957; LENT, 1966; SEED, 1973) is opposite to Rao’s (1953) results. Rao discarded all “gravid” animals, possibly biasing his samples toward juveniles. This bias can give high shell/tissue weight ratios (SEED, op. cit.). Further complications arise from indiscrete, disagreed upon or even year-round spawning in Mytilus californi- anus (ibid.), with M. edulis giving a 100% annual tissue weight variance (Dare & Epwarbs, 1975). Future studies may do well to include averages of year-round collec- tions (Fox & CoE, 1943). An interesting contradiction has arisen between the results of various studies of mussels. Allometric plots that vary more, judging from the visual appearance of the plots, in slope than Y-intercept (Mytilus californi- anus: Fox & Cor, 1943; present study; M. edulis: Srp, 1968, 1973) usually represent phenotypically different, genotypically similar samples, while those varying more in Y-intercept than slope represent genotypic variance (Goutp, 1966). Thus the former type of variance is the accumulating effect through time of differing environ- mental regimes (SEED, 1968). However, several workers report inter-site (A/. californianus), and inter- and intra- site (M. edulis) genetic variance via enzyme electropho- resis (LEVINTON & KoEHN, 1976). This apparent contra- diction awaits resolution. Page 56 THE VELIGER Vol. 22; No. 1 SUMMARY 1. Mytilus californianus were sampled in a random stratified fashion from the California rocky intertidal zone. 2. Shell width and weight relative to length and shell width relative to height had a positive and significant correlation, tissue weight relative to shell weight had a negative and significant correlation, and shell height relative to length had no correlation with tidal height. 3. Emersion percentage time and average emersion du- ration had a positive correlation, wave impact had a partial positive correlation, and population density had a negative correlation with tidal height. 4. Cryogenic preparations reveal a retracted outer man- tle fold during emersion. This is used to explain change in shell shape with the emersion differential. 5. The findings of this report are compatible with ex- planations elsewhere of spatial demographic shift and partial ontogenetic attenuation of growth, but are in conflict with explanations involving population density, overall growth rate, and wave stress. 6. Year-round collection is recommended for tissue- shell weight ratio determination. 7. Genotypic variance findings in this report conflict with recent biochemical studies elsewhere. ACKNOWLEDGMENTS For guidance and assistance, I would like to thank my colleagues at the University of California, Santa Cruz, especially Drs. M. W. Silver, A. T: Newberry, and J. S. Pearse. Also, my sincere thanks to W. C. Thompson, U. S. Naval Postgraduate School, Monterey, California, for providing the NOAA data and advice; to Drs. D. L. Fox, Scripps Institution of Oceanography, R. Seed, University Colle of North Wales, J. J. Gonor, Oregon State Univer- sity, and W. Lee, California Academy of Sciences, for critical review of the manuscript; and to P J. Dare and G. Davies, Ministry of Agriculture, Fisheries and Food, Conwy, Wales, for gracious provision of unpublished data. Illustrations are by G. M. Breed, O.S. U.S. D. G. Literature Cited Bairp, R. H. 2 R. E. Drinnan 1957. The ratio of shell to meat in Mytilus as a function of tidal expo- sure to air. Journ. Cons. Exp. Mer. 22: 229 - 236 Cocuran, W. G. 1963. Sampling techniques. 413 Pp. John Wiley & Sons, Inc. New York, Cor, Westey RosweLi 1945. Nutrition and growth of the California bay mussel (Mytilus edulis diegensis). Journ. Exp. Zool. 99 (1): 1-14; 2 text figs. (June 1945) Coz, WESLEY ROswELL & Denis L. Fox 1942. Biology of the California sea mussel (Mytilus californianus). I. Influence of temperature, food supply, sex and age on the rate of growth. Journ. Exp. Zool. g0 (1): 1-30; 6 text figs. (5 June 1942) Coreman, Nog. 1973. Water loss from aerially exposed mussels. Biol. Ecol. 12: 145 - 155 Dame, RicHarp EF 1972. Comparison of various allometric relationships in intertidal and subtidal American oysters. Fish. Bull. 70 (4): 1121-1126 Dare, D. J. « D. B. Epwarps 1975. Seasonal changes in flesh weight and biochemical composition of mussels (Mytilus edulis L.) in the Conwy estuary, North Wales. Journ. Exp. Mar. Biol. Ecol. 18: 89 - 97 Doty, MaxwELL STANFORD 1946. Critical tide factors that are correlated with the vertical distribu- tion of marine algae and other organisms along the Pacific coast. Ecology 27 (4): 315-328 Journ. Exp. Mar. Exuiotr, J. M. 1921. Some methods for the statistical analysis of samples of benthic invertebrates. The Ferry House, Ambleside, England, 148 pp. Fox, DENIS L. & WESLEY ROSWELL CoE 1943. Biology of the Californian sea mussel Mytilus californianus. II. Nutrition, metabolism, growth and calcium deposition. Journ. Exp. Zool. 93: 205 - 249 Giynn, PETER W. 1965. | Community composition, structure and interrelationships in the marine intertidal Endocladia muricata-Balanus glandula association in Monterey Bay, California. Beaufortia 12 (148): 1-97 Goutp, STEPHEN Jay 1966. Allometry and size in ontogeny and phylogeny. 41: 587-640 Harcer, Joun Rosin E. 1970. Effect of wave impact on some aspects of the biology of sea mussels. The Veliger 12 (4): 401-414; 9 text figs. (1 April 1970) Hayamyi, ITaru «& AxiHIKO MaTsuKUMA 1970. Variation of bivariate characters from the standpoint of allo- metry. Palaeont. 13 (4): 588 - 605 IMBRIE, JOHN 1956. | Biometrical methods in the study of invertebrate fossils. Bull. Amer. Mus. Nat. Hist. 108: 215 - 252 KENNEDY, V. S. 1976. Desiccation, higher temperatures and upper intertidal limits of three species of sea mussels (Mollusca: Bivalvia) in New Zealand. Mar. Biol. 35 (2): 127-138 Lent, Cuartes M. 1967. The effect of habitat on growth indices of the ribbed mussel Modiolus (Arcuatula) demissus. Chesapeake Sci. 8: 221 - 227 LEvVINTON, JEFFREY S. & R. K. KozHn 1976. Population genetics of mussels. In: Marine mussels: their ecology and physiology. B. L. Bayne (ed.), Cambridge Univ. Press: 357 - 384 Mizuts, FE C. 1955- Statistical methods. 842 pp. PAINE, ROBERT TREAT 1976. Size-limited predation: an observational and experimental ap- proach with the Mytilus-Pisaster interaction. Ecology 57: 858 - 873 PANNELLA, Gioroio & CopELAND MacC.intTock 1968. Biological and environmental rhythms reflected in molluscan shell growth. Journ. Paleontol. 42 (5, Suppl.) :64 - 80 Rao, K. PAMPAPATHI 1953. Shell weight as a function of intertidal height in a littoral popu- lation of pelecypods. Experientia 9: 465 - 466 Riepi, R. 1971. Water movement. In: Marine Ecology, vol. 1, Environmental factors. O. Kinne (ed.). John Wiley & Sons, Ltd. London: 1085 - 1156 Szzp, Ray 1968. Factors influencing shell shape in the mussel Mytilus edulis. Journ. Mar. Biol. Assoc. U. K. 48: 561 - 584 1973. Absolute and allometric growth in the mussel Mytilus edulis L. (Mollusca: Bivalvia). Proc. Malacol. Soc. London 40: 343 - 357 1976. Ecology. In: Marine mussels: their ecology and physiology. B. L. Bayne (ed.), Cambridge Univ. Press: 13 - 65 Woonin, SarAaH ANN ; : 1974. Polychaete abundance patterns in a marine soft-sediment envi- ronment: the importance of biological interactions. Ecol. Monogr. 44: 171 - 187 Biol. Rev. Henry Holt # Co., New York, 3r4 ed., Vol. 22; No. 1 THE VELIGER Page 57 Problacmaea moskalevi Golikov & Kussakin, a New Addition to the Eastern Pacific Limpet Fauna (Archaeogastropoda : Acmaeidae) DAVID R. LINDBERG Center for Coastal Marine Studies, University of California, Santa Cruz, CA (lifornia) 95064 and Department of Invertebrate Zoology, California Academy of Sciences, San Francisco, (CA) 94118 (2 Plates; 3 Text figures) THE ACMAED GENUS Problacmaea Golikov « Kussakin, 1972, is represented by 2 extant species in the boreal Pacific: Problacmaea sybaritica (Dall, 1871) and P mos- kalevi Golikov & Kussakin, 1972. Members of the genus differ from other acmaeids in possessing a penis-like struc- ture under the right cephalic tentacle. The radula is typi- cal of species that feed on encrusting algae, having 3 lateral teeth approximately equal in size and shape and arranged in a chevron-like pattern. The shells are white or reddish with or without darker rays, and are sculp- tured with concentric growth lines and in some, radial riblets. Problacmaea sybaritica has an extensive distribution in the northern Pacific: Hakodate, Japan (43°N, 141°E), Plover Bay, Chukotskiy Polvostrov, U.S.S.R. (64°N, 173°W), Pribilof Islands (57°N, 170°W), Aleutian Is- lands to Chirikof Island, Alaska (56°N, 156°W) (Mc- Lean, 1966: 47), but the type of the genus, P moskalevi, has been previously reported only from localities in the seas of Okhotsk and Japan, U.S.S.R. Recently, Dr. James H. McLean, Los Angeles County Museum of Natural History (LACM), noted that a specimen lot of Lepeta concentrica (Middendorff, 1847) from Alaska contained a single specimen that was different; it had smaller limpets attached to its dried body. Dr. McLean brought this specimen to my attention, and after exam- ining it I have identified it as P moskalevi. The specimen, re-catalogued as United States National Museum of Natural History (USNM) No. 749078, was collected on stones at a depth of 6.5m in Amkaknak Straits, Captain’s Bay, Unalaska, Aleutian Islands, Alas- ka (53°52’N, 166°34’W) by W. H. Dall in the early 1870’s. This represents a new distributional record over 2500 km east of the previous localities and suggests that this species may be present throughout the Aleutian Island arc. It is ironic that Dall, who contributed so extensively to acmaeid systematics and anatomy, overlooked this unique specimen, and thus it remained undescribed for approximately 100 years. The original description of Problacmaea moskalevi, translated from French, is given here and followed with additional remarks based on the Dall specimen. Problacmaea moskalevi “The shell is fine, rather fragile with a rounded oval base and a raised subcentral apex. The shell is white. The sculpture is described simply as concentric growth lines set off by annual rings. The inner sur- face of the shell has a small grey spot near the apex. The radula is typical of the genus. Height of the holotype: 4.7mm, length 10.5mm, width 9.5mm. Type locality— shallow basin in the middle horizon of the mesolittoral at Ostrov Bol’shoy Shantar [55°N, 137°42'E], Sea of Okhotsk (collected by M. B. Ivan- ova, 10 October 1966). The holotype is deposited in the systematic collections of the Institute of Zoology, Academy of Sciences of the U. S.S. R. [Leningrad]. The species is named in honor of Dr. L. Moskalev, a well-known specialist in limpets.” (GoLIkov & KUSSAKIN, 1972: 290) Golikov « Kussakin compared Problacmaea moskalevi Page 58 THE VELIGER Vol. 22; No. 1 to Acmaea apicina Dall, 1879, noting that the 2 species were superficially similar but differed in their respective length/height and width/height ratios, P moskalevi being consistently lower than A. apicina. Also, they considered P. moskalevi, like P. sybaritica, to be ovoviviparous and a protandric hermaphrodite. A paratype P moskalevi from the type locality contained a large number of embryos with shells in its nuchal cavity. The embryonic shells were approximately 0.3mm in length. Other specimens from Tartarskiy Proliv, Sea of Japan (50°N, 141°E), collected at a depth of 35 - 37m from sand and mud substrata, had nuchal cavities filled with large eggs, approximately 0.115 Shell Interior 1 Additional localities represented in Golikov and Kussakin’s mate- ria] are: Simushir (46°58’N, 152°02’E), Urup (46°N, 150°E) and Iturup (44°54’N, 147°30’E), Kuril Skiye Ostrova, U.S. S. R.; 5-15m on sand and stones (A. N. Golikov, in litt. March, 1978) mm in diameter. Golikov & Kussakin examined all to- gether 11 specimens of P moskalevi from 3 localities. * The Dall specimen (Figure 1) is 6.6mm in length, 5.7 mm in width, and 2.8mm in height. The length/height ratio is 2.4; the width/height ratio 2.0, both being slight- ly greater (+0.1) than the ratios given by Golikov « Kussakin for Asiatic specimens of Problacmaea moskalevi. The shell is white and encrusted with coralline algae. The anterior slope is slightly concave, the apex anteriorly di- rected. The posterior slope is convex and the lateral slopes straight. Sculpture of concentric growth lines is visible only at the shell margin. Viewed in profile the shell shows Approximate Schematic Position Structure Equivalent I~VY/Y{WYJVJJYJVJYVJJYJY/0 Mrernannennned cteetudtaltultult EY Nennnnnnnnee Mrarurnoannnned ornare weeararararararenenints hee AORN secs: Waatrarnnnnnne PKAXRXRXRX AY - M Myostracum Muscle Scar Concentric ia M+1 Crossed- Intermediate lamellar Area N \ M+e Complex- Interior N prismatic Margin Figure 2 Shell structure of the genus Problacmaea illustrating relationship of structure groups to terms used in acmaeid systematics. M — my- ostracum; m-1 — first structure group interior of myostracum; m-+1 — first structure group exterior of myostracum, etc. Explanation of Figures za to 1c, 4 to 8 Figure 1: Problacmaea moskalevi Golikov « Kussakin, 1972. ALASKA: Aleutian Islands, Unalaska (USNM No. 749078) ; shell length: 6.6mm a — dorsal; b — ventral; e — profile Figure 4: Dried body of Problacmaea moskalevi (USNM No. 749078). Body length: 3.9mm. NC -— remnant of nuchal cavity containing young limpets; V — viscera; IL — intestinal loops Figures 5 to 8: Protoconch of Problacmaea moskalevi (USNM No. 749078). Figure 5: dorsal view, X 340; Figure 6: anterior view, X 300; Figure 7: lateral view, X 340; Figure 8: posterior view, X 290 Tue VELIGER, Vol. 22, No. 1 [LinpBerc] Figures ra to rc, 4 to 8 Figure ra Figure 1b Figure 4 ee Figure 1C Tigure 6 ee ee Figure 5 Figure 7 Vol. 22; No. 1 THE VELIGER Page 59 unequal growth increments that result in uneven slopes [= Golikov « Kussakin’s “annual rings’?]. The interior is whitish with a slight yellow cast near the margin. The apical area is covered by the dried mantle, to which several young limpets are attached. The gray spot men- tioned by Golikov « Kussakin, if present, is not visible through the dried mantle. The shell structure (Figure 2), determined from the concentric bands visible on the inner surface, appears to be identical to that of P sybaritica, except that the inner margin (complex prismatic layer) of P moskalevi is much narrower than in P. sybaritica. The radula of the Dall specimen was not examined. Golikov « Kussakin’s figure of the radula of P moskalevi is given here in Figure 3. SLT TLT Figure 3 Radula of Problacmaea moskalevi (redrawn from Gotikov « Kus- SAKIN, 1972). FLT — first lateral teeth; SLT — second lateral teeth; TLT — third lateral teeth; scale unknown The dried body of the specimen (Figure 4) is 3.9mm in length and contains 24 small limpets in the nuchal cavity. Although contracted, large oral lappets are vis- ible on the muzzle. The foot is remarkably small, approx- imately 1.5mm in length. The nuchal cavity is large, ap- proximately 3.3mm in length or almost 85% of the ani- mal’s total length. Neither the ctenidium nor the nephridi- opores are visible because the specimen is dried and shriv- eled and the region is obscured by the juvenile limpets in the nuchal cavity. The visceral area is markedly posterior and ventral; the intestinal loopscontaina whitish material that appears to be calcium carbonate. The penis-like structure is not visible under the right cephalic tentacle. I do not know whether this is due to the specimen’s dried and contracted state, atrophy of the structure accompa- nying sex change, or degrees of both. Based upon the embryonic shells in the Dall specimen, the protoconch of Problacmaea moskalevi (Figures 5 - 8) is positioned in the center of the teleoconch (Figure 9). There is a slight asymmetry, the posterior portion of the protoconch being slightly to the left of the longitudinal axis of the teleoconch. The lateral edges of the protoconch are straight and approximately parallel. The posterior portion is rounded, with a small indentation on the postero-dorsal surface. The surface appears smooth, with no sculpturing. No coiling is visible, and the protoconch is presumed to becaecum-like. The posterior slope is strong- ly convex and ventrally appressed to the teleoconch (Fig- ures 7-8). Theanteriorslopeisalsoconvex although not as strongly as the posterior (Figure 7). The entire protoconch isseparated from the teleoconch bya finesuture (Figure 5). The initial teleoconch lacks sculpture and forms a slight depression around the posterior and lateral margins of the protoconch (Figure 8). After the initial growth, the teleo- conch is sculptured with approximately 40 sinuous, flat topped ribs (Figure 9). The interspaces between the ribs are divided by numerous ridges (Figure zo). Nearer the margin these ribs disappear, and the strong concentric growth lines typical of the species become apparent (Fig- ure II). DISCUSSION The shell structure of Problacmaea is identical to that of the western Pacific genus Patellouda Quoy « Gaimard, 1834, and unlike that of Acmaea, Collisella, and Noto- acmea (MacCiinTock, 1967). Although the genus Pa- telloida is characterized by 2 pairs of radular marginal teeth, the genus Problacmaea is simply derived by the loss of the marginal teeth as the radula became modified for feeding on encrusting algae. I consider the similarity of shell and radular morphologies to members of the eastern Pacific genus Acmaea Eschscholtz, 1833, to result from convergence rather than common ancestry. In temperate and tropical species acmaeid reproduc- tive strategy consists of external fertilization and pelagic development. THorsoN (1950) has shown that Arctic and near-Arctic prosobranchs tend to delete the pelagic phase and instead brood their young. In view of this model the apparent departure from typical acmaeid re- productive strategy by members of the genus Problacmaea is not surprising, and it is unlikely that this represents a distinct evolutionary tendency within the Acmaeidae wor- thy of subfamilial rank as proposed by Go.tkov « Kus- SAKIN (1972: 292). Page 60 THE VELIGER Vol. 22; No. 1 The protoconch of Problacmaea moskalevt is similar to that of P sybaritica (Figures 12, 13). 'The lateral pouches (Figure 14) described and figured by Morse (1910) and THompson (1912) in eastern Pacific and western At- lantic acmaeids are not present in either species of Probl- acmaea. Instead there is a simple suture between the apex and the lip of the protoconch of P sybaritica that is LP Bey Figure 14 Protoconch of Acmaea sp. (redrawn from Morsg, 1910). LP — lateral pouches; scale unknown visible under high illumination light microscopy. A ridge is also visible through the protoconch that seems to con- nect the anterior ends of the sutures. These sutures ‘are not visible in the protoconchs of P moskalevi, presumably because of the more advanced stage of development of these shells, the ventral portion of the protoconch being overlapped by the teleoconch. Specimens observed at an earlier stage of development would most likely show these sutures. The early teleoconch of Problacmaea moskalevi most closely resembles that of the Arcticspecies Acmaea rubella (Fabricius, 1780) (cf. THorson, 1935: 68; fig. 74), both species being sculptured with distinct radial riblets on the early teleoconch but having only concentric growth lines as adults. Specimens of P sybaritica collected by McLean in 1973 show that the initial teleoconch of P sybaritica is smooth but soon develops pits similar to those in P moskalevi. Because of the early stage of devel- opment of these shells I can not determine if these pits give rise to radial riblets as they do in P moskaleuvt. How- ever, as P sybaritica is sculptured with fine radial riblets as an adult, I suspect that this is the case. ACKNOWLEDGMENTS I wish to thank Dr. James H. McLean (LACM) for bringing the specimen of Problacmaea moskalevi to my attention, and also kindly reviewing the manuscript. Dr. Joseph Rosewater (USNM) supplied additional locality data on the Dall specimen. Dr. Alexandre N. Golikov (Academy of Sciences, Leningrad) supplied additional locality data on the western Pacific specimens. Special thanks to Ms. Jan Nowell (University of California Santa Cruz) for her assistance and patience with the scanning electron microscopy portion of this study. Literature Cited Go.utKkov, ALEXANDRE N. & O_ec G. KussakINn 1972. Sur la biologie de la reproduction des patelles de la famille Tec- turidae (Gastropoda: Docoglossa) et sur la position systématique de ses subdivisions. Malacologia 11 (2): 287-294; 7 text figs. (June 1972) MacCuinTocx, CopgLAND 1967. Shell structure of patelloid and bellerophontoid gastropods (Mol- lusca). Peabody Mus. Nat. Hist. Yale Univ. Bull. 22: ix+140 pp.; 128 text figs.; 31 plts. (21 February 1967) McLgan, James HamMILtTon 1966. West American prosobranch Gastropoda: superfamilies Patella- cea, Pleurotomariacea, and Fissurellacea. Ph. D. dissert., Stan- ford Univ.; x+262 pp.; 7 plts. Morse, Epwarp SYLVESTER 1910. An early stage of Acmaea. 313-323; 8 text figs. TuHompson, WILL FE 1912. The protoconch of Acmaea. 64: 540-544; 6 text figs. Txorson, GUNNAR Proc. Bost, Soc. Nat. Hist. 34: Proc. Acad. Nat. Sci. Phila. 1935. Studies on the egg capsules and development of Arctic marine prosobranchs. Medd. Grol. 100 (5): 1-71; 75 text figs. 1950. Reproductive and larval ecology of marine bottom invertebrates. Biol. Rev. Camb. Philos. Soc. 25 (1): 1-45; 6 text figs. Explanation of Figures 9 to 13 Figures 9 to 11: Early teleoconch of Problacmaea moskalevi (US NM No. 749078), shell length o.gmm. Figure g: dorsal view, X 80; Figure ro: radial sculpture, X 660; Figure 11: interface of radial sculpture with adult sculpturing, Vol. 22; No. 1 THE VELIGER Page 63 lighter in intensity of color. The sculpture is of definite nodes. The shell is thin and delicate. Terebra petiveriana Deshayes has a thicker subsutural band, usually of a definite contrast in color. Though often noded, frequently the sculpture is of rough axial ribs. Its shell is heavier and broader with coarser sculpture and with 2 folds on the columella. Although there appear to be 2 syntypes of Terebra glauca in the British Museum type collection, the word “unique” on the back of the mount seems to imply that originally there was only 1 specimen, the other being added later. The larger, more centrally mounted speci- men closely matches Hind’s measurements, so it must be considered the holotype. Terebra petiveriana Deshayes, 1857 (Figure 4) Terebra aspera Hinps, 1844: 154 [non Bosc, 1801] [not fig- ured]; Hinps, 1845: 174, plt. 43, fig. 44; CaTLow «& REEVE, 1845: 288; REEVE, 1860, sp. 40, plt. 10; VREDEN- BURG, 1921: 344, no. 18 Terebra radula Hinps, 1844: 155 [non GravENHORST, 1807] [not figured]; Hinps, 1845: 74, plt. 44, fig. 95; CaTLow «& REEVE, 1845: 290; REEVE, 1860, sp. 68, plt. 15 Terebra petiveriana DEsHAYES, 1857: 85, fig. 10; TRYON, 1885: 14, plt. 3, no. 31; TOMLIN, 1944: 14, no. I41 Terebra glauca Hinps, 1844: 155; ReEve, 1860: sp. 40, plt. 10; TRYON, 1885: 14, plt. 1, fig. 19 [exclude from synon- ymy of T: variegata Gray, 1834]; PaETEL, 1888: 251; TOMLIN, 1944: 14, no. 142 Description of Species: Size medium, to 46mm; color usually gray to brown, with whitish subsutural band, oc- casionally monochromatic flesh to beige; outline of whorls convex, with moderately wide convex subsutural band; protoconch of 3 to 34 slender, glassy whorls; early whorls of teleoconch sculptured with elongate nodes on subsu- tural band and straight, sharp axial ribs; suture well defined; spiral grooves appearing about 4" whorl, some- times developing small nodes upon crossing axial ribs about 7 whorl; sculpture coarse and extremely variable, of axial ribs with narrow spiral grooves or noded with broad grooves; subsutural band thick, convex, noded, well marked by suture and deep subsutural groove; body whorl with rough sculpture often continuing to siphonal fasciole ; outer lip moderately sturdy, columella recurved, with 2 basal folds; siphonal fasciole striate, with keel. Type Locality: Panama. Type Specimens: Holotype and 1 paratype, British Mu- seum (Natural History). Remarks: It is interesting to note that after Reeve fig- ured a specimen of Terebra petiveriana as T. glauca, subse- quent reviewers followed his lead and figured T. petiveri- ana as T: glauca or synonymized T: glauca with T. aspera or T: radula, both synonyms of T: petiveriana. Terebra carolae Bratcher, spec. nov. (Figures 5, 7, 9, 11) Synopsis: Shell medium-large with beige background marked with brown, flat sided except for convex subsu- tural band. Description: Shell moderately large, 52.3 X 11.7mm; color dull brownish-beige with darker brown between nodes of subsutural band and with light band at periphery of body whorl; outline of whorls flat, with convex sub- sutural band; protoconch missing from holotype and all mature specimens examined, but protoconch of immature paratype has 14 smooth mamillate embryonic whorls; axial sculpture of early whorls of teleoconch of narrow ribs; spiral sculpture of subsutural groove cutting through ribs to form noded subsutural band, with 2 additional spiral grooves crossing ribs; axial sculpture of later whorls of weak ribs, 14 on penultimate whorl; spiral sculpture consisting of subsutural groove cutting through axial ribs and 4 rows of cords crossing over ribs to form indistinct nodes; spiral cords becoming obsolete on body whorl with 3 spiral grooves anterior to periphery, with numerous axial striae between periphery and siphonal fasciole; sutures deep; aperture long semi-quadrate; columella recurved, with weak plication; siphonal fasciole striate, keel very sharp, strong. Type Locality: Santa Maria Bay, off Hughes Point, Baja California del Sur, Mexico (24°45'05’N; 112°19’W); 54m, shell bottom. Type Specimens: Holotype: LACM-AHF 1178. Para- types, all from west coast of Baja California, Mexico: LACM-AHF 1184 from type lot (13) ; LACM 71-163 off Rompiente Point, 38 - 25m (8); LACM A375 San Bar- tolomé Bay (21); LACM-AHF 71-164 off Rompiente Point, 50m (g); LACM 71-178 San Pablo Point, 23 - 30 m (1); LACM 71-180 Point Pequena, San Juanico Bay, 10m, fine sand (8); AMNH 18661 San Bartolomé Bay, 18m (1); ANSP 345789 Cedros Island, 15m (1) ; Brat- cher collection Santa Maria Bay, 15m (6); San Ignacio Lagoon (14); San Bartolomé Bay, 36-54m (7); CAS 59672 Santa Maria Bay, 15m (1); Cernohorsky collec- tion Santa Maria Bay, 15m (2); DuShane collection USIVM 7&2S71 anita | Page 64 THE VELIGER Vol. 22; No. 1 Cedros Island (2) ; MCZ 28805 San Bartolomé Bay (1) ; YPM 17717 Santa Maria Bay (1); Skoglund collection off Cedros Island (20) ; SBMNH 51671 Santa Maria Bay (1); SDMNH 70581 Santa Maria Bay (1); BM(NH) Santa Maria Bay (1). Discussion: ‘This species most closely resembles Terebra variegata Gray, 1834, but it bears some resemblance also to T. armillata Hinds, 1844, both of which have slender, multistriate protoconchs. Terebra carolae has a pauci- spiral mamillate protoconch and a consistently wider apical angle than either of the 2 species mentioned above. The plications on the exterior columella are less pro- nounced, and the interior columella shows an even greater difference. That of T: variegata has 2 closely spaced, ex- tremely sharp plications, while T. carolae has 2 wider spaced and weaker plications. Terebra carolae should be compared with T. tzare/la Deshayes, 1859, which is smal- ler and monochromatic. This species is named in honor of Carol Skoglund, who first brought it to my attention. ABBREVIATIONS Abbreviations have been used for a number of institutional collections cited in this paper. They are: ; AHF —- Allan Hancock Foundation (material on loan to Los Angeles County Museum of Natural History) AMNH -— American Museum of Natural History ANSP — Academy of Natural Sciences of Philadelphia BM(NH) - British Museum (Natural History) CAS —- California Academy of Sciences LACM - Los Angeles County Museum of Natural His- tory MCZ - Museum of Comparative Zoology, Harvard Uni- versity SBMNH - Santa Barbara Museum of Natural History SDMNH - San Diego Museum of Natural History USNM - United States National Museum YPM —- Yale University Peabody Museum of Natural History ACKNOWLEDGMENTS I wish to express my thanks to Dr. A. Myra Keen for reading and evaluating the manuscript of this paper. Literature Cited Avams, Cartes BAKER 1852. Catalogue of shells of Panama; with notes on their synonymy, station and geographical distribution. Ann. Lyc. Nat. Hist. New York 5: i- viii+334 pp. BraTCHER, Twita & RoBerT DoNALD BuRCH 1970. Five new species of Terebra from the eastern Pacific. The Veliger 12(3): 295-300; plt. 44 (1 January 1970) 1971. In: Keen, Sea shells of tropical west America, 27d ed., Stanford Univ. Press, Terebra, pp. 680 - 686; figs. 1520 - 1573 (Sept. 1971) CatLow, AcNness & Lovett REEVE 1845. The conchologists’s nomenclator. Str., Strand, London DesHayes, GERARD PAUL 1857. Description d’espéces nouvelles du genre Terebra. Journ. de Conchyl. 6: 65-102; plts. 3-5 (July 1857) 1859. A general review of the genus Terebra and a description of new species. Proc. Zool. Soc. London for 1859: 270 - 321 (between July and October 1859) Grant, Utyssgs Simpson, IV « Hoyt Ropney Gate 1931. Catalogue of the marine Pliocene and Pleistocene Mollusca of California and adjacent regions. San Diego Soc. Nat. Hist. Mem. I: 1 - 1036; 32 plts.; 15 text figs. (3 November 1961) Gray, JoHN EDwarp 1834. (Untitled; Terebra] 50 - 63 Hinps, RicHarp BriINsLeEY 1844. Descriptions of new shells, collected during the voyage of the Sulphur and in Mr. Cuming’s late visit to the Philippines. Proc. Zool. Soc. London for 1843 (XI): 149 - 168 (June 1844) 1845. Monograph of the genus Terebra. In G. B. Sowerby, The- saurus conchyliorum. 1: 147(bis) - 190(bis) ; plts. 41-45 (January) Keen, A. MYRA 1958. Sea shells of tropical west America; marine mollusks from Lower California to Colombia. i-xi+624 pp.; 10 colored plts.; 1700 text figs. Stanford Univ. Press, Stanford, Calif. (5 Dec. 1958) 1966. West American mollusk types in the British Museum (Natural History). II. Species described by R. B. Hinds. The Veliger 8 (4): 265 - 275; plts. 46, 47; 6 text figs. (1 April 1966) PAETEL, FRIEDRICH 1888. Catalog der Conchyliensammlung. Mit Hinzufiigung der bis jetzt publizierten Recenten Arten, sowie der ermittelten Synonyma; Fam. Terebridae. Gebr. Paetel, Berlin; pp. 247 - 255 Pitspry, Henry Aucustus & HerBert NELSoN Lowe 1932. West Mexican and Central American mollusks collected by H. N. Lowe 1929-31. Proc. Acad. Nat. Sci. Phila. 84: 33 - 144; 6 figs.; plts. 1-17; 2 photographs (21 May 1932) Reeve, Lovett 1860. A commentary on M. Deshayes’s revision of the genus Terebra. Proc. Zool. Soc. London 1860 (III): 448-450 (iss. betw. Aug. 1860 and March 1861) Reeve Bros., King William Proc. Zool. Soc. London for 1834 (2): SmiTtH, Epcar ALBERT 1873. Remarks on a few species belonging to the family Terebridae and descriptions of several new forms in the collection of the British Museum. Ann. Mag. Nat. Hist. (4) 11: 262-271 (April 1873) 1880. Descriptions of twelve new species of shells. Proc. Zool. Soc. London for 1880 (III): 478-485; plt. 46 (October 1880) VREDENBURG, E. 1921. Classification of the recent and fossil Terebridae. of India 51 (4):. Geol. Surv. Vol. 22; No. 1 THE VELIGER Page 65 A New Indo-Pacific Terebrid TWILA BRATCHER 8121 Mulholland Terrace, Hollywood, CA(lifornia) 90046 (1 Text figure) In 1969 AN UNUSUAL terebrid was dredged from 300m in Hawaii by E. R. Cross. Though subsequent research showed it to be a new species, there was a hesitancy in describing it on the basis of a single specimen, particularly as the protoconch was missing. Since that time other specimens of the same species have been sent to me for identification from several areas of the tropical Indo- Pacific. Because E. R. Cross was the instructor of my first course in underwater safety and is a longtime friend, be- cause of his years as editor of the Hawaiian Shell News, and because he discovered the first specimen of this new species, I now take pleasure in naming it in his honor. TEREBRDAE Morch, 1885 Terebra Bruguiére, 1789 Terebra Bruguiére, Encycl. Méth. Hist. Nat. Vers 1: xv. Type species by SD (Lamarck, 1799) Buccinum subulat- um Linnaeus, 1767. Recent; Indo-Pacific Terebra elliscrossi Bratcher, spec. nov. Diagnosis: A medium to large sized white Terebra with small fulvous dots, cancellate sculpture, and a double sub- sutural band. Description of Holotype: Shell size medium, color white with small round fulvous dots, usually in pairs, scattered at random; outline of whorls slightly concave with double convex subsutural band, anterior one being smaller; proto- conch missing, but protoconch of paratype having 34 slightly convex embryonic whorls; sculpture of early whorls of teleoconch consisting of narrow subsutural band with small nodes, followed by slightly curved axial ribs; spiral sculpture developing about 3% whorl; posterior end of ribs swelling into nodes, forming second subsutural band about 5" whorl; sculpture of later whorls cancel- late, with spiral and axial cords of about equal strength, forming small nodes at intersections, axial cords beginning at nodes of anterior band; double band occupying about half of whorl; cancellate sculpture continuing on body whorl to row of slightly enlarged nodes at periphery; spiral cords continuing anterior to periphery, axial sculp- ture becoming obsolete; aperture quadrate; columella re- curved, with moderate parietal callus and scarcely visible plication; siphonal fasciole striate, with moderate keel. Dimensions: Holotype 42.0 X 7.2mm. Paratypes from 21.4 X 5.4mm to 82.9 X 13.1mm Figure 1 Holotype of Terebra elliscrossi Bratcher, spec. nov. Page 66 THE VELIGER Vol. 22; No. 1 Type Locality: Honolulu side of Pearl Harbor entrance, Honolulu, Hawaii, 21°17’N; 157°56’ W at 300m, sand and coral rubble bottom; Jeg. E. R. Cross, 10 May 1960 Type Material: Holotype Los Angeles County Museum of Natural History no. 1257. Paratypes: Australian Muse- um no. c111658 (1); British Museum (Natural History) (1); Bratcher collection (2) ; E. R. Cross collection (1) ; Western Australian Museum (1); R. Schoening collection (1) [all above paratypes from 21 to 37.5m, Guadalcanal, Solomon Islands.] B. Parkinson collection (1), New Guin- ea; R. Schelling collection (1), Okinawa at 45m; U. S. National Museum no. 71899 (1), 104 km SW of Cap St. André, Madagascar, 150 - 300m. Discussion: Some individuals of this species have finer sculpture, and the intersections of axial and spiral sculp- ture may be less likely to form nodes at the intersections. The larger specimens tend to become less coarsely sculp- tured in later whorls. Some specimens have many tiny fulvous dots; others have few, scarcely noticeable ones. Terebra elliscrossi should be compared with several other Indo-Pacific species. Terebra waikikiensis Pilsbry, 1920, an endemic Hawaiian species, is also shiny white with pairs of small fulvous dots, but it has a turreted out- line and is smaller, to 35 mm. The dots, always in pairs, are placed at regular intervals. Terebra elliscrossi has a concave outline with convex subsutural band, is larger, to 82.9mm, and the dots, paired or individual, are scat- tered at random. Terebra insalli Bratcher « Burch, 1967, bears some resemblance to T: elliscrosst but has a smaller, more slender beige shell without the fulvous dots. Terebra trisertata Gray, 1834, has a much more slender shell, and that of 7: cumingi Deshayes, 1834, has more numerous and shorter whorls, neither showing fulvous dots. Terebra amanda, also without dots, is longer whorled and has a wider apical angle. Terebra floridana Dall, 1889, a western Atlantic species, has a shell remarkably similar to that of T. elliscrossi, ex- cept that it has more numerous and shorter whorls with no dots, and is beige instead of white. ACKNOWLEDGMENTS I wish to express my gratitude to Brian Bailey of Guadal- canal, Solomon Islands, who collected most of the para- types. I also want to thank E. R. Cross, Brian Parkinson, Ed Schelling, and Bob Schoening for the loan of type material for study, and Dr. A. Myra Keen for reading and evaluating this manuscript, and Bert Draper for taking the holotype photo. Literature Cited BratcHer, Twita & Ropert Donatp Burcu 1967. A new terebrid species with check list of Terebridae from the Red Sea (Mollusca : Gastropoda). The Veliger 10 (1): 7-9; plt. 2 (1 July 1967) Dai, Witt1am HEALEY 1889. Reports on the results of dredging under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-78) and in the Carib- bean Sea (1879-80), by the U.S. Coast Survey steamer “Blake’’, etc. XXIV. Report on the Mollusca—Part II. Gastropoda and Scapho- poda. Bull. Mus. Comp. Zool. 18: 1 - 492; plts. 1-40 (January to June 1889) Desuayes, Gfrarp Paut 1857. Description d’espéces nouvelles du genre Terebra. Journ. de Conchyl. 6: 65 - 102; plts. 3-5 (July 1857) 1834. Enumeration of the species of the genus Terebra with charac- ters of many hitherto undescribed. Proc. Zool. Soc. London for 1834 (xix): 59-63 (25 November 1834) Hinps, Ricuarp BrinsLey 1844. Descriptions of new shells, collected during the voyage of the Sulphur and in Mr. Cuming’s late visit to the Philippines. Proc. Zool. Soc. London for 1843 (XI): 149 - 168 (June 1844) Pirspry, Henry Aucustus 1921. Marine mollusks of Hawaii, VIII-XIII. Proc. Acad. Nat. Sci. Philadelphia 72: 296 - 328; plt. 12; figs. 1-11 (5 January 1921) Vol. 22; No. 1 THE VELIGER Page 67 Lolliguncula panamensis (Cephalopoda : Loliginidae) from the Pacific Coast of Colombia BY H. J. SQUIRES | ann J. H. BARRAGAN? UNDP/FAO Proyecto para el Desarrollo de la Pesca Maritima en Colombia, Bogota (5 Text figures) INTRODUCTION Lolliguncula panamensis Berry, 1911 is strictly a repre- sentative of the Panamic zoogeographical Province (OLs- SON, 1961) and is found only from Baja California to northern Peri. Loliolopsis diomedeae appears to be sim- ilar in this respect although it is reported from deeper water than the former (Voss, 1971). They are both taken in approximately the same depths where. shrimp fishing is carried on off the coast of Colombia and Ecua- i dor. Squids of the genus Lolliguncula are shallow-living ee warm water species (DRacovicH & KELLY, 1967). Water a : temperatures for L. panamensis on the Colombian Pacific ~ Cabo Corrientes coast were 21°-27°C at depths of 5-70m (SgQuIREs o i et al., 1970; 1971), while for L. brevis on the Caribbean = coast of Colombia they were 26° - 29° C at 10 - 35 m (Lo- A, PEZ, 1972). Where L. panamensis is taken the coastal area has fringing mangroves and many long estuaries with dis- | x charges of muddy water from rivers of the tropical rain Anas Juan forest. Secchi disc readings were 0.2 - 4.5m in 5-10m Canes deep, and 3 - 12m in 10 - 70m deep, and salinities about Bahia Malaga \_A Buena- 15 - 23 ppm (Sgumes et al., 1975). Ss aS (adjacent column —) Figure 1 Distribution of squid, Lolliguncula panamensis, along the Pacific coast of Colombia, and names used in the text I Y 9° ———————— Ecuador’ a Present addresses: (Gj ! Peri 3 ' 122 University Avenue, St. John’s, Newfoundland, Canada 2 Instituto Nacional de Pesca, Casillo de Correos 5918, Guayaquil, Ecuador Page 68 Both Lolliguncula panamensis and L. diomedeae were caught incidentally in shrimp trawls during survey fishing for shrimps (Sgumres et al., 1970; 1971). However, only small numbers of L. diomedeae were taken and are not included in this study. Commercially, only the largest of the squids, mainly L. panamensis, are selected during shrimp fishing, and packed and frozen at the processing companies for export to Europe. Total catch reported at Buenaventura in 1970 was 37 tons. During the survey for commercial shrimps and fishes, a chartered 22.5m shrimp trawler used paired trawls, each with an effective opening of about 20m wide and 2m high. Stretched meshes were 40 - 43 mm in the wings and 34 -41mm in the codend of the trawls when wet. Towing speeds were about 3 knots. A series of 5 trawling stations in depths of 10, 20, 35, 55 and 70m, respectively, were done at each of 16 shrimping areas along the coast. Stations north (or south) of Buenaventura were done in alternate months (Sourres ef al., 1970; 1971) during 1969 and part of 1970. In the process of separating shrimps, fishes and various invertebrates in the catches, the few squids were counted, weighed on a commercial balance and preserved in 10% formalin in sea water. After about 18 months in preservative, the squids were examined over a period of about 4 months in the labora- tory at Buenaventura. Each sample was first soaked for about 4 hour in fresh water, and lengths of mantle and tail fins measured on a measuring board marked in mm, or with vernier calipers. Total weight of each was taken on a precision balance (accuracy 0.1g) after draining off excess liquid. Mantles were cut open and head and vis- cera removed. Mantles, gonads and nidamental glands were weighed separately on the balance. Ovaries were examined and ova diameters measured on a mm grid under 10 X magnification. Stomachs were opened in a watch glass and contents examined under magnification of 10 - 30X. IDENTIFICATION The 2 species were separated by the following main characters in mature squids: A, Body short and thick; tail fins forming an ellipse in outline and reaching about 4 the length of the mantle. Only the left ventral arm hectocotylized in adult males. Suckers of arms with 5 teeth and of tentacular clubs Qi = QE MCC CUM ee ceases tctrtsteset Lolliguncula panamensis THE VELIGER Vol. 22; No. 1 A, Body slender; tail fins forming roughly a semicircle in outline and reaching about 4 the length of the mantle. Both ventral arms hectocotylized in adult males. Suckers of arms with 10 - 11 blunt teeth and of ten- tacular clubs 24 squared teeth... Loliolopsis diomedeae BIOLOGY Length-weight Relationships: Females of Lolliguncula panamensis were substantially larger than males (Figure 2) and greater in total weight for their length (Table 1). Average mantle weights were also slightly greater in fe- males than in males at given mantle lengths. Regression equations where L = mantle length were: Mantle Weight = 0.00794 L?®%” in males (N = 93); and Mantle Weight = 0.008667 L?’*> in females (N = 547). Also Total Weight = 0.00288 L? in males (N = 92) ; and Total Weight = 0.004284 L?7 in females ( (BarRAGAN, 1972). Mantle weights were slightly more than 55% of the total weight in males (55-57%) and about 50% of the total weight in females (59 - 50%; Table 1). This differ- ence can be explained by the large ovaries and nidamen- tal glands in females contributing more to the total weight than testes and spermatophore glands in males. Fin length (FL) increased slightly in proportion to mantle length (ML) with increase in size. Regression equations were as follows: FL = 0.521 ML — 0.94 in males (N = 94); and FL = 0.555 ML — 1.14 in females (N = 552) (Figure 3) (BaRRAGAN, 1972) = 550) Growth Estimates: Growth increments in females were estimated from apparent shifts in modes of mantle lengths in frequencies within months and between months from samples taken during January-October, 1969 (Table 2; Figure 4). Average increment estimated within months is 19 mm and between months, 16mm. Three modal clas- ses were apparent in most months with differences of 10 - 36mm (Table 2). Identifying modal shifts between months was based on most of the possibilities between successive months or every second month. Very small or very large apparent shifts were not included (Figure 4; Table 2). Data for males were too few to give definitive results. Vol. 22; No. 1 THE VELIGER Page 69 Table 1 Average Total and Mantle weights of Lolliguncula panamensis and percent mantle weight of total weight at given mantle length (96 males; 549 females). Percent Mantle weight Frequencies Ay. Total weights Ay. Mantle weights of Total weight ML M 18 M F M Je M 8 g g g g te te 34 1 2 pil 3.4 TET 1.7 55 50 38 3 3 4.2 4.6 2.3 2.3 55 50 42 15 6 5.4 6.0 3.0 3.0 56 50 46 23 30 Hall 7.8 3.7 3.9 52 50 30 19 33 8.3 9.7 4.7 4.8 57 49 54 7 53 10.3 12.0 5.8 6.0 56 30 38 8 4] 22S 14.6 Tl 12 57 50 62 ) 32 15.1 17.5 8.5 8.9 56 51 66 3 43 17.5 20.7 10.0 10.3 57 50 70 1 33 21.2 24.3 11.7 12.1 55 50 74 1 39 24.1 28.2 13.6 14.0 56 50 78 38 32.6 16.2 50 82 27 37.3 18.6 50 86 4] 42.5 21.1 50 90 38 48.1 23.9 50 94 46 54.1 26.9 50 98 26 60.6 30.1 50 102 9 67.5 33.6 50 106 8 75.0 35.3 50 110 1 82.9 41.3 50 Table 2 Estimates of increments in mantle length between modes in length frequencies (Fig. 4) of female Lolliguncula panamensis from January-October, 1969. Month Modal No. of Increments Increments Intervals 1969 classes females within groups between months between months mm mm mm Jan 44, 64, 78, 88 63 20, 14, 10 30, 20, 28 Jan-Feb, Jan-Mar Feb 44, 80, 94 27 36, 14 20, 48, 28 Feb-Mar, Feb-Apr Mar 52, 64, 92 138 12, 20 28, 12, 36 Mar-Apr, Mar-May Apr 44, 68, 92 35 24, 24 20, 32, 12 Apr-May, Apr-Jun May 48, 64, 88 59 16, 24 12, 24, 40 May-Jun, May-Jul Jun 76 21 — 12 Jun-Jul Jul 48, 72, 88 83 24, 16 12, 24, 36 Jul-Aug, Jul-Sep Aug 84 47 — = — Sep 52, 76, 92 55 20, 12 24, 16 Sep-Oct Oct 52, 76, 92 24 24, 16 = = Total females 552 Av. 19 mm Av. 16 mm Overall average 17 mm Page 70 THE VELIGER Vol. 22; No. 1 Maturity of Females: Four phases of maturity are de- age of 59% of all females in samples were immature. fined based on ova diameter, weights of ovaries and nida- Size of first mature was 80mm in mantle length. mental glands, and color of ova in preservative (Table According to proportions of Mature or Spent-and-recov- 3). The phases are categorized as Immature or Juvenile, ering in monthly samples, spawning occurred throughout Maturing, Mature, and Spent-and-recovering. An aver- the 10 months during which samples were collected. Ap- pearances of ovaries also indicate that females spawn more than once after maturing (Table 3). The propor- 60 tion of mature females ready to spawn (Phase III) or 50 60 40 $ g 50 Females 5 30 19 £0 4 = 1) 20 ai Males ig ae 5 10 10 oO nee a Coping 36 44 52 60 68 76 84 92 100 32 40 40 5 4 72 go 104 Manlte 1 - Manlte length - mm eee cam Figure 2 Figure 3 Length frequency histogram of 96 male and 552 female Lolligun- Regression of fin length (FL) on mantle length (M) of Lolligun- cula panamensis taken from January to October, 1969, on the cula panamensis. FL = 0.521 ML — 0.94 in 93 males; FL = 0.555 Pacific coast of Colombia f ML - 1.14 in 550 females Table 3 Phases of maturity of female Lolliguncula panamensis from the Pacific coast of Colombia. Mantle Weights of Weights Greater Color Presence length of nidamental of diameter of of Phase Category range glands Ovaries of ova ova oocytes Remarks mm g g g I Immature Pyriform or 32-75 0.1-0.5 0.3-1.5 0.1-0.3 Opaque Yes oocytes and juvenile white small developing opaque ova II Maturing 74-79 0.7-1.8 1.8-3.2 0.5-1.0 Opaque Yes attached toa yellowish dorsal (germinal) strand with III Mature 80-110 1.8-4.0 3.0-8.5 1.0-1.8 Translucent Yes branched yellowish stromae. IV Spent and 80-110 0.3-1.9 0.9-2.5 0.1-1.8 Opaque Yes A few large resid- recovering white but ual ova translu- some cent yellowish yellowish but degenerating. Vol. 22; No. 1 recently spawned (Phase IV) averaged 73% in each month (range 63 -87% with one exception of 41% in May; Table 4). Maturity of Males: In the samples examined all males were maturing or mature (with spermatophores ready for transferral). All had the left ventral arm hectocotylized. The smallest was 34mm in mantle length. Stomach Contents: Most of the squid (81%) with food in their stomachs had remains of fishes: scales, spines, vertebrae, dark integument, muscle tissues, etc. These ap- peared to be fragments of small pelagic fishes common in the area, 7. e., engraulids (Cetengraulis spp.) and clupe- ids (Opzsthonema spp.). Also present (in 15% of the stomachs) were remains of crustaceans, possibly small shrimps of the species Xiphopenaeus riveti and others. As noted by DracovicH « Ketty (1967) for small Lolhigunculus brevis in Florida, BARRAGAN (1972) men- THE VELIGER Page 71 tions that most (70%) of the immature squid examined had empty stomachs, while only a few (24%) of the mature squid were empty. He also found that more (70%) of the early mature or maturing squid had food in stomachs than late matures (40 and 60%, respectively). Males and females of the same size were not significantly different. DISCUSSION Estimation of Stock Potential: Squids caught during survey fishing for shrimps and fishes on the Pacific coast of Colombia, were found throughout the area, but mostly in depths of 5 - 30m (Sgumes et al., 1970; 1971). Ina few parts of the area, such as off headlands, untrawlable grounds may form sanctuaries for squids and some species Table 4 Percent of adult female Lolliiguncula panamensis in maturity Phases III and IV (ready to spawn or recently spawned: Table 3) each month from January-October, 1969, on the Pacific Coast of Colombia. Total No. Total No. of adults Percent in Phases Month examined (Phases II, III & IV) HI& IV Jan 81 34 74% Feb 26 13 87 Mar 141 61 85 Apr 36 18 88 May 63 16 4] June 26 15 87 July 87 22 63 Aug 48 30 83 Sept 59 17 74 Oct 24 11 73 Totals 591 237 Average % 73 Table 5 Incidence of principal food items in stomachs of Lolliguncula panamensis (% of stomachs containing food item) from the Pacific coast of Colombia. Total examined, 468; empty, 280. Principal food items Yo Small pelagic fishes such as engraulids and clupaeids (fragments of scales, vertebrae, dark pigmented integument, 81 muscle tissue, otoliths, etc.) Crustaceans (fragments of mouth parts, limbs, integument) 15 THE VELIGER Vol. 22; No. 1 “) January (63) uo June (21) 10 10 5 rs) mt oe July (83) February (27) 15 15 10 10 5 5 20 March (138) 20 August (47) 15 15 10 10 5 5 20 204 15 April (35) 15 September (55) 10 10 2 5 20 20 15 May (59) is October (24) 10 10 ) 5 32 48 64 80 94 32 48 64 80 94 Mantle Length - mm Page 72 Numbers Figure 4 Monthly frequencies of mantle lengths of Lolliguncula panamensis from the Pacific coast of Colombia, January to October, 1969 Vol. 22; No. 1 THE VELIGER Page 73 x x = , Ne! 70 ROL PRPS] KOSH POSER 60 DSK POLS PSOPS HK E so Kedroperkeord RSKXFSAKHRS BF SOR SBSRSY — PSOSISRAOPN BPSK PRSEIRLPRLIKOSKE Epo PEAR PSO PPR PS 40 PASSO OE hoof PO DoS PSP ERPS RASC P eR PSPSPS Eh oes noe oaea Seen abet sent eens Sons fatal goce PSEA OPH PON SISA POR ISOSOCF 20RD OPP OP PRPS OPP Re Poew OSI OOPO PCRS TOP POP SRO P OPA ooe RRR PSF PRP OR hoot Nee MA MO Se Aly SO 1969 Figure 5 Percentages of ready-to-spawn and recently-spawned female Lollig- uncula panamensis of mature females from January to October, 1969, on the Pacific coast of Colombia of fishes. The total area where squids could be taken ap- proximates 8500 km’. Catches of squids by the chartered shrimp vessel aver- aged 0.5 kg per 1-hour haul, the sweep of the nets being about 0.115km’, and catch per km’ therefore averaged 4.3 kg. If other shrimp vessels caught squids at a similar rate, it should be possible to calculate their total catch from the number of hours they fished in a year. In fact, the fleet of 87 shrimp vessels fished approximately 300 000 hours in 1970, and at a rate of 0.5 kg per hour the total catch would be 150000 kg (150 metric tons; Squires et al., 1971). Assuming that the vessels brought in catches of squid for the last 4 days of a 12 day trip (because of the tenden- cy of squid to spoil on ice), it may be assumed that they landed only 4 of their catch. The reported landings of squid at Buenaventura was 37 tons in 1970. A smaller part of the fleet (4) landed at Tumaco where it could be assumed another 5 tons were landed, giving a total land- ing of 42 tons. On the other hand, calculating the catch of squid for the total area of 8500km? from catch per km’ of 4.3 kg by the survey vessel would give an amount of 36550kg (37 tons). This estimate would assume that 100% of the squid in the path of the trawl were caught. This would be undoubtedly incorrect, since some squids could escape the slow movement of the trawl or through its meshes, and some could be schooling in the water above the nets when preying on pelagic fishes. It is likely that catch- ability of the trawls is much less than 100% for many species, including squids and fishes. Since an estimate of the total catch for the area by the commercial fleet is available (150 tons), it may be pos- sible to estimate the catchability by the following method: Estimated Annual Catch Estimated per Total Area Catchability tons % 37 100 74 5° 148 (150) 25 It may be assumed, therefore, that the catchability of this species by the shrimp trawls used by the fleet and the sur- vey vessel was about 25%. From the latter assumption it is further possible to estimate the biomass of squids in the area using GUL- LAND’s (1970) equation as follows: 1.25 X0.5 X0.6 Xsquid biomass = 150 tons Therefore, the squid biomass = 400 tons The potential annual yield is here assumed to be 150 tons, the natural mortality 0.6 and the catchability 25%. An increase in the rate of fishing for squids could not be rec- ommended on the basis of these estimations, although, if the squid could be frozen immediately when caught, the landings could be increased if the fishing rate remained the same. Growth: Estimates of growth in squids from modal clas- ses, averages, ranges, etc., of mantle length frequencies by various authors have been reviewed and compared by SumMMERs (1971). Estimates for Loligo pealei were 13-15 mm and for Illex illecebrosus 20 - 30 mm per month. Also growth of laboratory-reared squids has been documented by CuoeE (1966) for Sepioteuthis lessoniana and by La- Roe (1971) for S. sepioidea, giving average monthly in- crements of about 21mm in mantle length. In smaller squids (Sepiolinae) estimates were as much as 5mm per month (Bo.etzky et al., 1971). Ina larger squid, Dory- teuthis plei, estimates of 15 - 25mm per month for mantle length increments were obtained in laboratory-reared an- imals (LARoE, op. cit.). In view of the estimates of monthly growth in mantle length for various squids, our estimate from modal clas- Page 74 THE VELIGER Vol. 22; No. 1 ses of an average of 17 mm per month for female Lolligun- cula panamensis appears reasonable between mantle lengths of 40 and 94mm (Table 2; Figure 4). Reproductive Potential: The presence of more than one size of ova in ovaries and an area of oocyte proliferation (germinal strand) suggest that more than 1 batch of eggs is produced by individual females. Also the appearance of the ovaries full of translucent large eggs or spent and recovering with predominantly small maturing ova sug- gests that the eggs are laid in batches. Synchronization in spawning such as reported by TINBERGEN & VERWEY (1945) and LaRoe (1971) is indicated in this species by bimonthly peaks (Figure 4), although samples and dif- ferences are small. An average of 73% ready-to-spawn or recently-spawned (Phases III and IV, Table 3) in each of 1o monthly samples would indicate spawning every 14 to 2 months (Squires, 1973). The uniformly high temperatures throughout the year would most likely influence the rate of oogenesis as they affect the rate of embryogenesis (Mc- Manon & SUMMERS, 1971). Stomach Contents as Evidence of Feeding Behavior: Fish fragments occurred in most stomachs indicating that the main prey is small pelagic fishes. The latter are very abundant, although only lightly exploited by industry (about 1000 tons annually are made into fishmeal or canned for food; Oscar Arroyo, personal communication). Species of Opisthonema and Cetengraulis are apparently the most abundant, but several species of anchovies, silver- sides, etc., are also common. Some pelagic fishes are regu- larly taken in shrimp trawls, especially in shallow water, but they school mostly near the surface and great numbers tend to escape the trawl. Similarly, their predators, such as the squids, would also escape bottom trawls to a large extent. Squid stomachs had little evidence of bottom feed- ing such as mud particles reported for Lolliguncula brevis by DracovicH & KELLY (1966). Size Differences in Males and Females: Differences in mantle lengths and weights between males and females even before maturity reflect early dimorphism in this spe- cies. The few large females at the end of the size range (Table 1; Figure 2) may be caused by increased mortality at these sizes since the decrease in numbers is abrupt. In males, on the other hand, the decrease is noticeably more attenuate (Figure 2), perhaps as a result of increasing ability to escape the trawls. However, it is apparent that they do not grow any larger than samples indicate. They appear to mature at a much smaller size than females. ACKNOWLEDGMENTS We appreciate the assistance of Dr. Gilbert Voss of the University of Miami School of Marine and Atmospheric Science in identifying Loliolopsis diomedeae. We are also indebted to many of our colleagues in Colombia and the captains and crews of survey vessels for assisting in col- lections of squids at sea, and to INDERENA for labora- tory facilities at Buenaventura. Literature Cited Barracan, J. H. 1972. Contribucion al conocimiento biologico del calamar del Pacifico colombiano, Lolliguncula panamensis Berry. Tesis Licenc., Funda- cion Univ. Bogota (JTL): 61 pp. BoLetzky, SicurD voN, MaRIA VERENA VON Botetzky, D. FroscH & V. GAtzi 1971. Laboratory rearing of Sepiolinae (Mollusca: Cephalopoda). Mar. Biol. 8: 82 - 87 Cuog, S. 1966. On the eggs, rearing, habits of fry and growth of some cephalo- pods. Bull. Mar. Sci. 16: 330 - 348 DracovicH, A. & J. A. Keriy 1967. A biological study and some economic aspects of squid in Tampa Bay, Florida. Bull. Mar. Sci. 17: 87 - 102 Gutianp, J. A. 1970. The fish resources of the ocean. FAO Fish. Tech. Paper 97: 425 pp. LaRog, E. T. 1971. The culture and maintenance of the loliginid squids Sepioteuthis sepioidea and Doryteuthis plei. Mar. Biol. 9: 9 - 25 Lopez, N. 1972. Loliginidae del Caribe colombiano. Tesis Lic., Fundac. Univ. Bogot4é (TJL): 99 pp. McManon, J. J. W. C. Summers 1971. Temperature effects on the developmental rate of squid (Loligo pealet) embryos. Biol. Bull. 141: 561 - 567 Otsson, Axe, ADOLF 1961. Mollusks of the tropical eastern Pacific, particularly from the southern part of the Panamic-Pacific faunal province (Panama to Peru). Panamic-Pacific Pelecypoda. Paleont. Res. Inst. Ithaca, N. Y, 574 pp., 86 pits. Squires, H. J., A. Ben-Tuvia, O. Mora, O. Barona & O. ARROYO 1970. Preliminary results of cruises 6901-06 of the chartered shrimper “Cacique.” Prodepesca Estud. Investig. 2: 97 pp. Squires, H. J., M. Estevez, O. Barona & O. Mora 1975- Mangrove cockles, Anadara spp. (Mollusca : Bivalvia) of the Pacific coast of Colombia. The Veliger 18 (1): 57-68; 14 text figs. (1 July 1975) Squires, H. J., O. Mora, O. Barona & O. ARRoYO 1971. Results of cruises 6907-11 and 7001 of the chartered vessel “Ca- cique” on the Pacific coast of Colombia. Prodepesca Estud. Invest. 5: 41 pp. Summers, W.C. 1971. Age and growth of Loligo pealei, a population study of the com- mon Atlantic coast squid. Biol. Bull. 141: 189 - 201 TINBERGEN, Lukas & J. VERWEY 1945. Zur Biologie von Loligo vulgaris Lam. 7 (1, 2): 213-286 Voss, GILBERT LINCOLN 1971. Biological results of the University of Miami deep-sea expedi- tions. Cephalopoda collected by the R/V “John Elliott Pilsbury’ in the Gulf of Panama in 1967. Bull. Mar. Sci. 21: 1-34 Arch. Neerland. Zool. Vol. 22; No. 1 THE VELIGER Page 75 An Attachment Structure in an Epiparasitic Gastropod BY DANIEL L. HOFFMAN Department of Biology, Bucknell University, Lewisburg, Pennsylvania 17837 (1 Text figure) AN ECTOPARASITIC LIFE STYLE generally involves the de- velopment of specialized organs of attachment, as in the case of flatworms and arthropods where an array of simple to elaborate hooks, suckers, and spines has been well doc- umented. However, among gastropod mollusks, not only is the ectoparasitic habit rare, but the development of specialized attachment structures has only been reported in one species of prosobranch mollusk (VERRILL, 1897). No attachment organ with the possible exception of the foot has ever been noted in any of the ectoparasitic gastro- pods. The opisthobranch gastropod family Pyramidellidae consists of ectoparasites that feed on the body fluids of other mollusks and polychaete worms using a stylet on the end of a protrusible proboscis (FRETTER & GRAHAM, 1949). CLARK (1971) reports that the pyramidellan, Odostomia columbiana (Dall « Bartsch, 1909), responds to the surface texture of its host, the mesogastropod, Trich- otropis cancellata (Hinds, 1848). Also, the heavily spiny periostracum is highly attractive to this parasite, and has positive adaptive value in that it prevents the parasite from being dislodged by strong currents. Yet the para- sites are seen to move freely over all parts of the shell and also are able to move from one host to another. While in residence at the Friday Harbor Laboratories during the spring of 1978, I had the opportunity to study host-parasite relationships between these 2 species of gas- tropods. The snails were collected in what is called the “potato patch” of East Sound, off Orcas Island, Washing- ton, in approximately 30m of water. Approximately 52 specimens of Trichotropis cancellata were dredged and 46% were found to be infected with Odostomia columbi- ana. The mean number of parasites per host equaled 1.62 + 0.64 S. D. The snails were maintained in the lab- oratory in a 30 gallon [108 L] capacity plexiglass aquari- um that was supplied with running sea water. In the first attempts to remove some of these small pyramidellans from the surface of their host, one slipped from the grasp of the forceps. This was a most fortuitous accident, for the Odostomia did not fall to the bottom of the sea water-filled bowl, but was caught on the edge of the dorsal lip of the host’s shell, approximately 7mm from where it originally rested. It appeared to be held in place by a thread that was still attached to its former position on the surface of the shell. Removing the Tricho- tropis from the water, the attachment thread of the para- site became apparent under the dissecting microscope; so it appears that the thread has an optical density similar to that of sea water, since it is relatively invisible while under water. The attachment thread seems to be quite elastic and resilient, for when the Odostomia was care- fully nudged away from the edge of the shell of Tiricho- tropis with the tip of the forceps, it snapped back to its original position on the surface of its host. Stretching tends to reduce the elasticity of the thread, and if it is stretched more than 15 mm, the thread will break and the parasite will lose contact with its host. To test whether the para- sites could re-establish their attachment threads, the fol- lowing experiment was undertaken. Twelve Odostomia were removed from the surfaces of a number of Tricho- tropis, thus severing the attachment threads, and then placed on the surfaces of new hosts. By the following day, g of the parasites had re-attached, forming new threads. An aqueous saturated solution of neutral red selectively stains the attachment threads, making them readily vis- ible. Staining a shell of Trichotropis from which 2 Odo- stomia had previously been removed demonstrated at least 6 attached fragments. Since more attached fragments were visible than the number of parasites removed, it appears that the parasites are able to sever their own threads and produce new ones as they move about their hosts in a fashion similar to mussels utilizing their byssal fibers. The thread originates from the extreme posterior ven- tral surface of the foot (Figure 1). The margin of the posterior region is plicate, and there appears to be a median central pore from which the thread emerges. In addition, there is a ciliated median groove running up the antero-posterior axis of the foot terminating near the tip of the propodium. The function or role of this groove Page 76 was difficult to determine and cannot be answered here. La FotteTtrE (1977) has noted in dead and dried Chrysallida cincta, also a pyramidellid, on Norrisia nor- risii that the snails remain attached to their hosts. They may have been held in place by dried attachment threads. Normally, in gastropods, the foot is the major adhesive organ. What adaptive function might be served by pos- sessing an additional attachment structure? This question is best answered by looking at the habitat of the host and the feeding behavior of the parasite. YONGE (1962) re- cords that Trichotropis cancellata is restricted to hard bottomed subtidal substrates, consisting mostly of dead bivalve shells, that are repeatedly buffeted by strong tidal currents. A suspension feeder, it is relatively sedentary by habit in that it moves as high as possible on a vertical surface and stays there. When feeding, Odostomia colum- biana stations itself near the very edge of the aperture of the host’s shell in order to reach the soft mantle tissues with its long proboscis. This appears to be a precarious position, even on a relatively sedentary host with a spiny periostracum, given the force of the turbulent water and the vertical orientation of the host. The development and evolution of the attachment thread as an additional “life line” has positive adaptive values for O. columbiana. Such attachment threads in gastropods may be more common than first realized. VERRIL (1897) first noted threads of adhesive mucus formed by the foot glands of many land slugs and certain marine gastropods as well. He reports that the sargassum snail, Litizopa bombyx (Rang, 1829), attaches itself to its floating algal home by a thread of adhesive mucus. This was later verified by WALLER (1975) in L. melanostoma, a synonym of L. bom- byx (Asspott, 1974). Cadet Hand (personal communi- cation) has observed the dove shell, Mitrella carinata (Carpenter, 1865), that inhabits the rocky intertidal zone of the Pacific coast of the United States attaching it- self to the substrate by a thread of mucus secreted by the foot. In addition, I have observed on at least 7 different occasions while snorkeling in the Florida Keys the bubble Shell, Bulla striata (Bruguiére, 1792), hanging from a mucus thread after having dislodged them from a verti- cal facing wall. Dr. Hand feels that such forms of attach- ment may be size-related in that all the gastropods that demonstrate this phenomenon are quite small, generally less than 12mm in length. VERRILL (op. cit.) has hypo- thesized an evolutionary relationship between mucus threads and the byssal threads of the bivalve mollusks. He suggests that the earliest form of attachment was temporary, perhaps being aided by mucus secretion from the surface of the foot. He notes that “such a mode of adhesion to objects is common among planarians, small nemerteans, annelids, and the young forms of many groups at the present time” ; also, “from such a primitive adhesive THE VELIGER Vol. 22; No. 1 foot the transition to a larger foot with more specialized cells situated in a groove for the secretion of stronger byssus-like threads of mucus would have been easy.” Thanks are in order to Dr. A. O. D. Willows, Director of the Friday Harbor Laboratories, for the many facilities afforded me while in residence at the laboratories. Figure 1 Ventral and lateral perspectives of overturned specimens of Odostomia columbiana showing relationships of attachment thread (at) to the foot o — operculum t — ear-like tentacle Snails are approximately 2mm in length Vol. 22; No. 1 THE VELIGER Page 77 Literature Cited Assott, Rosert Tucker 1974. | American seashells. end ed., 663 pp.; 24 col. pits. Van Nostrand Reinhold, New York Crark, Kirsten 1971. Host texture preference of an ectoparasitic opisthobranch, Odo- stomia columbiana Dall « Bartsch, 1909. The Veliger 14 (1): 54-56; 1 pit. (1 July 1971) FRETTER, VERA & ALASTAIR GRAHAM 1949. ‘The structure and mode of life of the Pyramidellidae, parasitic opisthobranchs. Journ. Mar. Biol. Assoc. U. K. 28: 493 - 532 La Fotiette, Patrick I. 1977. Inbreeding and intraspecific variation in Chrysallida Carpenter, 1857 (Gastropoda: Pyramidellidae) . Ann. Reprt. West. Soc. Malac. 10: 18- 23 (14 December 1977) VERRILL, ADDISON EMERY 1897. A study of the family Pectinidae, with a revision of the genera and subgenera. Trans. Connect. Acad. Arts & Sci. 10: 41-95 Wa tter, THomas R. 1975. The behavior and tentacle morphology of pteriomorphian bi- valves: a motion picture study. Bull. Amer. Malacol. Union 7-13 Yonoe, CHartes Maurice 1962. On the biology of the mesogastropod Trichotropis cancellata Hinds, a benthic indicator species. Biol. Bull. 122: 160-181 Page 78 THE VELIGER Vol. 22; No. 1 Predation upon Crassostrea virginica (Gmelin) Larvae by Two Invertebrate Species Common to Chesapeake Bay Oyster Bars’ BY P D. STEINBERG* anv V. S. KENNEDY University of Maryland, Center for Environmental and Estuarine Studies Hom Point Environmental Laboratories, Cambridge, Maryland 21613, U.S.A. INTRODUCTION THORSON (1950, 1966) pDIscUSSED the high mortality of marine planktotrophic larvae and the dominant role played by predators in this mortality. MiLerkovsky (1974) expanded on this topic, concluding that the ecological significance of such predation was still unclear with more reliable quantitative data necessary. KorRINGA (1941) found that predation was the major cause of mortality in larvae of the European oyster (Ostrea edulis Linnaeus, 1758). Mature larvae of the American oyster, Crassostrea virginica (Gmelin, 1791), tend to be concentrated on or near the bottom of estuaries during ebb tide and slack water periods (CARRIKER, 1967; Woop & HarcIs, 1971). Further, settlement of pediveliger larvae of oysters has been shown to be gregarious in nature (Hu, 1969; Hipu et al., 1970), presumably resulting in aggregations of lar- vae over suitable substrate in response to a water-borne pheromone (VerrcH & Hu, 1971). Such gregarious be- havior should increase mortality rate of larvae due to predation by benthic organisms, especially those inverte- brates inhabiting shell surfaces of oyster bars which are major areas of oyster larval settlement (Crisp, 1967). In order to begin to establish significance of predation by such benthic invertebrates in Chesapeake Bay, in the summer of 1977 we investigated 2 common species, name- ly a sea anemone, Diadumene leucolena (Verrill, 1866) and a barnacle, Balanus improvisus Darwin, 1854. Diadumene leucolena is found on oyster beds through- out Chesapeake Bay, occasionally occupying 15-25% of t Contribution No. 944 from Center for Environmental and Estu- arine Studies, University of Maryland 2 Present address: Department of Biology, University of Califor- nia at Santa Cruz, Santa Cruz, CA 95064 the surface area of live oyster shells (ConEs & HavEN, 1969). Predation by sea anemones may be restricted mainly by their ability to seize and swallow prey, as annelids, mol- lusks and crustaceans can all be ingested (STEPHENSON, 1928). WiLtiaMs (1972) found the main components of gut contents of Diadumene luciae (Verrill) to be amphi- pods, isopods and copepods. MacKenzie (1977) recently reported preliminary observations that D. leucolena from Chesapeake Bay will feed on mature oyster larvae. We wished to quantify predation intensity, feeding rates and digestion times for this sea anemone. SouTHWARD (1955), BARNES (1959) and Crisp (1964) indicated that barnacles feed on a wide variety of plank- tonic organisms ranging in size from flagellates to small crustacea. Maturing oyster larvae may range in size up to 300 um (GALTSOFF, 1964; CHANLEY & ANDREWS, 1971), placing them within the reported size range of ingested material. Balanus improvisus is very common in Chesa- peake Bay (Wass, 1972) often settling thickly on oyster shell. It should have every opportunity to prey on oyster larvae as they occur over oyster beds. We wished to test the assumption that B. improvisus will ingest mature oyster larvae. We report here the results of our study of predation on mature Crassostrea virginica larvae by these 2 common invertebrate inhabitants of Chesapeake Bay oyster bars. MATERIALS anp METHODS Adults of both species were collected by oyster dredge in the Choptank River, Maryland (approximately 38°40’ N; 76°10’ W) during the summer of 1977. Pieces of shell carrying Diadumene leucolena or Balanus improvisus were chipped off oysters. Only sea anemones which remained Vol. 22; No. 1 THE VELIGER Page 79 attached to a shell fragment were used in experiments. This was generally true for B. improvisus as well, although occasionally barnacles not attached to a shell fragment but retaining their calcareous basis were used. All animals were maintained in aerated Choptank River water in the laboratory with collection and experimental salinities ranging from g to 11%, over the summer. Experimental water temperatures were kept at 21° to 22°C, a few degrees Celsius below collection temperatures. Oyster larvae were obtained from our laboratory’s shellfish hatchery. Pediveliger or “eyed” larvae were collected on a 177 um mesh screen and the younger umbo larvae on an 88 um mesh screen. Water in which larvae were held was first filtered through a 1oum filter. Over long holding periods, oyster larvae were fed cultured algae. During our experiments larval numbers were estimated as follows: Immediately after the water in their con- tainers was mixed thoroughly with a non-rotary move- ment, a 1mL sample was taken and the larvae therein were counted in a Sedgewick-Rafter cell. This was re- peated 10 times and a mean value was determined and used to extrapolate to the number of larvae in the volume of water being sampled. When different prey densities were needed, they were obtained by concentrating larvae in a volume of water to a known larval density and then pipetting appropriate quantities of this “stock solution” (after brisk agitation) to filtered water to provide for the experimental densities desired. Observations were made on individuals of each of the invertebrate species to observe their behavior in the pres- ence of oyster larvae. In all experiments, individuals used were not fed in the 24h period from time of collection until feeding observations began. Each invertebrate was placed in the experimental containers (11 to 20cm dia- meter glass bowls) at least 6 to 18h before larvae were introduced. Observations were made using a dissecting microscope at a magnification of 15 to 20. Experiments were made on each of the species as follows: Diadumene leucolena Ingestion of Larvae: Observations were made over 15 to 60 min on feeding behavior of individual sea anemones in the presence of umbo and pediveliger oyster larvae and details were noted. During experimentation, 3 arbitrary size classes of Diadumene leucolena were used, based on pedal diameter of the attached animal: small (2 - 4mm), medium (5 -8mm) and large (g- 11mm). Two experiments were performed to study larval in- gestion using varied densities of predator and prey. The first experiment lasted 48h and involved pediveliger den- sities of approximately 0.1, 0.4 and 1.6 larvae mL" and the presence of either 2 medium sea anemones or 1 small, 1 large and 3 medium sea anemones. Controls contained no sea anemones. The second experiment lasted 24h with larval densities of approximately 1.6, 3.2 and 6.4 larvae mL" and predator densities of 1 or 2 medium sea anem- ones. At the end of each experiment, numbers of surviving larvae were determined by subsampling and in addition, the shell fragments and walls and floors of the containers were examined carefully for settled larvae (none were found). Feeding Rates: Umbo or pediveliger larvae were added to 200 mL of water in 11 cm bowls containing single Dza- dumene leucolena which were then observed during feed- ing. Number of larvae ingested by sea anemones was counted over a 10min period, beginning immediately after 1 larva was seen to be eaten. Ingestion was deter- mined to occur when a larva travelled far enough down a sea anemone’s pharynx to become lost from view. Size of sea anemone, larval density and larval stage were varied in these experiments. Digestion Times: Sixteen individual sea anemones of varying sizes were allowed to ingest at least 10 pediveligers while under observation. They were then placed in water in clean bowls which were checked every 30min for empty pediveliger shells. Time to appearance of shells was noted in each case. Egestion and Gut Content Analysis: Forty-five sea anemones freshly dredged from the Choptank River were isolated on oyster-shell fragments, rinsed gently and then placed in individual clean glass bowls. Six to 12h later, the shell fragments and bowls were examined for the presence of larval shells as evidence of predation on bi- valve larvae in the field. In addition, gut contents of 1 lab- oratory-fed and 10 additional freshly dredged sea anem- ones were examined after dissection. Balanus improvisus Ingestion of Larvae: Two arbitrary size classes of bar- nacles were established based on greatest opercular dia- meter: small (2-4mm) and large (5-8mm). Known densities of umbo or pediveliger larvae were added to 11cm bowls containing single barnacles which were beating their cirri. Control bowls contained no barnacles. After 18h, the barnacles were removed, their shells and bowl surfaces were searched for settled oyster larvae and numbers of larvae surviving were determined by sub- sampling. Page 80 Egestion and Gut Content Analysis: Twenty-five freshly dredged barnacles were rinsed and placed in individual clean glass bowls which were examined 10 to 12h later for shells of bivalve larvae. If feces were present, they were examined for larval shell fragments resulting from mastication by the barnacles (Nicot, 1967). If no feces were present, the barnacles’ gut contents were examined after dissection. An additional 10 barnacles were allowed to sit and beat their cirri in a container containing pedi- veligers with a density of 20 larvae mL” and their feces or gut contents, or both, were examined at intervals over a Io to 12h period. Field Population Samples: We wished to determine densities of Diadumene leucolena and Balanus improvisus on oyster bars in Maryland’s portion of Chesapeake Bay to estimate the potential for predation in nature. Sixteen oyster bars were sampled using a towed oyster dredge. Of the dredged material collected from each oyster bar, 30 L were shoveled into a container and the contents of the con- tainer (live oysters, shell, shell fragments) were then pro- cessed immediately by close examination of all surfaces of the dredged material. Total numbers of D. leucolena were readily obtained. Because of high densities of small barnacles, no attempts were made to differentiate between B. improvisus and B. eburneus Gould, 1841, another bar- nacle found in some parts of the Bay that we sampled; our data are counts of barnacles in general. RESULTS Diadumene leucolena Feeding Behavior: Introduction of oyster larvae elicited pre-feeding behavior in Diadumene leucolena similar to that described by McFar.an_e (1970) in Tealia felina. In D. leucolena, this behavior consisted of widening of the oral disc, raising and spreading of tentacles, and protru- sion and opening of the actinopharynx. This pre-feeding response was quite stereotyped except for actinopharynx protrusion, which did not occur in some animals. Feeding behavior in Diadumene leucolena was similar to that of D. luciae as described by WituiaMs (1972), and his terminology is used here. The feeding response consists of 3 distinct actions: snatch, tentacular response, and oral response. The snatch occurs as a larva contacts and adheres to a tentacle which contracts quickly, al- though not completely. There is often a movement by the distal end of the tentacle to encircle the larva. A snatch response to a single larva never involved more than 2 THE VELIGER Vol. 22; No. 1 tentacles. The tentacular response follows the snatch and consists of the tentacle bringing the larva to the oral disc, which begins to expand. The oral response occurs as the larva is transferred from the tentacle to the lips of the actinopharyngeal region, with the latter expanding out- ward toward the tentacle as the mouth opens. The larva is transported inside the anemone with the transport sys- tem almost certainly being the pharyngeal cilia, although these were too small to be seen. An egestion response was observed in Diadumene leuco- lena that was similar to the oral response, with the direc- tion of transport reversed. The mouth opened, although to a smaller extent than in feeding, and the egested larval shells were transported to the edge of the oral disc where they dropped to the substrate. Whenever more than 1 shell was egested at one time, the shells clung together in a clump. Ingestion of Larvae: Table 1 contains results of the 2 ingestion experiments. Sea anemones fed heavily on pedi- veliger larvae as few larvae survived in the bowls with sea anemones whereas survival in control bowls was always much higher, usually by an order of magnitude. Decreased larval numbers in controls were presumably due to natural mortality and inevitable errors in sampling when adding larvae at the start of the experiments and counting sur- vivors at the end. Variations in larval density and sea anemone size or numbers did not affect the results. Feeding Rates: Results of feeding rate experiments are contained in Table 2. About 25% of the sea anemones initially used did not feed and were not considered when computing feeding rates. Rate variations were large, as would be expected for an individualistic process such as feeding. Rates were compared using Students’ t-test after we had determined that variances were homogeneous as indicated by results of the Fmax-test (SokaL & RoHLF, 1969). Feeding rate increased as larval density increased, with the differences statistically significant for medium individuals feeding on umbo larvae (P< 0.05; experi- ments 3 and 4) and pediveligers P< 0.01; experiments 7 and 8). Medium-sized individuals generally fed at a higher average rate than did smaller sea anemones, except in the presence of low densities of umbo larvae (experi- ments 1 and 3) ; however, these rate differences with size were statistically significant only for sea anemones feeding on high densities of pediveligers (P < 0.05; experiments 6 and 8). Small sea anemones ate fewer umbos min™ than pediveligers min* whereas medium sea anemones ate fewer umbos min™ than pediveligers min™; these differ- ences were not statistically significant (P > 0.05). Vol. 22; No. 1 THE VELIGER Page 81 Table 1 Crassostrea virginica and Diadumene leucolena. Ingestion of pediveliger larvae under varying conditions of predator and prey density. Controls—no sea anemones. I. 48 h experiment. Water volume = 1.75 litres. Numbers of sea anemones per container: A—Two medium (5-8 mm pedal diameter); B—One small (2-4 mm), one large (9-11 mm) and three medium. II. 24 h experiment. Water volume = 0.25 litre. A—one medium sea anemone; B—‘Iwo medium sea anemones. Larval numbers i Replicate 1 Replicate 2 Approximate Anemone pediveliger density density Start Finish Start Finish I. 0.1 ml—! A 174 20 = — B 174 3 = = Control 174 118 — a 0.4 ml—! A 740 2 702 1 B 740 4 702 1 Control 740 697 702 102 1.6 ml—! A 2775 6 2808 69 B 2775 6 2808 20 Control 2775 2147 2808 1006 II. 1.6 ml—! A 400 13 400 3 B 400 1 400 2 Control 400 131 400 118 3.2 ml—! A 799 ~ 16 799 0 B 799 77 799 7 Control 799 665 799 432 6.4 ml—! A 1598 il 1598 38 B 1598 2 1598 25 Control 1598 826 1598 812 Table 2 Crassostrea virginica and Diadumene leucolena. Feeding rates of two sizes of sea anemone on two stages of oyster larvae. Single anemones in 200 ml of estuarine water were observed for 10 min after feeding began. N = number of replications. Feeding rate = average number of larvae eaten per minute + | standard deviation. Anemone Pedal Larval density Feeding rate Experiment diameter (mm) N Larval stage (ml—?!) (min—!) 1 2-4 3 umbo 1 1.0 £0.8 2 2-4 4 umbo 4 2.4+2.9 3 5-8 8 umbo 1 0.7 = 0.6 4 5-8 7 umbo 4 2:8)==n1k9 5 2-4 3 pediveliger J 0.6 = 0.7 6 2-4 4 pediveliger 4 Hel ss31N 7 5-8 8 pediveliger 1 ie 22 2 8 5-8 7 pediveliger 4 4.9 + 2.6 Page 82 THE VELIGER Vol. 22; No. 1 Digestion Times: Digestion time was considered to be the interval from the end of feeding to the first appear- ance in the experimental container of empty larval shell(s}. A mean digestion time of about 4h resulted. Results were generally consistent, with 8 out of 16 meas- urements falling in the 3.5 to 4.5h range. The shortest interval of appearance of empty shells was 2.5 - 3.0h; the longest was 5.0-5.5h. To check for digestion, the egested shells were exam- ined. Of 163 pairs of hinged, intact shells examined, 133 (82%) were completely clear and empty. Of the re- maining 30 pairs which retained some internal color or texture, none appeared to contain whole larvae. Gut Content and Egestion Analysis: The gut of the 1 laboratory-fed sea anemone examined contained 50 pedi- veligers. Gut contents of 10 sea anemones freshly dredged from the Choptank River contained no bivalve larvae. Egested material from another 45 freshly dredged speci- mens collected at a later date yielded 1 complete pair of bivalve shells (comparable in size to those of oyster pedi- veligers) from each of 3 sea anemones. Balanus improvisus Ingestion Experiments: Results of these experiments are contained in Table 3. Compared with controls, larval Table 3 Crassostrea virginica and Balanus improvisus. Effects of two sizes of barnacles on numbers of two stages of oyster larvae. Single barnacles left in presence of larvae in 200 ml of estuarine water for 18 h. Control bowls contained no barnacles. N = number of replications. Larval numbers at end of experiment are reported as mean = | standard deviation. Larval numbers Barnacle opercular Larval Experiment diameter (mm) stage 1 2-4 umbo 2 5-8 umbo. 3 Control umbo 4 2-4 pediveliger 5 5-8 pediveliger 6 Control pediveliger N Start Finish 8 404 74 = 38 8 404 51 = 60 8 404 130 = 62 8 394 166 +102 8 394 == 7 8 394 273 = 39 Table 4 Diadumene leucolena and barnacles. Relative abundances on selected Chesapeake Bay oyster bars. Sample size per bar = 30 litres of oyster shell and associated material. Ovster bar Approximate location Diadumene leucolena Barnacles Swan Point 38°08'N: 76° 18'W 1 3583, Buoy Rock 39°00’: 76° 13’ 3 15943 Hood 38° 56’: 76°14’ 91 1289 Hollicutt Noose 38°51": 76°21’ 282 463 Cook Point 38° 39’: 76°17' 377 78 Deep Neck 38° 44": 76°15’ 250 782 Double Mills 38° 44’: 76°08’ 52 133 Horn Point 38° 36’: 76° 08’ 75 378 Green Marsh 38°35’: 76° 04' 48 458 Norman 38° 15'; 76°07' 235 547 Middleground 38°14': 75°55’ 495 1049 Georges 38°08": 75°50! 244 64 Marumsco 37°57": 75° 44’ 287 749 Sandy Pt. North 39°01’: 76° 23' 345 4282 Saunders 38° 53’: 76° 29' 36 1599 Cornfield Harbor 38° 03': 76° 20’ 153 873 Vol. 22; No. 1 numbers decreased markedly in bowls containing bar- nacles. Homogeneity of variances was established as before and t-tests indicated that large barnacles had a statisti- cally significant effect on both umbo larvae (P< 0.05) and pediveliger larvae (P< 0.001) compared with con- trol values. Small barnacles significantly affected only pediveliger numbers (P < 0.05). However, upon compar- ing effects of barnacle size on survival of umbo or pedi- veliger larvae (experiment 1 vs. 2; 4 vs. 5), no statistical differences were noted (P > 0.05). Egestion and Gut Content Analysis: Examination of gut content and feces of 25 freshly dredged Balanus im- provisus revealed no evidence of bivalve larvae ingestion, nor were any bivalve shells noted in the bowls. For the 1o barnacles active in the presence of pediveligers in the laboratory for up to 12h, we noted the following: more than 30 semidigested larvae in the gut of a large barnacle examined 1 h after initial exposure; no evidence of larvae in the gut of a small barnacle examined 1h after initial exposure; no evidence of larvae in the feces and gut contents of the 7 barnacles examined 10 to 12h after initial exposure; and 2 dead pediveligers inside the shell plates of a large barnacle examined 30 min after initial ex- posure to larvae. Field Population Samples: ‘Table 4 contains the results of our survey of 16 oyster bars conducted on July 19 and 20, and August 8, 1977. Numbers of sea anemones present varied from 1 to 495 per 30 L sample of dredged material. In general, barnacles predominated, averaging 92% of the animals counted. On some oyster bars (e. g., Buoy Rock Bar), barnacles were extremely abundant, covering most of the available shell substrate. On other oyster bars (e. g. Cook Point, Georges) their numbers were relatively low. DISCUSSION Few reports exist regarding field densities of late-stage Crassostrea virginica larvae, and their gregarious nature in setting makes estimates ambiguous. Hipu « Haskin (1971) reported a 200L sample taken in Delaware Bay to contain 4786 “eyed” larvae (0.024mL™"). NELson (1924) reported densities in his samples of up to 250 mature larvae L* (0.25mL"). While these reported densities are comparable to the lowest larval densities used in our ingestion experiments with Diadumene leuco- lena, it was shown in our experiments that sea anemones would almost completely eliminate larvae over the range of 0.1 to 6.4 larvae mL" (Table 1). Larvae are used as food as evidenced by egestion of empty shells. We ex- pect that, as oyster pediveligers aggregate (whether pas- sively or actively) over oyster shell preparatory to setting, THE VELIGER Page 83 sea anemones exact a toll, just as they did at the densities used in our experiments. Balanus improvisus (and pre- sumably other species of barnacles in Chesapeake Bay) would appear to have a similar effect. Presence of semi- digested larvae in one large barnacle exposed to pedi- veligers would indicate that the decline in larval numbers in the presence of barnacles (Table 3) is not just due to physical damage caused by contact with beating cirri but also involves feeding. The few findings of oyster larvae in the guts of freshly dredged specimens of the invertebrate species cannot be construed as evidence that only limited ingestion of oyster larvae occurs in nature. We did not measure densities of oyster larvae in the water column over the oyster beds at the time of collection and our field collections were limit- ed in number and over time. We expect that our laboratory observations demonstrating ingestion of oyster larvae by Diadumene leucolena and Balanus improvisus can be ex- trapolated to the field, although obviously these predators are exposed to a mixed food resource composed of more than just oyster larvae. There is no published evidence regarding selective feeding on oyster larvae in the pre- sence of such a mixed resource. Nevertheless, when one considers the gauntlet of waving tentacles of sea anem- ones, beating cirri of barnacles, and filtering currents of adult oysters and an additional fouling species, the hooked mussel Ischadium recurvum (Rafinesque, 1820) (ReEicH- ARDT, 1977) it becomes obvious that loss due to predation may be very high near or at the time of settlement by oyster larvae. SUMMARY Predation upon umbo and pediveliger larvae of Crassost- rea virginica (oyster) by Diadumene leucolena (sea anem- one) and Balanus improvisus (barnacle) was studied in the laboratory. Pre-feeding and feeding behavior of D. leucolena in the presence of oyster larvae was described. Few larvae survived in the presence of sea anemones. As larval density increased, sea anemones’ feeding rates in- ‘creased, with larger individuals generally feeding at a greater rate than smaller individuals. At 21° C to 22°C, larval shells were expelled about 4h after ingestion of liv- ing larvae by sea anemones had occurred, on average. In the presence of B. improvisus, numbers of surviving larvae decreased significantly. Pediveliger larvae were found in the gut of a barnacle which had been kept in their pre- sence for 1h. Examination of guts of very limited num- bers of freshly dredged individuals from the field provided evidence of bivalve larval shell in D. leucolena, but not in B. improvisus. Page 84 THE VELIGER Vol. 22; No. 1 ACKNOWLEDGMENTS This research was supported by the National Science Foundation, Undergraduate Research Participation Pro- gram, Grant Number SM176083577. We are grateful to Dr. G. E. Krantz, J. Perdue and D. Meritt for providing larvae and advice, to H. Reichardt for assistance in the field and for discussion, and to various reviewers for crit- ical reading of the manuscript. Literature Cited Barnes, Harotp 1959- Stomach contents and microfeeding of some common cirripedes. Canad. Journ. Zool. 37: 231 - 236 CarrikER, MELBOURNE ROMAINE 1967. Ecology of estuarine benthic invertebrates: A perspective. In: Estuaries: 442-487. G. H. Lauff, (ed.) Washington, D.C.: Amer. Assoc. Adv. Sci. Cuantey, Pau « Jay D. ANDREWS 1971. Aids for identification of bivalve larvae of Virginia. Malaco- logia 11 (1): 45-119 (8 October 1971) Cones, H.N., Jr. a Dexter Stearns Haven 1969. Distribution of Chrysaora quinquecirrha in the York River. Chesapeake Sci. 10: 75 - 84 Crisp, Dennis J. 1964. An assessment of plankton grazing by barnacles. In: Grazing in terrestrial and marine environments: 251 - 268. D. J. Crisp (ed.). Sympos. Brit. Ecol. Soc. no. 4 é 1967. | Chemical factors inducing settlement in Crassostrea virginica (Gmelin). Journ. Anim. Ecol. 36: 329 - 335 Ga.tsorF, Paut Simon 1964. The American oyster, Crassostrea virginica (Gmelin). Fish. Bull. Fish « Wildlife Serv., U. S. Bur. Comm. Fish. 64: 1-480; 400 figs.; 46 tables Hinu, HERBERT 1969. Gregarious setting in the American oyster Crassostrea virginica Gmelin. Chesapeake Sci. 10: 85 - 92 Hiovu, Hersert & Haron H. HasKin 1971. Setting of the American oyster related to environmental factors and larval behavior. Proc. Natl. Shellfish Assoc. 61: 35 - 50 Hipu, Herpert, FretcHer P Veitcu e« Patricia E. O’Brien 1970. Gregarious setting in the American oyster. Proc. Natl. Shell- fish. Assoc. Go: 4 Korrinca, PIrter 1941. | Experiments and observations on swarming, pelagic life and setting of the European flat oyster, Ostrea edulis. Arch. Neerl Zool. 10: I - 249 MacKenzie, Crypez Leonarp, Jr. 1977. Sea anemone predation on larval oysters in Chesapeake Bay (Maryland). Proc. Natl. Shellfish Assoc. 67: 113-117 McFar.ang, I. D. 1970. Control of the predatory feeding behavior in the sea anemone Tealia felina. Journ. Exp. Biol. 53: 211 - 221 MIverIKovsxy, Simon A. 1974. On predation of pelagic larvae and early juveniles of marine bottom invertebrates by adult benthic invertebrates and their passing alive through their predators. Mar. Biol. 26: g03 - 311 Nez.son, THurLow CHRISTIAN 1924. The attachment of oyster larvae. Nicax, J. A. CoLin 1967. The biology of marine animals. 204 Ed.; 669 pp. London, Sir Isaac Pitman e Sons ReicHarpt, Hucu F 1977. Influence of the hooked mussel Ischadium recurvum (Rafines- que) on settling larvae of the American oyster Crassostrea virginica (Gmelin). Univ. Maryl. Cent. Environm. Estuar. Stud. Ref. 77-144 HPEL Soxat, Ropert R. « FE. James Rouir 1969. Biometry. W.H. Freeman & Co., San Francisco, Calif 776 pp. SouTHwarp. A. J. 1955. Feeding of barnacles. STEPHENSON, T. A. 1928. The British sea anemones, vol. 1. London: Adlord & Son, Ltd. TuHorson, GUNNAR 1950. Reproductive and larval ecology of marine bottom invertebrates. Biol. Rev. 25: 1-45 1966. | Some factors influencing the recruitment and establishment of marine benthic communities. Neth. Journ. Sea Res. 3 (2): 267 - 293 VerrcH, FLETCHER P. e Hersert Hu 1971. | Gregarious setting in the American oyster Crassostrea virginica Gmelin: I. Properties of a partially purified “setting factor.” Chesa- peake Sci. 12: 173-178 Wass, Marvin L. 1972. A check list of the biota of lower Chesapeake Bay. Spec. Sci. Reprt. No. 65. 290 pp. Virg. Inst. Mar. Sci. WituuMs, R. B. 1972. | Chemical control of feeding in the sea anemone Diadumene luci- ae (Verrill). Comp. Biochem. Physiol. 41 A: 361 - 371 Woop, Lanciey & Wii1am Jennincs Harais, Jr. 1971. ‘Transport of bivalve larvae in a tidal estuary. Proc. Europ. mar. biol. Symp. 4: 29 - 44 Biol. Bull. 46: 143-151 Nature 17§: 1124-1125 Vol. 22; No. 1 THE VELIGER Page 85 NOTES & NEWS Range Extension for Penitella fitcht Turner, 1955 (Bivalvia : Pholadidae) BY E. C. HADERLIE Naval Postgraduate School, Monterey, California 93940 For THE PAST several years extensive dredging operations have been carried out in Monterey Bay in an attempt to determine the horizontal and bathymetric distribution of rock-boring bivalves. In addition, the intertidal sedi- mentary rocks of the northern part of Monterey Bay near Santa Cruz have been examined for borers and nestlers. Most of the bivalve borers found in this survey have been reported before from Monterey Bay, but one has not. ; Penitella fitcht was described by TURNER (1955) from specimens collected by J. E. Fitch from sedimentary rocks exposed at the north side of Bahia San Bartolomé, Baja California, Mexico. At the type locality P fitchi was fairly common along with about equal numbers of P penita (Conrad, 1837) and a smaller number of Chaceia ovo- idea (Gould, 1851). Subsequently a few other specimens have been found insouthern California at Redondo Beach, La Jolla, and San Diego, and from near Isla San Geroni- mo, Bahia San Cristobal, and at Punta Pequefia, Baja Ca- lifornia (KENNEDY, 1974). The specimen collected at La Jolla was a single dead shell found along with Penitella gabbu (Tryon, 1863), PR penita, Netastoma rostrata (Va- lenciennes, 1846), and Lithophaga plumula (Hanley, 1843) in a large intertidal zone mudstone cobble. KEn- NEDY (1974) also reported the rare occurrence of P fitchi in the fossil record. Specimens have been found in late Pleistocene Palos Verdes Sand above Upper Newport Bay, California, and from Pleistocene deposits of San Pedro, California, and Bahia San Bartolomé, Baja Cali- fornia. In the California Pleistocene deposits P fitchi was found associated with Chaceia ovoidea, Netastoma rost- rata, P. penita, and Barnea subtruncata (Sowerby, 1834). Of the several hundred pholads and boring mytilids collected in this survey by dredging sedimentary rock from the shallow water of Monterey Bay, 2 living Peni- tella fitchi and 1 set of valves from a dead specimen have been found. The 2 living animals were recovered from hard siliceous chert of the Monterey Formation dredged in April and August, 1977, from water 25m deep. The first animal was recovered from rock beneath the kelp bed off Del Monte Beach and the second from a reef outcrop east of Cannery Row, both sites near Monterey, California. In each case, P fitchi was associated with numerous living PR conradi Valenciennes, 1846, P gab- bu and Lithophaga plumula. Each P. fitchi had a shell 50mm long and 35.5 mm high and each was in the post- boring stage with a partial callum. The distinctive feature of this species of Penztella is the well-developed siphono- plax composed of numerous, flexible, over-lapping peri- ostracal leaves. One of the specimens lived in an artificial burrow in an aquarium with running sea water for several months until preserved. The third specimen recovered during this study consisted of a pair of valves 40mm long from a dead specimen in chert from water 15m deep off Del Monte Beach. No specimens of P fitchi have been found in sedimentary rocks from the intertidal zone in northern Monterey Bay. The present known range of Penitella fitchi is therefore extended northward to include the southern part of Mon- terey Bay, California. Literature Cited Kennepy, Groroe L. 1974. West American Cenozoic Pholadidae (Mollusca: Bivalvia) ~ San Diego Soc. Nat. Hist. Mem. 8: 1 - 128 Turner, RutH Drron 1955- The family Pholadidae in the western Atlantic and eastern Pacif- ic. Part Il — Martesiinae, Jouannetiinae and Xylophaginae. Johnsonia 3 (34): 65-160; pits. 35 - 93 Generous Donation by the San Diego Shell Club Shortly after our April issue was ready for mailing, we received from the San Diego Shell Club another generous donation to add to our Endowment Fund. We express our gratitude for the continued support by this organization. The income from the Endowment Fund materially assists and keeps on assisting in maintaining our very low mem- bership dues and subscription rates. Page 86 THE VELIGER Vol. 22; No. 1 A Correction On pages 408 - 409 of volume 21 we have published a review by Mr. Barry Roth of the “Catalogue of Molluscan Taxa Described by Tadashige Habe During 1939 - 1975.” Dr. Téru Inaba has written to us regarding several errors: for Karuro Oyama read KatsurA Oyama (the co- author of this work), and for Okinaebishu-no-kai read Okinaebisu-no-kai; add after this: Masatoyo Okamoto. Further, the price is stated by Dr. Inaba to be $30.00 (postage included). UNITAS MALACOLOGICA SEVENTH INTERNATIONAL MALACOLOGICAL CONGRESS 31 August to 6 September 1980 The Congress will be held at Perpignan and Banyuls-sur- Mer in southern France. All topics about living or fossil mollusks are welcomed. The presentations may be made in English, French, or German. Abstracts will be published ~ in “Malacologia.” For detailed information write to: Secrétariat du 7éme Congrés International de Malaco- logie, Laboratoire de B. I. M et Malacologie, 55 Rue de Buffon, F-75005, Paris, France. Provisional registra- tion by potential participants is requested. Forms for this purpose may be obtained from the Congress Secretary at the above address. It is recommended to do this as early as possible. The Unitas Malacologica was formerly the Unitas Malacologica Europaea, but the name was changed to indicate that there is no discrimination against the parti- cipation of non-European malacologists; also, active membership is now open to all malacologists. Sale of C. M. S. Publications: Effective January 1, 1978, all back volumes still in print, both paper covered and cloth bound, will be available only from Mr. Arthur C. West, P. O. Box 730, Oakhurst, CA (lifornia) 93644, at the prices indicated in our Notes and News section, plus postage and, where applicable, California State Sales Tax. The same will apply to the Supplements that are still in print, except for supplements to vol. 7 (Glossary) and 15 (Ovulidae), which are sold by The Shell Cabinet, P.O. Box 29, Falls Church, VI (rginia ) 22046; and supplement to volume 18 (Chitons) which is available from Hopkins Marine Station, Pacific Grove, CA (lifornia) 93950. Volumes 1 through 8 and 10 through 12 are out of print. Volume 9: $22.- — Volume 13: $24.- — Volume 14: $28.- Volume 15: $28.- Volume 16: $32.- Volumes 17 to 20: $34.- each; Vol. 21: $40.-. Postage and handling extra. There is a limited number of volumes 9, 11, 13, 14 to 20 available bound in full library buckram, black with gold title. 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This will necessarily result in delays. A Glossary of A Thousand-and-One Terms Used in Conchology by Winirrep H. ARNOLD originally published as a supplement to volume 7 of the Veliger has been reprinted and is now available from The Shell Cabinet, Post Office Box 29, Falls Church, Virginia 22046, U. S. A. The cost is US$ 3.50 postpaid if remittance is sent with the order. Supplement to Volume 15: Our stock is exhausted, but copies are still available from The Shell Cabinet, P O. Box 29, Falls Church, Virginia 22046. [A systematic Revision of the Recent Cypraeid Family Ovulidae by Crawrorp NEILL CaTE] Vol. 22; No. 1 Other supplements: [Growth Rates, Depth Preference and Ecological Succes- sion of Some Sessile Marine Invertebrates in Monterey Harbor by Dr. E. C. Haderlie] Supplement to Volume 17: Our stock of this supplement is exhausted. Copies may be obtained by applying to Dr. E. C. Haderlie, U. S. Naval Post-Graduate School, Mon- terey, CA (lifornia) 93940. 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Therefore we recommend either International Postal Money Orders or bank drafts on the Berkeley Branch of United California Bank in Berkeley, California. This institution has agreed to honor such drafts without charge. UNESCO coupons are NOT acceptable except as indicated elsewhere in this section. Page 88 THE VELIGER Vol. 22; No. 1 Publication Date of THE VELIGER THE PUBLICATION DATE of The Veliger is the date printed on the index page; this applies even if the date falls on a legal holiday or on a Saturday or Sunday, days when the U.S. Postal Service does not expedite second class mail matter. 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At a Regular Membership meeting of the Society in No- vember 1968 a policy was adopted which, it is hoped, will assist in building up the Endowment Fund of the Society. An issue of the journal will be designated as a Memorial Issue in honor of a person from whose estate the sum of $5000.- or more has been paid to the Veliger Endowment Fund. If the bequest is $25 o00.- or more, an entire volume will be dedicated to the memory of the decedent. CALIFORNIA MALACOZOOLOGICAL SOCIETY, Inc. is a non-profit educational corporation (Articles of In- corporation No. 463389 were filed January 6, 1964 in the office of the Secretary of State). The Society publishes a scientific quarterly, the VELIGER. Donations to the Society are used to pay a part of the production costs and thus to keep the subscription rate at a minimum. 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Please send for membership ap- plication forms to the Manager or the Editor. Membership renewals are due on or before April 15 each year. If renewal payments are made after April 15 Vol. 22; No. 1 THE VELIGER Page 89 but before March 15 of the following year, there will be a re-instatement fee of $1.-. Members whose dues pay- ments (including the re-instatement fee) have not been received by the latter date, will be dropped from the rolls of the Society. They may rejoin by paying a new initiation fee. The volume(s) published during the time a member was in arrears may be purchased, if still available, at the regular full volume price plus applicable handling charges. REGARDING POSTAL SERVICE We are much disturbed by the steadily increasing num- ber of premature claims for supposedly “missing”’ issues of our journal. Since we have announced on numerous occasions that our journal is mailed on the dates printed in the issues, 7.¢., number 1 on July 1, number 2 on October 1, number 3 on January 1 and number 4 on April 1 of each volume year, it is unreasonable to expect delivery of the issues earlier than at least one week after these dates; however, a much longer time must be allowed for delivery to addresses at various distances from Berkeley. Thus, for example, a two weeks lapse is not unusual for as short distances as 500km; and up to 3 and 4 months must be counted on for addresses in the Far East and in Africa. We are faced with the alternative of not replying to what we must consider premature claims or, if the trend con- tinues, we must increase our subscription rates to cover these additional expenses. Our past efforts at keeping the subscription rate as low as possible are, we believe, suffi- cient evidence that we simply cannot afford any other course of action. The postal service causes us enough financial losses. Therefore we urgently request that before a claim is made, the time schedule be carefully checked. We are grateful for the understanding of this difficult situation shown by many librarians and will be grateful to those who, heretofore being perhaps eager to make sure that the library receives what is coming to it, will exercise a little patience. Your harassed Editor. BOOKS, PERIODICALS, PAMPHLETS Biogeography and Adaptation: Patterns of Marine Life by GeeraT J. VERMEIJ. xi+332 pp.; illust. Harvard Uni- versity Press, Cambridge (Mass.) and London. $25.00. 15 December 1978 This is a thoughtful, provocative, and at times specula- tive book, based primarily on studies of molluscan distri- bution and shell form, in which a mature zoologist draws together extensive observations on the adaptations of marine organisms to their physico-chemical and _ biotic environments. Patterns in form, behavior, and physiology are among the most intriguing that the marine world has to offer, and their geographic distribution is as legitimate a concern of the biogeographer as are patterns treated primarily in taxonomic-evolutionary terms. In Part I (Patterns and Adaptations along Gradients), Professor Vermeij devotes Chapters 2 and 3 to gastro- pods and bivalves, respectively, and in Chapter 4 covers a variety of other slow-moving or sessile invertebrates, as well as plants. The author argues that physico-chemical conditions tend to provide the major selective forces in high and cold latitudes, but that biotic selective forces, especially predation pressure, result in greater defensive adaptations of prey species in more tropical waters. This is expressed in the evolution of such predation-resistant adaptations as armor, shell thickening, aperture narrow- ing, and shell sculpture in warmer seas. Further, Vermeij detects a comparable but micro-geographic trend in pred- ator effectiveness and defensive adaptations of prey spe- cies in low as compared with high intertidal situations, and similarly in shallow as compared with deep environ- ments. The author’s view that the biologically mediated com- ponent in the evolution of molluscan shell form is almost exclusively a response to predation, minimizing competi- tion as a selective force, may be, however, the most poten- tially controversial aspect of the book. For example, narrow apertures that predominate in tropical gastropods are explained as a response to increased predation pres- sure. This view neglects the strong correlation of shell form with feeding mode and taxonomy in marine gastro- pods. Elongate apertures have evolved predominately in neogastropods (and in many mesogastropods that are also carnivorous) engaging in what is often a highly com- petitive business. In forms adopting an active, predatory mode of life, the elongation may be viewed as serving the dual functions of streamlining, particularly when elonga- tion is coupled with detorsion, and of increasing the sep- aration of inhalent and exhalent currents. This is not to argue for competition as the major selective force, but rather to point out that both competition and predation have exercised profound influences on molluscan form, physiology, and behavior. In Part II (Interoceanic Patterns of Adaptation), Ver- meij in Chapter 5 generalizes that the biotic pressures of grazing and predation among gastropods on open surfaces are greater in the tropical Indo-Pacific than in the tropi- Page 90 THE VELIGER Vol. 22; No. 1 cal Atlantic, although this is not detectable in a number of other groups or on other substrates. Chapter 6 sum- marizes observations on the effects of present-day environ- mental factors such as water-movement, tides, salinity, temperature, and productivity on the establishment and maintenance of regional species diversity. Part III Geography and Evolution) becomes more gen- eral, as well as more varied in content than the title implies. In Chapter 7, Extinction and Speciation, Vermeij (without claiming originality) groups species into 3 cat- egories rather than the 2 of the commonly-used “r-K-ic” system (the author may be forgiven a few other equally bad puns): (1) weedy or opportunistic species (r-selec- ted); (2) stress-tolerant species adapted to variable or extreme environments;and (3) “biologically competent” species of stable and physiologically-favorable environ- ments, the latter 2 groups comprising species commonly called K-selected. This and Chapter 8, Patterns of Biotic History, deal with both Recent and fossil communities, and are so varied as to be difficult to characterize suc- cinctly. In the final Chapter 9, Barriers and Biotic Ex- change, Vermeij examines biotic contacts between marine biotas as well as the problem of introduced species. The book ends with a discussion of the possible faunistic con- sequences of the opening of a sea-level, interoceanic, Cen- tral American canal. In this, as in all preceding chapters, Vermeij generalizes freely and frankly, often admitting the inadequacy of his data, but posing questions deserving further study. Perhaps the most important and valuable aspect of this book is that it represents a synthesis of a truly great amount of observation by a perceptive ob- server, who does not hesitate to express his carefully-quali- fied intuitions. This is not a book in which one can easily look up facts, although it is reasonably indexed, and in- deed much of the discussion is difficult to follow if one is not broadly familiar with tropical molluscan and other genera. Many of Vermeij’s data consist of correlations and, as is always the case with such data, there are many possibil- ities for alternative interpretations and a lingering desire for substantiation. The tendency to speculation based on extensive observation has resulted in a book in which graduate students looking for research problems will find many a fascinating question. The text is very free of typo- graphical errors, but Harvard University Press should be chided for the low quality of some of the halftone figures. A 43-page list of references attests to the scholarly quality of the book, and constitutes an invaluable resource. It is refreshing to find a book of this sort so free of “models” and so full of good observations and ideas. Ralph I. Smith, Department of Zoology Carole S. Hickman, Dept. of Paleontology University of California, Berkeley The Abalone Book by Perer C. Howorrn. Naturegraph Publishers, Inc., Happy Camp, California 96039. $7.95 cloth; $3.50 paper. 1978 Since Keith Cox’s excellent “California Abalones, Fam- ily Haliotidae” (1962, Calif. Dept. Fish & Game, Fish Bull. 118) went out of print, I have been at a loss to recommend any available book as a ready source of infor- mation on eastern Pacific abalones, genus Halzotis. ‘This popularly written manual will undoubtedly fill many buy- ers’ needs. It contains a brief history of the abalone, from the fossil record to man’s present-day impact, a section on problems plaguing the resource and some possible solu- tions, basic abalone biology, a review of North American species, and advice on preparation and cooking. There is a key to identifying west coast abalone shells and another for underwater identification. Shells of California aba- lones, interior and exterior, are illustrated on brilliant color plates. The colored figures of each species’ epipodi- um are helpful for field identification by divers and sport fishermen. Northern California divers may take a second, hard look at the red abalone, Haliotis rufescens, in color plate I, for this appears to be a southern California form of the species, with reduced radial sculpture and elevated tremata — probably the result of introgressive hybridiza- tion with Haliotis sorenst. Hybrids between the two are particularly common in the Santa Barbara Channel area. The specimen illustrated as the Florida abalone, Haliotis pourtalesit (p. 65), differs in profile and sculpture from the original illustrations of that species. “The Abalone Book” is rich in firsthand observations, based on the author’s 24 years as a diver and 2 years managing an abalone processing plant. One can judge the depth of Howorth’s involvement with abalones from his comment regarding Haliotis cracherodu: “Blacks are the Bogarts of the abalone world.” Barry Roth THE VELIGER is open to original papers pertaining to any problem concerned with mollusks. This is meant to make facilities available for publication of original articles from a wide field of endeavor. Papers dealing with anatomical, cytological, distributional, ecological, histological, morphological, phys- iological, taxonomic, etc., aspects of marine, freshwater or terrestrial mollusks from any region, will be considered. Even topics only indi- rectly concerned with mollusks may be acceptable. In the unlikely event that space considerations make limitations necessary, papers dealing with mollusks from the Pacific region will be given priority. However, in this case the term “Pacific region” is to be most liberally interpreted. It is the editorial policy to preserve the individualistic writing style of the author; therefore any editorial changes in a manuscript will be sub- mitted to the author for his approval, before going to press. Short articles containing descriptions of new species or lesser taxa will be given preferential treatment in the speed of publication provided that arrangements have been made by the author for depositing the holotype with a recognized public Museum. Museum numbers of the type specimens must be included in the manuscript. Type localities must be defined as accurately as possible, with geographical longitudes and latitudes added. Short original papers, not exceeding 500 words, will be published in the column “NOTES & NEWS’; in this column will also appear notices of meetings of the American Malacological Union, as well as news items which are deemed of interest to our subscribers in general. Articles on “METHODS & TECHNIQUES” will be considered for publication in another column, provided that the information is complete and tech- niques and methods are capable of duplication by anyone carefully fol- lowing the description given. Such articles should be mainly original and deal with collecting, preparing, maintaining, studying, photo- graphing, etc., of mollusks or other invertebrates. A third column, en- titled “INFORMATION DESK,” will contain articles dealing with any problem pertaining to collecting, identifying, etc., in short, problems encountered by our readers. In contrast to other contributions, articles in this column do not necessarily contain new and original materials. Questions to the editor, which can be answered in this column, are in- vited. The column “BOOKS, PERIODICALS, PAMPHLETS” will attempt to bring reviews of new publications to the attention of our readers. Also, new timely articles may be listed by title only, if this is deemed expedient. Manuscripts should be typed in final form on a high grade white paper, 812” by 11”, double spaced and accompanied by a carbon copy. A pamphlet with detailed suggestions for preparing manuscripts intended for publication in THE VELIGER is available to authors upon request. A self-addressed envelope, sufficiently large to accom- modate the pamphlet (which measures 5/2” by 81/2”), with double first class postage, should be sent with the request to the Editor. EDITORIAL BOARD Dr. Donatp P. Azsort, Professor of Biology Hopkins Marine Station of Stanford University Dr. WarrEN O. AppicoTT, Research Geologist, U. S. Geological Survey, Menlo Park, California, and Consulting Professor of Paleontology, Stanford University Dr. Hans Berrscu, Curator of Marine Invertebrates San Diego Museum of Natural History Dr. Jerry DonounuE, Professor of Chemistry University of Pennsylvania, Philadelphia, and Research Associate in the Allan Hancock Foundation University of Southern California, Los Angeles Dr. J. Wyatr DuruaM, Professor of Paleontology Emeritus University of California, Berkeley, California Dr. Capet Hann, Professor of Zoology and Director, Bodega Marine Laboratory University of California, Berkeley, California Dr. Carote S. Hickman, Assistant Professor of Paleontology University of California, Berkeley, California Dr. A. Myra KEEN, Professor of Paleontology and Curator of Malacology, Emeritus Stanford University, Stanford, California Dr. Victor Loosanorr, Senior Biologist, Emeritus U.S. National Marine Fisheries Service EDITOR-IN-CHIEF Dr. Rupoir Srouter, Research Zoologist, Emeritus University of California, Berkeley, California Dr. Joun McGowan, Professor of Oceanography Scripps Institution of Oceanography, La Jolla University of California at San Diego Dr. FRANK A. PiTELKA, Professor of Zoology University of California, Berkeley, California Dr. Rosert Rosertson, Pilsbry Chair of Malacology Department of Malacology Academy of Natural Sciences of Philadelphia Dr. Peter U. Roppa, Chairman and Curator, Department of Geology California Academy of Sciences, San Francisco Dr. CiypeE F. E. Roper, Curator Department of Invertebrate Zoology (Mollusca) National Museum of Natural History Washington, D. C. Dr. JupirH Terry Smiru, Visiting Scholar Department of Geology, Stanford University Stanford, California Dr. Rap I. Smiru, Professor of Zoology University of California, Berkeley, California Dr. Cuar.es R. STASEK, Bodega Bay Institute Bodega Bay, California Dr. T. E. Tuompson, Reader in Zoology University of Bristol, England ASSOCIATE EDITOR Mrs. Jean M. Cate Rancho Santa Fe, California } - /, mee TAM Fd. if Divis) iON OF MD! tise THE VELIGER fy SS kat) LATIN >) din A Quantrly published by CAUFORNIA MALACOZOOLOGICAL SOCIETY, INC. Beskoley, Callfernia VOLUME 22 OcTOBER I, 1979 ContTENTS The Family Epitoniidae (Mollusca : Gastropoda) in the Northeastern Pacific. (6 Plates; 3 Text figures) ARLEN SOU SHANE yer oy rere Miri ras As sere eS gS Ot On the Growth Stages of Conus fergusoni Sowerby, 1873, the Reinstatement of Conus xanthicus Dall, 1910, and a New Species of Conus from the Galapagos Islands. (3 Plates; 4 Text figures) James H. McLean «& James W.NYBAKKEN . . .. ; SS Pek ries . 135 The Familiy Lepidopleuridae (Mollusca : Le in the Eastern Pacific. (4 Plates; 8 Text figures) ANTONIO J. FERREIRA. . Sides ten Cosa Benes iret ec hg ly A Statistical Study of Care tigris in the Central Pacific. (2 Text figures) RicHarp E. Brock . . . eeu eet geen geal) oe hOO Pholadomya candida ne The Last Cadaver Unearthed. (1 Plate; 1 Text figure) Bruce RUNNEGAR .. . : - 171 The Ultrastructure and elas Gihne s of the Ocelli in the Larva of Katharina tunicata (Mollusca : Polyplacophora). (3 Plates; 2 Text figures) Marc Davp Rosen, CHarztes R. Stasex a Corin O. HermaNS .. . 5 iG} New Information Concerning Humboldtiana taylori Drake, 1951 (Gisepea Pulmonata : Helminthoglyptidae) . (1 Plate; 1 Text figure) Arnie L:oMEtcatr © Davip H: RISKIND - 2). 0. 2s) es te es 179 [Continued on Inside Front Cover] Distributed free to Members of the California Malacozoological Society, Inc. Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc. Volume 22: $30.- plus $1.50 for postage (U.S.A. only) For ALL foreign countries: Swiss Francs 60.- plus SF 7.- for postage Single copies this issue $22.00. Postage additional. Send subscription orders to California Malacozoological Society, Inc. 1584 Milvia Street, Berkeley, CA 94709, U.S.A. Address all other correspondence to Dr. R. Stoner, Editor, Department of Zoology University of California, Berkeley, CA 94720 Second Clan Postage Paid at Berkelsy, Oalifornia NAL ieeiay ISSN 0042-3411 Contents — Continued Examination of a Reproductive Cycle of Protothaca staminea Using Histology, Wet Weight-Dry Weight Ratios, and Condition Indices. (4 Text figures) Howarp M. Fever, James C. Henpez, Pat Hotmes, Georce J. MuELLER & A. J. Paut Life-Cycle Completion of the Freshwater Clam Lasmigona compressa (Bivalvia: Unionidae) in an Experimental Host, Lebistes reticulatus. ALEx S. Tompa all gay. Soi, 1680! aly og det Wey, We SRR ay Ae Reena cM pak el Philip pia (Psilaxis) radiata: Another Indo-West-Pacific Architectonicid Newly Found in the Eastern Pacific (Colombia) . (1 Plate; 1 Map) Rosert ROBERTSON . ea a PSR ON, mes Le Vein Vota atoms: toes The Packaging of Ova in the Egg Cases of Aplysia californica. (5 Text figures) Paut KANDEL & THomas R. Capo . Modiolus aurum Osorio, spec. nov., from Juan Fernandez Archipelago, Chile. (Mollusca : Bivalvia : Mytilidae) . (g Text figures) Cercii1a Osorio R. ee ee mame he PET Slr gcd Range Extensions of Mollusk Species Found on the Tropical Coast of El Salvador. Marco ANTONIO HERNANDEZ A Re-Evaluation of the Northwestern Range of the Melongena corona Complex. (3 Text figures) IsaBeL C. Levy Rediscovery of the Holotype of Cymatium (Cymatium) ranzanii (Bianconi, 1850). (1 Plate) BruNo SABELLI, Marco TaviANi & STEFANO TOMMASINI . BOOKS, PERIODICALS & PAMPHLETS . 182 . 188 . 191 - 194 - 199 . 204 . 206 = 2110 - 212 Note: The various taxa above species are indicated by the use of different type styles as shown by the following examples, and by increasing indentation. ORDER, Suborder, DIVISION, Subdivision, SECTION, SupPERFAMILY, Famity, Subfamily, Genus, (Subgenus) New Taxa Vol. 22; No. 2 THE VELIGER Page g! The Family Epitoniidae (Mollusca : Gastropoda) in the Northeastern Pacific HELEN DuSHANE 15012 El Soneto Drive, Whittier, California 90605 (6 Plates; 3 Text figures) INTRODUCTION THIS PAPER IS A REVIEW Of the late Neogene and Quater- nary species of Epitoniidae (Mollusca: Gastropoda) which occur in the littoral and sublittoral zones of the north- eastern Pacific, from Point Barrow, Alaska (171° 20’ N) (including the Aleutian Islands) south to Cedros Island, Baja California Norte, Mexico (28° 12’ N). Species occur- ring south of Cedros Island were treated previously by DuSHANE (1974). This treatment is an attempt to characterize epitoniid species, discuss populational variation, summarize data on the ecology of the group. record the fossil occurrences, and to revise the nomenclature. The nomenclatural revisions herein have resulted in synonymizing some nominal taxa which hopefully will provide a better understanding of identity of other taxa. Species of Epitoniidae display a confusing degree of morphological variability. Some early authors separated specimens on very trivial characters without realizing how widely dispersed epitoniids are. That populational varia- bility exists has been established by Dat (1917), STRONG (1930, 1945), and others. The epitoniids of the northeast- ern Pacific region show considerable variation. In working with this family it soon became obvious that previous syn- onymies were incorrect, figures were inadequate, and very little was known of the infraspecific variation and distribu- tional limits of most species. The synonymies here are in- tended to be complete, except for those treated previously by DuSHaNneE (1974). Figures of some earlier type material do not exist, and some types have subsequently been lost. Many published geographic ranges cannot now be substan- tiated with referred material. The ranges given are based on material examined for this study. Useful references for the species in this family are: SOWERBY (1844); CARPENTER (1857, 1864); Nyst (1871); DaLL (1917, 1921); DEBoury (1919); STRONG (1930, 1937); GranT & GALE (1931); DuRHAM (1937); CLENCH & Tur- NER (1950, 1951, 1952); PALMER (1958). Morphology: Of the supraspecific taxa recognized by CLENCH & TURNER (1950, 1951, 1952) only the genera and subgenera Acirsa, Amaea, Asperiscala, Boreoscala, Depres- siscala, Nitidiscala, Opalia, Sthenorytis, and (Scalina) are used herein. Epztonium s.s. (type species Turbo scalaris Linnaeus, 1767), is characterized by convex, unattached whorls and does not seem to have typical members in the eastern Pacific. Certain morphological differences from those of Epitonium s.s. suggest that Nitidiscala and Asper- wscala should be given full generic rank, which has been done herein. Epitoniid genera can be characterized in the following ways: Asperiscala: Usually turreted with a consistent pattern of cording or fine threads on the whorls, sometimes obso- lete on later whorls; costae usually recurved, sometimes with a shoulder spine. In the eastern Pacific 22 species of Asperiscala are recognized from the tropical Panamic re- gion (DUSHANE, 1974), and 6 from the temperate north- eastern Pacific. Nitidiscala: Not always turreted; lacking spiral sculp- ture between whorls; costae bladelike, reflected, or over- lapping as in N. catalinae, with a sharp shoulder spine or extension, not continuous from whorl to whorl; basal disk lacking. From the tropical eastern Pacific 16 Nitzdiscala species are recognized (DUSHANE, 1974); and 7 occur in more temperate waters. In Opalia, the nominal subgenera Nodiscala and Denti- scala of De Boury have characters that are ill-defined and difficult to apply. They are not used herein. All Opalia have a heavy calcareous outer coat (intritacalx), that, when worn, removes the fine pitting characteristic of them, leav- Page 92 THE VELIGER Vol. 22; No. 2 ing the shell with a shiny appearance. The purpose of the punctae is not yet known. The remaining genera and sub- genera have diagnostic characters that make them readily recognizable and have been covered by CLENCH & TURNER (1950, 1951, 1952). Brief summaries of each are given under the separate headings. The length and diameter of a shell for a given number of whorls and costae are probably the best characters for identifying species of Epitonium. The limits of specific variation in the number of costae are greater in those spe- cies having more costae. ANKEL (1938b) suggested that the costae are a defense against boring naticids, a suggestion that parallels my own observations. Few epitoniids show damage from boring predators. Many type specimens, un- fortunately are beach-worn specimens and are too abraded to be of scientific value. Characters that are useful for sep- arating species, are: size of shell, length and diameter of shell, number of whorls, number of nuclear whorls in rela- tion to the remainder, sculpture and shape of early whorls, number and placement of costae on the whorls, absence or presence of spiral sculpture, presence of punctae (as in Opalia), absence or presence of a basal disk or cord, depth of suture, presence or absence of an umbilicus, shape and placement of aperture, sculpture of operculum, and geo- graphical and bathymetric origin. The internal anatomy of the epitoniids was partially described by THIELE (1928). The head includes the ten- tacles and eyes, the foot provides the animal with a means of locomotion, and the coiled visceral mass contains the internal organs. The foot is extended in front, grooved behind, ostensibly to help grip the surface, and witha thin, horny operculum. The long proboscis, when protruded, displays two specialized stylets alongside the mouth, which is at the base of the proboscis. The stylets, surrounded by a sheath of muscle, have openings for the salivary glands. There are no oesphageal glands. The hypobranchial gland produces a purple secretion. Other than the fact that it is a protein-like substance, we do not yet know the chemical constituents of the purple dye. ANKEL (1938a), THORSON (1958), and RoBERTSON (1963) suggested that this gland introduces an anesthetizing agent into its prey. Additional tests (SALO, 1977) suggested that Epitonium toxin inhibits some part of the neuromuscular system of the anemone on which it feeds. The structure of the buccal mass suggests that epitoniids can swallow their prey whole (FRETTER & GRAHAM, 1962). PERRAN (1978), in experiments with Boreoscala greenlandica (Perry, 1811), found this species could swallow whole the small anemone Gonactinia pro- lifera. Epitoniids have two sharp-edged jaws near the buccal mass. HOCHBERG (1971), DUSHANE (1974), C. R. Smiru (1977), SALO (1977), and PERRAN (1978) have shown that epitoniid radular teeth are different from all other proso- branch radulae with the exception of Bulla and Ianthina, which they most resemble. The numerous, evenly spaced, lateral teeth are arranged in regular rows forming obtuse angles. There are no central hooks as in other families of gastropods. The animal has an abnormally long acrem- bolic proboscis, which led FRETTER & GRAHAM (1962) to theorize that perhaps epitoniids have a method of feeding on either annelid worms or nemerteans that would require anesthetization before being seized by the radula. Subse- quent observations have not borne this out (HOCHBERG, 1971, C. R. SMirH, 1977, SALO, 1977, PERRAN, 1978). The radula consists of a ribbon of cuticle with rows of teeth that lie flat within the buccal mass. As the radula slips for- ward, the teeth become almost vertically erect and ready to rasp. As the radula returns, a reverse action takes place. This action constitutes the food gathering habit of the epitoniids. A detailed account of this process is given in FRETTER & GRAHAM (1962). HocHuBERG (1971: 22) noted that in southern California, Nitidiscala tincta lives in specific association with the small aggregate anemone, Anthopleura elegantissima, through- out its post-larval life. “This so-called micropredator is active twice a day during periods of high water when the anemone beds are covered and the polyps expanded. In order to feed, the snail everts a long acrembolic proboscis and slips it over the tip of a tentacle. The tentacle is held in place by the jaws and radula in combination with a mus- cular buccal bulb. Two stylets at the end of the proboscis inject a salivary toxin or anesthetizing agent. The tip of the tentacle is cut or torn off and pulled into the digestive system upon retraction of the proboscis.’ Hochberg’s ob- servations are corroborated by his excellent photographs. C. R. Smitu (1977) conducted experiments on the chem- ical recognition of prey of the same species. As a result, she also demonstrated that this gastropod is a micropredator on the anemone, Anthopleura elegantissima, its preferred prey, but will also eat the tentacles of A. xanthogrammica. The epitoniid also ingested tissue from two other Acti- niidae, Tealia lofotensis and Epiactis prolifera, but less commonly. SALO (1977) corroborated the findings of both Hocuserc (1971) and C. R. Situ (op. cit.). SALo (op. cit.) also tested the feeding behavior of Nitidiscala indianorum and found that Tealia crassicornis and T. lofotensis were most commonly accepted. THORSON (1958) observed Opalia funiculata [= O. cre- nimarginata] sucking on the body-cylinder of Anthopleura at mid-tide level. My observations in the field have been that O. funiculata and O. borealis attach themselves at the base of the anemone and not in the cup where the nema- Vol. 22; No. 2 THE VELIGER Page 93 tocysts are located. Also, the anemone is partially con- tracted at this time. All the observations were made in the intertidal zone and in daylight. PERRAN (1978) found that unlike Nitidiscala tincta, Boreoscala greenlandica fed on the column of the large anemone Metridium senile, and swallowed whole the small anemone Gonactinia prolifera. He also found that over short distances epitoniids are able to locate their prey by chemotaxis. Epitoniids are consecutive hermaphrodites, laying their eggs in small clusters, each cluster affixed to the next by a mucus thread. Those species with a habitat in or near sandy substrates have sand-agglutinated egg capsules, each capsule containing thousands of unhatched veligers. (Coral associated species do not have sand-agglutinated egg cap- sules.) As the eggs hatch, the veligers develop a smooth protoconch that is followed by whorls with costae or other characteristic sculpture. METHODS anp FORMAT The following outline has been adopted for each species: Synonymy: Original species name, author, date of pub- lication, page number(s), and figure number(s), if any, followed by like information for each author using the identical name. Original Description: Even if the description is not in English, it is given verbatim. Additional Description: Supplementary descriptions are given for all species. Most descriptive terms are sub- jective; however size (length) is defined herein as “small; o to 5mm; “medium;’ 5 to 10mm; and “large, 10mm and over. The number of costae and range for each species is recorded in whole numbers, as is the number of nuclear whorls and post nuclear whorls. Type Material: The repository of the holotype and any paratypes is listed with the type numbers. The same information is provided for each synonym, if any. Type Locality: This is given, when reported by the orig- inal author, together with the collector’s name, if known. The type locality is also given for any syn- onyms. Distribution, Ecology and BathymetricRange: Extremes of geographic and bathymetric range are given. Geologic Record: (Contributed by George L. Kennedy.) Cccurrences cited in this section are essentially only those of the Section of Invertebrate Paleontology of the Los Angeles County Museum of Natural History (LACMIP). Additional published records are not included herein be- cause it was not possible to examine the actual specimens cited. Literature citations, however, are included in the synonymy of each species, but the reader should be aware that these are unconfirmed occurrences. The specimens cited herein were identified by the author and arranged stratigraphically by G. L. Kennedy. Localities are arranged from north to south within each time-stratigraphic inter- val (lower Pliocene to upper Pleistocene). Although it is becoming increasingly apparent that there is a great variation of ages within coastal Pleistocene de- posits, it has not been possible to integrate this new chro- nology into this paper because of the lack of precise ages for many of the localities. ‘Thus some classical ‘Pliocene’ formations in Oregon and “‘lower Pleistocene” ones in southern California (Los Angeles basin) which are actually middle Pleistocene in age, are still cited herein as lower Pleistocene (in a two-fold division) for the sake of consist- ency with existing literature. Following any fossil occurrences are cited a number of representative recent stations, also based on collections of the Section of Malacology (LACM), as well as documen- tation of southern and northern-most occurrences. Discussion: Related information which is not applicable elsewhere is placed here. No attempt has been made to list all the locality data for each Recent species, as some records cannot be documented and some taxa are known from numerous records that would be cum- bersome to report. In cases where the species is rare, all locality data are used; otherwise only the north and south extremes of the known range, together with literature documentation are listed. Type Photographs: Photographs are of type specimens. If the type is badly worn, a photograph of a live-taken specimen is included. ABBREVIATIONS The following abbreviations are for institutions whose material has been used in this study: ANSP Academy of Natural Sciences of Philadelphia, Penn- sylvania AMNH American Museum of Natural History, New York City, New York BM(NH) British Museum (Natural History), London, England CAS(GTC) California Academy of Sciences, San Francisco, Cal- ifornia (Geologic Type Collection) Page 94 THE VELIGER Vol. 22; No. 2 LACM Los Angeles County Museum of Natural History, Section of Malacology, Los Angeles, California LACM-AHF Allan Hancock Foundation Collection, University of Southern California, now at the Los Angéles County Museum of Natural History LACMIP Los Angeles County Museum of Natural History, Section of Invertebrate Paleontology, Los Angeles, California MCZ Museum of Comparative Zoology, Harvard Univer- sity, Cambridge, Massachusetts PRI Paleontological Research Institute, Ithaca, New York RM Redpath Museum, McGill University, Montreal, Quebec, Canada NMC National Museum of Canada, Ottawa, Ontario, Canada SBMNH Santa Barbara Museum of Natural History, Santa Barbara, California SDNHM San Diego Natural History Museum, San Diego, California SU Stanford University Collection, Palo Alto, Califor- nia, now at the California Academy of Sciences UCMP University of California, Museum of Paleontology, Berkeley, California USNM United States National Museum of Natural His- tory, Washington, D.C. ACKNOWLEDGMENTS For the loan of type specimens and for valuable suggestions I would like to thank Frederick J. Collier and Joseph Rose- water (USNM), William K. Emerson (AMNH), Carole S. Hickman (UCMP), F. Eric Hochberg (SBMNH), George E. Radwint (SDNHM), Robert Robertson (ANSP), and Barry Roth (CAS). The loan of types was greatly facili- tated by numerous letters written in my behalf by Gale G. Sphon (LACM). The following collectors have also generously loaned specimens or provided relevant collecting data: Jean and Crawford Cate, Emery P. Chace, Steven Newswanger, and Donald R. Shasky. James T: Carlton (University Califor- nia, Davis) provided ecological data for a number of epi- toniids. This work is based mainly on the epitoniid collections, both Recent and fossil, at the Los Angeles County Museum of Natural History. Access and unlimited use of these col- lections were kindly allowed by James H. McLean, Curator of Malacology, and Edward C. Wilson, Curator of Inverte- brate Paleontology, respectively. Initial segregation of the fossils was done mainly by Philip G. Owen (LACMIP). George L. Kennedy (LACMIP, now U.S. Geological Sur- vey, Menlo Park) assembled the paleontological literature, prepared the fossil lists herein, and read several drafts of the manuscript. + deceased Personnel of the Museum library and that of the USC Allan Hancock Foundation have been most helpful in ob- taining rare or otherwise obscure references. Photographs used herein are those of the staff or associates of the Mu- seum, notably: Bertram C. Draper, James H. McLean, and Armando Solis. Their work is greatly appreciated. Radular extractions and drawings used in this study were done by F. Eric Hochberg (SBMNH) and George E. Radwin (SDNHM). The preliminary manuscript was read by William K. Emerson (AMNH), James H. McLean (LACM), and Ed- ward C. Wilson (LACMIP). To all of them I am grateful for their help. I should especially like to thank Professor Myra Keen (Stanford University, Emeritus) for her continued help and encouragement with various aspects of this work. Her suggestions on a number of taxonomic problems, reading of the manuscript, and loan of photographs of the Carpen- ter types in the British Museum (Natural History) are greatly appreciated. Dr. Keen also suggested the docu- mentation for the authorship of the family-group names for the Ep1TONIACEA and the Eprronmpae for which I am most grateful. SYSTEMATIC TREATMENT EpiTONIAGCEA Lamarck, 1822 [?1812] [Nom. correct. et nom. transl., BERRY, 1910, ex Scalariana Lam.] EprronnpAE Lamarck, 1822 [?1812] [Nom. correct. BRoDERIP, 1839, as Scalaridae, and BErry, 1910, ex Scalariana Lamarck] Epitonium Roding, 1798 [Scala of authors; Scalaria Lamarck, 1801] Shells usually colorless, somewhat turreted, sometimes um- bilicate, axial sculpture of heavy or slender costae, some- times recurved; whorls numerous, with varying degrees of convexity, coiled either loosely or tightly; spiral sculpture present or absent; aperture round or oval; operculum thin, horny, paucispiral. Acirsa Morch, 1857 [Type species: (M) Scalaria borealis Lyell, 1841] Outer lip thin; axial and spiral sculpture of low ribs; basal keel indistinct. Vol. 22; No. 2 mE VEEIGER Table 1 Chart showing geographical ranges and fossil records reported herein. Acirsa borealis (Lyell, 1841) Acirsa cerralvoensis (DuShane, 1970) Amaea (Scalina) brunneopicta (Dall, 1908) Amaea (Scalina) effiae (Willett, 1939) Asperiscala bellastriata (Carpenter, 1864) Asperiscala cookeana (Dall, 1917) Asperiscala hemphilli (Dall, 1878) Asperiscala lowei (Dall, 1906) Asperiscala minuticosta (DeBoury, 1912) Asperiscala tinctoria (Dall, 1919) Boreoscala greenlandica (Perry, 1811) Depressiscala polita (Sowerby, 1844) Nitidiscala caamanoi (Dall and Bartsch, 1910) Nitidiscala californica (Dall, 1917) Nitidiscala catalinae (Dall, 1908) Nitisdiscala catalinensis (Dall, 1917) Nitidiscala hindsti (Carpenter, 1856) Nitisdiscala indianorum (Carpenter, 1865) Mitisdiscala sawinae (Dall, 1903) Nitidiscala tincta (Carpenter, 1865) Sthenorytis stearnsit (Dall, 1892) Opalia borealis Keep, 1881 Opatia funiculata (Carpenter, 1857) Opalia infrequens (C. B. Adams, 1852) Opala montereyensis (Dall, 1907) Opalia spongiosa Carpenter, 1866 Opalia varicostata Stearns, 1875 Circumboreal 23°N - 28°N 10°N - 28°N 32°N - 37°N 24°N - 32°N 24°N - 34°N 1°S - 28°N 24°-N- 8°N 24°N - 138°N 50°N - 71°N 163°E - 172°W 1°N - 33°N 48°N 98°N - 33°N 28°N - 54°N 31°N - 50°N 14°S - 54°N 31°N - 54°N 24°N - 54°N 24°N - 54°N 28°N 31°N - 54°N 3°S - 34°N 8°N - 34°N 25°N - 54°N 1°S - 36°N Pliocene, San Diego County, California Pliocene, San Diego County, California Pleistocene, San Pedro, California Pliocene, Los Angeles and Orange Counties, California Pleistocene, same as above [Upper] Pliocene, San Diego County, California Pliocene, San Diego County, California Pleistocene, same as above Pliocene, southern California, Cedros Is., B.Cfa. Norte Pleistocene, Galapagos Islands, Ecuador Upper Pleistocene, Orange County, California, Punta Pequena, B. Cfa. Sur, Mexico Pleistocene, Alaska Pleistocene, Aleutian Islands Pliocene, Galapagos Islands, Ecuador Pleistocene, San Pedro, California Pliocene, Los Angeles and San Diego Counties, California Pleistocene, Los Angeles County, California Pleistocene, Los Angeles County, California Pliocene, southern California Pliocene, Santa Barbara County = Pleistocene, Santa Barbara County, California and Turtle Bay, B. Cfa. Sur, Mexico Pleistocene, from Santa Barbara south to San Diego Counties, California Pliocene, Orange, San Diego Counties, California Pleistocene, Oregon and California Pliocene, Los Angeles and San Diego Counties, California Pleistocene, Oregon and California Pliocene, Los Angeles south to San Diego Counties, California Pleistocene, Humboldt south to San Diego Counties, California Pliocene, Pacific Beach, San Diego County, California Pliocene, Los Angeles and San Diego Counties, California Pleistocene, Los Angeles County, California Pleistocene, Santa Barbara and Los Angeles Counties, California Pleistocene, Santa Barbara County, California Pliocene, Los Angeles and San Diego Counties, California Pleistocene, Santa Barbara, Los Angeles, San Diego Counties, California Pliocene, San Diego County, California Pleistocene, Los Angeles County, California Pliocene, Washington, California (San Diego, Orange Counties) and Baja California Norte, Mexico Page 96 THE VELIGER Vol. 22; No. 2 Acirsa borealis (Lyell, 1841) (Figure 7) Scalaria borealis LyE.1, 1841: based on Philos. Trans. (1835) pit. 2, figs, 11, 12. Nyst, 1871: 95 [as Aczrsa ‘Beck’. Tryon, 1887: 80. DALL, 1889: 308 [“Beck,’ as a synonym of Scala costulata (Mighels)]. DALL, 1917: 474 [“Beck’’]. KEEN, (in Burch) 1945: 20. MACPHERSON, 1971: 40 [‘“Beck’’]. Epitoneum (Acirsa) borealis “(Beck)” (Lyell): DAL, 1921: 114; plt. 13, fig. 10. I. S. OLDRoyD, 1927: 54; plt. 31, fig. 4 [as boreale “(Beck)” (Lyell) ]. Epitonium (Acirsa) borealis “Gould”: KEEP, 1935: 178; fig. 152 (description is for Opalia borealis Gould). Epitonium boreale (Lyell): KEEN, 1937: 35. BURCH, 1945: 20. Morris, 1952: 87; plt. 24, fig. 6. Turritella costulata MIGHELS & ADAMS, 1842: 50; plt. 4, fig. 20, (non Borson, 1825). CLENCH & TURNER, 1950: 229 to 231; plt. 99, figs. 1-3 (as synonym of Acirsa costulata (Mighels « Adams, 1842)). Scalaria costulata: TRYON, 1887: 80; plt. 16, figs. 10, 12, 18. MacPHERSON, 1971: 40 (as Acirsa). ABBOTT, 1974: 114 (as Acirsa and synonym of A. borealis). Scalaria ochotensis MIDDENDORFF, 1849: 98; plt. 12, figs. 12-14. CARPENTER, 1857: 216, 220, 223. TRYON, 1887: 76; plt. 15, fig.87. Day, 1917: 474. I.S. OLDRoyD, 1927: 51 as Scala. BURCH, 1945: 20. Epitonium (Acirsa) ochotensis: DALL, 1921: 114. f Acirsa ochotensis: ABBOTT, 1974: 114 (as synonym of A. bore- alis). Description of Aczrsa borealis (Lyell): “‘Whitish or grey- ish fulvous, thick; whorls 8, slopingly, slightly convex, with a spiral rib below, upper portion more or less distinctly longitudinally ribbed, gradually fading until the body whorl is smooth or nearly so. Length 18-22 mill.’ (from Tryon, 1887: 80). Additional Description: Shell brownish in color, thin, elongate, turreted; nuclear whorls 2, smooth; remaining whorls g or 10; suture definite but not impressed; axial sculpture of low ribs which continue to the suture but do not join the ones above, sometimes obsolete from the pe- nultimate whorl down to the basal disk; spiral sculpture of about 15 striae, sometimes brown in color; basal disk strong, elevated, beyond which the axial costae do not ex- tend; aperture round, slightly patulous anteriorly; oper- culum chitinous, paucispiral. Length, 19 to 60 mm; width, 6 to 20mm. Type Material and Type Localities: Scalaria borealis Lyell: Holotype from Uddevalla, Sweden, lost. Hypotypes MCZ no. 187118, from Greenland. Turritella costulata Mighels and Adams: Holotype from Mas- sachusetts, lost (R. I. Johnson, 1949): Neotype MCZ no. 165598, from Georges Bank, Massachusetts. Scalaria ochotensis Middendorff: Holotype in Museum of St. Petersburg (Leningrad), Russia; from south coast of Sea of Okhotsk. Distribution, Ecology and Bathymetric Range: Circum- boreal; in the Atlantic, from Massachusetts north to Nova Scotia, and Greenland; in the northern Pacific, Sea of Okhotsk and the Aleutian Islands, Alaska. Little is known of the ecology of this species. It occurs at least from low water to a depth of approximately gom. [For a complete summary of stations on the Atlantic seaboard see CLENCH & TURNER (1950: 230, 231).] Geologic Record: Pleistocene to Recent. Pleistocene oc- currences of this boreal species are few, at least in the northeastern Pacific. ALLISON (1973: 20) has reported it from the Pleistocene of Amchitka Island in the Bering Sea. Discussion: LYELL (1835) figured an unnamed Turri- tella sp. ina list of fossils from Uddevalla, Sweden. In 1841 he validated the specific name Scalaria borealis when he cited the previous figure of Turritella sp. [See ICZN Code, Art. 16 (vii)]. There has been confusion over the exact date of Lyell’s paper because the Transactions of the Geolog- ical Society of London were published in two parts; Part 1, containing the paper read by Lyell in 1839, was published in 1841, Part 2 was published in 1842. Beck has some- times been given credit for the name, but did not publish it. LYELL (1841: 136) gave him credit for supplying a num- ber of names. Turritella costulata is not only a synonym of Lyell’s taxon but also a homonym of Borson, 1825. Scalaria ochotensis is another synonym of Acirsa bore- alis, although because of its large size (60mm), Midden- dorff thought it to be distinct. Acirsa cerralvoensts (DuShane, 1970) (Figure 2) Epitonium (Acirsa) cerralvoensis DuShane, 1970: 3; fig. 3. Sphon, 1970: 10. Acirsa cerralvoensis: Keen, 1971: 436; fig. 666. DuShane, 1974: 49, 50; fig. 60. Epitonium (Asperiscala) cerralvoensis: Abbott, 1974: 123 (“an Acirsa?”’). Original Description: ‘‘Shell of medium size, tall and slender, thin but strong, light brown in color, with 10-12 gradually enlarging, rounded whorls, two opaque nuclear whorls; suture moderately impressed; axial ridges pale brown, 16 on the fourth whorl, gradually becoming obso- lete on succeeding whorls, spiral ridges wider than the interspaces, 19 on last whorl; peritreme incomplete, lip Vol. 22; No.*2 THE VELIGER Page 97 thin, columella arched; no basal ridge; operculum horn colored, paucispiral. Dimensions (in mm): length 11.8; width 3.7 (holotype):’ (DUSHANE, 1970: 3). Additional Description: A species with a slender, pale brown shell, differing from other Panamic-Galapagan epi- toniids in having reduced axial costae and unevenly spaced spiral striae. Length, 11 to 16mm; width, 3.7 to 4.2 mm. Type Material and Type Locality: Epitonium (Acirsa) cerraluoensis DuShane: Holotype LACM No. 1200; paratype LACM no. 1201; west side of Cerralvo Island, Baja California Sur, Gulf of California, Mexico. Distribution, Ecology and Bathymetric Range: Westside of the Gulf of California south to Cape San Lucas, Baja California Sur, Mexico, occurs on sand substrate in from 7 to 38m. Geologic Record: Pliocene, Recent. Upper Pliocene: San Diego Formation, hill southwest of Goat Canyon, southwestern-most San Diego County, LACMIP loc. gos, [1]. Recent: Specimens in the LACM Collection are all from the Gulf of California, Mexico; 1 specimen off Puerto Refugio, Angel de la Guarda Island (AHF 1051-40) in 38m; 2 specimens west side, Cerralvo Island (LACM Nos. 1200 and 1201) in 13m; 1 specimen off Rancho Palmilla (LACM 66-17), in from 18 to 36m; 1 specimen Pulmo Bay (LACM 66-20), in 7 m; 1 spec- imen off Punta Arena (LACM 66-21), in from 18 to 36 m; 1 specimen off Cape San Lucas (LACM 66-14), in from 18 to 36m. Discussion: Modern occurrences of this species are rare. Lengths of known specimens vary from 11 to 16mm and widths from 3.7 to 4.2 mm. As a fossil this species is known only from the upper Pliocene, in the southwestern-most portion of San Diego County, California. Amaea Adams & Adams, 1853 [Type species (SD, DEBoury, 1909): Scalaria magnifica Sowerby, 1844] Shells white, yellow-brown or dark brown; whorls joined; axial sculpture strong or weak, usually forming rectangles within which are extremely fine threads; sculpture below basal ridge similar to that above. Subgenus Scalina Conrad, 1865; synonym Ferminoscala Dall, 1908 [Type species (SD, PALMER, 1937): Scalaria staminea Con- rad, 1865] Sculpture below basal ridge or keel different from that above. The northern limit in the eastern Pacific of the genus Amaea and subgenus Scalina, previously thought to be the Holocene of Cedros Island, Baja California Norte, Mex- 1cO, is extended to the Pleistocene of southern California (see A. (S.) effiae). Amaea (Scalina) brunneopicta (Dall, 1908) (Figures 3, 4) Epitonium (Ferminoscala) brunneopictum DALL, 1908: 316; pit. 8, fig. 10. Boss, et al, 1968: 51. Epitonium brunneopictum: Dati, 1917: 474. STRONG, 1945: 21. CLENCH & TURNER, 1950: 287, as (Scalina). Scalina ferminiana brunneopicta: KEEN, 1958: 278. Amaea (Scalina) brunneopicta: KEEN, 1971: 436; fig. 670. DUSHANE, 1974: 53, 54; figs. 63, 64. ABBOTT, 1974: 117. ? Eglisia nebulosa DALL, 1919: 348. Boss et al, 1968: 218. Eglisia (?) nebulosa: KEEN, 1958: 277; fig. 155. Eglisia nebulosa: KEEN, 1971: 436; fig. 670 (as synonym of A. brunneopicta). DUSHANE, 1974: 53; fig. 64 (as synonym of A. brunneopicta). Englisia [sic] nebulosa: ABBOTT, 1974: 117 (as synonym of A. brunneopicta). Original Description: ‘‘Shell slender, acute, pale brown- ish, with broad peripheral band and basal disk of darker brown, and about 11 whorls, exclusive of the (lost) nucleus; sculpture of the same type as in the preceding species [Epitonium (Ferminoscala) ferminianum Dall, 1908] with, between the sutures, 4 primary and about 6 secondary spi- rals beside the spiral striae; the axial lamellae are very small and sharp, regularly spaced, little raised and about 36 on the penultimate whorl, they appear as whitish lines on a brown background; basal disk sharply spirally threaded, little raised; aperture as figured, when fully grown probably with a thick varix. Length of shell, 37 [mm], of last whorl, 14 [mm]; of aperture, 8 [mm]; diam- eter of basal disk, 9 [mm]; max. diameter 10mm.’ (DAL, 1908: 316). Additional Description: Shell size medium to large; slender, acute, color brown; whorls 13 to 14, convex, rounded in all stages; nuclear whorls lacking; first 2 whorls crenulated by the axial and spiral sculpture, third whorl with one strong cord at the periphery, fourth and fifth whorls with one strong cord at the periphery with a weaker one on either side, sixth whorl with g strong cords at the periphery with 2 weaker cords on either side, from there Page 98 THE VELIGER Vol. 22; No. 2 on down the cords increase in number until, on the last whorl, there are g rounded, broader cords; sutures mod- erately deep; costae 25 to 40 on the last whorl, white, thin, sharp on early whorls, less sharp on later whorls; the cords intersected by the costae forming small, unevenly spaced rhomboids within which there are fine axial and spiral threads, with the axial threads crossing over the spiral cords forming the rhomboid; basal area brown, defined by a somewhat thickened cord, sculpture same as on the body whorl but fainter; lip white, thin, slightly crenulated by the spiral ribs crossing the last costa, patulous; peritreme incomplete, brownish color of the base can be seen as a brown spot anteriorly on the inside of the aperture; oper- culum dark brown, multispiral. Length 11 to 44mm; width 3 to 12mm. Type Material and Type Localities: Epitonium (Ferminoscala) brunneopictum Dau: Holo- type USNM 97084; Cedros Island, Baja California Norte, Mexico. ? Eglisia nebulosa Dall: Holotype USNM 120702; Cape San Lucas, Baja California, Mexico. Distribution, Ecology and Bathymetric Range: From Cedros Island, Baja California Norte, Mexico (approxi- mately 28°07’ N, 115°11’ W), south throughout the Gulf of California, and south on the west Mexican coast to Costa Rica (10°53'45” N, 85°57'45” W). Occurs on a mud sub- strate, in from g to 72 m. Geologic Record: Pliocene to Recent. Upper Pliocene: San Diego Formation, southernmost San Diego County, California, LACMIP locs. 305 [1697], 305-A [12], 305-C [73], and 319 [23]. Recent Records: This is the only living species of Scalina taken as far north as Cedros Island, Baja California, Mexico. Cedros Island, Baja California Norte, Mexico (ap- proximately 28°047’N, 115°11’W), (type locality), Albatross station 2835, in 10m, one specimen (USNM 97084). Off Cabo Haro, Guaymas, Mexico (27°54’ N, 110°53’ W), Ariel expedition, several specimens, in 36 to 72m, August 1960. Playa Blanca, Costa Rica (10°53'45” N, 85°57'45” W), Velerolll, in 45 m, one specimen (LACM-AHF 463-35) February 1935. Discussion: DALL (1908b: 316) erred when he surmised that Amaea (Scalina) brunneopicta, when fully adult, probably had a thick varix on the aperture. DALL’s (1919: 348) Eglisia nebulosa from off Cape San Lucas, Baja Cali- fornia Sur, Mexico, was based on a worn specimen of A. (S.) brunneopicta. Amaea (Scalina) effiae (Willett, 1939) (Figure 8) Alabina effiae WILLETT, 1939: 202; plt. 54, fig. 1. WILSON & BING, 1970: 6. Alabina (?) aff. A. effiiae: KEEN & BENTSON, 1944: 13; table 1. Original Description: “Shell elongate-conic, white. Nu- clear whorls 2, globular, smooth and shining. Post-nuclear whorls well rounded; sutures deep. Early post-nuclear whorls apparently smooth; later ones with very obscure, raised, irregularly spaced spiral lines, visible only under a lens, and varying in strength in the different specimens examined. Base of last whorl rounded and somewhat wrin- kled at the columella. Imperforate. Aperture broadly Explanation of Figures 1 to 12 Figure 1: Scalaria borealis Lyell, 1841. From Datt, 1921: plt. 13, fig. 10; length, 19 mm; width, 6.5 mm X 2.9 Figure 2: Acirsa cerralvoensis (DuShane, 1970). Holotype, LACM 1200; length, 11.8mm; width, 3.7mm X 3.6 Figure 3: Epitonium (Ferminoscala) brunneopicta Dall, 1908. Hol- otype, USNM 97054; length, 37mm; width,iomm X 1.3 Figure 4: ? Eglisia nebulosa Dall, 1919. Holotype, USNM 120702; length, 19mm, width, 6mm X 2.7 Figure 5: Scalaria bellastriata Carpenter, 1864. Holotype lost; lec- totype by Parmer, 1958, USNM 14831b; length, 9mm; width, 7mm X 3.55 Figure 6: Epitonium (Asperiscala) cookeanum Dall, 1917. Holo- type USNM a2rro19; length, 9.5mm; width, 4mm x6 Figure 7: Scalaria hemphilli Dall, 1878a. Holotype, USNM 7991; length, “about an inch,” Dati, 1878a; Pliocene ca. X 2.2 Figure 8: Alabina effiae Willett, 1939. Holotype, LACMIP Type Collection 1061; length, 12mm, width, 4.5mm; Pleisto- cene X 4.5 Figure 9: Scala lowei Dall, 1906. Holotype lost. Paratype, USNM 191548; length, 7mm; width, 4mm X 5.1 Figure 10: Epitonium (Asperoscala) tinctorium Dall, 1919. Lecto- type by DuSHane (1974: 25), USNM 218100; length, 7 mm; width, 3mm X 5.2 Figure 11: Epitonium (Scala) caamanoi Dall « Bartsch, 1910. Type material lost. Figure after DaLt & BaRTscH, 1910: plt. 1, fig. 1; length, 7mm; width, 4mm xX5 Figure 12: Nitidiscala caamanoi (Dall & Bartsch, 1910). Specimen, LACM 36519; length, 16mm; width, 8mm x3 Tue VELIGER, Vol. 22, No. 2 [DuSHANE] Figures 7 to 12 Figure 2 Figure 4 Figure 5’ Figure 8 Figure 9 Figure 10 Figure 11 Figure 12 Vol. 22; No. 2 ovate, without trace of canal; outer lip thin, slightly re- flected. Columella strongly curved; parietal wall appar- ently not calloused. Length, 12mm; diameter, 4.5 mm<-’ (WILLETT, 1939: 202). Additional Description: Shell medium to large in size, nuclear whorls 2, smooth, remaining whorls 7 to g, con- vex, rapidly increasing in size, sutures moderately deep; spiral sculpture of fine threads varying in size, irregularly spaced; costae on the final whorl weak, irregularly spaced, but forming rhomboids with the threads of the spiral sculpture, axials crossing over the spiral threads forming the rhomboids; basal sculpture apparent, but fainter; aperture oval; outer lip thin, patulous, slightly crenulated by the spiral threads crossing the last costae; peritreme incomplete. Length 12 to 18mm; width 4.5 to 7.5 mm. Type Material and Type Locality: Amaea (Scalina) effiae (Willett): Holotype LACMIP 1061; Pleistocene, “Lower San Pedro series” [= Lomita Marl], Hilltop Quarry, San Pedro, California [LACMIP loc. 64]. Paratypes LACMIP 1729, 1730; Timm’s Point Silt, Timms Point, San Pedro, California [LACMIP loc. 62]. Distribution: Known only as a fossil. Geologic Record: Pleistocene. Lower Pleistocene: Santa Barbara Formation, Packerds [Beacon] Hill, Santa Barbara (Keen and Bentson, 1944: 13; as aff.), Lomita Marl, below Park Western Drive and Host Place, San Pedro, LACMIP loc. 435 [3], and at old Hilltop Quarry [type locality], LACMIP loc. 64 [1], Timms Point Silt, Timms Point, San Pedro, California, LACMIP loc. 62 [2]. Discussion: WILLETT (1939:202) questioned the place- ment of his new species in Alabina because of its large size. His specimens, although worn, indicate a close affinity to Amaea (Scalina) brunneopicta (Dall), having the same fine spiral and axial sculpture and a thin lip. This species ex- tends the known range of Amaea (Scalina) as a Pleisto- cene fossil north into southern California. J. W. Durham (personal comm., 1976) agrees with the placement of effiae in the genus Amaea. Asperiscala DeBoury, 1909 [Type species: (OD) Scalaria bellastriata Carpenter, 1864] Shells white or pink to dark gray or brown; costae usually recurved, sometimes with a spine on the whorl shoulder; sculpture of heavy cords to fine striations, sometimes obso- lete on the later whorls. THE VELIGER Page 99 Asperiscala bellastriata (Carpenter, 1864) (Figure 5) Scalaria bellastriata CaRPENTER, 1864a: 613, 660 [Reprinted, 1872: 99, 146]. CARPENTER, 1866: 221. ORCUTT, 1885: 539. TRYON, 1887: 84. Cooper, 1888: 263. DALL, 1917: 475. Scala bellastriata: ARNOLD, 1903: 269; plt. 9, fig. 17. ARNOLD, 1906: 36. KELSEY, 1907: 49 (as S. ballastriata). CossMAN, 1912: 28; plt. 2, figs. 42, 50. DEBourY, 1919: 34. Epitonium bellastriatum: KEEN, 1937: 35. ABBOTT, 1974: 121; fig. 1246. KENNEDY, 1975: 27. Epitonium (Asperoscala) [sic] bellastriata: DALL, 1921: 114. I. S. OLpRoyp, 1927: 55. Epitonium (Asperiscala) bellastriatum: GRANT & GALE, 1931: 857. KEEP, 1935: 179. DURHAM, 1937: 489. WILLETT, 1937b: 401. STRONG, 1945: 22 (as E. (A.) bellastriata). BuRCH, 1945: 23. SMITH & GORDON, 1948: 190. Epitonium bellastriatum: KEEN, 1937: 35. ABBOTT, 1974: 121; fig. 1246. KENNEDY, 1975: 27. Epitonium (Asperiscala) bellistriatum [sic]: PALMER, 1958: 189; plt. 20, figs. 25, 26 (lectotype). Original Description of Asperiscala bellastriata (Carpen- ter, 1864a): “Shape like pretiosa, jun.: ribs very close, spinous at shoulder, crossed by spiral riblets:’ Later Description: ‘‘S. t. tenuissima, albida; anfr. vii, valde tumentibus, haud nisi per costas attingentibus; costis circ. Xvil, acutis extantibus reflexis, haud semper in spira attengentibus, postice angulatis, parum spinosis; intersti- tils circa spiram basimque dense spiraliter lirulatis, lirulis rotundatis, super varicum angulum obsoletis; apertura ovata, umbilico magno. Long. 0.78, long. spir. 0.55, lat. 0.38; div. 40.” (CARPENTER, 1866: 221). Additional Description: Shell turreted, with elevated spire; nuclear whorls 3, smooth, opaque white, fourth whorl showing axial and spiral sculpture, remaining 5 to 6 whorls rapidly enlarging, very convex; suture deep; costae 15-17, reflected and with a sharp spine at the shoulder, dipping into the suture and joining the next costae above; spiral sculpture clearly defined by 20-25 ribs, interspaces slightly narrower; umbilicus large, aperture oval with re- flected margins resting on 2 costae posteriorly, with a spine at the posterior margin; operculum, very dark brown, horny, paucispiral. Length 3 to 20mm; width 2.33 to 11mm. Type Material and Type Locality: Scalaria bellastriata (Carpenter): Syntypes from southern California lost (fide PALMER, 1958: 190). Lectotype by PALMER, 1958 (USNM 14831b); Monterey, California. Distribution, Ecology and Bathymetric Range: Middle Farallon Island, San Francisco County, California (37° Page 100 THE VELIGER Vol. 22; No. 2 48’ N, 122°59/ W), south to the U.S.-Mexican border (32° 30’N, 117°09’W). Apparently feeds on sea anemones. Dredged in from 18 to 103m, on mud or sand substrate. Geologic Record: Pliocene, Pleistocene, Recent. Upper Pliocene: Fernando Formation, Sixth and Hope Streets, downtown Los Angeles, LACMIP loc. 466 [3 £.], Cherry Avenue and Interstate 40%, Long Beach, LACMIP loc. 423 [25] (KENNEDY, 1975: 27). Lower Pleistocene: Lomita Marl, below Park Western Drive and Host Place, San Pedro, LACMIP loc. 435 [1]. Timms Point Silt, Timms Point, San Pedro, LACMIP loc. 130-7 [1]. Upper Pleistocene: Palos Verdes Sand, Lincoln Avenue, northeast of Playa del Rey (Los Angeles), LACMIP locs. 59 [343] (WILLETT, 1937: 401), and 61 [1]. Dead- man Island, San Pedro, LACMIP loc. 2 [2]. East bluff above Upper Newport Bay, Newport Beach, LACMIP locs. 66-2 [10] (KANAKOFF & EMERSON, 1959: 27), and 66-1 [2]. Newport Mesa, west side of ‘‘middle’”’ New- port Bay, LACMIP loc. 68-B [specimens missing] (KANAKOFF & EMERSON, 1959: 27). Recent Records: Northeast of Middle Farallon Island, San Francisco County, California (37°48’ N, 122°59’ W), 67m, on sand, August 1938, one live specimen (LACM-AHF 887-38). Abalone Point, Laguna Beach, Orange County, California, 97 to 103 m, in mud, one specimen (LACM-AHF 1131-40). Off the U.S.-Mexi- can border (32°30’ N, 117°09’ W), 41m, March 1957, one live specimen (LACM-AHF 4910-57). Most of the recent specimens are from San Pedro to Santa Cata- lina Island, California, perhaps because more dredg- ing has been done in the past from off this limited locality than from any other area. Discussion: ‘The types of Carpenter’s species are lost (PALMER, 1958: 190). The lectotype of PALMER (1958) (USNM 14831b) from Monterey, California is a frag- mented specimen of 2 remaining whorls with the outer lip partially broken, with enough remaining to show that the shell is turreted, with a reflected portion of the upper lip, deep umbilicus and clean-cut spiral sculpture; length 9 mm; width 7mm (Figure 5). Except for CARPENTER’S (1866) Latin description, no complete description or diag- nosis has been published. Off the southern California coast some interesting ob- servations have been made among the artificial reefs made up of old street cars and automobiles. An Associate Marine Biologist for the State of California, the late Charles H. Turner (personal communication, 1968), SCUBA diving offshore from Santa Monica and Hermosa Beaches, Cali- fornia, among the man-made reefs, at 18 m depth, photo- graphed purple pile perch (Rhacochilus vacca). On exam- ining the stomach contents he discovered that the fish had been feeding on Asperiscala bellastriata. All of the fish ex- amined had absorbed the snail’s purple dye into their flesh, effecting a violet-purple hue to the fish, readily observable under water. This is the first reported instance of the pene- trating influence of the snail’s dye into the flesh of a fish, apparently with no ill effects upon the fish. Asperiscala cookeana (Dall, 1917) (Figure 6) Epitonium (Asperiscala) cookeanum DALL, 1917: 475. BAKER, HANNA & STRONG, 1930: 51; plt. 3, fig. 3. BURCH, 1945: 23. KEEN, 1958: 271. KEEN, 1971: 424; fig. 165. DUSHANE, 1974: 11, 12; fig. 8. ABBOTT, 1974: 123. Epitonium (Asperoscala) [sic] cookeana: DALL, 1921: 114. Epitonium cookeanum: I. S. OLDROYD, 1927: 56. KEEN, 1937: 35. Boss, et al, 1968: 89. Epitonium (Asperiscala) cookeana: STRONG, 1945: 22. Original Description: “Shell small, pink, solid, acute, im- perforate, the nucleus lost, with eight well rounded subse- quent whorls; with 10 rather solid, smooth, continuous white varices making less than half a turn round the spire; spiral sculpture of extremely fine uniform threads cover- ing the whorl between the varices; the terminal varix thicker than the others; all the varices broader at the inter- section with the suture but not spinose. Length, 9.5 mm; diameter, 4mm.” (DALL, 1917: 475) Additional Description: Shell small, white, sometimes with a pinkish cast between the costae; nuclear whorls 3, small, brown, glassy; later whorls 8, rounded; costae 10 to 12, solid, white, smooth, heavier at the suture, not re- flected; spiral threads between costae fine; suture deep; umbilicus lacking; aperture round; outer lip heavy, thin- ner parietally. Length 7 to 10mm; width 3 to 5 mm. Type Material and Type Locality: Epitonium (Asperiscala) cookeanum Da.i: Holotype USNM 211019. Named in honor of Miss J. M. Cooke of San Diego on the basis of one specimen obtained for her, presumably from La Paz, by Capt. Porter. Distribution, Ecology and Bathymetric Range: La Jolla and San Diego, California (approximately 32°40’ N, 117° 20’ W), to La Paz, Baja California Sur (24°15’ N, 110915’ W), and Espiritu Santo Island, Gulf of California, Mexico (24°30’ N, 110°15’ W). Nothing is known of the habitat of this species. Geologic Record: Pliocene, Recent. Upper Pliocene: San Diego Formation, in southwestern- Vol. 22; No. 2 THE VELIGER Page 101 most San Diego County, California, LACMIP loc. go5-C [1]. Recent Records: La Jolla, California, one specimen (DuShane Collection). Ocean Beach, San Diego County, California, one specimen (SDNHM 43661). Magdalena Bay, Baja California Sur, Mexico, col- lected by Orcutt, 1917, eight specimens, depository unknown. Cape San Lucas, Baja California Sur, Mex- ico, collected by McLean, 1966, one specimen (LACM 66-13). Espiritu Santo Island, Gulf of California, Mex- ico, one specimen (CAS). Discussion: ‘This is a rare species about which we know little. All of the modern records are of dead specimens, taken intertidally. Asperiscala hemphilli (Dall, 1878) (Figure 7) Scalaria hemphilli Dax, 1878a: 12, 16. DALL, 1878b: 29. Cooper, 1888: 263. ARNOLD, 1903: 28, 44, 264. DEBoury, 1919: 35. Boss, et al, 1968: 155. Scala hemphilli: ARNOLD, 1906: 36. Epitonium hemphilli: DALL, 1917: 472. Epitonium (Boreoscala) hemphilli: GRANT & GALE, 1931: 856. Epitonium (Asperiscala) hemphilli: DURHAM, 1937: 481, 490; pit. 56, fig. 856. Original Description: ‘‘Shell in general resembling a ro- bust specimen of S. indianorum, having from nine to twelve varices on the last whorl, coronated behind near the suture, wholly pure white; surface of the whorls beneath the varices longitudinally delicately sculptured, with alter- nate riblets and grooves. Length about an inch; apical angle about 30°. This species has the sculpture of S. bellastriata, but the shape of S. indzanorum, and is the only grooved species, except the former, which has yet been reported from this region. All the specimens are decollate:’ (DALL, 1878: 16.) Additional Description: Shell large in size, heavy look- ing; nuclear whorls lost, remaining whorls about 6, with fine spiral sculpture and equally fine interspaces; costae 10 to 11, varying from narrow to wide, strongly reflected with an angulation at the shoulder; suture not deep; non-umbil- icate; aperture oval, reflected, resting on the costae which continue to the base; outer lip thickened by the last costa. Length 13 to 22 mm; width 7 to iomm. Type Material and Type Locality: Scalaria hemphilli Dall: Holotype USNM 7991; San Diego Formation, Balboa Park, San Diego, California (Plio- cene). Distribution and Ecology: Extinct; known only from southern California as a fossil. Geologic Record: Pliocene; Pleistocene (?). Upper Pliocene: Niguel Formation, Via La Mirada, San Juan Capistrano, LACMIP loc. 4922 [1]. San Diego Formation, Pacific Beach, LACMIP loc. 4847 [1], Pacific Beach, at end of Arroyo Drive, LACMIP loc. 107 [g], and in southwesternmost San Diego County, LACMIP loc. 305, [4]. Discussion: ‘This species was described at an early date (Dat, 1878a: 16) but was not figured (see Figure 7). Dall compared Asperiscala hemphilli to a robust Nitidiscala indianorum with the spiral sculpture of Asperiscala bella- striata, each of which has a different shell. DALL (1917: 472) later compared his species to Boreoscala greenlandica, which it does not resemble. Asperiscala hemphilli has been reported in the Pleisto- cene upper San Pedro Series [= Palos Verdes Sand] (ARNOLD, 1903: 44, 264), and Pliocene San Diego Forma- tion (ARNOLD, 1906: 36). Asperiscala hemphilli seems to be restricted to the southern California area, with few specimens extant. DurHAM (1937: 490) suggested that A. hemphilli, A. clarki, and A. bellastriata are closely related. DUSHANE (1974: 20, 21) synonymized A. clark: with A. minuticosta (DeBoury). Asperiscala hemphilli differs from A. bellastri- ata by having a more slender shell, fewer costae (10 or 11), finer spiral sculpture, no umbilicus and from A. minutz- costa by having less swollen whorls, heavier costae with a blunter spine at the shoulders, no umbilicus, and a shal- lower suture. Asperiscala lowe: (Dall, 1906) (Figure 9) Scala lowei DALL, 1906: 44. KELSEY, 1907: 49. Boss, et al., 1968: 191. Epitonium lowet: DALL, 1917: 475. I. S. OLDROYD, 1927: 56; plt. 31, fig. 1. KEEN, 1937: 35. Epitonium (Asperoscala) [sic] lowei: DALL, 1921: 114; plt. 6, fig. 11. ipsa (Asperiscala) lowet: STRONG, 1945: 22. KEEN, 1971: 426; fig. 623. ABBOTT, 1974: 123; fig. 1290. DUSHANE, 1974: 18-20; figs. 42, 157. Epitonium (Asperiscala) cf. E. (A.) lowei: Berry, 1956: 153; fig. 1. Original Description: ‘‘Shell small, conic, with five or more rapidly increasing whorls after the (lost) nucleus; color white, whorls very convex, with deep sutures and a small spiral umbilicus; there is no basal disk or cord; sculp- Page 102 THE VELIGER Vol. 22; No. 2 ture of about twenty-seven rather thick, strongly reflected, smooth, close-set varices, and very close, fine spiral threads, covering the whorl between the varices, and separated by about equal sulci; aperture sub-circular, slightly higher than wide, the reflected margin wide at the outer lip, pat- ulous at the inner base, narrow between the shoulder and the preceding whorl, and at the shoulder produced into a short, rather stout spine which, repeated on successive varices, coronates the whorls. Length (without nucleus), 7.0mm, diameter, 4.0mm.’ (DALL, 1906: 44). Additional Description: Shell medium in size, white, conic; nuclear whorls 4, white, in most cases the fourth nu- clear whorl showing fine axial costae, with here and there aheavierone; teleoconch of 5 tog whorls, rapidly enlarging, convex; costae 25 to 32, strongly reflected, with occasion- ally a heavier one, with a spine at the whorl shoulder on all costae; raised spiral ridges between costae, unevenly spaced, forming rectangles within which are 2 fine hori- zontal threads (magnification x 40); sulci twice the width of the ridges; suture deep; umbilicus deep, spiral; aperture oval; lip thin, patulous at base, reflected, with a pointed (adjacent column —) Figure 69 Asperiscala lowei (Dall, 1906) Specimen showing about 20 compressed conical uncini in the row (DuShane Collection). a: area of attachment; b-c-d: tooth re- volved from lateral to ? view; e: tooth seen from attachment area; f: tooth from central area of ribbon; g: tooth from outer area of ribbon; h: orientation of tooth uncertain spine at the shoulder; operculum dark brown, horny, thin, and paucispiral. Length, 3 to 25 mm; width, 11%4 to 9mm. The radula is rectangular, with compressed, conical, horn- like teeth, about 20 in one row (Figure 69). The radular slide is at the SDNHM;; the specimen from which it was dah ia Explanation of Figures 13 to 24 Figure 13: Scala (Viciniscala) minuticosta DeBoury, 1912. Holo- type: Zool. Museum Berlin, East Germany, no. 2578 (fide R. Kilias. Univ. Berlin). Photograph after DEBoury, 1912, Journ. Conchyl. 60 (2): plt. 7, fig. 1; length, 35 mm; width 17mm X 2.4 Figure 14: Epitonium cedrosensis Jordan & Hertlein, 1926. Hol- otype, CAS(GTC) 2116, Pliocene; length, 8.5mm; width, 3.75 mm X 6.5 Figure 15: Epitonium dallas: Jordan «& Hertlein, 1926. Holotype, CAS(GTC) 2122; Pliocene; length, 7mm; width, 4mm X 9 Figure 16: Epitonium clarki T. S. Oldroyd, 1921. Holotype, SU 5948 [now at CAS(GTC)]; Pleistocene; length, 19mm; width, 8mm XK 2.5 Figure 17: Epitonium densiclathratum Dall, 1917. Holotype, US NM 111209; length, 17 mm; width, 7.5 mm x 2.8 Figure 178: Boreoscala greenlandica (Perry, 1811). Greenland. From Perry, 1811: plt. 28, fig. 8 Figure 19: Boreoscala greenlandica (Perry, 1811). Queen Char- lotte Islands, B. C., Canada, from 146 m, rock and sand sub- strate, collected by Bernard and Quayle, October, 1966; length, 39 mm; width, 14mm (DuShane Collection) Xg Figure 20: Boreoscala greenlandica smithi (MacNeill, 1943). Hol- otype, USNM 499037; length, 14.6mm; width, 8mm; Pliocene X3 Figure 21: Scalaria polita Sowerby, 1844 (after SowerBy, 1844). Holotype presumably lost; not found at BM(NH) by M. Keen in 1965. No dimensions given Figure 22: Epitonium (Nitidiscala) pedroanum Willett, 1932. Hol- otype, LACM 1036; length, 11.7mm; width, 3.7mm _ X 4.7 Figure 23: Epitonium implicatum Dall & Ochsner, 1928. Holo- type, CAS(GTC) 2932; length, 18mm; width,6mm X 3.5 Figure 24: Epitonium appressicostatum Dall, 1917. Syntype, US NM 59334; length, 14mm; width, 4.5mm x 4.5 Tue VELIGER, Vol. 22, No. 2 [DuSHANE] Figures 13 to 24 Figure 20 Figure 17 Figure 27 Figure 22 Figure 23 Figure 24 x % Vol. 22; No. 2 THE VELIGER Page 103 taken is in the DuShane Collection, from Bay of Los An- geles, Gulf of California, Mexico. Type Material and Type Locality: Asperiscala lowei (Dall): Holotype lost (Lowe Collection). Paratype, USNM 191548; dredged off Avalon, Santa Catalina Island, California. Distribution, Ecology and Bathymetric Range: From Anacapa Island, California (34°02’N, 119°21’ W), south to Magdalena Bay, Baja California Sur, Mexico (24°32’ 30” N, 112°01’10” W) with many intermediate stations; sparingly into the Gulf of California, Mexico south to Pan- ama. Dredged from gravel, shale, sand and rock substrate in 25 to 171 m depths. Geologic Record: Recent. Recent Records: Northeast of Anacapa Island, Califor- nia (34°02’N, 119°21’W), dredged by Velero III (1938), 81m, on rock, one specimen (LACM-AHF 876-38). Rompiente Point, Baja California Sur, Mex- ico, collected by McLean and LaFollette (Searcher 265), 1971, 90m, shale and gravel substrate, two speci- mens (LACM 71-168). Magdalena Bay, Baja Califor- nia Sur, Mexico (24°32’30” N, 112°01’10” W), bot- tom sample by Velero III (1936), 31m, one minute specimen (LACM-AHF 235). Three additional rec- ords indicate it ranges sparingly into the Gulf of Cali- fornia and south to Panama: San Pedro Martir Island, in 25m, sand substrate (length 21mm; width 9.25 mm (Skoglund Collection); Bay of Los Angeles, in 72 m, live-taken specimen (length 17mm; width 9mm) (DuShane Collection); Panama, live-taken specimen (length 5 mm; width 214 mm) (Shasky Collection). Discussion: ‘The holotype of Asperiscala lowei was re- turned to Lowe after a figure had been drawn of it, but is now missing. It is not at the San Diego Natural History Museum where the Lowe Collection is housed (teste Rap- WIN, 1972). A second specimen, with the same measure- ments, was sent by Lowe (DALL, 1906: 44), also from the same type locality (Avalon, Catalina Island, California), which became a paratype (Figure 9). This taxon varies considerably in size (length 3 to 21 mm; width 114 to gmm). It resembles Asperiscala bella- striata (Carpenter, 1864), but has more reflected and nu- merous costae, with finer spiral sculpture, spaced farther apart, between costae. Both species occur offshore. Asperiscala minuticosta (DeBoury, 1912) (Figures 23, 74, 15, 16) Scala (Viciniscala) minuticosta DEBourY, 1912: 87; plt. 7, fig. 1. DEBourRY, 1919: 40. Epitonium (Asperiscala) minuticosta: DUSHANE, 1974: 20-22; figs. 20-26 [contains complete synonymy]. Epitonium clarki T. S. OvpRoyD, 1921: 115; plt. 5, fig. 13 [not Dentiscala clarki Olsson and Smith, 1951]. GRANT & GALE, 1931: 857 [as E. (Asperiscala)]. DURHAM, 1937: 4go [as E. (Asperiscala)]. WILLETT, 1937b: 401 [as E. (Asperiscala)]. VALENTINE & MEADE, 1961: 12. Epitonium minuticosta: KENNEDY, 1975: 30. Epitonium cedrosensis JORDAN & HERTLEIN, 1926: 446, plt. 30, fig. 3. Epitonium (Aspertscala) cedrosense: GRANT & GALE, 1931: 857. DURHAM, 1937: 489. Epitonium dallasi JoRDAN & HERTLEIN, 1926: 447; plt. 30. fig. 2. Epitonium (Asperiscala) dallast: GRANT & GALE, 1931: 857. DuRHAM, 1937: 489. Original Description of Asperiscala minuticosta (De- Boury): “‘S. testa alba, tenuis, paulum translucens, sat fra- gilis, elongata-conica, anguste sed profunde umbilicata, sutura sat obliqua, angusta et omnino disjuncta; anfract. summis costarum tantum commissi. Anfract. embryonales deficientes. Sequentes 6, valde convexi costis mediocriter obliquis, sat distantibus, tenuibus, acutis, vix prominuis ornuli. Transversim nonnullae striae spirales, sat crebre dis- positae adsunt. Ult. anfract costis 21 munitus. Basis con- vexa, funiculo circumbasati omninodestituts. Columella funiculo nullo firmata. Apertura rotundata. Peristoma in- ternum continuum et foliaceum. Peristoma externum tenue. Long. 35 mm, diam. maj. 17mm. Habitat.—West Colombia” (DEBoury, 1912: 87). Additional Description: Shell medium to large, horn color or white, nuclear whorls 3, brown, glassy, conic; body whorls 7 to 8, well-rounded; suture deep; costae 13 to 21, white, reflected, with wide interspaces, true varices at irregular intervals, short, sharp spine in front of suture; spiral sculpture of about 20 sharp threads on the last whorl, with wider interspaces in which are about 8 axial striae crossing the threads between the costae and about 5, hori- zontal striulae within each rectangle; with a smooth space in front having no sculpture other than the costae and the axial striae, repeated around the narrow but deep umbili- cus; aperture oval; reflected lip continuous, free from the costae, with a sharp spine; operculum horny, dark brown, paucispiral. Length 3 to 35 mm; width 114 to 17mm. Page 104 THE VELIGER Vol. 22; No. 2 Type Material and Type Localities: Scala (Viciniscala?) minuticosta De Boury: Holotype, Zoolo- gisches Mus. Berlin, East Germany, no. 2578 (fide R. Kilias pers. comm. 1978), Univ. Berlin: West Colombia. Epitonium clarki T. S. Oldroyd: Holotype, SU 5948 (now at CAS); Santa Monica, California. Pleistocene. Epitonium cedrosensis Jordan and Hertlein: Holotype CAS (GTC) 2116. Paratypes CAS (GTC) 2117-2120; Bern- stein’s Abalone Camp, Cedros Island, Baja California Norte, Mexico. Pliocene. Epitonium dallasi Jordan and Hertlein: Holotype CAS (GTC) 2122; Turtle Bay, Baja California Sur, Mexico. Pliocene. Epitonium nesioticum Dall « Ochsner. Holotype CAS (GTC) 2928; Paratypes CAS (GTC) 2929, 2930, 2931; Isla Isa- bela (Albemarle Island), Galapagos Islands, Ecuador. Pleistocene. Distribution, Ecology and Bathymetric Range: Southern California (fossil), Cedros Island, Baja California Norte, Mexico (fossil and recent), south to Magdalena Bay (re- cent), east side Gulf of California (recent), west coast of Mexico (recent), Galapagos Islands, Ecuador (recent and fossil). Dredged on sand and broken shell substrate in 18 to 137m. Geologic Record: Pliocene to Recent. Upper Pliocene: Niguel Formation, Via La Mirada, San Juan Capistrano, LACMIP locs. 4923 [2 f.], and 4925 [6 f.]. San Diego Formation, in San Diego, at end of Arroyo Drive, Pacific Beach, LACMIP loc. 107 [9], and at Euclid Avenue and Market Street, LACMIP loc. 485 [6], in southwesternmost San Diego County, LACMIP locs. 305 [483], 305-A [230], 305-B [y], 305-C [487], 318 [9], and 319 [14]; and in Baja Cali- fornia Norte, 414 miles south of International border, on old coast highway, LACMIP loc. 449 [4]. Salada Formation, south side of Bahia Tortugas, Baja Cali- fornia Sur, LACMIP loc. g62 [1 f.]. Lower Pleistocene: Lomita Marl, below Host Place and Park Western Drive, San Pedro, LACMIP loc. 435 [a]. Upper Pleistocene: Long Wharf Canyon [= Potrero Canyon], near Santa Monica, LACMIP loc. 2670 [3]. Palos Verdes Sand, Lincoln Avenue, northeast of Playa del Rey (Los Angeles), LACMIP locs. 59 [240] (WILLETT, 1937b: 401; as E.(A.) clarki),and 5171 [1], Vermont Avenue and Sepulveda Boulevard, west of Carson, LACMIP loc. 147 [1]. Formation uncertain, in San Pedro, on Gaffey Avenue, LACMIP loc. 227 [46], and at Pacific Avenue and Oliver Street, LACMIP loc. 131 [6]. Long Beach, west of Interstate 405 and Cherry Avenue, LACMIP loc. 424 [3]. Palos Verdes Sand, on Pacific Coast Highway between Sev- enth and Colorado Streets, LACMIP loc. 4568 [1 f.] (KENNEDY, 1975: 30). East Bluff above Upper New- port Bay, Newport Beach, LACMIP locs. 66-2 [27] (KANAKOFF & EMERSON, 1959: 27; as E. clarki), 66-1 [go]. Newport Mesa, west side of Newport Bay, LACMIP loc. 68-B [“36,” specimen missing] (Kana- KOFF & EMERSON, 1959:27;as E.clarki). Pacific Coast Highway, below Newport Heights, Newport Beach, LACMIP loc. 241 [1]. Discussion: For a complete synonymy of Panamic-Gala- pagan occurrences see DUSHANE (1974: 20-22, figs. 20-26). This is a commonly dredged species in the Panamic-Gala- pagan area. It has been named and renamed, both as a fossil and as a Recent species, but DeBoury is the earliest author. T. S. OLpRoypD (1921: 115) named and described a Pleistocene fossil from Santa Monica, California as Epitonium clark: (length 19mm; width 8mm) (Figure 16). His description includes the characteristic fading of sculpture adjacent to the sutures, the short, sharp spines on the costae and the deep umbilicus. JORDAN & HERTLEIN (1926: 446, 447) described two Pliocene fossils from Baja California Norte (outer coast) as E. cedrosensis (length 814mm; width 334mm) (Figure 14), from Bernstein’s Abalone Camp, Cedros Island; and £. dallasi (length 7 mm; width 4mm) (Figure 75), from 1.6km southeast of Turtle Bay. Four whorls remain of each of the holotypes. They show the rounded whorls, with definite spiral sculp- ture, reflected costae, although badly worn, and the pointed spine just short of the suture, the typical umbilicus, and the heavy outer lip with the spine at the top of the aperture. Both E. cedrosensis and E. dallas: are well within the range of variability of Asperiscala minuticosta. Fossil occurrences of A. minuticosta north of Cedros Island, Baja California Norte, are extralimital for the Pleistocene and Pliocene of southern California. One live specimen of this taxon was taken by the Allan Hancock Expeditions (Velero III), off Cedros Island (LACM-AHF 1705-49). Asperiscala tinctoria (Dall, 1919) (Figure zo) Epitonium (Asperoscala) [sic] tinctortum DALL, 1919: 340. LowE, 1932: 113; plt. 9, fig. 6. Boss, et al, 1968: 320. Epitonium (Asperiscala) tinctorium. STRONG, 1945: 22, 23. KEEN, 1958: 272; fig. 97. KEEN, 1971: 428; fig. 629. Du- SHANE, 1974: 24; figs. 12, 18. Original Description: “Shell small, white, with a narrow purple-brown spiral line in front of the suture, of six or more whorls excluding the (lost) nucleus; the whorls ad- jacent, the axis imperforate, with 11 to 12 varices which Vol. 22; No. 2 THE VELIGER Page 105 about half encircle the spire; the varices smooth, slightly reflected, wider where they cross the rather deep suture; there are no varical angles or spines; the whorls are uni- formly finely striated; aperture rounded; there is no basal disk or cord; no operculum was taken with the specimen; height of shell, 7; of last whorl, 3; diameter, 3 mm.” (DALL, 1919: 340). Additional Description: Shell small, yellowish-brown, with white costae; nuclear whorls 3, white, glassy; post nuclear whorls 8 or g, rapidly enlarging; costae 8 to 12, slightly reflected, showing growth lines, thicker on the body whorl; spiral threads unevenly spaced between whorls; suture fairly deep; with a purple-brown band on the whorls near the suture; aperture round; shell imper- forate; lip slightly patulous at bottom; operculum brown, horny, thin. Length, 2 to 8.5 mm; width, 1 to 4.5 mm. Type Material and Type Locality: Epitonium (Asperoscala) [sic] tinctorium Dall: Lectotype, USNM 218100 by DUSHANE (1974: 25); four paralecto- types, USNM 698126 by DuSHANE (1974: 25); Magda- lena Bay, Baja California Sur, Mexico. Distribution, Ecology and Bathymetric Range: Magda- lena Bay, Baja California Sur, Mexico, throughout the Gulf of California, south on the west coast of Mexico, pos- sibly to Panama. Usually found on rubble reefs from the intertidal zone down to approximately 6 m. Geologic Record: Pleistocene to Recent. Upper Pleistocene: East bluff above Upper Newport Bay, Newport Beach, LACMIP loc. 136 [3]. South- east of Punta Pequena, outer coast of Baja California Sur, LACMIP loc. 2719 [28]. Discussion: Live-taken shells have white costae and yel- lowish-brown whorls, but beach specimens are white. The brown band between the costae is almost on the outer periphery of the whorl. The costae show growth lines, the spiral striations are fine and irregularly spaced between the costae, but not on them, and the lip is fairly heavy. Fresh shells are easily identified but dead shells present a problem unless these points are noted. When DaLt (1919: 340) described Asperiscala tinctoria he failed to choose a holotype from the 5 specimens in the type lot. DUSHANE (1974: 25) selected a lectotype (USNM 218100) that most nearly fitted Dall’s description and measurements. Although collected by C. R. Orcutt many years ago, it still retains the brown band between the cos- tae, and the fine spiral striations; the aperture is slightly damaged. The remaining 4 specimens are paralectotypes (USNM 698126). Essentially a Panamic species, Asperiscala tinctoria is most frequently taken from around rocks on rubble reefs and under rocks in tide pools. On the outer coast of Baja California it is taken as far north as Magdalena Bay. Pos- sibly because of the inaccessibility of the coast from Mag- dalena Bay north to Scammon’s Lagoon there seem to be no records for this species, although it is quite possible for it to occur there in a shallow water habitat. Boreoscala Kobelt, 1902 [Type species: Turbo clathrus groenlandicus Chemnitz, 1795 (non-binomial) (=Scalaria greenlandica Perry, 1811: OD Kobelt, 1902] Shells usually off-white, costae straplike, strongly sculp- tured continuing to the aperture. Whorls convex, attached, with spiral sculpture of strong ridges with deep depressions between; suture definite, but not deep; basal disk present. Aperture oval, operculum horny, black. Boreoscala greenlandica (Perry, 1811) (Figures 18, 19, 20) Turbo clathrus groenlandicus CHEMNITZ, 1795: 155; plt. 195a, figs. 1878, 1879 (non-binomial). Scalaria greenlandica PERRY, 1811: app., plt. 28, fig. 8. GouLp, 1841: 249, fig. 107 [as S. Groenlandica]. G. B. SowErsy II, 1844a: plt. 34, figs. 105, 110. CARPENTER, 1857: 216, 223, 336 [as S. groenlandica]. DAL, 1873: 58 [as S. grénlandi- cum]. DALL, 1874: 251 [as S. grénlandica]. TRYon, 1887: 76; pit. 16, fig. 91 [as S. grénlandica]. DAL, 1921: 114. DALL, 1925: 6; plt. 22, fig. 2. I. S. OLDRoyD, 1927: 349 [as Scala grénlandica]. Epitonium (Arctoscala) greenlandicum: KEEP, 1910: 184. Epitonium (Boreoscala) greenlandicum: DALL, 1917: 472. GRANT & GALE, 1931: 856. KEEP, 1935: 179 [as E. (B.) groénlandicum]. Epitonium greenlandicum: KEEN, 1937: 35. MORRIS, 1952: 87; plt. 24, fig. 2. CLENCH & TURNER, 1952: 320; plt. 131, fig. 2, plt. 154, figs. 1-3. MACGINITIE, 1959: 83; plt. 5, figs. 2, 3. COWAN, 1964: 111. BERNARD, 1970: 81. ALLISON, 1973: 20. ABBOTT, 1974: 121, 122; fig. 1247. Boreoscala greenlandica: HABE & ITO, 1965: 29; plt. 7, fig. 24. Boreoscala greenlandica ‘(Chemnitz)’: GoLikov & SCARLATO, 1967: 47; fig. 35. Epitonium (Boreoscala) greenlandicum smitht MAcNEI (in MacNet1, MertTie & Pinssry, 1943), 82; pit. 11, fig. 2. Original Description of Boreoscala greenlandica (Perry): “Shell of a pale brown invested with thong-shaped ribs; the four upper folds of the spire smooth; the mouth round. A native of Greenland.” (PERRY, 1811: accompanying plate 28, fig. 8). Page 106 Additional Description: Shell large, pale brown or off- white; nuclear whorls 2 to 4, smooth; post-nuclear whorls 11 to 12, moderately convex, suture impressed, not deep; costae strong, curving into the suture, sometimes nearly covering the intercostal spaces, 9 to 12 on the body whorl. riding over the basal ridge, continuing to the aperture; spiral sculpture of strong ribs, 5 times the width of the deep grooves; basal ridge usually present, with about 9 regularly spaced ribs above the ridge and g to 5 inconspic- uous ribs below; lip usually thickened, aperture subovate; operculum black, horny, with a narrow brown border. Length 11.5 mm; width 4 to 19.5 mm. Type Material and Type Localities: Scalaria greenlandicum Perry: Holotype lost (CLENCH & TURNER, 1952: 320); type locality restricted to Godthaab, southwest Greenland (CLENCH & TURNER, loc. cit.). Epitonium (Boreoscala) greenlandicum smithi MacNeil: Holotype USNM 499037; Inner Submarine Beach, Nome, Alaska (“Pliocene” = Pleistocene). Distribution, Ecology and Bathymetric Range: Circum- boreal, Arctic Ocean near Point Barrow, Alaska, south to British Columbia, Canada; Siberia and northern Japan; taken from depths of 38 m to 163 m. Geologic Record: Pleistocene to Recent. Pleistocene: The species has been reported from the Pleistocene of Amchitka Island in the Aleutians by ALLISON (1973: 20), and the “Inner Submarine Beach” at Nome, Alaska by MacNei (in MacNeEt, MERTIE & PILsBRY, 1943: as Epitonium (Boreoscala) greenlandicum smithi, new subspecies; as “Plio- cene’’); length 14.6 mm; width 8 mm (USNM 499037) (Figure 20). MacNeil’s specimen appears to be an eroded example of Boreoscala greenlandica, with a THE VELIGER Vol. 22; No. 2 broader and flatter base, and with weaker axial costae than is normal for the species. Recent: Cowan (1964: 111), stated this species is known south to Wrangell, Alaska and off the north end of Graham Island, Queen Charlotte Islands, British Co- lumbia, Canada; one specimen (LACM 365,18) from off Point Barrow, Alaska (collected by G. E. MacGin- itie, 1948); one specimen taken off the west coast of Queen Charlotte Islands, British Columbia, from 146 m, rock and sand substrate (collected by F. R. Bernard and D. Quayle, 1966) (DuShane Collection) (Figure 19). Discussion: ‘The earlysynonymy of thiscircumboreal spe- cies in the Atlantic Ocean is well covered by CLENCH & ‘TURNER (1952: 320) and need not be repeated. An account of the biology of this species is given by PERRAN (1978). Depressicala DeBoury, 1909 [Type species: (OD) Scala aurita Sowerby, 1844; synonym: Pictoscala Dall, 1917: type species (OD) Scalaria lineata Say, 1822 (not Réding, 1798; not Kiener, 1838)]. Shells purple-brown with low white costae and fine spi- ral striae that do not form a pattern, aperture oval. Depressiscala polita (Sowerby II, 1844) (Figures 271, 22, 23, 24) Scalaria polita SowERBY II, 1844b: 30. SowErsy II, 1844a: 100; plt. 34, fig. 99. REEVE, 1873: plt. 10, fig. 77. TRYON, 1887: 69; plt. 14, fig. 43. DALL, 1917: 488. Scala politas ADAMS & ADAMS, 1853: 222. Explanation of Figures 25 to 35 Figure 25: Epitonium californicum Dall, 1917. Holotype, USNM 201202; length, 1omm; width, 4mm X5 Figure 26: Epitonium orcuttianum Dall, 1917. Holotype, USNM 273998; length, 6.5mm; width, 3 mm x 7.6 Figure 27: Epitonium diegense Dall, 1917. Holotype, USNM a11- 904; length, 5mm; width, 2mm x4 Figure 28: Epitonium sawinae, variety ? catalinense Dall, 1917. Holotype, USNM 109502; length, 13.5 mm; width, 6mm X 3.6 Figure 29: Epitonium (Crisposcala) catalinae Dall, 1908. Holo- type, USNM 198628; length, 12 mm; width, 4.5 mm X 3 Figure 30: Epitonium (Nitidoscala) tabulatum Dall, 1917. Holo- type, USNM 109569; length, 18 mm; width,7.5mm X 2.2 Figure 31: Epitonium (Nitidoscala) regum Dall, 1917. Holotype, USNM 206596; length, 9mm; width, 4mm X 4.4 Figure 32: Epitonium (Nitidiscala) contrerasi Jordan « Hertlein, 1926. Holotype, CAS(GTC) 2121; length, 22mm; width, gmm; Pliocene X 3 Figure 33: Scalaria hinds Carpenter, 1856. Syntypes (2) BM (NH) 1963.21; left hand specimen designated as lectotype herein; length, 25mm; width, 1omm X 2.6 Figure 34: Epitonium persuturum Dall, 1917. Holotype, USNM 211021; length, 15.5 mm; width, 6mm X 3.2 Figure 35: Epitonium (Nitidiscala) cooperi Strong, 1930. Holo- type, SDMNH 345; length, 20mm; width, 8mm X 2.4 Vol No. 2 [DuSuHANE] Vigures 25 to 3: Tue VELIGER, Vol. 22, : Figure 28 Figure 27 i Figure 31 Figure 29 Figure 30 gure 3 Figure 32 Figure 34 Figure 33 Vol. 22; No. 2 THE VELIGER Page 107 Epitonium (Nitidiscala) politum: KEEN, 1971: 432; fig. 655. DUSHANE, 1974: 35, 39; figs. 89-93, 96. ABBOTT, 1974: 123 (as E. (Nidiscala) [stc]). Epitonium (Nitidoscala) [sic] appressicostatum DALL, 1917: 482. Epitonium (Nitidiscala) appressicostatum: STRONG, 1945: 26, 27. KEEN, 1958: 272. Epitonium appressicostatum: Boss et al., 1968: 31. ABBOTT, 1974: 123. Epitonium implicatum DALL & OCHSNER, 1928: 111; plt. 6, fig. 1. Epitonium (Nitidiscala) pedroanum WILLETT, 1932: 88; plt. 5, fig. 3. STRONG, 1945: 27. BURCH, 1945: 30, 31. Epitonium pedroanum: KEEN, 1937: 35. SPHON, 1971: 11. ABBOTT, 1974: 123. Original Description of Scalaria polita Sowerby: “‘T. tenui, elongata, laevi, imperforata; anfractibus numerosis, vix prominentibus; varicibus sub-numerosis, tenuis, in medis anfractuum obsoletis; colore pallide griseo. The whorls are numerous, and not very prominent. The varices appear as if worn away in the centre of the whorls. Taken by Mr. Cuming at Xipixappi, West Columbia, in sandy mud, ten fathoms deep.’ (SoweErRBy II, 1844a: 30). Additional Description: Shell medium in size, thin, pol- ished, yellowish brown in color; nuclear whorls 3 to 5, smooth, glassy, amber colored; post nuclear whorls 7 to 12, rounded; costae g to 15, white, nearly obsolete on the body whorls, slightly angulate at shoulder, continuous over suture; suture moderately deep; base rounded, non-umbil- icate; aperture oval, margin slightly angular in front. Length 9 to 16mm; width 314 to 5 mm (Figure 22). Type Material and Type Localities: Scalaria polita Sowerby II: Holotype presumably lost; not found at BM (NH) by M. Keen (pers. comm.) in 1965; West Colombia. Epitonium (Nitidoscala) appressicostatum Dall: Lectotype USNM 678703 herein, length 14mm; width 4.5 mm; paralectotype USNM 678703 herein, length 12 mm; width 4 mm; Acapulco, Mexico. Epitonium implicatum Dall and Ochsner: Holotype CAS [GTC] 2932; Galapagos Islands, Ecuador. Pliocene (?). Epitonium (Nitidiscala) pedroanum Willett: Holotype LACM 1036; paratype (1) in J. Q. Burch Collection; San Pedro, California. Distribution, Ecology and Bathymetric Range: Catalina Island, California (33°27'59” N, 118°36’15” W) south to the Galapagos Islands and Ecuador (approximately 1°35’ 10” S, 80°51’55” W). Dredged in from 11 to 393 m, rocks, nullipores, broken shells, and sandy mud substrate. Geologic Record: Pliocene (?) to Recent. Pliocene (?): Indefatigable Island, Galapagos Islands, Ecuador (1°35/10”S, 80°51’55” W), collected by Ochsner and Slevin, from zone “D;’ November 1905, one specimen, as Epitonium implicatum (Holotype CAS [GTC] 2932). Recent Records: West end, Catalina Island, California (33°27'59” N, 118°36’15” W), Velero III, 40m, gray mud, August 1957, one specimen (LACM-AHF 5240- 57). Off San Pedro, California, collected by Willett, 36 m, two specimens as Epitonium pedroanum (Holo- type LACM 1036). Off Agua Chale, Gulf of Cali- fornia, Mexico, Chamizal, 11 m, out of starfish stom- ach, June 1968, one specimen (DuShane Collection). Manzanillo, Mexico, dredged by Shy, 39 m, December 1965, three specimens (DuShane Collection). Off Aca- pulco, Mexico (DALL, 1917: 482), 2 specimens, as Epitonium appressicostatum (USNM 59334).Off La Plata Island, Ecuador, Velero III, February 1934, 18 m, one specimen (LACM-AHF 213-34). Discussion: This subtidal species has a wide geographic range although few records have been obtained. It is easily recognized, being the only species along the north-temper- ate coast to have a yellowish brown shell with white costae and a slender outline. Fresh specimens are glassy, some- times brownish in color, the costae being mere threads. Spiral striulae mark succeeding whorls after the first 2 on some specimens. Occasionally the costae are discontinuous. Nitidiscala DeBoury, 1909 [Type species: (OD) Scalaria unifasciata Sowerby II, 1844] Spiral sculpture between whorls absent. Nitidiscala caamanoi (Dall & Bartsch, 1910) (Figures 12, 12,17) Epitonium (Scala) caamanoi DALL & BARTSCH, 1910: 13; plt. 1, fig. 1. Boss, et al., 1968: 53. Epitonium caamanot: DALL, 1917: 482. STRONG, 1945: 24. T. S. OLDROYD, 1925: 13. I. S. OLDROYD, 1927: 62; plt. 31, fig. 3. KEEN, 1937: 35. BERNARD, 1970: 81. Epitonium (Nitidoscala) [sic] caamanot: DALL, 1921: 116. I. S. OLpRoyD, 1924: 107; plt. 12, fig. 1. Epitonium (Nitidiscala) caamanoi: GRANT & GALE, 1931: 858. STRONG, 1945: 24, 26. BURCH, 1945: 31. Epitonium densiclathratum DALL, 1917: 478. I. S. OLpRoyp, 1927: 61. KEEN, 1937: 35. BURCH, 1945: 29. Boss, et al., 1968: 102. Epitonium (Nitidoscala) [sic] densiclathratum: DALL, 1921: 115. I. S. OLDROYD, 1924, 107. Epitonium (Nitidiscala) densiclathratum: STRONG, 1945: 24. Page 108 THE VELIGER Vol. 22; No. 2 Original Description of Epitoniwm (Scala) caamanoi Dall and Bartsch: ‘‘Shell small, rather conic, white, with thir- teen broadly reflexed, axially conspicuously striated var- ices; nucleus? (lost); whorls, more than six, varices contin- uous up the spire, narrow near the suture, more than doubling in width at the shoulder, where they are provided with a small spine or prominent angulation, then contin- uing to the base, where they are again narrowly contracted; there is no basal disk or cord, the umbilicus is closed; the surface of the whorls between the varices is smooth; the whorls are evenly rounded, and the aperture, if perfect, would probably be nearly circular. Length of six whorls (decollate), 9.5; diameter at base, 5.0, at decollation, 0.7; of aperture, 2.0mm.’ “Collected by Mr. John Macoun, No. 77a (in part), with four other species of the genus, and two dead specimens, in from 7 to 10 fathoms” (DALL & BARTSCH, IgI0: 13). Additional Description: Shell large, white, often worn looking; nuclear whorls eroded; post nuclear whorls 6 to 8, well rounded; costae 11 to 13, broadly reflected, axially striated, solid, thick, continuous from whorl to whorl, nar- row near the suture, expanding on the shoulder to a small spine, remainder of costae doubling in width over the whorl, narrow at the base; surface between whorls smooth; suture distinct; aperture round, lip reflected over the cos- tae; non-umbilicate; operculum horny, round, paucispiral. Length 9.5 to 17 mm; width 5.5 to 7.5mm. Type Material and Type Localities: Epitonium (Scala) caamanoi Dall and Bartsch: Type mate- rial lost; not at Geological Survey Museum, Ottawa, Can- ada (pers. comm. Arthur H. Clarke, 1975); Barkley Sound, Vancouver Island, British Columbia, Canada. Epitonium densiclathratum Dall: Holotype USNM 111209;_ ~ Puget Sound, Washington, taken by the U.S. Fish Com- mission, station 3068, 243 m, green mud. Distribution, Ecology and Bathymetric Range: Barkley Sound, Vancouver Island, British Columbia (50° N, 125° 20’ W) Neah Bay and Puget Sound, Washington (48° N, 122° W). This is a subtidal species and good specimens are taken only by dredging in from 12 to 243m. Geologic Record: Pleistocene to Recent. Upper Pleistocene: Second and Orizaba [now Beacon] Streets, San Pedro, LACMIP loc. 142 [46]. East bluff above Upper Newport Bay, Newport Beach, LACMIP locs. 66-1 [1], 66-2 [5], and 136 [11]. The four Pleis- tocene lots indicate a southern range in earlier times than is apparent from collections of recent material. Recent Records: Ucluclet to Amphitrite Point, Van- couver Island, British Columbia, collected by C. H. Young and W. Spreadborough, 1909, one badly dam- aged specimen (NMC 1415) 6% whorls remain with 12 costae, length 10mm; width 5 mm. Departure Bay, Vancouver Island, British Columbia (49°12’N, 123° 58’ W) one specimen (LACM 36519) with 8 whorls, 11 costae, length 16mm; width 8mm. Seiku, Wash- ington, collected DuShane, one beach specimen, with 7 whorls, 11 costae, length 11mm; width 514mm (DuShane Collection). Discussion: ‘The description of Nitidiscala densiclathra- tum (Dall), ex Carpenter MS (DALL, 1917: 478) agrees with that for Nitidiscala caamanoi and the 2 names are con- sidered synonymous herein. The holotype of N. densi- clathratum (USNM 11 1209)has 7 whorls, 11 costae, length 17mm; width 7.5mm. The only other lot of N. densi- clathratum (USNM 23353) from Neah Bay, Washington, collected by J. G. Swan, has 3 broken specimens. DALL (1921: 115) gave a range from Neah Bay, Washington to San Diego, California but there are no literature citations to substantiate his southern range. Nitidiscala caamanoi differs from other epitoniids within the genus by having a broader shell, inflated whorls, reflected costae, narrow at the distinct suture. Nitzdiscala californica (Dall, 1917) (Figures 25, 26, 27) Epitonium californicum Dati, 1917: 482. I. S. OLpRoyp, 1927: 62. KEEN, 1937: 35. STRONG, 1945: 25. BURCH, 1945: 31. KANAKOFF & EMERSON, 1959: 27. Boss, et al., 1968: 56. Epitonium (Nitidoscala) [sic] californica: DALL, 1921: 116. Epitonium (Nitidiscala) californicum: STRONG, 1930: 195; plt. 20, figs. 11, 12a, 12b. KEEN, 1958: 273. WILSON & KEN- NEDY, 1967: 251. Epitonium (Epitonium) californicum: ABBOTT, 1974: 122. Epitonium diegense DALL, 1917: 483. KEEN, 1937: 35. BURCH, 1945: 32. Boss, et al., 1968: 104. Epitonium (Nitidoscala) [sic] diegensis: DALL, 1921: 116. Epitonium diegensis: I. S. OLDROYD, 1927: 63. STRONG, 1945: 26. Epitonium (Nittdiscala) diegense: KEEN, 1958: 274. Epitonium (Epitonium) diegense: ABBOTT, 1974: 123. Epitonium orcuttianum DALL, 1917: 484. I. S. OLDRoyD, 1927: 66. KEEN, 1937: 35. BURCH, 1945: 24. Boss, et al., 1968: 232. Epitonium (Nitidiscala) orcuttiana: STRONG, 1945: 24. Epitonium (Epitonium) orcuttianum: ABBOTT, 1974: 123. Original Description of Nitidiscala californica (Dall): “Shell small, white, solid, with seven whorls and a very small brownish nucleus of a whorl and a half; varices 9 or 10, low, sharp, reflected, anteriorly axially striated, with a very small sharp spine at the shoulder; suture deep, the Vol. 22; No. 2 THE VELIGER Page 109 whorls almost separated, rounded; base rounded, imper- forate; aperture short-ovate, its inner margin resting on the preceding varices, not touching the body whorl. Length, 10.5; diameter, 4mm. Range, San Miguel Island, California, to the Gulf of California:’ (DALL, 1917: 482). Additional Description: Shell small to large in size, nu- clear whorls 3, brown, glassy, the third whorl down show- ing faint axial sculpture; post nuclear whorls 6 to 7, well rounded, with very faint axial striations; suture deep; cos- tae g to 12, reflected, with some axial growth lines crossing the whorls diagonally and before entering the very deep suture rising to a point, becoming very narrow as they enter the suture, overlapping the costa above; non-umbili- cate, aperture slightly oval, lip reflected, resting on the preceding costae, with a small spine posteriorly, slightly patulous anteriorly, operculum horny, thin, brown. Aver- age dimensions are: length 4 to 11 mm; width g to 4.5 mm. Type Material and Type Localities: Epitonium californicum Dall: Holotype USNM 201202; paratype (1) USNM 635569; San Diego, California. Epitonium diegense Dall: Holotype USNM 211904; para- type (1 plus fragments) USNM 635561; San Diego, Calli- fornia. Epitonium orcuttianum Dall: Holotype USNM 273998; paratypes (many) USNM 635563; San Diego, California. Distribution, Ecology and Bathymetric Range: Nitidis- cala californica appears to be restricted in range from New- port Bay, Orange County, California (33°34’15” N, 117° 30’00 W) south to Cedros Island, Baja California Norte, Mexico (28°20'25” N, 115°11’20” W). Occurs in muddy back bays, intertidally, and sublittorally to depths of 36 m. Geologic Record: Pliocene to Recent. Upper Pliocene: Fernando Formation, Sixth and Hope Streets, downtown Los Angeles, LACMIP loc. 466 [10]. San Diego Formation, end of Arroyo Drive, San Diego, LACMIP loc. 107 [2], southwesternmost San Diego County, LACMIP loc. 305-C [26]. Lower Pleistocene: Lomita Marl, below Park Western Drive and Host Place, San Pedro, LACMIP loc. 435 [4]. Upper Pleistocene: Palos Verdes Sand, Lincoln Avenue, northeast of Playa del Rey (Los Angeles), LACMIP loc. 59 [85]. Corona del Mar, LACMIP loc. 74 [1]. East bluff above Upper Newport Beach, LACMIP loc. 136 [10]. Recent Records: Newport Bay, Orange County, Cali- fornia, collected by A. M. Strong, five specimens (LACM 36520). Todos Santos Bay, Baja California Norte, collected by E. P. Chace, January 1926, one specimen (DuShane Collection). Cedros Island, Baja California Norte, Mexico, collected McLean, et al., RV Searcher, October 1971, two specimens (LACM 71-152). Discussion: The holotype of Nitidiscala diegense DALL (1917: 483), length 5mm; width 2mm (Figure 27), and Nitidiscala orcuttiana Datu (1917: 484), length 6.5mm; width 3mm (Figure 26), have the same characteristics as Nitidiscala californica. All 3 species have sharp spines, in- flated whorls, and the same number of costae. All have San Diego, California as their type localities. DaLL’s (1917: 483) range for N. diegense from San Diego to La Paz, is in error since the specimen (USNM 211507) from La Paz, Baja California Sur, Mexico, on which he based his south- ern record, has a more slender shell and is not related. Datv’s (1917: 482) range for N. californica, from San Mi- guel Island, California to the Gulf of California, cannot be substantiated by southern specimens. It is odd that STRONG (1930), in comparing members of the genus Nitidiscala on the California coast, did not include a discussion of N. die- gense and N. orcuttiana. Nitidiscala californica differs from Nitidiscala tincta, with which it has often been confused, by having a more slender shell, more reflected costae, and with a spine on the shoulders of the whorls and one on the last costa forming the lip. In addition, the costae are of a different shape. They become very narrow before entering the suture, whereas in N. tincta the costae are broad as they enter the suture. Nitidiscala catalinae (Dall, 1908) (Figures 29, 30, 31) Epitonium (Crisposcala) catalinae DAL, 1908a: 252. DALL, 1917: 484. DaLL, 1921: 116 (as (Nitidoscala)) [sic]. I. S. OLpDROYD, 1927: 65. BURCH, 1945: 32. SMITH & GORDON, 1948: 191. Boss, et al., 1968: 67. Epitonium catalinae: STRONG, 1930: 191. KEEN, 1937: 35. WILLETT, 19372: 64. STRONG, 1945: 24. COWAN, 1964: 111. ABBOTT, 1974: 123 (as E. (Epitonium)). Epitonium (Nitidoscala) [sic] tabulatum Datt, 1917: 482. Epitonium (Crisposcala) tabulata: DALL, 1921: 116. I. S. OLp- ROYD, 1927: 66. BURCH, 1945: 33. SMITH & GoRDON, 1948: 191. Epitonium tabulatum: KEEN, 1937: 35. STRONG, 1945: 24. VALENTINE, 1961: 408. Boss, et al., 1968: 313. ABBOTT, 1974: 123 (as E. (Epitonium)). Epitonium (Nitidiscala) [sic] regum Da.x, 1917: 484. Epitonium (Crisposcala) regum: DALL, 1921: 116. I. S. OLD ROYD, 1927: 64. BURCH, 1945: 32. SMITH & GORDON, 1948: 191. tee regum: KEEN, 1937: 35. VALENTINE & MEADE, 1961: 12, 23. Boss, et al., 1968: 275. ABBOTT, 1974: 123 (as E. (Epitonium)). Nitidiscala rega: STRONG, 1945: 24. Page 110 THE VELIGER Vol. 22; No. 2 Original Description of WNitidiscala catalinae (Dall): “Shell slender, white, turreted, imperforate, with more than 7 adherent whorls; nucleus (lost); suture distinct, closed; varices (on the last whorl 14) continuous, making nearly one revolution around the axis in ascending the spire; they are flatly reflected, axially striate, subspinose at the shoulder, giving a tabulate aspect to the rounded whorls. There is no basal disk on the whorl, but on the basal part of each reflection of the varices there is a smooth area over which the suture travels, and which, taken collectively, gives the effect of segments of a disk imposed on the varices but not on the whorl; below the shoulder the varices are widely reflected, extending for a space to the angle of re- flection of the preceding varix, where it would seem these extensions are normally attached, covering a hollow space between them and the whorl, but in the type specimen most of these extensions are broken away; aperture subcircular. Height of (decollate) six whorls, 12.0; of last whorl, 6.0; of aperture, 2.7; maximum diameter, 4.5 mm.” (DALL, 1908a: 252). Additional Description: off-white in color; nuclear whorls 21/2 to 3, brownish; post- nuclear whorls 7 to 12; suture distinct; costae 12 to 20, re- flected, and overlapping the next costae, axially striated, often appearing chalky, angular above, tabulating the pro- file of the spire; lip entire, with a small spine at the shoulder formed by the last costa; non-umbilicate; operculum horny, thin, light brown, slightly oval in shape. Length 7.9 to 18 mm; width 4 to 7.5mm. Shell medium to large in size; Type Material and Type Localities: Epitonium (Crisposcala) catalinae Dall: Holotype, USNM 198628; Catalina Island, California. Epitonium (Nitidoscala) tabulatum Dall: Holotype, USNM 109569: Los Coronados Islands, Baja California Norte, Mexico. Epitonium (Nitidoscala) regum Dall: Holotype, USNM 206596: Point Reyes, Marin County, California. Geographical Distribution, Ecology and Bathymetric Range: From Forrester Island, Alaska (54°50’N, 133° 32’ W) south to at least Cedros Island, Baja California Norte, Mexico (28°o1’ N, 115°29’ W). Occurs on sand sub- strate in from 29 to 110m depths. Geologic Record: Pleistocene to Recent. Lower Pleistocene: Lomita Marl, below Host Place and Park Western Drive, San Pedro, LACMIP loc. 435 [64]. Timms Point Silt, in San Pedro, at Third and Mesa Streets, LACMIP loc. 99 [3], and at Timms Point, LACMIP locs. 62 [2], 130-7 [85], and 4805 [8]. Upper Pleistocene: Palos Verdes Sand, Lincoln Avenue, northeast of Playa del Rey (Los Angeles), LACMIP loc. 59 [2]. Recent Records: Forrester Island, Alaska (54°50'N, 133°32' W), dredged by Willett (1914-1917), 36-72 m, one specimen. La Perouse Bank, off Barkley Sound, Vancouver Island, British Columbia, dredged by Cowan (1964), 68 to 72m, one specimen. Northwest coast of Vancouver Island, British Columbia, dredged by Bernard (1967), 75m, from gravel and mud, one specimen (DuShane Collection). Queen Charlotte Sound, British Columbia (51°54’N, 128°51’ W), dredged by D. B. Quayle on G. B. Reed, 106m, two specimens, one live-taken (LACM 69-71). Point Reyes, Marin County, California, dredged by U.S. Fish Com- mission, station 4309, 110m (USNM 206596), as Niti- discala regum (Dall). Coronados Islands, Baja Cali- fornia Norte, Mexico, dredged by U.S. Fish Commis- sion, station 2931, 61m (USNM 1095609), as Nitidis- cala tabulata (Dall). Eight miles southwest of Cedros Island, Baja California Norte, Mexico (28°01/23’N, 115°29'37” W), dredged by Velero III (1950), three specimens (LACM-AHF 1948-50). Discussion: Although Dall placed Nitidiscala catalinae (Figure 29), in the subgenus Crisposcala DuRHAM (1937: 483) states, “C'risposcala has a basal disk; it is distinguished by anetwork of crossed spiral lines that produce a decussate appearance on the varices. Living in New Caledonia (De- Boury) Eocene to Recent.” As seen in well-preserved speci- mens, the species is most unusual because of the overlapping costae. Most specimens have suffered damage and are some- times difficult to identify. Nitidiscala tabulata (USNM 109569) (Figure 30) the holotype of which has 16 costae, 5% remaining whorls, is here considered a synonym of N. catalinae. The type of Nitz- discala regum (USNM 206596) (Figure 37) with 19 costae and 5 remaining whorls is a badly damaged specimen of N. catalinae. Nitidiscala catalinense (Dall, 1917) (Figure 28) Epitonium sawinae, variety ? catalinense DALL, 1917: 481. STRONG, 1930: 191; plt. 20, fig. 10. Epitonium (Nitidoscala) [sic] catalinensis: DALL, 1921: 116. Epitonium catalinensis: WILLETT, 1937b: 401. BURGH, 1945: 30. ABBOTT, 1974: 119. Epitonium catalinense: I. S. OLDROYD, 1927: 61. KEEN, 1937: 35- Epitonium sawinae catalinense: Boss, et al., 1968: 68. Original Description: ‘Shell with a small three-whorled nucleus, smooth and white, and seven and a half subsequent whorls; varices 22 to 24, not spinose or angular, not regu- larly continuous over the suture, with the anterior faces of Vol. 22; No. 2 THE VELIGER Page 111 the varices finely lamellose or deeply striated. Base rounded with a minute umbilical perforation in the adult; aperture nearly circular. Length, 13.5; diameter, 6mm. Range, off Catalina Island, California. It differs from sawinae by the absence of angularity on and the greater number of varices, and by the umbilical perforation.” (Dat, 1917: 481). Shell white, thin, nuclear whorls 3, smooth, amber colored; post nuclear whorls 9 to 10, very inflated, glossy; costae 19 to 30 or more, thin, with a few axial striations on the faces; sutures definite but not deep; base rounded; aperture oval, with reflected face rest- ing on the anterior costae; peritreme complete but very thin and easily broken, barely covering a minute umbilicus. On older specimens, a lime deposit covers all but the upper 4 of each nuclear whorl giving the appearance of a dark, cir- cular band at the top of each whorl. On the fourth whorl down sculpture of fine costae covers half of the whorl, the remainder being amber in color. Length 3 to 22 mm; width .75 to8mm. Additional Description: Type Material and Type Locality: Epitonium sawinae, variety ? catalinense Dall: Holotype, USNM 109502; off Catalina Island, California. Distribution, Ecology and Bathymetric Range: Van- couver Island, British Columbia (50°40’05” N, 128°31’9” W), south to Todos Santos Island, Baja California Norte (31°53 20 N, 116°48'15” W). Occurs in mud, gray-green sand or gravel substrate in from 20 to 360m depths. Geologic Record: Pliocene to Recent. Upper Pliocene: San Diego Formation, southwestern- most San Diego County, LACMIP locs. 305 [82], 318 [9], and 319 [4]. Lower Pleistocene: Lomita Marl, in San Pedro, at old Hilltop Quarry, LACMIP loc. 64 [1], and below Park Western Drive and Host Place, LACMIP loc. 435 [13]. Upper Pleistocene: Palos Verdes Sand, Lincoln Avenue, northeast of Playa del Rey (Los Angeles), LACMIP loc. 59 [2] (WILLETT, 1937b: 401). Recent Records: NW tip of Vancouver Island, British Columbia (50°40'5”N, 128°31’9” W), dredged by D. B. Quayle on A. P. Knight, FRB Station 68-10, 110m, 1968, two specimens (LACM 68-177). Pacific Grove, Monterey Bay, California, collected by Mc- | Lean, in 36m, 1960, two specimens (LACM 60-23). Southwest side of Santa Rosa Island, Santa Barbara County, California, dredged by Velero III, 1941, 77- 81m, on green mud, one specimen (LACM-AHF 1390-41). East side of Santa Barbara Island, Califor- nia, collected McLean and Margetts, diving at 30 to 70 feet, 1972, one specimen (LACM 72-97). Four miles north of Todos Santos Island, Baja California Norte, Mexico, dredged by Velero III, 1941, in 74m, from shell, mud, gray sand substrate, one specimen (LACM- AHF 1245-41). Discussion: Although named as a variety of Nitidiscala sawinae by Datu (1917: 481), Nitidiscala catalinense is dis- tinct (Figure 28). It differs by having a more tabulate out- line, with more inflated (convex) whorls, a deeper suture, more costae, and a minute umbilicus partially hidden by the thin, reflected lip. This species is not rare but is usually en- countered in from 20 to 360m off the islands of the Santa Barbara Channel group. WiLLETT (in BurcH 1945: 30) states, “Specimens taken in 20-40 fms [36 to 72m], (San Pedro, Catalina), vary greatly, some shoulders being rounded, others angled and varices numbering from 19-27 . .. However, 12 specimens taken at 200 fms [360m] off Catalina Island all have shoulders rounded and varices are more in number, 25 to 32.” Nitidiscala hindsii (Carpenter, 1856) (Figures 32 to 35) [For a complete synonymy on this species in the southern part of its range (Panamic Province), see DUSHANE (1974: 34)]. Scalaria hindsit CARPENTER, 1856: 165. CARPENTER, 1857a: 284, 336. Scala hindsii: ARNOLD, 1903: 264. Epitonium (Nitidiscala) hindsii: PALMER, 1963: 331; plt. 67, figs. 3-6. DUSHANE, 1974: 15, 34-35; figs. 79-81, 83-84, 87-88. Not Epitontum or Scalaria hindsvi of California authors (e. g., KEEP, 1911: 183, fig. 174 = E. tinctum (Carpenter) q. v. Epitonium (Nitidoscala) [sic] fallaciosum DALL, 1917: 478 (nomen nudum). WoopRING, 1931: 31. Boss, et al., 1968: 126 (“new name”). [Invalid, lacking indication.] Not E. (N.) fallactosum DALL, 1921 = E. tinctum (Carpenter), q.v. Epitonium persuturum DALL, 1917: 478. OLDROYD, 1927: 59. Epitonium (Nitidoscala) [sic] persutura: DALL, 1921: 115. Epitonium (Nitidiscala) contrerasi JORDAN & HERTLEIN, 1926: 446; plt. 30, fig. 4. GRANT & GALE, 1931: 858. DURHAM, 1937: 488. Epitonium (Nitidiscala) cooper STRONG, 1930: 194; plt. 20, figs. 6-8. WiLson & KENNEDY, 1967: 251. Epitonium (Epitonium) cooperi: ABBOTT, 1974: 119; fig. 1236. Original Description of Nitidiscala hinds (Carpenter): “S. testa ‘S. Cumingii’ simili, sed magis elongata, majore, anfr. x.haud profunde separatis; varicibus acutis viil., acutis coronatis, lineis regularibus, ad marginem alteram spirae Page 112 parallelis, ascendentibus. Long. 1.04, long. spir. .79, lat. .4, div. 25°.” (CARPENTER, 1856: 165). Additional Description: Shell medium to large in size, white, nuclear whorls three; opaque, postnuclear whorls seven to eleven, loosely coiled; costae eight to fourteen, sharp, high, thin, with an angle or sharp spine at the shoul- der, free of the reflected outer lip; suture deep and at an angle, giving the whorls a slanted appearance; aperture oval; operculum horny, thin, paucispiral. Length 3 to 26 mm; width 11/2 to 10mm. Type Material and Type Localities: Scalaria hindsii Carpenter: Syntypes (2), BM(NH) 1963. 21; Panama. Lectotype designated herein, BM(NH) No. 1963.21, larger of the two syntypes (length 25 mm; width 10 mm). Epitonium fallaciosum Dall: California (no locality; fig- ured specimen of Keep, present repository undeter- mined). Epitonium persuturum Dall: Holotype USNM 211021; San Diego, California. Epitonium (Nitidiscala) contrerasi Jordan and Hertlein: Holotype CAS (GTC) 2121: Turtle Bay, Baja California Sur, Mexico; CAS loc. 945, Pliocene. Epitonium (Nitzdiscala) cooperi Strong: Holotype SDNHM 345; paratypes SDNHM 3386-3391; USNM 46222, and 56052; San Pedro, California. Distribution, Ecology and Bathymetric Range: From Forrester Island, Alaska (54°50’ N, 133°32’ W), south along the north temperate coast including the outer coast of Baja California, the Gulf of California, and south to Peru (14° THE VELIGER Vol. 22; No. 2 14/08” S, 76°08’30” W). Intertidal to 195 m, on mud, silty sand, or gravel bottoms. Geologic Record: Pliocene to Recent. Upper Pliocene: Fernando Formation, downtown Los Angeles, Fifth and Hope Streets, LACMIP loc. 27 [1, 4f.], and Sixth and Hope Streets, LACMIP loc. 466 [6]. San Diego Formation, southwesternmost San Diego County, LACMIP loc. 305 [12]. Turtle Bay, Baja California Sur, Mexico, CAS loc. 945 (Jordan and Hertlein, 1926: as E. contrerasi). Lower Pleistocene: Santa Barbara Formation, east of Santa Barbara General Hospital, LACMIP loc. 183-A [1]. Upper Pleistocene: Isla Vista, near Goleta, LACMIP loc. 416 [2]. Palos Verdes Sand, Lincoln Avenue, near Playa del Rey (Los Angeles), LACMIP locs. 59 [11], and 4745 [103], near Vermont Avenue and Sepulveda Boulevard, west of Carson, LACMIP loc. 147 [3], south of Union Oil Refinery, northern San Pedro, LACMIP loc. 1210 [25]. Formation uncertain, in San Pedro, at San Pedro and Wilmington Road, LACMIP loc. 299 [7], Second and Orizaba [now Beacon] Streets, LACMIP loc. 142 [7], below old Nob Hill, LACMIP loc. 300 [18], and at Arnold’s lumberyard exposure, LACMIP loc. 76 [4]. West of Interstate 405 and Cherry Avenue, Long Beach, LACMIP loc. 424 [3]. East bluff above Newport Bay, Newport Beach, LACMIP locs. 66-2 [141] (KANAKOFF & EMERSON, 1959: 27 as N. indianorum, in part), 66-1 [7], 66-9 [1], Explanation of Figures 36 to 48 Figure 36: Epitonium (Nitidiscala) indianorum (Carpenter, 1865). Lectotype of Stronc (1930: plt. 20, fig. 1) USNM 15521, “Neeah Bay - J. G. Swan;” length, 26mm; width, 9mm x 1.88 Figure 37: Epitonium columbianum Dall, 1917. Holotype, USNM 111211; length, 21 mm; width, 7.5 mm XK 2.1 Figure 38: Epitonium montereyense Dall, 1917. Holotype, USNM 111217; length, 6 mm; width, 2.5 mm X 4.8 Figure 39: Scala rectilaminata Dall, 1907. Holotype, USNM 110- 430; length, 3.25 mm; width, 1.6mm x2 Figure 4o: Scala berry: Dall, 1907. Holotype, USNM_ 107724; length, 3.5mm; width, 1.75 mm X2 Figure 41: Nitidiscala tincta (Carpenter, 1865). Laguna de San José, Baja California Norte, Mexico, specimens with egg clusters, in tide pools with large sea anemones (DuShane Collection) Figure 42: Nitidiscala sawinae (Dall, 1903). Santa Barbara Island, California; length, 19mm; width, 7mm (DuShane Collec- tion) X 2.5 Figure 43: Epitonium (Crisposcala) acrostephanus Dall, 1908. Holotype, USNM 110638; length, 20mm; width, 6.5mm X7 Figure 44: Scala sawinae Dall, 1903. Holotype, USNM 109309; length, 10.5 mm; width, 4mm X 7-5 Figure 45: Scalaria subcoronata Carpenter, 1866. Holotype, USN M 14830b; length, 13mm; width, 7mm X 3.8 Figure 46: Epitonium tinctum var. bormanni Strong, 1941. Para- type, DuShane Collection; length, 4.5mm; width, 2.5mm X 117 Figure 47: Epitonium continuatum T. S. Oldroyd, 1925. Holotype, USNM 352383 (type lost since 1952, fide Rosewater, 1975) 5 length, 8mm; width, 3.6mm. Figure from T. S. O.tproyp (1925: plt. 2, fig. 10) X 6.5 Figure 48: Scalaria (? Indianorum, var.) tincta Carpenter, 1865. Lectotype of Patmer, (1958: 189); USNM 19510; length, 10.5 mm; width, 4.5 mm X 4.5 THE VELIGER, Vol. 22, No. 2 [DuSHANE] Figures 36 to 48 Figure 39 Figure 40 Figure 36 Figure 37 Figure 38 Figure 42 Figure 43 Figure 44 Figure 45 Figure 46 Figure 47 Figure 48 F Vol. 22; No. 2 and 136 [43]. Pacific Coast Highway, below Newport Heights, Newport Beach, LACMIP loc. 241 [1]. Near MacArthur Boulevard and San Joaquin Hills Road, Newport Beach, LACMIP loc. 480 [1]. Bay Point Formation, old Spanish Bight, Coronado Island (San Diego), LACMIP loc. 336 [2], and on Pomona Street, Glorietta Bay, Coronado, LACMIP loc. 2658 [1]. Recent Records: Nitidiscala hindsu ranges from For- rester Island, Alaska (54°50’ N, 133°32’ W), south to Independencia Bay, Peru (14°14’08” S, 76°08’30” W), with documentation of many stations throughout its range. Forrester Island, Alaska, ex Antoniss Jay Col- lection [53 specimens, dead. Intertidal ?] (LACM 13448). Venado Island, Canal Zone, Panama, col- lected McLean, 1970, one specimen (LACM 70-15). Independencia Bay, Peru, Velero III, 1935, two speci- mens (LACM-AHF 380-35). Discussion: CARPENTER (1856: 165) described Scalaria hindsu from Panama. He later recognized the species in the California fauna, but because some of the material so iden- tified turned out to be Epitonium tinctum, there has been confusion. Dall attempted to recognize the California form as distinct, but unfortunately in proposing the name E. fal- laciosum for it in 1917 he gave no real diagnosis, cited no type specimen, and did not select a type locality. In 1921 he remedied the oversight by citing a figure, based on KEEP (1911: 183, fig. 174), but unfortunately one that authors agree is of E. tinctum. Because there is a geographically consistent pattern of variation within Nitzdiscala hindsii, it would be convenient to recognize two geographic sub- species (drawing the boundary line between them, how- ever, might be difficult)—the southern or Panamic sub- species having a larger shell, with fewer costae (8 to 9) the northern variant tending to be smaller with up to 14 costae. The name for the latter, given by Strong, Epitonium coopert, has been accepted by many authors as taking the place intended by Dall for his EF. fallactosum. Wooprinc’s attempt (1931) to supply an “indication” for the latter by designating a “lectotype’”’ was futile, for Dall had in 1921 supplied the indication when he cited a figure, though that turns out to represent another species. Against adoption of E. cooperi stands the barrier of two prior synonyms: E. per- suturum Dall, 1917, based on a beachworn shell, and E. contrerast Jordan and Hertlein, 1926, based on a fossil. Neither has received much currency. Because of this matter of priority, it seems better to regard the shift in morpho- logical characters from south to north as clinal and to con- sider that the true range of N. hindsi is from Alaska to Panama. DUSHANE (1974: 34-35) synonymized 6 nominal species from the Panamic-Galapagan region with Nitidiscala hind- THE VELIGER Page 113 sw. In addition, Epitontum persuturum Dall, and Epito- nium cooperi Strong are synonymized herein. According to Burcu (1945: 29), Nitidiscala hindsii is commonly dredged off the southern California coast in from 65 to 195m, on a gravel substrate. It is often washed in by the hundreds on beaches after a storm. Common as dead shells are, I have seen only one live-taken specimen, dredged in 11 to 15m off Puertecitos, Gulf of California, Mexico, December, 1964 (DuShane Collection). Nitidiscala indianorum (Carpenter, 1865) (Figures 36, 37, 38) Scalaria indianorum CARPENTER, 1864: 613, 628, 660, 683 [reprinted, 1872: 99, 114, 146, 149]. CARPENTER, 1865: 10 [Reprinted, 1872: 244]. Cooper, 1888: 263. SowERBY, in REEVE, 1873: plt. 11, fig. 81. DALL, 1878a: 27, 28. DALL, 1878b: 12. ORCUTT, 1885: 539. TRYON, 1887: 70; plt. 14, figs. 48, 49. Scala indianorum: ARNOLD, 1903: 15, 19, 22, 24, 25, 28, 31, 45, 264. ARNOLD, 1906: 31, 36. Epitonium indianorum: KEEP, 1911: 183. Moopy, 1916: 43. T. S. OLpRoyp, 1925: 13. I. S. OLDROYD, 1927: 57, 58. KEEN, 1937: 35. KEEN, 1943: 37. WILLETT, 1937b: 32. Morris, 1952: 87; plt. 24, fig. 4. EMERSON & ADDICOTT, 1953: 439. VALENTINE, 1961: 360. AppICOTT & EMERSON, 1959: 15, 21. KANAKOFF & EMERSON, 1959: 27. VALENTINE & Meapg, 1961: 12, 17, 23. ADpicoTT, 1966: 4; pit. 2, fig. 29. BERNARD, 1970: 81. ABBOTT, 1974: 119. Epitonium (Nitidoscala) [sic] indianorum: DALL, 1917: 477. DALL, 1921: 115. I. S. OLDROYD, 1924: 107. I. S. OLDRoYD, 1927: 57. STRONG, 1930: 186, 192; plt. 20, figs. 1, 2a-b. GRANT & GALE, 1931: 859. : Epitonium (Nitidiscala) indianorum: T. S. OLDROYD, 1925: 13. DURHAM, 1937: 481, 487; plt. 56, fig. 14. WILLETT, 1937b: 401. KEEN & BENTSON, 1944: 155. BURCH, 1945: 24, 27. A. G. SMITH & GorDON, 1948: 191. BERRY, 1956: 153. PALMER, 1958: 186; plt. 20, figs. 23, 24. GLEN, 1959: 185; fig. 5. WILSON & KENNEDY, 1967: 252. Moore, 1968; 58; plt. 27, fig. g. Epitonium (Gyroscala) indianorum: ABBOTT: 1974: 165. Epitonium columbianum Datt, 1917: 481. I. S. OLDRoyD, 1927: 60. KEEN, 1937: 35. BURCH, 1945: 30. Boss ef al., 1968: 85. Epitonium (Nitidoscala) [sic] columbiana: DALL, 1921: 115. Epitonium (Epitonium) indianorum: ABBOTT, 1974: 122. Epitonium montereyense DAL, 1917: 481 [not Scala (Cirso- trema) montereyensis DALL, 1907]. I S. OLDROYD, 1927: 57. Boss, et al., 1968: 213. Scalaria regiomontana DaLt in DEBoury, 1919: 39 (new name for E. montereyense Dall, 1917). Scalaria (Nitidoscala) [sic] regiomontana ‘Dall, 1917’ [sic]: DALL, 1921: 116 [new name for montereyense Dall, 1917]. Boss, et al., 1968: 275. Epitonium regiomontanum: I. S. OLDROYD, 1927: 64. KEEN, 1937: 35. BURCH, 1945: 31. SMITH & GoRDON, 1948: 191. Page 114 Epitonium (Epitonium) regiomontanum: ABBOTT, 1974: 122. Original Description of Nitidiscala indianorum (CARPEN- TER, 1865): “Scalaria testa gracili, turrita, alba: anfr. circ. X., rotundatis, parum separatis, laevibus; basi sim- plici, haud umbilicata; costis viii.-xv. (plerumque xii), acu- tioribus, subreflexis, interdum latis, plerumque lineis irre- gularibus margini spirae recto parallelis ascendentibus, rarius juxta suturam subnodosis; apertura ovata. Long. 1.05, long. spir. .8, lat. .36, div. 28. Hab. Neeah Bay (Swan). Strung as ornaments by the Indian children. Intermediate between S. communis and S. Turtonts, and scarcely differs from ‘S. Georgettina, Kien.,’ Mus. Cum. No. 34, Brazil.” (CARPENTER, 1865: 10). Additional Description: whorls 8 to 10, convex, with longitudinal growth lines; costae 10 to 17, low, rounded, somewhat striated, contin- uous over the suture which is shallow; at the suture each costa joining the one above and extending along the suture to the next costa on the left; costae on the lower and middle portion somewhat reflected unless worn, when they appear to be thick, rarely with a small spine at the shoulder; costae continuous to the base and the columellar lip, to which they fuse; non-umbilicate, lip entire; operculum horny, pauci- spiral. Length 7.4 to 38mm; width 3.3 to 12mm. Shell large in size, white, solid; Type Material and Type Localities: Scalaria indianorum Carpenter: Lectotype USNM 15521 by STRONG (1930: plt. 20, fig. 1); paralectotypes (4) plus fragments (2) USNM 635564; Neah Bay, Washington. Epitonium (Nitidoscala) [sic] columbianum Dall: Holo- type USNM 111211; off the Columbia River, Oregon. Epitonium_montereyense Dall: Holotype USNM 111217 (rejected homonym of Scala (Cirsotrema) montereyensis Dall, 1907); Monterey Bay, California. Distribution, Ecology and Bathymetric Range: Forrester Island, Alaska (54°50’ N, 133°32’ W), south to Todos San- tos Bay, Baja California Norte, Mexico (31°53’ N, 116°48’ W); intertidal to 114m; sandy mud substrate. Geologic Record: ? Miocene, Pliocene to Recent. Upper Pliocene: Fernando Formation, near Interstate 405 and Cherry Avenue, Long Beach, LACMIP loc. 423 [1]. San Diego Formation, southwesternmost San Diego County, LACMIP loc. 305-C [1]. Lower Pleistocene: Port Orford Formation, southeast of Cape Blanco and north of old mouth of Elk River, Ore- gon, LACMIP locs. 3947 [12, 3 f.], 3949 [1], and 3960 [7]. Elk Head, north of Trinidad, LACMIP loc. 3941 [1]. Crannell Road and highway 101 frontage road, south of Trinidad, LACMIP loc. 3936 [19]. Santa Barbara Formation, Rincon Hill Road, southeast of Carpinteria, LACMIP loc. 473 [1]. Stockton Ranch, THE VELIGER Vol. 22; No. 2 Las Posas Hills (Ventura County), LACMIP loc. 161 [3]. Don Felipe Drive, Baldwin Hills, Los Angeles, LACMIP loc. 293 [1]. Lomita Marl, in San Pedro, at old Hilltop Quarry, LACMIP loc. 64 [9], and below Host Place and Park Western Drive, LACMIP loc. 435 [21]. Upper Pleistocene: Near Cayucos, LACMIP loc. 421 [5]. Isla Vista, near Goleta, LACMIP loc. 2694 [r]. San Pedro Sand, Second and Orizaba [now Beacon] Streets, San Pedro, LACMIP loc. 142-B [5]. Forma- tion uncertain, in San Pedro, at Pacific Avenue and Oliver Street, LACMIP loc. 131 [1], Third and Mesa Streets, LACMIP loc. 98 [3], and at Deadman Island, LACMIP loc. 2 [4, 1f.]. Palos Verdes Sand, Lincoln Avenue, Playa del Rey (Los Angeles), LACMIP locs. 59 [29] (Willett, 1937b: 401), and 4745 [14], Anaheim Street, east of Normandie Avenue, Harbor City (Los Angeles), LACMIP loc. 229 [6], San Pedro, LACMIP loc. 2695 [1 f.], Eighth and Palos Verdes Streets, San Pedro, LACMIP loc. 226 [3], “Crawfish Georges,” 22™¢ and Mesa Streets, San Pedro, LACMIP loc. 2697 [2], Pacific Coast Highway between Seventh and Col- orado Streets, Long Beach, LACMIP loc. 4568 [4] (KENNEDY, 1975:30; as cf.). East bluffs above Upper Newport Bay, Newport Beach, LACMIP loc. 66-2 [1] (KANAKOFF & EMERSON, 1959: 27; in part). Pa- cific Coast Highway, below Newport Heights, New- port Beach, LACMIP loc. 241 [1]. Recent Records: Forrester Island, Alaska, collected by Willett (1914-1917), 72m (USNM 216346). Neah Bay, Washington, collected by J. G. Swan, five speci- mens and fragments (USNM 15521 and 635564). Off the Columbia River, Oregon, taken by the U.S. Fish Commission, station 3065, 49 m, fine black sand, length 21mm; width 7.5mm, with nine whorls and 17 costae as Epitonium columbianum Dall, 1917 (US NM 111211), Monterey Bay, Monterey County, California, taken by the U. S. Fish Commission, station 4140, in 54m, length 6mm; width 2.5mm; with 6 whorls and 14 costae as Scalaria regiomon- tana Dall, 1921 (USNM 111217), Carmel Subma- rine Canyon, Monterey County, California, col- lected by McLean diving at 40 to 125 feet, one live- taken specimen, length 38mm; width 7mm, with 12 whorls and 13 costae (LACM 60-24). Todos Santos Bay, Baja California Norte, Mexico, taken by the U.S. Fish Commission, station 2936, in 108 m, juvenile spec- imens (USNM 209427). Discussion: The range of Nitidiscala indianorum is ex- tensive along the coast from Alaska south to Todos Santos Bay, Baja California Norte, Mexico. Southern specimens Vol. 22; No. 2 THE VELIGER. Page 115 are smaller, indicating the terminus of the range. Two nom- inal species, Nitidiscala columbiana and Nitidiscala regio- montana are placed in synonymy herein because no differ- ences in shell characters can be found. Each is within the geographical range for N. indianorum. DALL (1917: 481) based his southern record of N. columbiana (USNM 97027) dredged off Punta Abreojos, Baja California Sur, Mexico on I specimen that appears to be of another species. Nitidiscala indianorum is sometimes confused with N. tincta, but has a much larger shell and lacks the brown su- tural line sometimes found in N. tincta. Nitidiscala indiano- rum also occupies a more northern, although somewhat overlapping, geographical range. It is also found only sub- littorally and must be dredged or obtained by diving, whereas N. tincta occurs mainly intertidally. Although Carpenter mentioned Nitidiscala indianorum several times in 1864, he did not describe the species until 1865, indicating that it was intermediate between Scalaria communis and S. turtonis. When Pater (1958: 187; plt. 20, figs. 23, 24) chose a syntypic specimen of Nitidiscala indianorum as lectotype and a second specimen as a paratype she was unaware that Stronc (1930: plt. 20, fig. 1) had already chosen a lectotype (USNM 15521) (length 26mm; width 9mm), which had probably been segregated from the other syntypic speci- mens. All remaining specimens (USNM 635564),other than the lectotype chosen by Strong, became paralectotypes. Nitidiscala sawinae (Dall, 1903) (Figures 39, 40, 42, 43, 44) Scala sawinae DALL, 1903: 175. KELSEY, 1907: 49. DALL & BARTSCH, 1910: 21. Boss, et al., 1968: 287. Epitonium sawinae: DALL, 1907: 481. PACKARD, 1918: 319; pit. 36, fig. 13. STRONG, 1930: 191, 194; plt. 20, figs. g, 10. KEEN, 1937: 35. WILLETT, 19374: 64. BURCH, 1945: 30. ABBOTT, 1974: 119; fig. 1237. DUSHANE, 1974: 39. Epitonium (Nitidoscala) [sic] sawinae: DALL, 1921: 115; plt. 6, fig. 12. Epitonium (Nitidiscala) sawinae: STRONG, 1945: 24, 27. SMITH & GORDON, 1948: 191. Scala berryi DALL, 1907: 127. BERRY, 1907: 42. Boss, et al., 1968: 46. Epitonium berryi: DALL, 1917: 483. KEEN, 1937: 35. STRONG, 1945: 27. BURCH, 1945: 31, 32. Epitonium (Nitidoscala) [sic] berryi: Datu, 1921: 116. I. S. OLDbROYD, 1927: 63. Epitonium (Nitidiscala) berryi: STRONG, 1945: 25. SMITH & GorDON, 1948: 190. Epitonium (Epitonium) berryt: ABBOTT, 1974: 123. Scala rectilaminata DALL, 1907: 127. BERRY, 1907: 42. Boss, et al., 1968: 274. Epitonium rectilaminatum DALL, 1917: 482 (lapsus for Scala rectilaminata DALL, 1907: 127). I. S. OLDRoYD, 1927: 62. KEEN, 1937: 35. STRONG, 1945: 27. BURCH, 1945: 31, 32. SMITH & GORDON, 1948: 191. KANAKOFF & EMERSON, 1959: 27. Epitonium (Epitonium) rectilaminatum: ABBOTT, 1974: 122. Epitonium (Crisposcala) acrostephanus Dati, 1908: 251. Burcu, 1945: 32. Boss, et al., 1968: 9. Epitonium acrostephanus: DALL, 1917: 478. KEEN, 1937: 35. VALENTINE & MEADE, 1961: 23. COWAN, 1964: 111. Epitonium (Nitidoscala) [sic] acrostephanus: DALt, 1921: 116. Epitonium (Nitidoscala) [sic] acrostephanum: T. S. OLpRoyp, 1925: 13. I. S. OLDRoyD, 1927: 65. Epitonium acrostephanum: KANAKOFF & EMERSON, 1959: 27. Epitonium (Nitidiscala) acrostephanum: GRanT & GALE, 1931: 858. WILLETT, 1937b: 401. Epitonium (Crisposcala) acrostephanum: SMITH & GORDON, 1948: 191. Epitonium (Epitonium) acrostephanus: ABBOTT, 1974: 123. Original Description of Nitidiscala sawinae (DALL, 1903): “Shell small, elongate, sub-acute, with ten or more whorls; nucleus of three smooth polished whorls; subsequent whorls smooth, with about 19 low, sharp, slightly reflected varices which entirely cross the whorl; at the shoulder these are slightly spinose; aperture rounded ovate, entire, with a small spine at the shoulder angle and a less conspicuous one at the inner base of the aperture; there is no trace of a basal cord or disk, and no spiral sculpture. Length, 10.5; diameter of aperture, 2.5; max. diameter of last whorl 4.0mm. A broken specimen with three more whorls seems to have measured 24mm in total length when perfect, and 8mm in diameter.” (DALL, 1903: 175). Additional Description: Shell medium to large; white; nuclear whorls 2 to 3, usually broken, sometimes with a small, round plug formed inside the last nuclear whorl; re- maining whorls 8 to 10, smooth, with occasional fine axial striations; suture deep; costae 14 to 21, reflected, with a sharp spine at the shoulder, slightly striate longitudinally, with a turn into the suture, crossing it and becoming con- nected with one of the costae of the preceding whorl, slightly flattened at the base; aperture ovate; peritreme thin, re- flected over the costae on which they rest; non-umbilicate ; operculum thin, horny, paucispiral, horn color. Length 3.5 to 24mm; width 1 to8mm. Type Material and Type Localities: Scala sawinae Dall: Holotype USNM 109309; Avalon, Cata- lina Island, California. Scala berryi Dall: Holotype USNM 107724; San Pedro Bay, California. Scala rectilaminata Dall: Holotype USNM 110430; Mon- terey Bay, California. Epitonium (Crisposcala) acrostephanus Dall: Holotype USNM 110638. Newport, California. Page 116 THE VELIGER Vol. 22; No. 2 Distribution, Ecology and Bathymetric Range: British Columbia, Canada (54°17’N, 132°37’ W), south to Mag- dalena Bay, Baja California Sur, Mexico (24°31’30”N, 112°o1'10” W). Occasionally taken by divers, but usually dredged in mud, sand and broken shell substrate in from 18 to 360m. Geologic Record: Pliocene to Recent. Upper Pliocene: Fernando Formation, downtown Los Angeles, at Fifth and Hope Streets, LACMIP loc. 19 [2], and at Sixth and Hope Streets, LACMIP loc. 466 [18]. San Diego Formation, Pacific Beach, LACMIP loc. 309 [1], Market Street, San Diego, LACMIP loc. 127 [1], southwesternmost San Diego County, LACMIP locs. 305 [109], 305-A [158], 305-C [18], 318 [4], and 319 [5]. Lower Pleistocene: Port Orford and Elk River Forma- tions, southeast of Cape Blanco and north of old mouth of Elk River, Oregon, LACMIP locs. 3947 [7] and 3960, Moonstone Beach, south of Trinidad, LACMIP loc. 3942 [13, 4f.], Santa Barbara Formation, east of Santa Barbara General Hospital, LACMIP locs. 183-A [15], and 183-B [22], Santa Barbara County Dump, LACMIP loc. 322 [4, 1 f.], bluff on Santa Bar- bara Yacht Harbor, LACMIP loc. 4965 [1], Rincon Hill Road, southeast of Carpinteria, LACMIP loc. 4890 [6]. Ondulando Highlands, Ventura, LACMIP loc. 441 [4]. Lomita Marl, in San Pedro, at old Hilltop Quarry, LACMIP loc. 64 [5], and below Host Place and Park Western Drive, LACMIP loc. 435 [130]. Timms Point Silt, Timms Point, San Pedro, LACMIP locs. 62 [9], and 130-7 [12]. Upper Pleistocene: Palos Verdes Sand, Lincoln Avenue, northeast of Playa del Rey (Los Angeles), LACMIP locs. 59 [105 | (WILLETT, 1937b:401; asE.(N.)sawinae and E. (N.) acrostephanum), and 4745 [2], Vermont Avenue and Sepulveda Boulevard, west of Carson, LACMIP loc. 147 [2], and Lomita Boulevard and Main Street, Wilmington (Los Angeles), LACMIP loc. 77 [27]. Terrace seven, Palos Verdes Hills, LACMIP loc. 1308 [2] (MarincovicH, 1976: 23; as E. sp.). Formation uncertain, in San Pedro, Pacific Avenue and Oliver Street, LACMIP loc. 131 [1], Se- condand Orizaba [now Beacon] Streets, LACMIP loc 142 [6], and Deadman Island, LACMIP loc. 2 [1]. Near Interstate 405 and Cherry Avenue, Long Beach, LACMIP loc. 424 [2]. East bluff above Upper New- port Bay, Newport Beach, LACMIP locs. 66-2 [26] (KANAKOFF & EMERSON, 1959: 27; as E. rectilamina- tum and E. acrostephanum), 66-1 [6], and 136 [4]. West bluff of “middle” Newport Bay, Newport Beach LACMIP locs. 68-A [1] (KANAKOFF & EMERSON, 1959: 27; as E. rectilaminatum) and 68-B [‘3”speci- mens missing] (KANAKOFF & EMERSON, 1959: 27; as E. acrostephanum). Recent Records: This species occurs off the Queen Char- lotte Islands, British Columbia, along the Oregon and southern California coasts, including the offshore is- lands of Santa Rosa, Santa Cruz, Catalina, San Mar- tin, Guadalupe, as far south as Magdalena Bay, Baja California Sur, Mexico, in from 18 to 360m. Off the northwest tip of Graham Island, Queen Char- lotte Islands, British Columbia, Canada, collected by F. Bernard (G. B. Reed, station 67-32), 179m, August 1967, one live but damaged specimen with operculum intact (LACM 67-30). La Perouse Bank, Barkley Sound, Vancouver Island, British Columbia, Canada, dredged by Cowan (1964: 111), one live specimen (no depository given), 72m. Monterey Bay, California, length 3.5mm; width 1mm, 22m, I specimen as Explanation of Figures 49 to 59 Figure 49: Opalia insculpta Carpenter, 1865. Holotype, PRI 7090 (formerly Cornell Univ. 4950); length, 14mm; width, 7 mm xX 4 Figure 50: Opalia (Dentiscala) crenimarginata Dall, 1917. Holo- type, USNM 111207; length, 17mm; width,7mm _ X 3.7 Figure 51: Opalia (Dentiscala) nesiotica Dall, 1917. Holotype, US NM 56900; length, 10.5 mm; width, 5 mm x6 Figure 52: Scalaria (Cirsotrema) funiculata Carpenter, 1857. Lec- totype of Keen (1968: 408; plt. 57, fig. 50a; text fig. 40) BM(NH); length, 21 mm X2 Figure 53: Opalia bullata Carpenter, 1865. Holotype, RM No. 76, Montreal, Canada; length, 8mm; width, 2.5 mm x7 Figure 54: Opalia tremperi Bartsch, 1927. Holotype, USNM 362- 454; length, 7.8mm; width, 2.5 mm X7 Figure 55: Rissoa infrequens C.B. Adams, 1852. Holotype, MCZ 186418; length, 6.5 mm; width, 2mm X 7.6 Figure 56: Opalia mazatlanica Dall, 1908. Holotype, USNM 168669; length, 12.5 mm; width, 4.5 mm x5 Figure 57: Epitonium (Nodiscala) ordenanum Lowe, 1932. Holo- type, ANSP 157988; length, 7mm; width, 2mm xX7 Figure 58: Scala (Cirostrema) [sic] montereyensis Dall, 1907. Holo- type, USNM 110431; length, 2.5mm; width,1.5mm X 4 Figure 59: Scala (Opalia) pluricostata Dall, 1917. Holotype, USN M 56054; length, 16mm; width, 5mm X 3.1 THE VELIGER, Vol. 22, No. 2 [DuSuaAneE] Figures 49 to 59 Ww Figure 49 Figure 52 Figure 50 Figure 51 Figure 55 Figure 54 Figure 53 Figure 56 Figure 57 Figure 58 Vigure 59 Vol. 22; No. 2 THE VELIGER Page 117 Scala rectilaminata (USNM 110430). Off Naples, Cal- ifornia, dredged by Pat Brophy from a kelp holdfast, 290m, one live specimen, November 1968 (Santa Bar- bara City College Collection). San Pedro, California, length 3.5mm; width 1.75mm, 360m, one specimen as Scala berryi (USNM 107724). The Hancock Expe- ditions 1931-1941 (Velero III) dredged it commonly from Anacapa Island, California south to the Baja California, Mexico outer coast islands of San Martin (station 1694-49), Guadalupe (station 1919-49), Ce- dros (station 1948-50), and from Magdalena Bay (24°31’30" N, 112°01'10" W). Discussion: Datt (1903: 175) described Nitidiscala saw- inae from off Catalina Island, California. The type has a medium sized shell of 10.5mm and so later workers neg- lected toassess the relationship to N.acrostephanus (DALt, 1908), which is normally 20 mm in length. When the type of N. sawinae is photographed and enlarged to a size com- parable with N. acroste phanus the synonymy is obvious (Fig- ures 43, 44). Both species have the same proportions, 14 to 21 costae, and a similar geographic and depth range. DALL (1908: 251), in his original description, gave the range of this species from Monterey, California to the Coronados Islands, Mexico, in from 29 to 61m, explaining that the species was common in the dredgings, but was usually smaller than the holotype of Nitidiscala acrostephanus dredged by H. N. Lowe off Newport, California. It is odd that STRONG (1922) listed N. acrostephanus, but neglected to discuss the species in his (1930) paper on Nitidiscala from the west coast of North America. The characteristics of Nitidiscala berryi and N. rectila- minata, both of Dall, are essentially the same as those for N. sawinae and the 2 are synonymized here (Figures 39, 40). Datu (1917: 483) likened N. berryi to Epitonium incon- spicuum (Sowerby, 1842), from the West Indies, but did not imply synonymy. In a general statement, Dat (1917: 481) suggested that Nitidiscala sawinae ranged into the Gulf of California, Mexico, but did not validate his suggestion with any spe- cific data. There seems to be a species group affinity among a number of nominal species in the genus Nitidiscala, namely: N. curvilineatum (Sowerby, 1844) from Central America, N.imbrex from Panama, N. willetti from Mazat- lan, Mexico, and N. sawinae from California. Unfortu- nately, the type of N. curvilineatum could not be located at the British Museum (Natural History) (teste A. M. Keen), so we have only the illustration in Sowersy (1844: plt. 33, fig. 59) by which to make an evaluation. All the species have many costae, thin, erect, somewhat flattened on the base with a fragile texture and swollen whorls. Perhaps all these taxa may prove to be synonymous with N. curvilineatum. Nitidiscala tincta (Carpenter, 1865) (Figures 41, 45, 46, 47, 48) Scalaria ? Indianorum var. CARPENTER, 1864a: 613 [Re printed, 1872: go]. Scalaria ? var. tincta: CARPENTER, 1864a: 660 [Reprinted, 1872: 146]. Scalaria (Ind. var.) tincta: CARPENTER, 1864a: 665 [Reprinted, 1872: 151]. Scalartia (? Indianorum, var.) tincta: CARPENTER, 1865: 31 [Reprinted, 1872: 244]. Cooper, 1870: 67. Scalaria tincta: Dax, 1878b: 12. Tryon, 1887: 70; plt. 14, fig. 49. CoopPER, 1888: 263. Epitonium (Nitidoscala) [sic] tinctum: Datt, 1917: 477. T. S. OLpROYD, 1925: 13. I. S. OLDRoYD, 1927: 58. Scala tincta: ARNOLD, 1903: 265; plt. 5, fig. 3. ARNOLD, 1906: 28, 36. DEBoury, 1919: 34. Epitonium tinctum: T. S. OLDROYD, 1925: 13. STRONG, 1930: 190. KEEP, 1935: 180; fig. 153. KEEN, 1937: 36. WILLETT, 1937b: 401. STRONG, 1941: 46. BurcH, 1945: 27, 28. STRONG, 1945: 26. Morris, 1952: 88; plt. 24, fig. 5. A. CLarK, 1957: plt. 2. THORSON, 1957: 55. PALMER, 1958: 188. VALENTINE, 1961: 360, 374. VALENTINE & MEADE, 1961: 10, 16. MCLEAN, 19609: 34; fig. 17.1. BERNARD, 1970: 81. HOCHBERG, 1971: 22, 23. C. R. SMITH, 1972: 47. ABBOTT, 1974: 119; fig. 1234. KENNEDY, 1975: 27. R. I. SMITH & CARLTON, 1975: 488. VEDDER & Moore, 1976: 1g0; plt. 2, figs. 4, 5. C. R. SMITH, 1977a: 331-340; figs. 1-4. C. R. SmitH, 1977b: 9. MCLEAN, 1978: 34; fig. 17.1. Epitonium (Nitidiscala) tinctum: STRONG, 1930: 193; plt. 20, figs. 3, 4, 5a-hb. GRANT & GALE, 1931: 859, 860. DURHAM, 1937: 488. WILLETT, 1937b: 401. A. G. SMITH & GORDON, 1948: 191. PALMER, 1958: 188. WILSON & KENNEDY, 1967: 252. Epitonium (Nitidiscala) tincta: STRONG, 1945: 24. Epitonium tinctium [sic]: KANAKOFF & EMERSON, 1959: 27. Scalaria subcoronata CARPENTER, 1864a: 613, 660 [Reprinted, 1872: 99, 146]. CARPENTER, 1866: 221. COOPER, 1870: 67. Dat, 1874: 297. Dati, 1878b: 29. Cooper, 1888: 263. DEBoury, 1919 (as Scala): 34. STRONG, 1930: 187, 192. PALMER, 1958: 188. Epitonium (Nitidoscala) [sic] subcoronatum: DALL, 1917: 478. I. S. OtpRoyp, 1924: 107. I. S. OLpRoyD, 1927: 58. Epitonium (Nitidoscala) [sic] subcoronata: DALL, 1921: 115. Epitonium subcoronatum: T. S. OLDROYD, 1925: 13. KEEN, 1937: 35. BURCH, 1945: 28. ABBOTT, 1974: 119 (as a syn- onym of E. tinctum). Epitonium continuatum T. S. OLDROYD, 1925: 13, 35; plt. 2, fig. 10. Epitonium (Nitidiscala) continuatum: GRANT & GALE, 1931: 858. Epitonium (Nitidiscala) eelense DURHAM, 1937: 488; plt. 56, fig. 7. Epitonium (Nitidoscala) [sic] tinctum var. bormanni STRONG, 1941: 47. BURCH, 1945: 28. WILSON & KENNEDY, 1967: 252. Epitonium bormanni: ABBOTT, 1974: 119, as a “form” of E. tinctum. Page 118 THE VELIGER Vol. 22; No. 2 Original Description of Nitidiscala tincta (CARPENTER, 1865): “S. ? Indianorum costis acutis, haud reflexis; an- fractibus postice fuscopurpureo tinctis. Hab. Cerros Island (Ayres) ; San Pedro (Cooper). The Lower-Californian shell may prove distinct. It is like S. regularis Cpr., but without the spiral sculpture.” (Car- PENTER, 1865: 31). Additional Description: Shell small to medium in size; nuclear whorls 3, rapidly increasing, opaque, solid looking; remaining whorls 4 to 8, sometimes showing a brown area at the sutures, rounded; costae 11 to 14, very slightly re- flected, and slightly shouldered on the whorls at times, reflected under the face of the aperture; suture deep; aper- ture oval, patulous anteriorly, operculum thin, horny, light in color. Length 4 to 15mm; width 1 to 5mm. Type Material and Type Localities: Scalaria (? Indianorum, var.) tincta Carpenter: Lectotype USNM 19510 by Palmer (1958: 189) (figured by Strong, 1930: plt. 20, fig. 3); San Pedro, California (collected by Cooper). Scalaria subcoronata Carpenter: Holotype USNM 14830b; Monterey, California. Epitonium continuatum T. S. OLDRoyD: Holotype USNM 352383 (type lost since 1952; fide Rosewater, 1975); Nob Hill Cut, San Pedro, California; Pleistocene (San Pedro Sand). Epitonium (Nitidiscala) eelense Durham: Holotype UCMP 30147; Eel River, Humboldt County, California; Pliocene. Epitonium tinctum var. bormanni Strong: Holotype LACM 1064; paratypes (4) SBMNH 08119; paratypes (8) SDNHM 454, 1521-1527; paratypes (13) DuShane Col- lection. Distribution, Ecology and Bathymetric Range: For- rester Island, Alaska (54°50’ N, 133°32’ W), south to Mag- dalena Bay, Baja California Sur, Mexico (24°18’N, 112° 12’ W). Nitidiscala tincta is a micropredator and occurs in- tertidally on the small colonial anemone Anthopleura ele- gantissima and less commonly on the large, solitary anem- one Anthopleura xanthogrammica. The most common habitats are in sand pockets and on open-faced boulders, in sand filled crevices, under kelp, and on rock ledges, from a zero tide down to 40m. It is capable of withstanding long periods of exposure between tides. During low tide N. tincta can be found clustered on the column of the sea anemone cup (the oral disk), but twice each day at high tide it feeds by snipping off the ends of the tentacles of the anemone. Dentition of the radula is in the form of long, bifid struc- tures, unlike any other epitoniid thus far reported. The largest teeth have a long, smooth blade without projections, while the smaller teeth almost always show projections in this area (Figures 70, 71). 20pm Figure 70 Nitidiscala subcoronata (Carpenter, 1864) (SBMNH 51501). The bifid structure of the radular teeth is unlike any other thus far reported, with little variation in length. The scale (20 um) gives an accurate comparison of teeth size and denti- tion sculpturing The radulae of Nitidiscala subcoronata and N. tincta were extracted by Dr. Eric Hochberg (SBMNH) from dried specimens by soaking in full strength Clorox bleach (sodium hypochlorite). Ribbons were mounted temporarily in glycerin and examined under oil immersion on a Zeiss compound microscope utilizing Nomarski differential in- terference illumination. Drawings were done with the aid of a drawing tube attached to the microscope. The radulae were per- manently mounted in Hoyers Mounting Medium. Figure 71 Nitidiscala tincta (Carpenter, 1865) (SBMNH 21450). Radular slides of Figure 70 and 71 are virtu- ally identical; the larger teeth having a long, smooth blade without projections, the smaller teeth almost always showing projections in this region. (Scale line = 20m) Vol. 22; No. 2 THE VELIGER Page 119 Geologic Record: Pliocene to Recent. Lower Pliocene: Towsley Formation, Sand Canyon, off of Santa Clara River Valley (Los Angeles County), LACMIP loc. 291 [4]. Upper Pliocene: Fernando Formation: Sixth and Hope Streets, downtown Los Angeles, LACMIP loc. 466 [106], near Interstate 405 and Cherry Avenue, Long Beach, LACMIP 423 [4] (KENNEDY, 1975: 27). Niguel Formation, Via La Mirada, San Juan Capistrano, LACMIP loc. 4923 [4]. San Diego Formation, Fifth Street, San Diego, LACMIP loc. 323 [5], southwest- ernmost San Diego County, LACMIP locs. 305 [629], 305-C [159], and 319 [42]. Lower Pleistocene: North of Trinidad at Omenoku Point, LACMIP loc. 3940 [3]. Santa Barbara Forma- tion, bluff on Santa Barbara Yacht Harbor, Santa Bar- bara, LACMIP loc. 4684 [48]. Rustic Canyon, Pacific Palisades, near Santa Monica, LACMIP loc. 2621 [23]. Baldwin Hills, Los Angeles area, LACMIP loc. 4668 [49], Lomita Marl, in San Pedro, at old Hilltop Quarry, LACMIP loc. 64 [2], and below Host Place and Park Western Drive, LACMIP loc. 435 [544]. Timms Point Silt, Timms Point, San Pedro, LACMIP loc. 62 [1]. Upper Pleistocene: Palos Verdes Sand, Lincoln Avenue, northeast of Playa del Rey (Los Angeles), LACMIP locs. 59 [448] (WILLETT, 1937b:401,and 4745 [5],near Vermont Avenue and Sepulveda Boulevard, west of Carson, LACMIP loc. 147 [21], Lomita Boulevard and Main Street, Wilmington (Los Angeles), LACMIP loc. 77 [9]. San Pedro Sand, Miraflores Avenue, San Pedro, LACMIP loc. 4565 [7]. Terrace twelve, Palos Verdes Hills, LACMIP loc. 1305 [1] (MarincovicH, 1976: 23; as E. sp.). Formation un- certain, Via Valmonte, Walteria (Torrance), LACMIP loc. 409 [2], and in San Pedro, Pacific Ave- nue and Oliver Street, LACMIP loc. 131 [1], San Pedro and Wilmington Road, LACMIP loc. 299 [5], and San Pedro waterfront area, LACMIP loc. 1 [6]. West of Interstate 405 and Cherry Avenue, Long Beach, LACMIP loc. 424 [26]. East bluff above Upper Newport Bay, Newport Beach, LACMIP locs. 66-2 [322] (Kanaxorr & EMERSON, 1959: 27;plus E. recti- laminatum), 66-1 [8], 66-9 [1], and 136 [133]. New- port Mesa, west side of “middle” Newport Bay, LACMIP loc. 68-A [specimen missing], and 68-B [1] (KANAKoFF & EMERSON, 1959: 27). Pacific Coast Highway, below Newport Heights, Newport Beach, LACMIP loc. 241 [2]. Calle Fortuna, Capistrano Beach, LACMIP locs. 58 [37] (WILLETT, 1937c [1938 a]: 107) and 137 [2]. Near San Clemente RR station, LACMIP loc. 452 [2]. Bay Point Formation, east side of Mission Bay, San Diego, LACMIP loc. 4562 [3]. South of San Juanico, Baja California Sur, LACMIP loc. 2719 [56]. Recent Records: Forrester Island, Alaska, collected by Willett (1914-1917), one live-taken specimen, Du- Shane Collection; off British Columbia (23-51° N), collected F. Bernard (1970: 81), intertidal to 20m; Cape Arago lighthouse, Coos County, Oregon (Velero III), among eel grass root masses, July 1942, one spec- imen (LACM-AHF 1491-42); Depoe Bay, Oregon, collected DuShane, intertidal, one specimen, August 1958, DuShane Collection; Fort Bragg, California, collected DuShane, intertidal in tide pools on anem- ones, 53 specimens live-taken, August 1965, DuShane Collection; Todos Santos Bay, Baja California Norte, Mexico, collected E. P. Chace, intertidally, 18 speci- mens live-taken, April 1937, DuShane Collection; Playa San Ramon, Baja California Norte, Mexico (105 miles south of Ensenada), collected DuShane, inter- tidally, 51 specimens, 1963 and 1967, DuShane Collec- tion; Laguna de San Jose, Baja California Norte, Mex- ico (Guerrero Negro area), collected DuShane, in tide pools with large anemones, 11 specimens with eggs, February, 1976, DuShane Collection; Magdalena Bay, Baja California Sur, Mexico, collected by Orcutt, March 1917 (USNM 217877). Discussion: CARPENTER (1864a: 613, 660, 665) referred to Nitidiscala tincta as a new species, but gave no descrip- tion of it until 1865 (p. 31). This species, living intertidally from 0 to 40m, with a continuous geographical range from Forrester Island, Alaska, collected by Wimtetr (1914- 1917), to Magdalena Bay, Baja California Sur, Mexico (USNM 217877), collected by Orcutt, is represented by hundreds of stations (LACM) along the California coast. Stronc (1941:46) believed that the specimens occurring north of Point Conception, Santa Barbara County, Cali- fornia were different enough to allow for a subspecific name, Nitidiscala tincta subcoronata (Carpenter). The lectotype of Nitidiscala tincta (PALMER, 1958: 189; USNM 19510), has 8’ whorls, 10 costae, and measures, length 10.5 mm; width 4.5mm (Figure 48, also figured by Srronc [1930: plt. 20, fig. 3]). The holotype of Nitidiscala subcoronata (USNM 14830b), has 11 nuclear whorls, plus 7 additional ones, 11 costae, and measures, length 13mm; width 7 mm (Figure 45, also figured by Strone [1930: plt. 20, fig. 4; incorrect USNM number)). Nitidiscala continu- ata (USNM 352383) is synonymous with N. tincta, length 8mm; width 3.6mm (Figure 47). Nitidiscala tincta var. bormanni is a small N. tincta (Figure 46), as is N. eelense. Radulae of N. tincta and N. subcoronata are virtually iden- Page 120 THE VELIGER Vol. 22; No. 2 tical. Specimens north of Point Conception, California have larger and heavier shells than those from the south. Extractions of radular teeth and drawings were made by Dr. Eric Hochberg (SBMNH). Shells and ribbon slides are at the Santa Barbara Museum of Natural History and are: Nitidiscala tincta (SBMNH 21450), Refugio Beach State Park, Santa Barbara County, California, collected by Gale Sphon, 1965; Nitidiscala subcoronata (SBMNH 51501), Fort Bragg, Mendocino County, California, collected by DuShane, 1965 (Figures 70, 71). Sthenorytis Conrad, 1862 [Type species (SD, De Boury, 1889): Scalaria expansa Con- rad, 1862] Whorls rapidly expanding, spire low; costae heavy; basal disk present. Sthenorytis stearnsit (Dall, 1892) (Figure 64) Scala (Sthenorytis) stearnsit DALL, 1892: 245; plt. 21, fig. 4. Boss, et al., 1968: 304. Scala stearnsi: STEARNS, 1898: 298. ARNOLD, 1906: 28, 100. Epitonium (Sthenorytis) stearnsii: Schuchert, Datu, et al., 1905: 583. DALL, 1908b: 317. GRANT & GALE, 1931: 855. DuRHAM, 1937: 482. DUSHANE, 1974: 46, 47. Original Description: “Shell large, stout, short conical, of about six whorls (the specimen having lost its apex, the exact number is doubtful) ; each whorl provided with about seven strong reflected varices, slightly angulated near the suture, where each varix is appressed against the interspace and corresponding varix of the antecedent whorl, forming a wave in the varical contour; varices very thick, sharp- edged, somewhat rugose from the prominence of the incre- mental lines, on the base overlapping one another and on the spire continuous over the whorls to the apex; surface smooth, except for lines of growth; a faint indication of a basal cingulum is perceptible in one or two of the inter- varical spaces; suture filled by the appressed varices; whorls very round, aperture nearly circular, except for the sutural wave. Lon. (of decollate shell 21% whorls) 26; max. lat. 25 mm” (DALL, 1892: 245). Additional Description: Shell large, heavy, thick; de- pressed-turbinate; whorls rapidly enlarging; costae 7, wide, reflected, triangular shaped, each costa flattening as it ap- proaches the suture, joining the costa immediately above, leaving a pit at the suture, broadly reflected on the base, fusing with the lip; suture deep; aperture round, oblique; lip heavy, reflected, continuous; operculum horny, black. Length, 27mm; width, 24mm. Type Material and Type Locality: Scala (Sthenorytis) stearnsii Dall: Holotype USNM 106904; Pacific Beach, San Diego, California. San Diego Forma- tion, Pliocene. Geologic Record: Pliocene (type locality). ° Discussion: In more than 80 years there have been no subsequent records of this species from southern California Pliocene deposits. Perhaps Stearns’ fossil is an ancestral Sthenorytis reaching as far north as San Diego during one of the warm-water phases of the Pliocene. The species is known from only one badly damaged specimen, collected by Stearns from the upper Pliocene San Diego Formation, in Pacific Beach, San Diego, California. However, enough remains to show its affinities to Panamic-Galapagan species of Sthenorytis (DUSHANE, 1966: 311; plt. 52, figs. 1-5. 1974: 45-47, figs. 113-117). Explanation of Figures 60 to 68 Figure 60: Ovpalia chacei Strong, 1937. Holotype, LACM 1045; length, 28.3 mm; width, 10.5 mm X 2.1 Figure 61: Opalia borealis Keep, 1881. Live-taken specimen, dredged at 36m by Willett, off Forrester Island, Alaska; length, 22 mm; width, 7mm (DuShane Collection) X 2.9 Figure 62: Opalia spongiosa Carpenter, 1866. Holotype, USNM 14830; length, 8.5 mm; width, 3 mm x8 Figure 63: Opalia retiporosa Carpenter, 1866. Holotype, USNM 11843; length, 7mm; width, 2.3 mm X7 Figure 64: Scala (Sthenorytis) stearnsii Dall, 1892. Holotype, US NM 106904; length, 27 mm; width, 24 mm xX 1.4 Figure 65: Opalia varicostata Stearns, 1875. Specimen collected by Henry Hemphill [CAS(GTC) 11860], length, 51 mm; width, 27mm Xi Figure 66: Opalia anomala Stearns, 1875. Specimen collected by Henry Hemphill [CAS(GTC) 11861], length, 37 mm; width, 14mm X 1.3 Figure 67: Opalia varicostata Stearns, 1875. Lectotype, USNM 214040; length, 55mm; width, 21 mm x1 Figure 68: Opalia varicostata Stearns, 1875. Five specimens (LAC MIP) showing gradation from ribbed form to smooth form. Specimens coated with magnesium chloride x1 Tue VELIGER, Vol. 22, No. 2 [DuSHANE] Figures 60 to 68 Figure 62 i Figure 6 eon e See Figure 66 Figure 67 Figure 68 Vol. 22; No. 2 THE VELIGER. Page 121 rss a eer Opalia Adams and Adams, 1853 [Type species: Scalaria australis Lamarck, 1822; (SD) De Boury, 1886] Shells white or light gray to brown in color, solid, imper- forate, with axial sculpture of strong ribs that may some- times be angulated, with or without basal ridge and with spiral sculpture of fine threads between each 2 of which is a row of small pits. Chalky outer layer (intritacalx) over entire shell easily abraded. Oval aperture set at an angle to axis of shell; lip thickened by last axial rib; operculum horny, thin, paucispiral. O palia borealis Keep, 1881 (Figures 60, 61) “Scalaria borealis Gould, 1852’: [207], of authors. CARPEN- TER, 1857: 212, 244. REEVE, 1937-1874: plt. 10, fig. 75. TrYON, 1887: 76; plt. 16, fig. 89. R. I. JOHNSON, 1964: 47. [Not of Lyell, 1841.] “Opalia borealis Gould” of authors: CARPENTER, 1864a: 532, 613, 628, 660 [Reprinted, 1872: 18, 19, 114, 146]. Car- PENTER, 1865: 31, 32 [Reprinted, 1872: 245]. Cooper, 1870: 67. KEEP, 1881: 28; plt. 6, fig. 5. CooPER, 1888: 255. KEEP, 1893: 49; fig. 30. ARNOLD, 1903: 266. ARNOLD, 1906: 36. Scala borealis: KEEP, 1904: 201; fig. 212. DEBOURY, 1919: 35. Scala (Opalia) borealis: BERRY, 1907: 42. BERRY, 1908: 38 [as Epitonium (O.)]. KEEp, 1911: 182; fig. 173 [as Epi- tonium]. Epitonium (Acirsa) borealis (“Gould”): KEEP, 1935 ed.: 178 (text only; figure, by error [fig. 52] of Acirsa borealis (Lyell). Scala Wroblewskyi MOrcH, 1875: 251 (replacement name for “S. borealis Gould” of authors). DEBoury, 1919: 35. Scalaria (Psychrosoma) gouldi TAPPARONE-CANEFRI, 1876: 152 (not S. gouldi Deshayes, 1861). STRONG, 1937: 4. Epitonium (Opalia) wroblewskii [sic]: Dati, 1917: 412. DALL, 1921: 113. I. S. OLDRoyD, 1924: 108. T. S.OLDRoyp, 1925: 13. I. S. OLDROYD, 1927: 51; plt. 31, fig. 5. BERNARD, 1970: 81. ABBOTT, 1974: 115; fig. 1195. Epitonium wroblewskyi: GRANT & GALE, 1931: 853 (as E. (Opalia)). KEEN, 1937, 36. WILLETT, 1937b: 401 (as E. (Opalia)). Opalia wroblewskyi: KEEN, 1937: 43. DURHAM, 1937: 501 (as O. (Opalia)). STRONG, 1937: 5; plt. 2, fig. 10 (as O. wro- blewskit [sic]). BERRY, 1948: 15-19. Morris, 1952: 86; plt. 24, fig. 1 (as O. wroblewskit [sic]). Opalia chacei STRONG, 1937: 5; plt. 2, fig. 9. KEEN, 1937: 43. Burcu, 1945: 17. A. G. SMITH & GORDON, 1948: 190. Cowan, 1964: 111. R. I. SMiTH & CARLTON, 1975: 489. Opalia wroblewskyi chacei: EMERSON, 1956b: 338. EMERSON & ADDICOTT, 1958: 9. ADDICOTT & EMERSON, 1959: 16, 21. KANAKOFF & EMERSON, 1959: 29. Opalia cliacei [sic]: ZINSMEISTER, 1970: 124. Original Description of Opalia borealis Kerr, 1881: “Figure 5, Pl. VI is about an inch long, white, strong, with fewer and blunter ribs than Scalaria. It also differs from it by not having a continuous peristome.” (KEEP, 1881: 28). Additional Description: Shell large, with calcareous outer coat (intritacalx), color off-white; nuclear whorls 1 i, usually eroded, with axial and spiral sculpture starting on the third whorl down; body whorls 8 to 11, attached at the suture, not rounded, but ascending rapidly, with an acumi- nate profile; interspaces wider than the ribs; varices usually 7, with here and there a heavier one, particularly on the last 2 whorls, often with a varix continuing beyond the heavy basal ridge onto the base, usually ascending in almost a straight line, one above the other, in older specimens be- coming obsolete on later whorls although the basal disk per- sists; suture distinct but not deep, with a remnant of the earlier basal disk as a small ridge just above the suture line; spiral sculpture of small punctations covering the entire sur- face of the chalky outer layer, sometimes obliterated on older specimens; lip patulous, peristome coalescing with the parietal wall; operculum brown, horny, with growth lines showing as small ridges on large specimens. Length 6.5 to 43mm; width 3 to 13mm. Type Material and Type Localities: Scalaria borealis Gould: Holotype missing, probably lost (fide R. I. Johnson, 1964: 47); Puget Sound, Washington. Scalaria wroblewskyi Mérch: Holotype would be that of Scalaria borealis Gould. Opalia borealis Keep: No definite locality given; figured specimen of Keep, present repository undetermined. Opalia chacei Strong: Holotype LACM, Section of Mala- cology 1045; paratypes (6) DuShane Collection; Crescent City, California. Distribution, Ecology and Bathymetric Range: For- rester Island, Alaska (54°50’N, 133°32’ W), with a con- tinuous distribution south to just north of Todos Santos Island, Baja California Norte, Mexico (31°53'20” N, 116° 48'15’ W). Intertidal to 180m, usually associated with Antho pleura elegantissima, among rocks and in tide pools. Geologic Record: Miocene (?), Pliocene to Recent. Upper Pliocene: Fernando Formation, Fifth and Hope Streets, downtown Los Angeles, LACMIP loc. 27 [2], east bluff above Upper Newport Bay, Newport Beach, LACMIP loc. 471 [1] (ZiNSMEISTER, 1970 [1971]: 124; as O. cliacei). San Diego Formation, Market Street, San Diego, LACMIP loc. 127 [2], southwest- ernmost San Diego County, LACMIP locs. 305 [29, 14 others possibly O. varicostata], and 305-A [1 f]. Lower Pleistocene: Santa Barbara Formation, east of Santa Barbara General Hospital, LACMIP loc. 183-A Page 122 [1]. Lomita Marl, in San Pedro, at old Hilltop Quarry, LACMIP loc. 64 [1], and below Host Place and Park Western Drive, LACMIP loc. 435 [34]. Timms Point Silt, Timms Point, San Pedro, LACMIP loc. 130-7 [1]. Discussion: In a Danish paper published early in 1875 (p. 251), MOrcx proposed a replacement name Opalia wroblewskyi for “Scalaria borealis Gould (not Beck)” 2.e., presumably, Goutp, 1852: 207. Whether Morch regarded Gould’s usage as a misidentification or a homonym is im- material, for the name was preoccupied by S. borealis Lyell, 1841. Mérch’s name has priority over Scalaria (Psy- chrosoma) gouldi proposed by TAPPARONE-CANEFRI (1876: 152), which was in turn preoccupied by S. gouldi Deshayes, 1861. According to Morch, Gould thought that Scalaria australis Lamarck, 1822, was so similar a species that he withdrew his name. Scalaria borealis Gould, 1852 was, in any case, a nomen nudum. The replacement name O palia wroblewskyi proposed by Morch, 1875 was not accom- panied by any description or figure; it was based only on the nomen nudum of Gould. The first author to give an adequate indication is KEEp, 1881 (p. 178, fig. 152) who used the name Opalia borealis for this species [not to be confused with Acirsa borealis (Lyell, 1841)]. Opalia bore- alis, as Gould thought, may be similar to S. australis, but because of the geographical difference it seems wise to re- tain Keep’s species as distinct, one with a wide distribution in the eastern Pacific, from Forrester Island, Alaska, to near Todos Santos Island, Baja California Norte, Mexico. No essential differences can be observed between O palia borealis and O. chacet. Specimens inhabiting colder waters seem to be longer and slimmer with a more acuminate spire, while those from warmer waters tend to be chunkier, par- ticularly near the basal whorls, ribs slightly wider and rounder and occasionally with one continuous over the low disk, although the relationship of ribs to interspaces is the same. Specimens from off northern California intergrade within the same population. Berry (1948: 16), considered Opalia borealis Keep and O. montereyensis Dall to be closely related. Berry’s view seems to be logical even though O. borealis has a more acu- minate outline, less rounded ribs ascending the spire with more regularity and symmetry, sometimes a heavier rib being carried onto and beyond the basal disk; spaces be- tween ribs twice the width of the ribs. O palia borealis often clusters at the base of the oral disk of the sea anemone Anthopleura elegantissima (Brandt), everting its proboscis into the soft base and sucking the body fluids. THE VELIGER Vol. 22; No. 2 Opalia funiculata (Carpenter, 1857) (Figures 49, 50, 51, 52) Scalaria (Cirsotrema) funiculata CARPENTER, 18574: 260, 284, 336. CARPENTER, 1857b: 447 [Reprinted, 1967: 447]. HERTLEIN & STRONG, 1951: 89. PALMER, 1951: 62. BRANN, 1966: 17. KEEN, 1968: 408; plt. 57, figs. 50a, b; text fig. 40 (= Opalia diadema funiculata). Opalia funiculata: CARPENTER, 1864c: 547, 551 (as Scalaria sp.) [Reprinted, 1872: 33, 37]. CARPENTER, 1865: 31 [Reprinted, 1872: 244]. McLEAN, 1969: 34; fig. 2. ABBOTT, 1974: 115; fig. 1198. ABBOTT, 1974: 116; fig. 1202 [Opalia (Cylindriscala) watsoni (DeBoury, 1911) as syn- onym of O. funiculata]. McLEAN, 1978: 34; fig. 17.2. . Not Scalaria funiculata Watson, 1883: 609; ? = Cylindriscala watsont (DEBourY, 1911). [Deep-water, Brazil.] Scala funiculata: ARNOLD, 1903: 267. DEBouRY, 1919: 34, 36. Epitonium (Cirsotrema) funiculatum: KEEN, 1958: 272. Opalia diadema funiculata: KEEN, 1968: 408; plt. 57, figs. 50a-b [fig. 50a: lectotype of Opalia funiculata]. Opalia (Dentiscala) funiculata: KEEN, 1971: 440; fig. 680. DUSHANE, 1974: 61-63; figs. 122 to 125 and fig. 162. Opalia (? crenatoides, var.) insculpta CARPENTER 1864c: 539, 619, 660 [reprinted, 1872: 25, 105, 146]. CARPENTER, 1866: 275,277 [Reprinted, 1872: 322, 324; also reprinted in DALL, 1909: 189]. CooPER, 1888: 255. ARNOLD, 1903: 267. PALMER, 1958: 19, 20. Opalia insculpta: CARPENTER, 1864-65: 10 [Reprinted, 1872: 244]. VAN WINKLE (PALMER), 1921: 4; plt. 1, figs. 10, 11. VALENTINE & MEADE, 1961: 10. ABBOTT, 1974: 115. Opalia (Dentiscala) insculpta: DALL, 1917: 473. STRONG, 1945: 18. PALMER, 1958: 28, 50, 191; plt. 22, figs. 4, 5. Epitonium (Dentiscala) insculpta: BAKER, HANNA & STRONG, 1930: 48. GRANT & GALE, 1931: 855 (as E. (D.) insculp- tum). WILLETT, 1938: 10 (as E. insculptum). Dentiscala crenimarginata DALL, 1917: 473. DALL, 1921: 114 (as E. (Dentiscala)). KEEN, 1958: 278; fig. 157. PALMER, 1958: 191. MCLEAN, 1961: 464. Boss, et al., 1968: 94. Scala crenimarginata: DEBouRY, 1919: 36. Epitonium (Dentiscala) crenimarginatum: 1. S. OLDRoYD, 1927: 53. BAKER, HANNA & STRONG, 1930: 47; plt. 2, fig. 6. KEEP, 1935: 180. A. G. SMITH & GORDON, 1948: 190. Epitonium crenimarginatum: KEEN, 1937: 35. WILLETT, 1938: 10. STRONG, 1945; 18. Opalia crenimarginata: THORSON, 1957: 55. ABBOTT, 1974: 115. Dentiscala nesiotica DALL, 1917: 473 [not Epitonium nestott- cum Dall and Ochsner, 1928]. DALL, 1921: 114 (as E. (D.)). STRONG, 1945: 18. Boss, et al., 1968: 219. Scala nesiotica: DEBouRY, 1919: 37. Epitonium (Dentiscala) nesioticum: I. S. OLDROYD, 1927: 53. Epitonium nesioticum: KEEN, 1937: 35- Opalia nesiotica: ABBOTT, 1974: 115. Vol. 22; No. 2 Original Description of Opalia funiculata (Carpenter): “C. t. subelongata, graciliore, alba; marginibus spirae rec- tis; anfr. ix. prope suturam valde impressum acute angu- latis; costibus variantibus (xv.-xx.) haud acutis, angustiori- bus, ad angulam coronatis, supra basin vix continuis; anfr. ult. costis plerumque obsoletis, varicibus paucis validad; costa rotundata, suturam continuante, circa basin tumente, a costis radiantibus nodosa; costa spirali altera labio adjici- ente; tota superficie minutissime decussata; labro a costis sinuato. “This shell agrees with C. diadema in almost every par- ticular down to the minute decussation of the surface. The remarkable pupiform growth of the Peruvian shells how- ever, contrasted with the very regular spire (with one more whirl in proportion) of the Panama and Mazatlan speci- mens, is thought by Mr. Cuming sufficient to separate the species. The Mazatlan shells are not so large as those in Mr. Cuming’s collection, which measure long. ’7, long. spir.’47, oO 953 lat. 34, div. 37°.” (CARPENTER, 1857b: 447). Additional Description: Shell medium in size, off-white in color; nuclear whorls 3 to 4, usually eroded into a blunt apex; remaining whorls 5 to 7, enlarging rapidly, somewhat rounded, dipping abruptly into a deep suture, ribs 12-16, rounded, following contour of whorl, with an occasional large varix continuing onto the base, strongly developed on top of whorl, appressed to antecedent whorl giving appear- ance of nodes; ribs almost obsolete on later whorls; inter- spaces as wide as ribs; punctations fine, between equally fine spiral threads over entire shell, continuing onto base, where they are stronger just below a wide, elevated basal disk, returning to normal as they approach elliptical aper- ture; lip thick, entire, outer surface concentrically striated, inner lip surface smooth; operculum horny, brown, pauci- spiral, with about 7 to 9 ridges emanating from nucleus. Length 5 to 17mm; width 3 to 8mm. Type Material and Type Localities: Scalaria (Cirsotrema) funiculata Carpenter: Lectotype by Keen, 1968 (Panama); paralectotypes (2) (Panama; hy- polectotype by Keen, 1968 (Mazatlan); all at BM(NH). Opalia (? crenatoides, var.) insculpta Carpenter: Holotype PRI 7090 [formerly Cornell University 4950]; Santa Bar- bara, California. Pleistocene, Santa Barbara Formation. Dentiscala crenimarginata Dall: Holotype USNM 111207; La Paz, Baja California Sur, Mexico. Dentiscala nesiotica Dall: Holotype USNM 56900; Cata- lina Island, California. Distribution, Ecology and Bathymetric Range: From Refugio Beach State Park, Santa Barbara County, Cali- fornia (34°26’ N, 120°4’ W), with a continuous distribution to the Galapagos Islands, Ecuador, and Tumbes Province, THE VELIGER Page 123 Pert (3°54’S, 80°53’ W). Occurs intertidally down to 30 m; ectoparasitic with sea anemones. Geologic Record: Pleistocene to Recent. Upper Pleistocene: Palos Verdes Sand, Lincoln Avenue, northeast of Playa del Rey (Los Angeles), LACMIP loc. 59 [1]. East bluff above Upper Newport Bay, New- port Beach, LACMIP locs. 66-2 [7] (KANAKOFF & EMERSON, 1959: 29; O. insculpia), and 136 [4]. Recent Records: O palia funiculata is a common species from California to Peru,so only the most northern and southern points are reported herein: Refugio Beach State Park, Santa Barbara County, California (34°26’ N, 120°4’ W), col- lected by Gale Sphon intertidally, four specimens, Feb- ruary 1965 (DuShane Collection). El Rubio and Punta Moro, Tumbes Province, Peru (3°54’S, 80°53’ W), intertidally by McLean, Shasky, and Pena, April 1972 (LACM 72-85). Discussion: throughout the years, partly because early workers did not realize the wide geographic range of the species. CARPEN- TER (1864a: 539) originally considered O palia inscul pta, a Pleistocene fossil from Santa Barbara, California, a variety of Opalia crenatoides Carpenter. Later he thought O. in- sculpta to be a southern form, but contradicted himself (p.660) when he stated the species is found only as a fossil. He (1866: 275) validated O. insculpta by describing it from the “Pleistocene” [= lower Pliocene], explaining that “Opalia crenatoides is a recent species with fewer varices than insculptum and appears to be quite distinct, though no doubt related.” Dati (1917: 473) described O palia crenimarginata from La Paz, Baja California Sur, Mexico (length 16mm). Some later workers (see WILLETT, 1938: 10) considered it “iden- tical with Carpenter’s earlier znsculptum and thought the latter name should be used for our recent California species.” Datu (1917: 473) also described Opalia nesiotica, dredged in 29 m off Catalina Island, California (length 10.5 mm). Dall separated O. nestotica from other Opalia by the sharp spiral sculpture that covered the surface. However, this sculpture is only on the intritacalx, easily abraded and worn off by wave action and rubbing against the substrate. Baker, HANNA & STRONG (1930: 47) and A. G. SmirH & GorDoN (1948: 190) thought O palia insculpta Carpenter was the fossil form of Opalia crenimarginata Dall. The ex- istence of O. insculpta at Monterey, California was doubted by Smit & Gorpon (op. cit.). PALMER (1958: 191) re- ported, “The exterior of O. insculpta has been worn away Opalia funiculata has had many names Page 124 THE VELIGER Vol. 22; No. 2 except in the sutural region, [so] the exact character of the longitudinal ribs cannot be described. However, prominent impressions of each reveal 14 ribs on the last whorl.” She further questioned Carpenter’s reasons for stating there was no spiral sculpture. Comparisons of the types of these 4 taxa show them to be conspecific (Figure 49, 50, 51, 52). Some of the types are so badly worn that their true relationship to one another was not previously suspected. CARPENTER (1864: 551) stated that the Scalaria sp. no. c. reported by C. B. Apams (1852) from Panama is O pala funiculata. KeeEn (1968: 408) chose the largest (length 21 mm) of the syntypic specimens of O palia funiculata, from Panama, as lectotype. A specimen with immature lip from Mazatlan (length 16mm), was selected as hypolectotype. The radula of Opalia funiculata has blunt teeth, short and with a rounded cusp at the center of each tooth in addi- tion to the terminal ones. The broad outline and blunt cusps suggest a carnivorous mode of feeding. Opalia funiculata is known to feed on the sea anemone Anthopleura elegan- tissima. The specimen from which the radula was obtained (collected at Palos Verdes Peninsula, California by Du- Shane), and the radular slide are at the SDMNH. Opalia infrequens (C. B. Adams, 1852) (Figures 53, 54; 55,59, 57) Rissoa infrequens C. B. ApAMs, 1852: 403 [reprinted 1852: 179]. TURNER, 1956: 57; plt. 12, fig. 2. DUSHANE, 1974: 67; figs. 134, 135, 143, 145, 147. Opalia (Nodiscala) infrequens: DUSHANE, 1974: 67; figs. 134, Epitonium (Pliciscala) infrequens: BARTSCH, 1915: 34. Rissoina infrequens: KEEN, 1971: 375, 903. Opalia bullata CARPENTER, 1864c [not Scalaria bullata Sow- erby, 1844]: 537, 660 [reprinted, 1872: 23, 146]. CARPEN- TER, 1865: 397 [reprinted, 1872: 287]. PALMER, 1945: 100. Scalaria (Psychrosoma) crosseana 'TAPPARONE-CANEFRI, 1876: 154 [replacement name for Opalia bullata Carpenter, 1865]. TRYON, 1887: 84. Opalia (Nodiscala) bullata: PALMER, 1958: 20, 23, 50, 192; pit. 22, fig. 6. KEEN, 1971: 440; fig. 681. ABBOTT, 1974: 117. Opalia (Dentiscala) mazatlanica DALL, 1908: 250. DALL, 1917: 474. STRONG, 1945: 19, 21 (O. (Nodiscala)). KEEN, 1958: 278 (as O. (Nodiscala)). Boss et al., 1968: 201. Epitonium (Nodiscala) mazatlanica. BAKER, HANNA & STRONG, 1930: 44. Scala (Nodiscala) mazaltanica [sic]: DEBouRY, 1919: 37. Opalia tremperi BARTSCH, 1927: 3; plt. 1, fig. 8. I. S. OLDRoyD, 1927: 279. STRONG, 1930: 42. STRONG, 1937a: 7. KEEN, 1946: 8. Epitonium (Opalia) trempert: GRANT & GALE, 1931: 854. Epitonium tremperi: KEEN, 1937: 36. Epitonium (Nodiscala) tremperi STRONG, 1945: 19. BURCH, 1945: 19. Epitonium (Nodiscala) ordenanum LoweE, 1932: 114; plt. 9, figs. 3, 3a. Opalia (Nodiscala) ordenana: STRONG, 1945: 21. KEEN, 1958: 278; fig. 164. Original Description of Opalia infrequens C. B. Adams: “Shell very long, ovate conic; white; with, on each whorl, sixteen very obtuse slightly elevated indistinct ribs, which are separated only by striae, and a few microscopic revolv- ing striae; apex subacute; spire with the outlines nearly rectilinear; whorls seven, contracted above, otherwise con- vex or subangular with a moderately impressed suture; last whorl long, subangular; aperture oblique, subovate; scarcely effuse ; labrum much excurved, well thickened; um- bilicus wanting. Length .24 inch; breadth .075 inch.” (C. B. ADAMS, 1852: 403). Additional Description: Shell small, slim, conic, brown, varying in shading, dead shells white; with soft, calcareous outer coat; nuclear whorls 2, glassy, brown; body whorls 8, evenly punctate between spiral sculpture of small raised threads; suture well defined; ribs 15 to 20, rounded, coales- cing at the suture, overlapping at the edge of the preceding whorls to form cusps, with an occasional heavier rib; aper- ture oval, large; lip strongly expanded and thickened, com- plete, with the same sculpture as on the whorls; outer lip rounded, inner lip set at an angle against the body whorl; operculum dark, oval, horny, paucispiral, with the nucleus apical. Length 7 to 12mm; width 2 to 4mm. Type Material and Type Localities: Rissoa infrequens C. B. Adams: Holotype MCZ Harvard 186418; Panama. Opalia bullata Carpenter: Holotype RM 76, Montreal, Canada; Santa Barbara, California; Pleistocene (Santa Barbara Formation). Opalia (Dentiscala) mazatlanica Dall: Holotype USNM 168669; Mazatlan, Mexico. Opalia tremperi Bartsch: Holotype USNM 362454: San Clemente Island, California. Epitonium (Nodiscala) ordenanum Lowe: Holotype ANSP 157988; San Juan del Sur, Nicaragua. Distribution, Ecology and Bathymetric Range: From Santa Cruz Island, California (34°03’ N, 119°28’ W), along the southern California coast to Mazatlan, Mexico, south to Panama. Occurs intertidally to 36m depths. Geologic Record: Pleistocene, Recent. Pleistocene: Santa Barbara Formation, Santa Barbara, California (type locality) [1]. Recent Records: Santa Cruz Island, California (34°03’ N, 119°28’ W), dredged inshallow water (teste KEEN, 1946: 8), SU specimen (now at CAS). South end of Vol. 22; No. 2 THE VELIGER Page 125 Pulmo Bay, Baja California Sur, Mexico, coarse sand at 20 feet, collected McLean and Oringer, April 1966, one specimen (LACM 66-20). Guatulco Bay, Oaxaca, Mexico, collected Willett, 72-126m, March 1938, one specimen (LACM A.375). San Juan del Sur, Nica- ragua, collected by H. N. Lowe (1931), 36m, one spec- imen (as Epitontum ordenanum) holotype ANSP 157988. Bahia Herradura, Puntarenas Province, Costa Rica (9°37'58’ N, 84°40’30” W), collected McLean, 70 feet, March 1972, one specimen (LACM 72-53). Panama, collected C. B. Adams, one specimen, holo- type of Opalia infrequens, MCZ 186418. Discussion: Although C. B. Adams placed his species in the Rissoidae, which later workers followed, BartTscu (1915: 34) recognized that it belonged in the Epitoniidae and reallocated it in the subgenus Pliciscala without com- menting on his decision. The type of Pliciscala DeBoury, 1887 (Eocene of Europe) has a basal disk and strong axial sculpture that bears little resemblance to the species named by C. B. Adams. One Pleistocene specimen collected at Santa Barbara by Col. Jewett was first referred to as Opalia bullata by Car- PENTER (1864: 537, 660), although he did not describe it until 1865 (p. 397). Palmer erroneously thought O. bullata, O. spongiosa and O. retiporosa were conspecific. The latter 2 species, which are probably inseparable, differ from O. bullata by having stronger shouldered whorls, ribs that coalesce near the aperture into the basal disk near the aper- ture, and a less oval aperture. TAPPARONE-CANEFRI (1876: 154), believing incorrectly that O. bullata was a homonym of Scalaria bullata Sowerby, 1844, renamed Carpenter’s species Scalaria (Psychrosoma) crosseana. TRYON (1887: 84) correctly reinstated O. bullata. The type specimen of O. mazatlanica Dat (1908: 250) is the largest of the con- specific types; length 12 mm; width 4mm. No specific diag- nostic characters can be found which would differentiate it from O palia infrequens, length 6.5mm; width 2mm. Opa- lia tremperi BARTSCH (1927: 3) is here considered to be a synonym of O. infrequens. The type is 7.8mm long and 2.5 mm wide. Epitonium (Nodiscala) ordenanum Lowe (1932: 114), has a brown shell with 15 slender ribs; length 7mm; width 2 mm and is here placed in synonymy. Opalia montereyensis (Dall, 1907) (Figures 58, 59) Scala (Cirostrema) [sic] montereyensis DALL, 1907: 128. BERRY, 1907: 42. [Not preoccupied by‘‘Scalaria hellenica montereyensis DeBoury, 1889” of Palmer, 1958: 190-191, a non-existent name based on a lapsus of DeBoury.] Epitonium (Cirsotrema ?) montereyensis: DALL, 1917: 477 [not p. 481]. DALL, 1921: 115. I. S. OLDRoyD, 1927: 355. GRANT & GALE, 1931: 190. Opalia montereyensis: STRONG, 1937: 6; plt. 2, fig. 12. KEEN, 1937: 43. BURCH, 1945: 17. BERRY, 1948: 15-19; figs. 1-5. VALENTINE & MEADE, 1961: 10-17. MCLEAN, 1969: 34; fig. 17.3. BERNARD, 1970: 81. R. I. SMITH & CARLTON, 1975: 488. MARINCOVICH, 1976: 6, 12, 23. MCLEAN, 1978: 34; fig. 17.3. Opalia (Opalia) montereyensis: ABBOTT, 1974: 117. Scala (Opalia) pluricostata Carpenter MS label: BERRY, 1907: 42 [not Scala pluricostata DeBoury, 1913]. PALMER, 1958: 190. Opalia pluricostata DALL, 1917: 473 [fide Boss, et al., 1968: 259]. Scalaria pleurocostata [sic] ‘Carpenter’: STRONG, 1937: 36. Scala evicta DEBoury, 1919: 36 [new name for O. pluricostata Dall]. Epitonium (Opalia) evictum: DALL, 1921: 114. I. S. OLDRoyp, 1927: 350. BURCH, 1945: 17. Opalia evicta: KEEN, 1937: 43. STRONG, 1937: 6; plt. 2, fig. 11. BuRCH, 1945: 17. BERRY, 1948: 15. A. G. SMITH & GORDON, 1948: 190. PALMER, 1958: 190; plt. 20, fig. 22 [syn. O. plu- ricostata Dall]. ABBOTT, 1974: 116. Original Description of Opalia montereyensis Dall: “Shell small (probably not full grown), the nucleus lost but with five subsequent rapidly increasing whorls; shell sub- stance in two layers, the inner translucent white, solid, the outer opaque white, frothy, porous, with numerous punc- ticulations arranged in harmony with the incremental lines; varices low, solid, with a spongy surface, nine in number; basal disk conspicuous, slightly concave; bordered by a con- spicuous cord; aperture gibbous, patulous near the imper- forate axis. Long. 2.5, diam. 1.5mm.” “Dredged in 25 fms, mud, off Del Monte, in Monterey Bay, Cala., by S. S. Berry. U.S.N.M. 110431.” “This shell represented by two specimens, of which one isin Mr. Berry’s collection, is doubtless immature, but there isno other species known north of Cape St. Lucas belonging to this special group, and, though the characters require rather high magnification to see them clearly, it cannot be confounded with any other Californian species.” (Datu, 1907: 128). Additional Description: Shell small, with calcareous outer coat (intritacalx), color off-white; 11/2 nuclear whorls, usually eroded; body whorls 6 to 9, 7 being the usual num- ber, rounded, moderately deep depressions at the junction with the suture, axial ribs 8 to 10, rounded, sometimes dis- junct as they ascend the spire; basal disk heavy, joined by the extension of the axial ribs which rarely extend beyond the basal disk; spiral punctations cover the entire shell, about 35 on each whorl, evenly spaced; area below the basal disk concave, making the disk conspicuous; lip quadrate in Page 126 THE VELIGER Vol. 22; No. 2 outline, aperture round, lip extended anteriorly; operculum horny, brown, paucispiral. Length 2.5 to 16mm; width 1.5 to5 mm. Type Material and Type Localities: Scala montereyensis Dall: Holotype USNM 110431; para- type (1) S. S. Berry Collection; off Del Monte, Monterey Bay, California. Opalia pluricostata Dall: Holotype USNM 56054; Neah Bay, Washington. Distribution, Ecology and Bathymetric Range: From Forrester Island, Alaska (54°50’ N, 133°32’ W), south to off Vancouver Island, the coasts of Oregon, Washington and California to Cedros Island, Baja California Norte, Mexico (28°20'25” N, 115°11'20” W). This species occurs inter- tidally among rocks or in tide pools, and may be dredged down to 90m on rock or among kelp beds. Geologic Record: Pliocene to Recent. Upper Pliocene: Fernando Formation, Sixth and Hope Streets, downtown Los Angeles, LACMIP loc. 466 [4], east bluff above Upper Newport Bay, Newport Beach, LACMIP 4956 [1], San Diego Formation, Market Street, San Diego, LACMIP loc. 127 [32], and in southwesternmost corner of San Diego County, LAC- MIP locs. 305 [430], 305-A [8], 305-C [23], and 318 [3]. Lower Pleistocene: Santa Barbara Formation, Santa Barbara General Hospital, LACMIP locs. 183-A [1], and 183-B [cf. 1 f.]. Upper Pleistocene: Point Arena, LACMIP loc. 4816 [1 f.]. North side of Point Ano Nuevo, LACMIP loc. 5019 [2]. Terrace Seven, Palos Verdes Hills, LACMIP loc. 1307 [1] (Marincovich, 1976: 23). Palos Verdes Sand, Vermont Avenue and Sepulveda Boulevard, west of Carson, LACMIP loc. 147 [1]. East bluff above Upper Newport Bay, Newport Beach, LACMIP loc. 66-2 [1 f.] (KANAKOFF & EMERSON, 1959: 29; as O. wroblewskyi chacei). Calle Fortuna, Capistrano Beach, LACMIP loc. 58 [1 f.] (WILLETT, 1937[1938a]: 107; as O. evicta). Recent Records: Off Forrester Island, Alaska, collected by Willett (1914-1917) 90m (USNM 216347). En- trance to Bull Harbor, Hope Island, north end of Van- couver Island, British Columbia, collected by McLean, 15-40 feet, May 1963 (LACM 63-28). Off Cedros Is- land, Baja California Norte, Mexico, collected by Mc- Lean and LaFollette (Searcher 231) 15-40 feet, Octo- ber 1971 (LACM 71-151). Discussion: D£EBoury (1919: 39) stated that the specific name, ““montereyensis” was preoccupied, that Dall himself had used it in 1889 as a subspecies of Scala hellenica Forbes (1845: 189). DeBoury apparently misread Datw’s (1889: 320-322) S. hellenica var. moerchiana, an Atlantic species, as “montereyensis.” There is no usage of “montereyensis” by either author in 1889. Scala (Opalia) pluricostata ‘Carpenter,’ Dati, 1917 (p. 473), based on a Carpenter MS label name, is synony- mous with Opalia montereyensis. Opalia evicta DEBoURY (1919: 36, 40) is a needless new name for O. monterey- ensis. The syntypes of Opalia pluricostata have much larger shells than Opalia montereyensis (length 16mm; width 5mm) but have the same number of rounded ribs (8-10), punctate surface, and a conspicuous basal disk. For a more complete discussion see STRONG (1937: 6), BERRY (1948: 16), and Patmer (1958: 190). Opalia montereyensis Dall, 1907 should not be confused with Epitonium montereyense Dall, 1917, the latter now in the genus Nitidiscala. Berry (1948: 16) discussed the relationship between Opalia borealis Keep and O. montereyensis, being of the opinion that the 2 are extremely close allies and that there may exist a geographical gradient from one to the other, an opinion with which I concur. Opalia montereyensis has a smaller shell, broader in its proportions, with more com- pact coiling, a chunkier outline, more deeply and sharply cut relief and a stronger basal disk over which the ribs do not extend. O palia borealis has a more acuminate outline, less rounded ribs ascending the spire with more regularity and symmetry, with sometimes a heavier rib carried onto and beyond the basal disk, and with spaces between the ribs twice the width of the ribs. Opalia spongiosa Carpenter, 1866 (Figures 62, 63) Opalia spongiosa CARPENTER, 1864a: 613, 660 [Reprinted, 1872: 99, 146]. CARPENTER, 1865: 31 [Reprinted, 1872: 244]. CARPENTER, 1866: 222. Cooper, 1870: 67. ABBOTT, 1974: 116. Epitonium (Nodiscala) spongiosum: DALL, 1917: 474. DALL, 1921: 114 (as E. (N.) spongiosa). Epitonium spongiosum: BAKER, HANNA & STRONG, 1930: 44. KEEN, 1937: 35. Opalia (Nodiscala) spongiosa: STRONG, 1945: 19, 21. PALMER, 1958: 192, 193. KEEN, 1971: 442; fig. 686. DUSHANE, 1974: 68-71; figs. 138, 139. Opalia retiporosa CARPENTER, 1864a: 613, 660 [Reprinted, 1872: 244]. CARPENTER, 1865: 31 [Reprinted, 1872: 244]. CARPENTER, 1866: 222. DeBoury, 1919: 34 (as Scala). ABBOTT, 1974: 116; fig. 1201. DUSHANE, 1974: fig. 138. Epitonium (Opalia) retiporosa: Moony, 1916: 43. Vol. 22; No. 2 THE VELIGER Page 127 ge Epitonium (Nodiscala) retiporosum: DALL, 1917: 474. BAKER, HANNA & STRONG, 1930: 45; plt. 2, fig. 3. GRANT & GALE, 1931: 855. Epitonium (Nodiscala) retiporosa: 1. S. OLDROYD, 1927: 53. STRONG, 1945: 19, 21. A. G. SMITH & GORDON, 1948: 190. KEEN, 1958: 278; fig. 165. Epitonium (Opalia) retiporosum: WILLETT, 1937b: 401. Epitonium retiporosum: KEEN, 1937: 35. Opalia (Nodiscala) retiporosa: DURHAM, 1937: 505; plt. 57, fig. 19. KEEN, 1958: 278; fig. 165. Original Description of Opalia spongiosa Carpenter: “O. t. turrita, parva, albida, marginibus spirae rectis; anfr. ix. subplanatis suturis impressis; costis undulantibus circ. xiii., plerumque (nisi ad suturas crenulatus) obsoletis; tota superficie lineis punctorum creberrimis spiralibus, punctis creberrimis, minutis, altissimis; cerca basim imperforatum costa antica latissima, spirali; apertura ovata, valde callosa; operculo auratiaco, paucispirali. Long., 0.36; long. spir., 0.26; lat. 0.12; div. 20°.” (CARPENTER, 1866: 222). Additional Description: Shell brown when live-taken, slender-conic, medium in size; nuclear whorls 21/2, glassy, brown; post-nuclear whorls 8 to 9 (southern California specimens tending to have 7 post-nuclear whorls), some- what rounded; axial and spiral sculpture present over entire shell; ribs 10 to 15, sinuous, prominent, rounded, overlap- ping the sutures to leave deep pits between, with tubercles at the periphery, more prominent on the last whorl; inter- spaces as wide as the ribs; sutures deep; spiral rows of punc- tations over entire shell, about 28 rows on the last whorl, alternate rows smaller in size; basal disk joined by the exten- sions of the axial ribs to form depressions between; punctate sculpture continuing to the anterior extremity of the shell; aperture subpyriform, set at an oblique angle; lip heavy and reflected on mature specimens, punctate ; operculum brown, horny, paucispiral. Length 7 to 13mm; width 212 to 514 mm. Type Material and Type Localities: Opalia spongiosa Carpenter: Holotype USNM 14830; Mon- terey, California. Opalia retiporosa Carpenter: Holotype USNM 11843; Catalina Island, California. Distribution, Ecology and Bathymetric Range: Monte- rey, California (36°37’ N, 121°54’ W), south into the Gulf of California, and south along the west Mexican coast to the Galapagos Islands, Ecuador (approximately 1°35’S, 80°51’ W). Usually dredged on a sand substrate, in from 18 to 72m. Geologic Record: Pliocene to Recent. Upper Pliocene: San Diego Formation, on Market Street, San Diego, LACMIP loc. 127 [2], and in south- westernmost San Diego County, LACMIP locs. 305 [14], 305-A [4], and 305-C [1]. Lower Pleistocene: Lomita Marl, below Park Western Drive and Host Place, San Pedro, LACMIP loc. 435 [36]. Timms Point Silt, Timms Point, San Pedro, LACMIP locs. 62 [1], and 130-7 [4]. Upper Pleistocene: Palos Verdes Sand, Lincoln Avenue, northeast of Playa del Rey (Los Angeles), LACMIP loc. 59[2] (WILLETT, 1937b: 401; as (O.) retipor- osum). Recent Records: Off Hopkins Marine Station, Pacific Grove, Monterey, California (36°37.5’ N, 121°54’ W), collected by McLean, 36m, 1960, three specimens (LACM 60-23). Off Santa Cruz Island, California, Velero III, 67m, sand and shell substrate, September 1940, two specimens (LACM-AHF 1191 and 1303- 41). Coronados Islands, Baja California Norte, Mex- ico (Pacific side), Puritan Expedition, two specimens dredged (AMNH 145: 77775). Maria Madre Island, Tres Marias Islands, Mexico, Puritan Expedition, 30 m, one specimen (AMNH 59: 74791). Post Office Bay, Charles Island, Galapagos Islands, Ecuador (approx- imately 01°15 S, 90°27’ W), Velero III, bottom sam- ple from 16m, January 1933 (LACM-AHF 402). Discussion: Although CarPENTER (1864c: 613) sug- gested, by implication, the synonymy of Opalia spongiosa and O. retiporosa, 80 years passed before workers ac- cepted the similarities between the 2 taxa. Unfortunate- ly, Carpenter had very poor specimens from which to make his diagnoses. CARPENTER (1866: 222) states of O. retiporosa: “The texture has a rotten appearance; yet one of the specimens was stained with purple, and contained the dried remains of the animal, with its operculum. In the endeavor to extract this, the shell gave way.” Carpenter referred to the two species as new several times in his earlier publications before he finally described them in 1866. PALMER (1958: 194) met the “first reviser” rule (ICZN 24A) in discussing O. spon- giosa and O. retiporosa,citing both taxa, considering them to be the same species, relegating O. retiporosa to the syn- onymy of O. spongiosa. Opalia varicostata Stearns, 1875 (Figures 65, 66, 67, 68) Opalia varicostata STEARNS, 1875: 463; plt. 27, figs. 2-5. DALL, 1878a: 29. DALL, 1878b: 12. CoopeEr, 1888: 255. DALL, 1892: 245; plt. 21, fig. 4. ARNOLD, 1903: 63, 267. ARNOLD, 1906: 28, 100. DALL, 1917: 473. WOODRING, STEWART & RICHARDS, 1941: 38, 46, 54, 70; plt. 36, fig. 6. WoopRING & BRAMLETTE, 1951: 74, 103, 104; plt. 10, fig. 2. KERN, 1973: 85. Scala (Opalia) varicostata: DEBoury, 1919: 36. DURHAM, 1937 [as Opalia (O.)]: 502 Page 128 Epitonium (Opalia) varicostatum: GRANT & GALE, 1931: 853; plt. 24, fig. 20. WILSON, 1966: 112. Opalia anomala STEARNS, 1875: 464; plt. 27, fig. 1. DALL, 1878a: 29. DALL, 1878b: 12. COOPER, 1888: 255. DALL, 1892: 245. ARNOLD, 1903: 63, 266. ARNOLD, 1906: 28. DALL, 1917: 473. Scala (Crassiscala) anomala: DEBourRY, 1919: 36. Epitonium (Opalia) varicostatum var. anomalum: GRANT & GALE, 1931: 854. Opalia (Opalia) varicostata var. anomala: DURHAM, 1937: 502. Opalia (Opalia) varicostata var. granti DURHAM, 1937: 502; ple. 57, fig. 7. Opalia varicostata var. anomala: WoopRING & BRAMLETTE, 1950: 107. “Shell elon- gated-conical, turreted, tapering, solid, imperforate, aper- ture ovate, peristome continuous, thickened; dingy to clear white; suture well defined; whorls united, exceedingly var- iable in convexity and altitude; specimens all decollate, or truncated, equally solid, though varying in length from .75 to 2.45 inches, showing four and one-half whorls within the first measurement to five in the latter. Perfect specimens have probably from 8 to 12 whorls, or even more. Longi- tudinal ribs 9 to 12, varying in number, prominence, and regularity, as well as in obliquity, when compared with the axial line of the shell, and, in some specimens, irregularly thickened and distorted by the intrusion of a varical rib more or less conspicuously. In some individuals the termi- nation of the rib at the suture gives the upper part of the whorls a crenulated appearance, and the suture in all speci- mens is more or less waved, dependent upon the prominence of the ribs, which terminate anteriorly at and join a trans- verse rib at about the middle of the basal whorl. Number of specimens 22. Collected by Henry Hemp- hill’ (Stearns, 1875: 463). Original Description of Opalia varicostata: Additional Description: Shell heavy, solid, tapering, off- white in color; whorls 10 to 12, variable in convexity; suture well defined; ribs 9 to 12, variable in prominence, regularity and thickness, terminating at the prominent basal disk, with an occasional very heavy one riding over the basal disk and continuing to the oval aperture; peristome thick, contin- uous. Length 20 to 60 mm; width 14 to 20mm. Type Material and Type Localities: Opalia varicostata Stearns: Lectotype (USNM 2140402), selected herein; length, 55mm; width, 21mm. Paralec- totypes (3) (USNM 214040b), designated herein; Pacific Beach, San Diego, California (Pliocene, San Diego For- mation). Opalia anomala Stearns: Holotype USNM 214041; same locality as above. THE VELIGER Vol. 22; No. 2 Opalia varicostata var. granti Durham: Holotype UCMP 30166; Pacific Beach, California (Pliocene, San Diego Formation). Distribution, Ecology: Extinct species from Wishkah River, Washington, Zapata Creek, Fresno County, Cali- fornia, Los Angeles and San Diego Counties, California, and northern Baja California Norte, Mexico. Geologic Record: Note: The number of specimens of the variety anomala is followed by “a” in brackets follow- ing each locality. Lower Pliocene: Towsley Formation, Sand Canyon, off Santa Clara River Valley (Los Angeles County), LACMIP loc. 291 [2]. Upper Pliocene: Temescal Canyon, Pacific Palisades, LACMIP locs. 42 [1], and 4472 [1]. Niguel Forma- tion, Via La Mirada, San Juan Capistrano, LACMIP loc. 4923 [4a]. San Diego Formation, in San Diego, at end of Loring Street, Pacific Beach, LACMIP loc. 122 [180; 29a], Diamond Street, Pacific Beach, LACMIP loc. 4523 [8], end of Arroyo Drive, Pacific Beach, LACMIP loc. 107 [6; 2a], near Wabash Boulevard and Gateway Drive, LACMIP loc. 1187 [1], on Mar- ket Street, LACMIP loc. 127 [2], San Diego “area,” LACMIP loc. 4758 [4; 2a], southwesternmost San Diego County, LACMIP locs. 305 [28; 17, rf. al, 305-A [1; 22a], 305-C [2], 318 [2a], and 319 [5a]; in Baja California Norte, 41/2 miles south of U.S. border on old coast highway, LACMIP loc. 449 [2f.], La Joya turnoff on Mexico Highway 1-D, south of Ti- juana, LACMIP loc. 4755 [1]. Discussion: ‘This Pliocene fossil seems to be common in the San Diego, California area, having been reported by many authors. The most northerly occurrence is reported by DurHam (1937: 502) at Wishkah River, Washington from the upper Miocene or lower Pliocene, Montesano For- mation. O palia anomala, described by STEARNS (1875: 464) asa variety of O. varicostata grades directly into the latter through the gradual obsolescence of the longitudinal ribs, except on the spire, where axial sculpture is often present. As a variety it is of little value. DEBoury (1919: 36) erro- neously placed O. anomala in the subgenus Crassiscala, al- though he conceded that O. varicostata belonged in O palia. Duruam (1937: 502) named the variety O. varicostata var. granti, from the Pliocene San Diego Formation, in Pacific Beach, California, with 8 axial ribs, and a thin, high spired shell of 9 whorls. Vol. 22; No. 2 EPITONIIDAE or DOUBTFUL STATUS Epitonium (Nitidiscala) crebricostatum (Carpenter, 1864) Scalaria crebricostata CARPENTER, 1864: 613, 660 [Reprinted, 1872: 99, 146]. CARPENTER, 1866: 222. CooPER, 1870: 67. Keep, 1881: 28; plt. 6, fig. 4 (E. tinctum). Cooper, 1888: 263. KEEP, 1911: 184. Scala crebricostata: ARNOLD, 1903: 263. KEEP, 1904: 202. ARNOLD, 1906: 36. STRONG, 1930: 188. Epitonium crebricostatum: DALL, 1917: 478. T. S. OLDRoyD, 1925: 13. ABBOTT, 1974: 119. Epitonium (Nitidoscala) [sic] crebricostata: DALL, 1917: 478. DALL, 1921: 115. I. S. OLDRoyD, 1924: 108. I. S. OLDRoyp, 1927: 61. Epitonium (Nitidiscala) crebricostata: KEEP, 1935: 180. STRONG, 1945: 24. Epitonium (Nitidiscala) crebricostatum: GRANT & GALE, 1931: 858; plt. 22, fig. 11. KEEN, 1937: 35. STRONG, 1945: 26. Burcu, 1945: 29. A. G. SMITH & GORDON, 1948: 191. KEEN, 1958: 274. PALMER, 1958: 184; plt. 20, figs. 27-28. Description of Scalaria crebricostata CARPENTER (1864: 660): “= Mus. Cuming no. 32: 15 sharp, reflexed ribs, coro- nated against the sutures.” Description of Scalaria crebricostata CARPENTER (1866: 222): “S. t. gracili, tenui, alba; anf. x.rotundatis, haud at- tingentibus; costis circ. xv., acutis, reflexis, vix attingenti- bus, lineis irregulariter spiralibus ascendentibus; costis juxta suturam eleganter coronatis; sculptura spirali, nisi striulis interdum exillimis, nulla; aperture rotundata; um- bilico nullo; operculo normali, dense corneo. Long. 0.70; long. spir. 0.52, lat. 0.18, div. 26 [length 17.71 mm; width 4.55 mm]. Hab. Monterey, San Pedro, Cooper, common. = “Scalaria, unique” Mus. Cum. No. 32. Somewhat resembles S. tenuis, Sby., but is not so turrited.” Material and Localities: BM(NB), Registry 1950. 3. 29. 1, one specimen labelled “California,” 12 costae, length 9.7 mm; width 3.6 mm. USNM 46234 [ex 14831]. Recatalogued in 1885, labelled “Type-Monterey-Cooper,” 3 live-taken specimens: 11 costae, length 16 mm; width 8.5 mm 10 costae, length 9 mm; width 4 mm 4 costae, length 8.5 mm; width 4 mm Discussion: Through the years Scalaria crebricostata Carpenter has been an enigma to workers. None of the spec- imens extant at the BM(NH), and the USNM match the descriptions given by CARPENTER (1864: 660; 1866: 222). Inasmuch as Carpenter’s brief description (1864) and de- tailed description (1866) specify a shell of dimensions differ- ent from those of the one specimen in the BM(NH), one can only assume that the original specimen is lost. None of the THE VELIGER. Page 129 specimens at the USNM can qualify because their dimen- sions are far different from the descriptions given by Car- penter. Because of the uncertainty of the identity of Carpen- ter’s taxon it is here rejected as a species inquirenda. The specimens at the USNM labelled “crebricostata” are Niti- discala caamanoi (Dall and Bartsch, 1910). REJECTED RECORDS Epitonium (Asperoscala) sic] arnoldi Dall, 1917 The description given by DaLt (1917: 475), type locality San Pedro, California (length, 14 mm; width, 5.5 mm), par- allels that given by CLENCH & TURNER (1952: 292) for Epi- tonium (Asperiscala) multistriatum (Say, 1826). Compar- ison of the holotype of E. arnoldi Dall (USNM 106875), with photographs of the type of E. multistriatum (Say) con- firms the opinion that the description of E. arnold: is based on specimens of E. multistriatum (Say) with an erroneous locality label. On the Atlantic coast the range is from Buz- zards Bay, Massachusetts south to Cape Canaveral (Cape Kennedy), Florida. Epitonium (Nitidoscala) [sic] barbarinum Dall, 1919 The holotype of Epitonium (Nitidoscala) [sic] barbarinum Dall, 1919 (USNM 46229) from the Stearns collection, type locality, San Diego, California, is a specimen of Epitonium angulatum (Say, 1830), with a range on the Atlantic sea- board and in the Gulf of Mexico, from Long Island, New York south to Florida (excluding the Florida Keys) and west to Texas. DALL (1921: 116) gave the range for E. bar- barinum from San Diego, California to Panama. Moreover, Dall’s original measurements (length 19mm; width 6.5 mm) are in error. The type of E. barbarinum has been meas- ured several times by Joseph Rosewater (USNM) and Du- Shane: length 13.9mm; width 6mm, within the size range for E. angulatum. Thus, E. barbarinum, as a species, is elim- inated from the eastern Pacific fauna, and was renamed as a previously misidentified species from the upper Gulf of California, Mexico (DUSHANE, 1979: 379). Epitonium (Nitidiscala) hexagonum (Sowerby, 1844) Originally described by SowerBy (1844: 98) (four syn- types: BM(NH)), from Acapulco, Mexico, this species is not known north of Magdalena Bay, Baja California Sur (SDMNH Collection), collected by Orcutt, 1917. There are several records of beachworn shells, from California and Page 130 THE VELIGER ! Vol. 22; No. 2 Baja California, dating from the turn of the century; USNM 221848 and USNM 221849, Santa Cruz, Califor- nia (Button); USNM 211020, Ocean Beach, California (Cook); USNM 105506, Scammon’s Lagoon (Hemphill). It is possible that the species had a more extensive distribution at that time; however, until live-collected specimens are known from this region, the species is not regarded as a member of the fauna. The range is from Magdalena Bay, Baja California Sur, and the west side of the Gulf of Cali- fornia, Mexico; disjunct to Acapulco, Mexico, then south to Panama. Epitonium lagunarum Dall, 1917 Known only from the holotype (USNM 253024), type lo- cality Laguna Beach, California, there have been no sub- sequent reports of this taxon from the eastern Pacific. Datv’s (1917: 477) description matches well the descrip- tion of Epitonium rupicolum (Kurtz, 1860), and examina- tion of the type of E. Jagunarum confirms the opinion. Epi- tonium rupicolum is a common species from Massachusetts to Texas. Epitonium tiara (Carpenter, 1856) Datv’s record (1917: 480) of Epitonium tiara Carpenter, ranging from Catalina Island, California to Todos Santos Bay, Baja California Norte (repeated by I. S. O_prRoyp (1927), KEEN (1937),and Burcu (1945)) wasa misidenti- fication, as the taxon is synonymous with Asperiscala obtusa (Sowerby, 1844) (DUSHANE, 1974: 22), a species ranging from the Gulf of California south to Colombia. Epitonium zephyrium Dall, 1917 The United States National Museum has two specimens labelled “Epitonium zephyrium” (USNM 56056 and USNM 635572). Neither specimen matches the descrip- tion given by Dat (1917: 485) who stated, “without basal disk or cord.” Both USNM specimens have a basal cord and neither has the dimensions given by Dall in his original description. Dall did not figure the specimen. One can only conclude that the holotype is now lost. No other specimens of E. zephyrium seem to be in museum or private collec- tions. The two specimens at the USNM are Epitonium lamellosum (Lamarck, 1822), which has been reported along the shores of Australia, Japan, southern Europe, western, southern, eastern Africa, eastern coast of the United States from Florida and the Gulf of Mexico south to eastern Colombia. Literature Cited Aszott, Ropert TucKER 1974. American seashells. Nostrand Reinhold, New York Apams, CHarRLes BAKER 1852. Catalogue of shells collected at Panama with notes on synonymy, station and habitat. Ann. Lyc. Nat. Hist. New York 5: 229 - 296 (June); 297 - 549 (July) [reprinted: 1852,, R. 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Suppl. 11: 223 - 295 ARNOLD, RALPH 1903. ‘The paleontology and stratigraphy of the marine Pliocene and Pleistocene of San Pedro, California. Calif. Acad. Sci. Mem. 3: 420 pp.; 37 pits. (27 June 1903) [reprinted as: Stanford Univ., Contributions to Biology from the Hopkins Seaside Laboratory no. 31, same date and pagination] 1906. ‘Tertiary and Quaternary Pectens of California. U.S. Geol. Surv. Prof. Paper 47: 7-146; 53 plts.; 2 text figs. Baxer, Freperick, G Dattas HANNA & ARCHIBALD McC.iure STRONG 1930. Some Mollusca of the family Epitoniidae from the Gulf of Cali- fornia. Proc. Calif. Acad. Sci. (4) 19 (5): 41-56; plts. 2, 3 (15 July 1930) BartscH, Paut 1915. The Recent and fossil mollusks of the genus Rissoina from the west coast of America. Proc. U.S. Natl. Mus. 49 (2094): 33 - 62; pits. 28 - 33 (24 July 1915) 1927. New west American marine mollusks. Proc. U.S. Natl. Mus. 70 (11): 1-36; pits. 1-6 (8 April 1927) BERNARD, FRANK R. 1970. 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Museum Boltenianum sive catalogus cimeliorum e tribus regnis nature quae olim collegerat Joa. Fried. Bolten ... Pars secunda con- tinens conchylia sive testacea univalvia, bivalvia + multivalvia. Hamburgi, Johan Christi Trappii: i-viit+1-199 (post-10 September, 1798; facsimile repr. by Charles Davies Sherborn & Ernest Ruthven Sykes, March 1906) Sato, SicRip 1977. Observations on feeding, chemoreception, and toxins in two species of Epitonium. The Veliger 20 (2): 168-172; 1 plt.; 1 text fig. (1 October 1977) ScHucHErT, CHariges, WILLIAM HEALEY DALL, T. W. STANTON & R. S. Basser 1905. Catalogue of the type specimens of fossil invertebrates, in the Department of Geology, United States National Museum. U. S. Natl. Mus. Bull. 53 (1) (sec.1): 1-704 SmitrH, ALLYN Goopwin & MacKenzie Gorpon, Jr. 1948. The marine mollusks and brachiopods of Monterey Bay, Cali- fornia, and vicinity. Proc. Calif. Acad. Sci. (4) 26 (8): 147-245; pits. 3, 4; 4 text figs. (15 December 1948) Smitu, Carzy RescH 1977a. Chemical recognition of prey by the gastropod Epitonium tinc- tum (Carpenter, 1864). The Veliger 19 (3): 331-340; 4 text figs. (1 January 1977) 1977b. Chemical recognition of prey by the gastropod Epitonium tinc- tum. Ann. Rprt. West. Soc. Malacol. 10: 9 (14 Dec. 1977) SmitH, RatpH INcRAM & James T. Car.Ton (eds.) 1975. Light’s Manual. Intertidal invertebrates of the central California coast. gid ed., xvilit+716 pp.; 156 plts. (8 May 1975) SowErBy, GeorcE BretTINGHAM 2nd 1844a. Thesaurus conchyliorum, or monograph of genera of shells, ed- ited by G. B. Sowerby, Jr., completed by G. B. Sowerby 3rd, London, 1 (4): 83-146; plts. 32-40 (11 April 1844) 1844b. Descriptions of new species of Scalaria, collected by Hugh Cum- ing, Esq. Proc. Zool. Soc. London prt. 12: 1 - 38 (July 1844) SpHon, GALE G. 1966. Material contained in the molluscan type collection of the Santa Barbara Museum of Natural History. The Veliger 9 (2): 244 to 246 (1 October 1966) 1971. Type specimens of recent mollusks in the Los Angeles County Museum of Natural History. Los Angeles Cty. Mus. Contr. Sci. no. 213: 37 PP. (27 May 1971) STEARNS, RoperT Epwarps CaRTER : 1875. Descriptions of new fossil shells from the Tertiary of California. Proc. Acad. Nat. Sci. Philadelphia, 1875: 463 - 464; 1 plt. Strono, ARCHIBALD McCiure 1930. Notes on some species of Epitonium, subgenus Nitidiscala, from the west coast of North America Trans. San Diego Soc. Nat. Hist. 6 (7): 183 - 196; plt. 20 (go August 1930) 1931. Further notes on Epitonium fallaciosum. The Nautilus 45 (2): 70-71 (14 October 1931) 1937- Notes on west coast Epitoniidae. The Nautilus 51 (1): 4-8; part of plate 2 (3 July 1937) 1941. Notes on Epitonium (Nitidoscala) [sic] tinctum (Carpenter). The Nautilus 55 (2): 46-47 ; (24 October 1941) 1945. Key to subgenera of Epitonium. Minutes Conch. Club So. Calif. (John Q. Burch, ed.) 52: 16 (September 1945) TAPPARONE-CANEFRI, CESARE MARIA 1876. Rectifications dans la nomenclature de quelques espéces du genre Scalaria. Journ. de Conchyl. (3) 24 (2): 152-156 THIELE, JOHANNES 1928. Uber ptenoglosse Schnecken. 73 - 94 TuHorson, GUNNAR : : 1957- Parasitism in the marine gastropod family Scalidae. Medd. naturh. Foren. Kobenhagen 119: 55 - 58 Tryon, GzorcE WASHINGTON, Jr. : 1887. Manual of conchology; structural and systematic. (1) 9: 1 to 488; 71 plts. Philadelphia Zeitschr. wissensch. Zool. 19: Vedens. Page 134 THE VELIGER Vol. 22; No. 2 } WILLETT, GEORGE oe Nie pare Re eae eee Fish Bull. 1918. ; Notes on the Mollusca of Forrester Island, Alaska. The 146: 221 pp ; 74 text figs. Nautilus 32 (2): 65-69 } (22 October 1918) TAS, RaeR DIOR ee A ae Pistia from agree i eae oe ae ene 1956. The eastern Pacific marine mollusks described by C. B. Adams. eat a cad aah neste e ae April ue Mus. Comp. Zool., Harvard Univ., Occ. Pap. Mollusks 2 (20): 21 - 136; 1937a. Additions to our knowledge of the fossil invertebrate fauna of Vv a 57 Ia Ww (21 September 1956) California. Bull. South. Calif. Acad. Sci. 36 (2): 60-64; plts. 24, ALENTINE, JAMES WILLIAM a 0) September 1961. Paleoecologic molluscan geography of the Californian Pleisto- I Laas An upper Pleistocene fauna from the meee Hills, oa) cene. Univ. Calif. Publ. Geol. Sci. 34 (7): 309-442; 16 text geles County, California. Trans. San Diego Soc. Nat. Hist. 8 (30) : figs. (17 May 1961) 379 - 406; plts. 25, 26 (15 December 1937) 1966. Numerical analysis of marine molluscan ranges on the extra- 1938. Remarks on some west American mollusks. The Nautilus tropical northeastern Pacific shelf. Limnol. Oceanogr. 11 (2): 198 52 (1): 10-11 (22 July 1938) to 211; 7 figs. 1939. A new species of mollusk from the San Pedro Pleistocene. VALENTINE, JAMES WILLIAM & Rosert F MEADE Bull. South. Calif. Acad. Sci. 38 (3): 202-203; plt. 54 (December) 1961. Californian Pleistocene paleotemperatures. Univ. Calif. Witson, Epwarp C. « Donatp E. Binc Publ. Geol. Sci. 40 (1): 1-46; 4 text figs. (14 July 1961) 1970. Type specimens of fossil invertebrata in the Los Angeles County Van WINKLE, KaTHERINE EVANGELINE HILTON Museum of Natural History, exclusive of paleoentomology. Los [see also: Parmer, KaTHERINE E. V. W] Angeles County Mus. Contrib. Sci. no. 181: 20 pp. (27 Feb. 1970) 1921. Illustrations and descriptions of fossil Mollusca contained in the Witson, Epwarp C. & Grorce L. Kennepy : paleontological collection of Cornell University. Cornell Univ. Bull. 1967. Type specimens of Recent invertebrates (except Arachnida and Amer. Paleont. 8 (36): 3-6; 1 plt. Insecta) in the San Diego Natural History Museum. Trans. San Vepper, Joun G. & ELLEN J. Moore Diego Soc. Nat. Hist. 14 (19): 237-279 (17 November 1967) 1976. Paleoenvironmental implications of fossiliferous Miocene and Wooprine, WENDELL PHILLIPS : Pliocene strata on San Clemente Island, California. In: Aspects of 1931. Epitonium fallaciosum. The Nautilus 45 (1): 31 (13 July) the geological history of the California continental borderland, D. G. Wooprinc, Wenvext Puituips & Mitton NuNN BRAMLETTE irae arc Howell, ed. Pacific sect. Amer. Assoc. Petrol. Geol. Misc. Pub. 24: 1950. Geology and paleontology of the Santa Maria District, ip HEA : 6) fornia. U.S. Geol. Surv. Prof: Paper 222: 1 - 185; 23 plts.; 9 text 107-136; 4 plts.; 9 text figs. (15 June 197 fous @ seme Wooprino, WENDELL PuHixups, RaLpH STEWaRT & R. W. RICHARDS 1941. Geology of the Kettleman Hills oil field, California. U. S. Geol. Surv. Prof. Paper 195: i- v+167 pp.; 57 plts.; 15 text figs.; 2 inserts (7 June 1941) ZINSMEISTER, WILLIAM J. 1970 [1971]. A late Pliocene macrofossil fauna of Newport Beach, Orange County, California. Bull. So. Calif. Acad. Sci. 69 (3 & 4): 121-125 (October-December) Vol. 22; No. 2 THE VELIGER On the Growth Stages of Conus fergusont Sowerby, 1873, the Reinstatement of Conus xanthicus Dall, 1910, and a New Species of Conus from the Galapagos Islands BY JAMES H. McLEAN Natural History Museum of Los Angeles County, Los Angeles, California 90007 AND JAMES NYBAKKEN Moss Landing Marine Laboratories of the Central California State Universities and Colleges Moss Landing, California 95039 (3 Plates; 4 Text figures) Page 135 INTRODUCTION IN REVIEWS OF EASTERN PACIFIC Conus species, NYBAKKEN (1970: 25; 1971: 97) reported markedly different radula morphology between specimens considered adults and juve- niles of C’. fergusoni Sowerby, 1873. Nybakken suggested 3 possible explanations for this apparent anomaly: 1) di- morphism inC.fergusoni, 2) change inradula morphol- ogy with growth, 3) confusion between 2 species. Too few specimens were then available to allow resolution of the problem. Since that time additional specimens have en- abled us to reach new conclusions. Mature specimens of Conus fergusoni, largest of the trop- ical eastern Pacific species of Conus, have readily been rec- ognized by previous workers. The juvenile shell of the spe- cies has not been understood, however. Recent authors (HANNA & STRONG, 1949; KEEN, 1958; EMERSON & OLD, 1962; Hanna, 1963; NyBAKKEN, 1970, 1971; KEEN, 1971) have considered the taxon C. xanthicus Dall, 1910, to represent the immature form of C. fergusoni. We now show that C. xanthicus is separable from C. ferguson: on both shell and radula morphology. The true juvenile of C. fergusoni is described and figured for the first time. During this study, we found that some specimens from the Galapagos Islands thought to be Conus xanthicus had a radula unlike that of either C’. xanthicus or C. fergusoni. We describe these specimens as a new species. C. kohni McLean & Nybakken. In this account we give comparative descriptions of the 3 taxa and figure a number of specimens to illustrate growth stages and some of the variation in each species. We have examined all previously illustrated specimens in the Cali- fornia Academy of Sciences and the American Museum of Natural History collections. The figure citations in our syn- onymies are based upon new determinations of the identity of these specimens. Our radular descriptions employ the terminology used by NyBAKKEN (1970). The accounts of the radula are based on the following number of examined specimens: Conus fergusoni 5, C. xanthicus 10 (including the holotype); C. kohn 6 (including the holotype). Museum abbreviations used in the text are as follows: AHF, Allan Hancock Foundation, University of Southern California (collection housed at LACM); CAS, California Academy of Sciences, San Francisco; LACM, Los Angeles County Museum of Natural History; USNM, United States National Museum of Natural History, Washington. ACKNOWLEDGMENTS Weare especially grateful to Mr. Leroy H. Poorman, West- minster, California, who was much involved in the discus- sions that led to the recognition of the growth stages of these species. He has also donated specimens (Figures 6, 11, and 15) important to this work. Mr. Gerald Wellington, Uni- versity of California, Santa Barbara, contributed specimens Page 136 from the Galapagos Islands, including the type lot of the new species. Additional specimens have been loaned by Mr. Alex Kerstitch of Tucson, Arizona. We thank curators and staff at the American Museum of Natural History, Califor- nia Academy of Sciences, and the National Museum of Natural History for the loan of the specimens. William K. Emerson, A. Myra Keen, Alan J. Kohn, Pat- rick I. LaFollette, and Leroy H. Poorman critically read the manuscript and offered helpful suggestions. We are grateful to Dr. Kohn for allowing us to compare the new species with his collection of type photography, specimens, and radula slides. The photographs are the work of Mr. Bertram C. Draper, Los Angeles. SPECIES ACCOUNTS Conus fergusoni Sowerby III, 1873 (Figures 1, 2, 5-77) Conus fergusoni SowERBY III, 1873: 145; plt. 15, fig. 1 - 1887: 256; plt. 508, fig. 675 — DALL, 1910: 218 — HANNA & STRONG, 1949: 294; plt. 7, fig. 3 — KEEN, 1958: 485; fig. 938 — EMERSON & OLD, 1962: 26; fig. 14 — HANNA, 1963: 42; plt. 4, fig. 2; plt. 9, fig. 10 — NYBAKKEN, 1970: 13; figs. 18, 19 (radula), figs. 40, 41 — NYBAKKEN, 1971: 97; fig. 4 — KEEN, 1971: 667; fig. 1511 left. Diagnosis: Coronations persisting through 10th whorl, spire whorls slightly concave; large specimens white, small THE VELIGER Vol. 22; No. 2 specimens yellow-orange, with lighter, even banding; shells under 25mm in length with widely spaced spiral rows of dark brown spots; spire lacking color pattern. Description: Shell large (maximum length 153 mm, with 12 to 13 teleoconch whorls); spire low to moderately ele- vated; spire outline concave in small specimens to nearly straight in large specimens; shoulder sharply angulate in small specimens, less angulate in large specimens; the angu- lation with low coronations in small specimens, the corona- tions often persisting through a shell length of 50mm; coronations indistinct and undulating in large specimens; spire whorls slightly concave, spire sculptured with fine spiral striae and growth lines; suture produced at the shoulder, deeply and narrowly incised; whorl profile more or less straight except convex below the shoulder; surface smooth except for 10-15 spiral striae on lower third of whorl ; aperture moderately broad, of about the same width throughout and conforming to whorl profile. Protoconch homeostrophic, 3-whorled, smooth, dark; shoulder of early teleoconch whoris angulate; suture produced well below the angulation; suture rising to meet the shoulder by the 6th whorl; coronations on the shoulder angulation persist- ing to at least the 8-whorled stage. Color light yellow- orange, paler in medium-sized specimens, fading to white in large specimens; small specimens with a distinct lighter spiral band about the middle of the shell and usually a second light band at the shoulder; spire lacking color pat- tern; small specimens with spiral rows of dark brown dots on body whorl; aperture white within. Periostracum thin Explanation of Figures 5 to 12 (Figures are scaled to render a shell 60 mm in length at life size and a 15 mm shell at a length of 35 mm, with intermediate-sized shells proportionally scaled. Spire views are oriented perpendicular to a plane touching the apex and the shoulder and are enlarged to a diameter of 2/3 the length of the frontal view.) Conus fergusoni Sowerby III, 1873 Figure 5: AHF 395-35, 26-29 m, rocky, off Isla Lobos de Afuera, Peru. Radula verified specimen, length 61 mm, periostra- cum intact. Figure 6: LACM 35506, 73m, muddy, Bahia Bocochibampo, Guaymas, Sonora, Mexico. Length 52.6 mm, periostracum removed (spire with persistent coronations; largest speci- men examined showing juvenile spotted pattern) . Figure 7: LACM 72-73, 40-55 m, Golfo Dulce, Puntarenas Prov., Costa Rica. Length 52.4mm, periostracum intact (spire coronations relatively faint) . Figure 8: AHF 431-35, 82m, sand & gravel, off Rocas Octavia, Colombia. Radula verified specimen, length 43.5 mm, perio- stracum intact (spire coronations prominent). Figure 9: CAS 12310, 77 m, off Punta Judas, Costa Rica. Length 42.8 mm, periostracum removed from body, intact on spire (spotted specimen figured by Hanna, 1963, plt. 9, fig. 10). Figure zo LACM 72-12, 53-26 m, mud, Bahia Elena, Guanacaste Prov., Costa Rica. Radula verified specimen (see Figure 2), length 26.1 mm, periostracum removed (spotted juvenile; smallest radula-verified specimen). Figure 1z: LACM 36563, 20m, Bahia Santiago, Colima, Mexico. Length 14.0mm, dead specimen, periostracum worn away (spotted juvenile, smallest specimen examined). Conus virgatus Reeve, 1849 Figure 12: CAS 39174, vicinity of Guaymas, Sonora, Mexico (from shrimp boats). Length 41 mm, periostracum removed, sur- face eroded (example of banded form; note lack of corona- tions; specimen previously misidentified as C. fergusoni). Tue VELIGER, Vol. 22, No. 2 [McLean « NypakkeEn] Figures 5 to 12 ) Figure 8 — Figure 10 . 5 Ti Figure 11 Figure 12 Vol. 22; No. 2 THE VELIGER Page 137 and light colored in small specimens, thick and dark brown in large specimens, produced in closely set, concave ridges on the spire. Operculum bluntly unguiculate, about 4 times longer than wide. Radula (Figures 1, 2): Tooth from mature specimens Figure 1 Conus fergusoni, from 26.1 mm specimen (shell: Figure zo). Radular tooth (scale = 0.1mm) (Figure 2) long; serration extending half the length of the tooth, terminating proximally in a pointed or rounded cusp; barbs 2, one at the tip, the other on the opposite side; blade and waist lacking, base enlarged, rounded, bearing a pointed spur. Tooth from the smallest specimen exam- ined (Figure 1) (shell length 26.1 mm, Figurg¢ 70), propor- tionately shorter, the 1st barb weaker, 2nd barb undevel- oped; waist indistinct, the proximal portion of the shaft wider than the distal; base enlarged, rounded, with prom- inent spur. Radula preparations were made from 5 specimens, in- cluding those illustrated in Figures 5, 8, and ro. Except for the radula from the smallest specimen (Figure 10), there was no significant variation in tooth morphology. Distribution and occurrence: Bahia Tortuga, Baja Cali- fornia, Mexico (HANNA & STRONG, 1949), north in the Gulf of California to Guaymas, Sonora, south to Isla Lobos de Afuera, Peru; Galapagos Islands, Ecuador. Conus fer- gusoni is uncommon at the Galapagos Islands; we have received specimens collected by Gerald Wellington at Isla Isabela and HANNA & STRONG (1949) mentioned a speci- men of 128mm in length collected at Caleta Tagus, Isla Isabela. Near the northern and southern range extremes, the entrance to Bahia Magdalena, Baja California, Mex- SN rere way cOoKc cS ST SRee Figure 2 C. fergusoni, from 51.9 mm specimen (Gulf of Panama, Univ. of Miami, uncatalogued). Radular tooth (scale = 0.1mm) ico, in the north, and Isla Lobos de Tierra and Isla Lobos de Afuera, Peru, in the south. McLean observed C. fergu- soni on rocky bottoms at shallow subtidal depths. At local- ities in the central part of the range it is less common and is known only from dredged records. The species is not re- stricted to a rocky substrate; McLean has observed large numbers taken on soft bottoms on the shrimp fishing ground in the Gulf of Guayaquil off northernmost Peru. Comparisons: Immature specimens of Conus fergusoni differ from C. xanthicus and C. kohni in having prominent coronations on the early whorls. The coronations persist through at least the 8-whorled stage and may be observed in spire view on specimens of any size, including shells with Page 138 THE VELIGER Vol. 22; No. 2 intact periostraca. Similar persistent coronations charac- terize C’. patricius Hinds, 1843, but that species differs in having a rounded shoulder and a marked concavity of the lower body whorl profile, features particularly apparent in young shells. The color patterns of young shells of C. fer- gusoni differ from those of both C. xanthicus and C. kohni in lacking mottling on the spire and in having even band- ing, and in younger stages, rows of dark spots, instead of mottled banding with jagged edges. An uncommon color form of Conus virgatus Reeve, 1849, has also been a source of confusion. This form, an example of which was described as C. signae Bartsch, 1937, lacks axial flammules and may have a central band of lighter color intensity. However, such specimens of C. virgatus (Figure 12) lack coronations on the spire and the whorls are not as rapidly expanding as in C. fergusoni. Variability: Conus fergusoni is moderately variable in shell proportion and height of spire. For example, shells in Figures 6 and 7 are of about the same length but that in Figure 6 is lower spired, broader at the shoulder and has a more pronounced bulge below the shoulder. Young shells lose the typical coloration of banding and spotting at dif- ferent sizes. All of the 5 smallest shells examined (none ex- ceeding 26 mm in length) are spotted. Spots have been observed on one shell as large as 52.6 mm in length (Figure 6); the smallest shell observed on which spots are lacking is 43.5 mm in length (Figure 8). Remarks: The original illustration of Conus fergusont is an accurate rendition of a large, white-shelled specimen lacking the periostracum, 145mm in length, apparently life size, no dimensions being given. The description con- sisted of a brief Latin diagnosis. The source of the original material was given as follows: “Several specimens . . . col- lected at Panama by Mr. Ferguson.” The present location of type material is unknown; specimens have not been lo- cated in the British Museum (Alan J. Kohn, personal com- munication). Previous accounts of Conus ferguson have included C. xanthicus Dall, 1919, as a synonym, thereby attributing ex- cessive variability to the species. This confusion is no doubt due to scarcity of juvenile specimens of C’. fergusont. In all the museum and private collections examined by us, we have located only five specimens under 40mm in length. McLean can attest to the apparent absence of juveniles, having seen numbers of mature specimens at Isla Lobos de Afuera, Peru, but none under 55mm in length, despite having taken many gravel samples from crevices and under rocks. Hanna & STRONG (1949: 295) and Hanna (1963: 43) claimed that ‘“‘a magnificent series of growth stages has en- abled us to state with assurance that C. xanthicus is the young of C. ferguson.” We have examined all small speci- mens in the California Academy collection previously iden- tified as C. fergusoni and find that the smallest authentic specimen studied by Hanna is the 42.8 mm shell figured in Explanation of Figures 13 to 23 Conus xanthicus Dall, 1910 Figure 73: USNM 111236, Holotype of C. xanthicus, 130 m, sand, off Guaymas, Sonora, Mexico. Radula verified specimen, length 42.5 mm, periostracum removed. Figure z4: Berry Collection, Holotype of C. chrysocestus, 55-82 m, off Morro Colorado, Sonora, Mexico. Length 45.3 mm, periostracum removed (relatively low-spired specimen). Figure 75: LACM 11345, 73 m, off La Paz, Baja California, Mex- ico. Radula verified specimen, length 53.4 mm, periostracum removed (relatively low-spired specimen). Figure 76: AHF 1118-40, 108-126m, coarse gray sand, Banco Gorda, Baja California, Mexico. Length 40.0 mm, perio- stracum removed on ventral side, intact on dorsal side (rela- tively high-spired specimen). Figure 17: CAS 17809, 29 m, Bahia Chamela, Jalisco, Mexico. Length 41.9 mm, periostracum intact (relatively low-spired specimen; spiral sculpture on spire whorls especially prom- inent). Figure 18: AHF 300-34, 73 m, sand, Bahia Azufre, Isla Clarion, Revillagigedo Islands, Mexico. Radula verified specimen, length 24.3 mm, periostracum removed. Figure 79: AHF 431-35, 82 m, sand & gravel, off Rocas Octavia, Colombia. Radula verified specimen, length 39.1 mm, perio- stracum removed (relatively slender specimen). Figure 20: AHF 325-35, 146 m, rocky, Caleta Tagus, Isla Isabela, Galapagos Islands, Ecuador. Length 46.8 mm, periostracum removed (largest specimen examined from Galapagos Is- lands). Figure 27: AHF 792-38, 128-146 m, off Isla Daphne Chica, Gala- pagos Islands, Ecuador. Radula verified specimen, length 35-7 mm, periostracum intact. Figure 22: CAS 38975, 18-37 m, off Isla Rabida, Galapagos Is- lands, Ecuador. Length 32.2 mm, periostracum removed. Figure 23: AHF 324-35, 82m, rock, Caleta Tagus, Isla Isabela, Galapagos Islands, Ecuador. Radula verified specimen, length 14.3mm, periostracum partially intact (smallest specimen examined). THE V: Vol. 22, No. 2 [McLEAN & NyBAKKEN] Figures 13 to 23 HE VELIGER, Vol. 22, No. Figure 22 Figure 23 Figure 27 Vol. 22; No. 2 THE VELIGER Page 139 color (HANNA, 1963: plt. 9, fig. 10), our Figure 9. Other small specimens in the Academy collection are identified by us as C. xanthicus. There is no indication of intergrada- tion with C. fergusoni. EMERSON & OLD (1962: 27) consid- ered specimens resembling C. xanthicus “to be merely a highly colored deep water ecotype of C. fergusoni,” but did not document any depth-related distinctions. They noted coronations in the one authentic specimen of C. fergusoni they illustrated but did not question the lack of this char- acter in the other specimens so identified. Conus fergusoni is one of few species in which a young specimen of Conus has been shown to have a radular tooth morphology differing from that of the adult. We treat this further in the discussion section of this paper. Conus xanthicus Dall, 1910 (Figures 3, 13-23) Conus xanthicus DALL, 1910: 225 — HANNA & STRONG, 1949: 294 (as syn. of C. fergusont), plt. 7, figs. 1, 2, 4 (holotype) — KEEN, 1958: 485 (as syn. of C. fergusont) - EMERSON Oxp, 1962: 26 (as syn. of C. fergusoni), figs. 13 left 18 right, 15 left, 15 right — HANNA, 1963: 42 (as syn. of C. fergusoni), pit. 2, fig. 4, plt. 5, fig. 5 (holotype), plt. 7, fig. 9 - NYBAKKEN, 1970: 25 (status uncertain); fig. 20 (radula), figs. 42, 43 -— NYBAKKEN, 1971: 97 (as possibly valid species); fig. 5 — KEEN, 1971: 667 (as syn. of C. fer- gusont); fig. 1511 right (holotype of C. chrysocestus). Conus chrysocestus BERRY, 1968: 157 — NYBAKKEN, 1971: 99 (as syn. of C. fergusoni — KEEN, 1971: 667 (as syn. of C. fergusonz); fig. 1511 right (holotype). Diagnosis: Coronations present only at very early stage, spire whorls slightly concave, aperture narrow; color yel- low-orange, lighter color bands irregular, mottled; spire whorls mottled; spire color darker than that of body whorl. Description: Shell medium-sized (maximum length 54 mm, with 10 teleoconch whorls) ; spire varying from low to moderately elevated; spire outline straight to somewhat concave; shoulder distinctly ridged, not coronated in ma- ture stages; spire whorls flat to slightly concave, spire sculp- tured with faint growth lines and microscopic spiral striae; suture produced below the shoulder ridge, well de- fined but not deeply channeled; anal sinus of medium depth; body whor! profile with a faint convexity below the shoulder, basal profile variable from straight to convex or concave; surface smooth except for 10-15 spiral striae on base; aperture relatively narrow, of same width through- out and conforming to whorl profile. Protoconch homeo- strophic, 3-whorled, smooth, light colored; shoulder of early teleoconch whorls coronated, coronations lost by 3rd to 6th teleoconch whorl; early juvenile shells with 2-3 in- cised spiral striae below the shoulder. Color dark yellow to orange brown, with two irregular, often interrupted white bands; basal area variegated with white; white mottling at the shoulder produces light and dark radial markings on spire; ground color on spire of greater intensity than that of body whorl; aperture white within. Periostracum brown- ish, thin over body whorl, thicker on spire and produced in closely set, thin, arched ridges. Operculum bluntly unguic- ulate, about two times longer than wide. Radula (Figure 3): Tooth of the “Conus regularis type” (NYBAKKEN, 1970), with single barb at the tip, a well- developed opposite blade extending about 1/3 the length of the tooth; serration prominent and extending about the same distance as the blade; waist slight; base enlarged with a small spur on one side. Figure 3 C. xanthicus, from 35.7 mm specimen (shell, Figure 21). Radular tooth (scale = 0.1mm) Radula preparations were made from ten specimens, ranging in shell length from 14.2-54.3 mm, including those illustrated in Figures 13 (holotype), 15, 18, 79, 21, and 23. There was no significant variation in tooth morphology among the examined specimens. Distribution and occurrence: Morro Colorado, Sonora, Mexico (type locality of Conus chrysocestus), to Rocas Octavia, Colombia (6°47’N; AHF 431-35); Revillagigedo Islands, Mexico, and Galapagos Islands, Ecuador. All oc- Page 140 currences are well offshore, chiefly in the 50-140m depth range. We have examined 4 lots from Sonora, 14 lots from the southeastern side of Baja California from Isla Carmen to the Gorda Banks off Cape San Lucas, 12 lots from Isla Clarion, Revillagigedo Islands, 1 lot from southern Mex- ico, 1 lot from the Perlas Islands, Panama, 1 lot from Colombia (Figure 19) and 1o lots from the Galapagos Islands. The Rocas Octavia locality in Colombia is both the southernmost record known for C. xanthicus and the only station from which both C. xanthicus and C. fergu- soni have been collected together. Comparisons: Although Conus xanthicus could not be confused with large, fully mature specimens of C. fergu- soni, it has been confused with immature specimens of the latter. Mature C. xanthicus differ from immature C. fer- gusont in lacking coronations, producing the suture below (rather than at) the shoulder, having a narrower aperture, having an irregular, mottled (rather than even) banding pattern on the body whorl, and having a mottled (rather than unmarked) pattern on the spire. Juvenile C. xanthi- cus do not have the regular rows of spots of juvenile C. fer- gusoni. Conus xanthicus has a general resemblance to C. virgatus Reeve, 1849, but has a more angulate shoulder and does not have the axial color markings of that species. Like C. fergusont, C. virgatus also produces the suture at the shoulder. Comparisons with C. kohni are given under the treatment of that species. Variability: Conus xanthicus is highly variable in breadth and in spire profile. Too few specimens are avail- able to fully document the range of variation, but some generalizations may be made. Those seen from Sonora are relatively low spired, although the spire of the holotype of C. xanthicus (Figure 13) is higher than that of the holo- type of the synonymous C. chrysocestus (Figure 14). Those from the southeastern side of Baja California are similarly THE VELIGER Vol. 22; No. 2 variable, but some higher spired forms are known from the region (Figure 16). Specimens from Isla Clarion are rather small and uniform (Figure 18). The Colombian specimen (Figure 19) is rather narrow and high spired. Galapagan specimens (Figures 20-23) are rather uniformly narrow and moderately high spired. Remarks: The holotype of Conus xanthicus (USNM 111236, Figure 13), from 130m off Guaymas, Sonora, Mexico, was not originally figured. The taxon was appar- ently not discussed again until HANNA & STRONG (1949) figured the holotypes and placed the name in the synonymy of C. fergusont. The holotype of the synonymous C. chryso- cestus Berry remains in the private collection of S. Stillman Berry, Redlands, California (Figure 14). It is also from Sonora, Mexico: “trawled in 30 to 45 fms., off Morro Colo- ado, Sonora; Antonio Luna, Dec. 1965.” It is a large, brightly colored specimen which was not compared by Berry to C. xanthicus. He apparently did not question the long-held view of other authors concerning the validity of C. xanthicus. Conus kohni McLean & Nybakken, spec. nov. (Figures 4, 24-29) Diagnosis: Coronations present only at very early stage, spire whorls markedly concave, aperture relatively broad; color yellow-orange, with mottled, lighter colored band- ing; spire with radial markings of same intensity as that of rest of shell. Description: Shell medium-sized (maximum length 52.5 mm, with 9 teleoconch whorls) ; spire moderately elevated; spire outline slightly concave to straight; shoulder distinctly ridged, not coronated in mature specimens; spire whorls Explanation of Figures 24 to 29 Conus kohni McLean « Nybakken, spec. nov. Figure 24: LACM 1885, holotype, 18-37 m, Isla Isabela, Gala- pagos Islands, Ecuador. Radula verified specimen, length 35-3 mm, periostracum removed ventrally, intact on spire. Figure 25: AHF 788-38, 101m, coral & shell bottom, off Isla Daphne, Galapagos Islands, Ecuador. Length 52.3 mm, periostracum removed, lip broken back (largest specimen examined). Figure 26: AHF 788-38, same locality as Figure 25. Length 43.5 mim, periostracum intact. Figure 27: LACM 72-200, 40-45 m, coralline algal rubble, Bahia Academia, Isla Santa Cruz, Galapagos Islands, Ecuador. Radula verified specimen, length 40.0mm, periostracum removed. Figure 28: CAS 46379, Isla Santa Cruz, Galapagos Islands, Ecua- dor. Length 33.6 mm, periostracum removed. Figure 29: AHF 324-35, 82m, rock, Caleta Tagus, Isla Isabela, Galapagos Islands, Ecuador. Radula verified specimen, length 15.0mm, periostracum partially intact (smallest specimen examined). THE VELIGER, Vol. 22, No. 2 [McLean « NyBAKKEN] Figures 24 to 29 Figure 28 Figure 29 Vol. 22; No. 2 markedly concave, sculptured with faint growth lines; su- ture produced below the shoulder ridge, well defined but not deeply channeled; anal sinus of medium depth; outer lip thin, arcuate; body whorl profile with a convexity 1/8 the distance below the shoulder and a slight concavity at 1/3 the distance from the base; surface smooth except for 10-12 spiral lirae on the lower third of body whorl; aper- ture relatively wide, of same width throughout and con- forming to the body whorl profile. Protoconch homeo- strophic, 3-whorled, smooth, light colored; shoulder of first three teleoconch whorls weakly coronated, coronations lost by the 5th whorl. Color yellow to orange, with 2 more or less distinct spiral bands of white or less intense ground color, bands often narrow and bearing smaller white spots; the basal area may have some lighter mottling or less in- tense coloration; the shoulder bears another series of whit- ish spots that extend across the spire; ground color on the spire of equal intensity to that on the body whorl, aperture white within. Periostracum brownish, thin over body whorl, thicker on the spire and produced in closely set, thin, arched ridges. Operculum bluntly unguiculate, mar- gins unserrated, about 3 times longer than wide. Dimen- sions of holotype: length 35.3 mm, width 18.7 mm, length of operculum 6.5 mm. Radula (Figure 4): “Tooth with g barbs, one near the tip and 2 on the opposite side; the first 2 barbs sharply pointed, the 3rd more rounded; serration lacking; the narrowest part of the tooth posterior to the 3rd barb and marked by a more or less abrupt step or shelf; posterior to the waist the shaft diameter expands to a maximum and then con- Figure 4 C. kohni, from 35.3mm holotype specimen (shell: Figure 24) Radular tooth (scale = 0.1 mm) THE VELIGER Page 141 stricts slightly before the base; the base is large and bears a prominent spur. Radula preparations were made from 6 specimens, in- cluding those illustrated in Figures 24 (holotype), 27, and 29. No variation was noted. Type locality: 18-37m, mud-silt bottom, Caleta Tagus, Isla Isabela, Galapagos Islands, Ecuador (0°24.5’S, go° 23° W), 3 specimens collected by Gerald Wellington, 15 January 1975. Type material: Holotype, LACM 1885; 1 paratype, CAS 59690; 1 paratype, USNM 758900. Referred material (all Galapagos Islands, Ecuador): AHF 201-34, 46-64 m, Isla Espanola, 31 January 1934, 3 specimens; AHF 324-35, 82m, Isla Isabela, 10 December 1934, 1 specimen; AHF 788-38, 101m, Isla Daphne, 19 January 1938, 4 specimens; LACM 66-210, 34m, Isla San Salvador, 24 September 1966, 1 specimen; CAS 46379, 100m, Isla Santa Cruz, June 1969, 1 specimen; LACM 72-205, 40-45m, Isla Santa Cruz, 31 January 1972, 3 specimens. Distribution and occurrence: Galapagos Islands, Ecua- dor, on a variety of bottom types at depths of 18-100m. At stations AHF 324-35 and 788-38 it occurred with Conus xanthicus. It may eventually be shown to occur elsewhere in the tropical eastern Pacific, although this now seems un- likely, considering that the region has been well sampled. Etymology: We are pleased to name the species in honor of Dr. Alan J. Kohn of the University of Washington, who has contributed greatly to knowledge of the genus Conus over the past 20 years. Comparisons: Conus kohni is similar to C. xanthicus in size and color pattern but may be distinguished on radula and shell characters. The radula tooth of C. kohni (Figure 4) differs from that of both C. xanthicus (Figure 3) and C. fergusoni (Figures 1, 2) in completely lacking a serration and having 3 barbs. The chief shell difference is the broader aperture of C. kohni, which increases the rate of expansion and results in mature shells of similar length having one less whorl. Mature shells typically have 10 whorls in C. xanthicus and g whorls in C. kohni. Addition- ally, shells of C. kohni are broader, the lower portion of the body whorl is more concave, the lip is more arcuate, and the spire whorls are more concave. The ground color and the color patterns are remarkably similar, although the lighter bands of C. kohni tend to be narrower. Comparison of specimens with the periostracum removed shows the ground color on the spire to be darker than the ground color of the body whorl in C. xanthicus, whereas in C. Page 142 kohni the ground color on the spire and on the body whorl is identical. Variability: The 16 available specimens of C. kohni are uniform in proportions. Ground color varies from dark yellow to yellow-orange. Remarks: Although specimens of Conus kohni were first collected over 40 years ago, the species has escaped notice until now, having no doubt been dismissed along with C. xanthicus as representative of the juvenile stage of C. fer- gusont. Unlike C. xanthicus, however, misidentified speci- mens have not previously been illustrated. We were first alerted to its existence by discovery of an apparent dis- crepancy in radular characters among specimens thought to be C. xanthicus. Only after we had confirmed the pres- ence of two species on radular characters were we able to recognize the associated shell characters that separate the 2 species. Unfortunately we have not observed living examples of either Conus xanthicus or C. kohni, so we do not know if there are body color differences between the 2 species. Similarly we do not know if there are differences in habi- tat or bottom type preference. The bottom type data available for both species ranges from mud, sand, coral- line algal rubble, to rock, suggesting that both have a wide range of substrate occupancy. DISCUSSION Conus kohni is the only species of the genus at present considered endemic to the Galapagos Islands. Most mol- lusks known from the Galapagos Islands also occur in the tropical eastern Pacific, although there is a major endemic element and a small percentage of trans-Pacific migrants with Indo-Pacific faunal affinity. Two Indo-Pacific species of Conus are known from the Galapagos: Conus ebraeus Linnaeus, 1758, and C. chal- daeus (Réding, 1798). These two species have also been reported from the mainland (see references in EMERSON, 1978). Conus tessulatus Born, 1778 is an Indo-Pacific spe- cies known from the Revillagigedo Islands and the main- land (EMERSON, 1978). It is possible that C. kohni is in this category, but if this is so, it remains to be discovered in the Indo-Pacific. If Conus kohni is an eastern Pacific species, it may yet be found to occur elsewhere in the Panamic province, or it could represent a relict population surviving at the Gala- pagos Islands and extinct on the west American mainland. Few of the eastern Pacific Conidae, especially those that THE VELIGER Vol. 22; No. 2 occur offshore such as C. xanthicus, are known from the fossil record. We have no data as to the food of Conus fergusoni, C. xanthicus, or C. kohni, but would anticipate that prey items would be different in all 3, based on major differ- ences in tooth morphology. Conus xanthicus has a radula tooth morphology that is the most common in the genus, a type that NYBAKKEN (1970) termed “regularis.” At least nine other Panamic species have a similar tooth (NYBAK- KEN, Op. cit.). The food of these species, where known, consists of errant polychaetes (NYBAKKEN, 1979). Conus fergusont has a radula tooth similar to that of C. princeps Linnaeus, 1758, and C. patricius Hinds, 1843. Again, er- rant polychaetes should be the preferred food for species with a similar tooth (Nybakken, in press). The tooth of C. kohni is of an uncommon type for which no food data are available for any species with a similar morphology. It is most similar to that of the eastern Pacific species C. re- curvus Broderip, 1833, and C. arcuatus Broderip & Sow- erby, 1829. The similarity between the tooth of Conus kohni and C. recurvus leads us to compare the shell morphology of the 2: proportions are similar; both have the concave shoulder and a markedly arcuate lip, but a very different color pattern. If other evidence can be found to support the theory that C.kohni and C. recurvus are closely related, it will imply that the similarity of color pattern between C. xanthicus and C. kohni is coincidental and convergent and that tooth structure is a conservative character. An alternative possibility is that the similar color pattern of C. kohni and C. xanthicus is the conservative feature and that the tooth structure has diverged in response to changes in diet. Small specimens of C. fergusoni evidently have a differ- ent radula tooth morphology from that of large specimens. The tooth of a 26.0mm long specimen is shown in Figure 1. The next smallest specimen from which we have ob- tained a radula is 43.5 mm in length. Its tooth, as in others from shells of larger sizes we have prepared (shell lengths, 85.1, 61.0, 60.7, and 51.9 mm) is similar to that of Figure 2. The tooth of the smallest specimen is proportionately shorter, lacks the second barb and has the first barb very weakly developed. The fact that the serration is the same, running half the length of the tooth, enables us to consider the transition to be reasonable, in the absence of inter- mediate-sized examples. This is the second known species of Conus in which a change in radular tooth morphology has been demon- strated. NYBAKKEN (1970: 13) found that the radula tooth of a specimen of C. patricius 27.1 mm in length was Vol. 22; No. 2 THE VELIGER Page 143 proportionately shorter than that of the adult and entirely lacked the blade, barb, and serration. That, however, probably represented an earlier stage of tooth development than in our example with C. fergusoni. NyBAKKEN (1970) found that other Conus species examined did not show any significant change in tooth morphology between young and mature specimens. Both species reach large sizes — changes in tooth morphology with growth may prove to be a fea- ture of only those species that attain the largest sizes. In addition to large size and similar tooth morphology, both Conus fergusoni and C. patricius have coronations in immature stages that are not formed in fully mature stages. The subgenus Pyruconus Olsson, 1967, was pro- posed for C. patricius (type species) and fossil species with a similar pyriform shape (OLsson, 1967: 21). Although C. ferguson: is not similarly pyriform, it is evidently re- lated and may also be tentatively assigned to the subgenus. However it is premature to attempt assignment of the other species treated here. CONCLUSIONS Two of the 3 possible explanations originally suggested by NYBAKKEN (1970) to account for different radula tooth morphologies in the then understood Conus fergusoni have proven to be the case: 2 species were being confused, and tooth morphology in C. ferguson: does change with growth. Change of tooth morphology with growth is evidently un- usual in the genus and its occurrence in other species should be further documented. Recognition of Conus xanthicus has long been delayed because of confusion with immature specimens of C. fer- gusoni, due no doubt to the rarity of the earliest stages of C. fergusoni — for reasons not apparent. Previous authors had not realized that coronated early whorls are a feature of C. fergusoni and that small specimens have spiral rows of dark spots. Conus fergusoni is not highly variable, whereas C. xanthicus is unusually variable in elevation of the spire. Conus xanthicus and the herein described C. kohni have been confused because of a nearly identical color pattern. The two may be separated on radula and shell characters. It remains to be demonstrated whether they are closely re- lated or are more nearly related to other species having corresponding tooth morphologies. Despite a spate of recent papers on the systematics of Eastern Pacific Conus, none of these papers contain rig- orous comparative diagnoses of the shells, nor do we have information about the color of living animals or food habits for many of the species. Progress has been made with documentation of tooth morphologies. Future system- atic work on the genus should include a discussion of radula characters. NOTE ADDED IN PROOF After this paper was submitted, we received “Cone Shells: A Synopsis of the Living Conidae,” by Jerry G. Walls, ro11 pages, T. F.H. Publications, Inc., Neptune, N.J., published March 6, 1979. Our conclusions about the validity of Conus xanthicus, the synonymy of C. chrysocestus, and the spotted juveniles of C. fergusont were also reached by Walls. However, he included C. fulvocinctus Crosse, 1873, in the synonymy of C. fergusoni, an opinion not shared by us. The locality for C. fulvocinctus was originally given as West Africa, and the 75mm long specimen was said to have a thin periostracum. The original figure shows strong spiral cords extending from the base to the mid-whorl, spire whorls that are convex rather than concave, and no trace of coronations in early stages. We agree, however, that C. consanguineus E. A. Smith, 1880, locality un- known, is a probable junior synonym of C. fergusont. Literature Cited Berry, SAMUEL STILLMAN 1968. Notices of new eastern Pacific Mollusca. VII. Leafl. in Malacology 1 (25): 156-157 Dai, Wituiam Hearey 1910. Summary of the shells of the genus Conus from the Pacific coast of America in the United States National Museum. Proc. U. S. Natl. Mus. 38 (1741): 217-228 (6 June 1910) Emerson, WILLIAM KEITH 1978. Mollusks with Indo-Pacific faunal affinities in the eastern Pa- cific Ocean. The Nautilus 92 (2): 91 - 96 Emerson, Witi1aM Keitu & Witt1AM E_woop OLp, Jr. 1962. Results of the Puritan-American Museum of Natural History expedition to western Mexico, 16. The recent mollusks: Gastropoda, Conidae. Amer. Mus. Novitates no. 2112: 44 pp.; 20 figs. Hanna, G Datras 1963. West American mollusks of the genus Conus, IT. Occ. Papers Gal. Acad. Sci., no 35: 103 pp.; 11 plts.; 4 text figs. Hawna, G Darras & ArcHIBALD McCiure STRONG 1949. West American mollusks of the genus Conus. Proc. Cal. Acad. Sci. (4) 26: 247-322; plts. 5-10; 4 text figs. Kesen, A. Myra 1958. Sea shells of tropical west America; marine mollusks from Lower California to Colombia. Stanford Univ. Press, Stanford, Calif. i-xi + 624 pp.; 1700 text figs.; 10 colored pits. (5 Dec. 1958) 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru, 20d ed. Stanford Univ. Press, Stanford, Calif. i-xiv+ 1064 pp.; ca. 4000 text figs.; 22 col. plts. (21 September 1971) NYBAKKEN, JAMES WILLARD 1970. Radular anatomy and systematics of the west American Conidae (Mollusca, Gastropoda). Amer. Mus. Novitates no. 2414: 29 PP-; 45 figs. : a 1971. The Conidae of the Pillsbury Expedition to the Gulf of Panama. Bull. Mar. Sci. 21 (1): 93-110; figs. 1-26 Dace 1979. Population characteristics and food resource utilization of Conus in the Galapagos Islands. Pacif. Sci. 32 (3): 271-280 in press Population characteristics and food resource utilization of Conus in the Sea of Cortez and West Mexico. Journ. Mollusc. Stud. Page 144 THE VELIGER Vol. 22; No. 2 Ousson, Axer ADOLF 1967. Some Tertiary mollusks from South Florida and the Caribbean. Paleontol. Res. Inst., Ithaca, N. Y. 61 pp.; 9 pits. SowersBy, GzorcE BRETTINGHAM (3!) 1873. Descriptions of five new cones. Proc. Zool. Soc. London 1873: 145-146; plt. 15 1887. Thesaurus conchyliorum, or monographs of genera of shells. London, vol. 5: Conus suppl., 249 - 279; pits. 29-36 (507 - 512) Wo rae Vol. 22; No. 2 THE VELIGER Page 145 The Family Lepidopleuridae (Mollusca : Polyplacophora ) in the Eastern Pacific BY ANTONIO J. FERREIRA ' Research Associate, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118 (4 Plates; 8 Text figures) INTRODUCTION THE Taxonomy of the eastern Pacific lepidopleurids has long remained uncertain. These most primitive of Recent chitons are usually small in size, uniform in color, fragile, and scarce in numbers. They do not lend themselves easily to study given the simplicity of their anatomical features. In museum collections they are represented by few and small lots, which makes it difficult to draw firm conclusions about intraspecific variations and interspecific distinctions. The family Lepidopleuridae has never been reviewed. In the eastern Pacific, chiton workers have often suspected that far too many nominal species (Dall alone named 13) of lepidopleurids had been described. Insufficiently character- ized, unfigured, known only from the type material (often limited to 1 or 2 specimens), many such nominal species of eastern Pacific lepidopleurids have crowded the chiton no- menclature in the long wait for a taxonomic decision about their validity. It is the purpose of this paper to review the lepidopleurids of the eastern Pacific, and, following the study-plan of the late ALLYN G. Smiru (1977), to contribute to the rectifica- tion of the west coast chiton nomenclature. The present work draws mostly upon material in the col- lections of the California Academy of Sciences (CAS), Stan- ford University (SU) [at CAS], Natural History Museum of Los Angeles County (LACM), San Diego Museum of Natural History (SDNH), and Academy of Natural Sciences ' Mailing address for reprints: 2060 Clarmar Way, San Jose, CA (lifornia) 95128, U.S. A. of Philadelphia (ANSP). However, particular emphasis must be given to the contributions of Dr. Joseph Rosewater who graciously permitted me to examine and study critical type material in the repository of the National Museum of Natural History (USNM), and of Piet Kaas (The Nether- lands), Richard Van Belle (Belgium), and Dr. B. Sirenko (Leningrad, U.S.S.R.) who generously provided me with valuable specimens and data. This review of the family Lepidopleuridae is limited to the area known as eastern Pacific, which extends from the Bering Strait (65.5° N) to the northern tip of Chiloé Island, Chile (44°S), through the north cold temperate, north warm temperate, tropical, and south warm temperate re- gions of the west coast of the Americas. It excludes the poorly known (to the author) south cold temperate region (Magellanic Province) which starts at Chiloé Island. From this investigation 7 species are recognized as components of the lepidopleurid fauna of the eastern Pacific: Leptochiton rugatus (Pilsbry, 1892) Leptochiton nexus Carpenter, 1864 Leptochiton alveolus (Lovén, 1846) Leptochiton incongruus (Dall, 1908) Leptochiton albemarlensis Smith & Ferreira, 1977 Hanleyella oldroydi (Dall, 1919) Oldroydia percrassa (Dall, 1894) The following descriptions and synonymies of species are not intended to be exhaustive. Repetition of what is already easily available in the literature is avoided in favor of ex- tending the descriptions and differential diagnoses of the species involved. Page 146 SYSTEMATIC TREATMENT POLY PLACOPHORA de Blainville, 1816 Neoloricata Bergenhayn, 1955 LEPIDOPLEURINA Thiele, 1910 LEPMoPLEURDAE Pilsbry, 1892 Definition: ‘Relatively small [less than 5 cm long]; valves generally sculptured with granules set in lines or quincunx; valve coverage complete; girdle narrow, mi- nutely spiculose or scaly [or both]; insertion plates lacking, or if present, weak and unslit; articulamentum layer usually consisting only of weakly developed sutural laminae in valves ii to viii. Gills short, posterior.” (SmiTH, 196ob: 52). Type-genus: Lepidopleurus Risso, 1826 [not DALL, 1879] (Type species: Chiton cajetanus Poli, 1791, by SD, HERRMANNSEN, 1847). Leptochiton Gray, 1847 Definition: Valves rounded and thin. Tegmental surface with granules in lines or quincunx. Lateral areas low and inconspicuous. Girdle narrow, with minute scales or scale- like processes, often with interspersed small spines. Type Species: Chiton cinereus Montagu, 1803 (not Chi- ton cinereus Linnaeus, 1767) [== Chiton asellus Gmelin, 1791] by SD, Gray (1847, Nov.). Synonyms: Terenochiton Iredale, 1914b [ type species, Lepidopleurus (Terenochiton) subtropicalis Iredale, 1914, by OD]; Xtphi- ozona Berry (1919a) [type species, Lepidopleurus (Xiphi- ozona) heathi BERRY, 1919, by OD]. Remarks: Gray (1847a) defined Leptochiton on the ba- sis of the “valves rounded and thin” which clearly distin- guished its type species, Chiton cinereus Montagu, 1803, from Chiton cajetanus Poli, type species of Lepidopleurus Risso, 1826. By citing C. cinereus together with C. caje- tanus Gray introduced an element of uncertainty as to the validity of Leptochiton; however, this uncertainty was dis- pelled when Gray (1847b) listed C. cinereus as type of Leptochiton Gray, and C. cajetanus as type of Lepido- pleurus Risso. Dati (1879b) accepted Leptochiton, but Pitssry (1892) placed it in thesynonymy of Le pido pleurus. IREDALE, having at first recognized Leptochiton (19144), erected a THE VELIGER Vol. 22; No. 2 new genus, Terenochiton, for “the small Australian ‘Lepi- dopleurus’” which have “nothing in common save the ab- sence of insertion plates” with the type of Lepidopleurus (IREDALE, 1914b: 29). Berry (1917) rehabilitated Gray’s Leptochiton pointing out that “the relatively delicate and smooth shelled Lepidopleuridae of the west American coast seem only diffidently congeneric with L. cajetanus Poli, the thick rugose shell selected by Pilsbry as Risso’s type.” And although introducing Xiphiozona (19192), Berry decided “to retain” Leptochiton (1919b: 6, footnote) as the proper generic name to be used for most west Amer- ican lepidopleurids. ALLyYN G. SmrtH at first (1960a) considered Leptochiton as a subgenus of Lepidopleurus; but in the Treatise (1960b) SmitH regarded Leptochiton as a synonym of Lepido- pleurus Risso. However, in an unpublished revision of the chiton classification, on fileat CASG, Smith had reaccepted Leptochiton and conferred it generic rank. In the most re- cent classification of Polyplacophora, VAN BELLE (1975) also accepted Leptochiton, although treating it as a sub- genus of Lepidopleurus. Terenochiton Iredale, 1914, was placed in the synonymy of Leptochiton by Smrru (1960b), and VAN BELLE (1975). The examination of 66 specimens of Terenochiton inquina- tus (Reeve, 1847), collected in 9 localities on the east coast of North Island, New Zealand (AJF 382-385, 387-391) demonstrated that in all respects of shell, girdle, and radula, the species belongs quite properly in the genus Leptochiton. Leptochiton Gray is here given full generic rank in the recognition of Gray’s original description, and the uniform- ity of characteristics typified in Leptochiton asellus (Gmelin, 1791). Leptochiton rugatus (Pilsbry, 1892) (Figures 1, 2, 7, 33, and 34) Leptochiton internexus rugatus DALt, 1879b: 319 (nomen nudum) Lepidopleurus rugatus PILsBrY, 1892: 11-12; plt. 3, figs. 67-70. 1898: 287 — BERRY, 1907: 47 — CHACE, 1917: 44 - DALL, 1921: 186 (in section Leptochiton) — OLpRoyp, 1927: 849 (in section Leptochiton) — BuRcH, 1942: 7 — CHACE, 1958: 330 — ABBOTT, 1974: 393 Leptochiton rugatus (Pilsbry). THIELE, 1909: 12-13; plt. 1, figs. 41-50 — SMITH, 19474: 6. 1947b: 17 — SMITH & Gor- DON, 1948: 205 — PALMER, 1958: 263; plt. 28, fig. 7; plt. 35, fig. 3 — Linpsay, 1966: 348 — BuRGHARDT & Burc- HARDT, 1969: 26; plt. 2, fig. 35 — THORPE in KEEN, 1971: 863; Polyplacophora, fig. 1 — A. G. SMITH in R. I. SMITH & CARLSON, eds. (Light’s Manual, 3rd ed.), 1975: 262, 265. Leptochiton internexus DALL, 187gb: 319 (nomen nudum) Vol. 22; No. 2 Lepidopleurus internexus CARPENTER in PILSBRY, 1892: 12 — DALL, 1921: 186 (in section Leptochiton) - OLDROYD, 1927: 849 (in section Leptochiton) — La RocQuéE, 1953: 8 — BURGHARDT & BURGHARDT, 1969: 24; plt. 2, fig. 34 - ABBOTT, 1974: 393 Leptochiton internexus (Carpenter in Pilsbry). SMITH 1947a: 4. 1947b: 17 — PALMER, 1958: 262, plt. 28, figs. 3-6 Leptochiton cancellatus (Sowerby). DALL, 1879b: 315-317. 1886: 211 — BERRY, 1917: 232-233 ~ WILLETT, 1919: 27 — Berry, 1927: 160 — SMITH, 1947a: 4. 1947b: 17 - SMITH & GoRDON, 1948: 205 — BERRY, 1951: 218. [not Chiton cancellatus Sowerby, 1839] Lepidopleurus cancellatus (Sowerby). PiLssry, 1892: 3-5 (in part) — DALL, 1921: 186 — OLDROYD, 1924: 184. 1927: 848-849 — LELoupP, 1940: 6-7; figs. 10-11, 16-18 — Burcu, 1942: 7 — BURCH & BURCH, 1943: 5 — JAKOVLEVA, 1952: 46-47; fig. 14; plt. 1, fig. 2 (in part) — La RocQuE, 1953: 7 (in part) — TakI, 1964: 407 — ABBOTT, 1974: 392 (in part). [not Chiton cancellatus Sowerby, 1839] Lepidopleurus assimilis THIELE, 1909: 11-12, plt. 1, figs. 30-39 — TAK! & TAKI, 1929: 162 — TAKI, 1938: 328-331; plt. 14, fig. 2; plt. 16, figs. 5, 9-13, 15; plt. 17, figs. g-11 — JAKOVLEVA, 1952: 48-49; fig. 16; plt. 1, fig. 4 - Tamu, 1962: 31 — Taki, 1964: 408 Lepidopleurus alascensis THIELE, 1909: 13; plt. 1, figs. 51-60 - DALL, 1921: 187 — (in section Leptochiton) — As- BOTT, 1974: 393 Leptochiton alascensis (Thiele). SMITH, 1947a: 3. 1947b: 17 Type Material: Leptochiton rugatus (Pilsbry, 1892): Syntypes (ANSP 35586) [““two whole specimens plus completely disarticulated valves without girdle,” M. Miller, in litt., 27 March 1978]. Lepidopleurus internexus Carpenter in Pilsbry, 1892: Hol- otype (USNM 30750); two accompanying labels give locality as ‘Sta. Barbara,” California. Lepidopleurus assimilis Thiele, 1909: Types at the Zoolo- gisches Museum, Humboldt Universitat, Berlin [“r set of plates (a little dilapidated), locality Sachalin (it must be the holotype), and 1 dried specimen, locality Wladiwostok (marked as a type)” Dr. R. Kilias, in litt., 6 September 1978]. Lepidopleurus alascensis Thiele, 1909: Holotype at Zoolo- gisches Museum, Humboldt Universitat, Berlin [“(in alco- hol), locality Alaska,” Dr. R. Kilias, in litt., 6 September 1978]. Type Locality: ‘near S. Tomas River, Lower Califor- nia,” Mexico [31°32’N; 116°26’ W], as indicated in the label accompanying the syntypes [M. Miller, in litt., 27 March 1978]. Description: The original description of Pirssry (1892), supplemented in THIELE (1909), is very adequate to iden- tify the species. Color, a uniform creamy white; some speci- mens may be orange color as described for “internexus” by CaRPENTER in Pitspry (1892), and often seen in “assimilis” from Japan; and some may be blackened by fuliginous ma- terial. In size, specimens from Alaska tend to be appreciably smaller than those from the southern part of the range; THE VELIGER Page 147 largest specimen examined measures 15.8 mm in length, in- cluding girdle (LACM 62-8: “Intertidal, Franklin Point, San Mateo County, California, leg. J. H. McLean, June 1962-November, 1963’’). Based on 10 adult specimens, the width/length ratio is 0.53. Specimens of Leptochiton ruga- tus are somewhat variable in height; however, a constant diagnostic feature is the definitely rounded, i.e., not cari- nated, back (Figure 1). Tegmental sculpture consists of microgranules linearly juxtaposed to form subgranose rib- lets (Figure 2). These riblets are longitudinally disposed in the central areas (about 20-40 riblets per side), somewhat radially oriented in the end valves and lateral areas (about 10-15 per area). The strength of these sculptural features is quite variable, as is the degree of crowding of the riblets and their microgranulose appearance; the space between the riblets may appear, at first glance, pitted or cancellate. Figure 33 Leptochiton rugatus (Pilsbry, 1892) Girdle scales and spines of (A) the upper surface, and (B) the under surface. Specimen collected at Monterey Bay, California at a depth of 8m (AJF 50) Mucro, central to subcentral, often a bit pointed and ele- vated; postmucro, quite variable, convex to concave, often dropping abruptly. Lateral areas of intermediate valves usually inconspicuous; in many specimens they show con- Page 148 THE VELIGER Vol. 22; No. 2 centric, more or less coarse and irregular growth lines. Girdle, pebbly to gravelly in appearance, with no conspic- uous spicules or spines; in some specimens there may be a definitely spiculous appearance at the sutures. Girdle scales (Figure 33), mostly conical, irregularly striated, small (about 60-80 pm in height, 20-40 zm in diameter of base), close together but not imbricated. An occasional scale is elon- gated to the point it could be called a spinule, a feature more often seen at the sutures where, in some specimens, they form small ill-defined tufts. Girdle undersurface cov- ered with small triangular scales, about 40 wm x 20 pm, imbricated, pointing outward. Girdle scales display con- siderable degree of polymorphism in size and shape, not seemingly related to locality or depth. The radula of Leptochiton rugatus was first described by THIELE (1909). Consistently, in the specimens examined, it comprised over 100 rows (170 in 1 specimen) of mature teeth. A specimen (AJF 50: Monterey Bay, California, at 8m depth), measuring 10.3 mm in length, has a radula 3.9 mm long (i.e., 37% of the specimen’s length), with 146 rows of mature teeth. The central tooth (Figure 34) has a rela- tively narrow front end, about 18 jum in width, with a small 50 um Figure 34 Leptochiton rugatus (Pilsbry, 1892) Radula: Median, first lateral and head of second lateral tooth. Specimen collected at Monterey Bay, California at a depth of 8m (AJF 50) blade. F irst lateral teeth, aliform, wider in the front. Major lateral teeth with a very long (go um), unicuspid head. Outer marginal teeth about 22 uum in length, and 44 um in width; length/width ratio, 0.5. The examination of the radula of several specimens from different localities, from Alaska to Baja California and the Gulf of California, dis- closed no appreciable variation or discrepancy in these essential characteristics. A specimen of “asstmilis” from Vostock Bay, Sea of Japan (collected by B. Sirenko, July 26, 1971, at a depth of 5 m), generously donated by Dr. B. Sirenko, Zoological Institute, Academy of Sciences, Lenin- grad, U.S.S.R., measuring 5.2mm in length, has a radula 3.3mm long (53% of shell length) with about 170 rows of mature teeth; the median tooth is narrow in front where it measures 13 jum in width; major lateral teeth have very long (45 m), unicuspid heads; outer marginal teeth are 20 um in length, 38 4m in width (length/width ratio, 0.52). These observations, in the absence of significant concho- logical distinctions, are to be regarded as well within the expected variation in L. rugatus, particularly when differ- ences in geographical sites are considered. Distribution: Leptochiton rugatus has a seemingly con- tinuous distribution from Alaska to Magdalena Bay, Baja California, Mexico. The northernmost verified record is Cohen Island, Kachemak Bay, Kenai Peninsula, Alaska (§9°32.5 N; 151°28.5 W), leg. J. H. McLean & R. Baxter, July 31, 1973 (LACM 73-20); the southernmost record is at Magdalena Bay, “about 1 mile northeast of Pta. Belcher,” Baja California, Mexico (24°35’N; 112°05’ W), leg. J. H. McLean & Oringer, Dwyer Expedition, December 14, 1967 (LACM 67-70). Leptochiton rugatus has been collected in the Aleutian Islands (CASG 39538), and in the offshore islands: Heskett (LACM 73-23), Kodiak (LACM 75-32), Queen Charlotte (LACM 66-46; LACM 70-95), Van- couver (LACM A.4052; LACM-AHF 1151-40; LACM 63-62; LACM 66-36; LACM 73-40), Orcas (LACM M- 541; LACM M-544), Farallon (LACM 62-9), Anacapa (LACM-AHF 874-38), Santa Rosa (LACM-AHF 1385-41; LACM 73-10), Santa Cruz(LACM-AHF 1192-40; LACM- Explanation of Figures 1 to 8 Figure 1: Leptochiton rugatus (Pilsbry, 1892). 12.3 mm long. Mon- terey, California (LACM 59-12) Figure 2: Close-up of specimen in Figure 1 to show detail of central and lateral areas Figure 3: Lepidopleurus nexus Carpenter, 1864. Holotype (US NM 16270) Figure 4: Close-up of specimen in Figure 3 to show detail of central and lateral areas Figure 5: Lepidopleurus ambustus Dall, 1919. Paralectotype (here- in. designated) (USNM 274120) Figure 6: Lepidopleurus ambustus Dall, 1919. Lectotype (herein designated): Close-up of the lectotype to show detail of central area, lateral areas, and girdle (USNM 274120) Figure 7: Lepidopleurus internexus Carpenter in Pilsbry, 1892. Holotype (USNM 30750) Figure 8: Lepidopleurus lycurgus Dall, 1919. Holotype (USNM 274119) [FeRREIRA] Figures 7 to 8 Tue VELIGER, Vol. 22, No. 2 Vol. 22; No. 2 AHF 1196-40), Catalina (LACM-AHF 1149-40; LACM- AHF 1151-40; LACM-AHF 1323-41; LACM-AHF 1430- 41; LACM 65-6), and Guadalupe (LACM-AHF 1924-49). In what may be an isolated population, L. rugatus is also present throughout most of the Gulf of California, from “Puertecitos to Bahia de los Angeles, Baja California, and in the Guaymas region of Sonora, Mexico” (THORPE in KEEN, 1971), as verified by specimens collected at Puerte- citos (LACM 62-19), Bahia de San Francisquito (AJF 36), and San Augustine, Sonora (AJF, August, 1972). Leptochiton rugatus, identified as “assemilis” and “can- cellatus,” has been reported off the coasts of U.S.S.R., Sea of Japan, Sea of Okhotsk, and the Bering Sea. Bathymetrically, Leptochiton rugatus ranges from the intertidal zone to 458m (LACM 1058, off Catalina Island, California). Median depth, based on the lots examined, 14m. As reflected in the literature and museum la- bels, many serious students of chitons have held the view that the “rwgatus’’-like specimens of the eastern Pacific rep- resent two or more distinct species. However, the observa- tions made in the course of this study indicate that, at least on conchological grounds, all specimens examined should be regarded as conspecific. Radula studies did not contra- dict this first impression. But the question remained whether the eastern Pacific species, Leptochiton rugatus (Pilsbry, 1892) was conspecific with L. cancellatus (Sowerby, 1839) from the North Atlan- tic. From the original descriptions of L. rugatus and L. can- cellatus it seemed impossible to differentiate the two popu- lations, Pacific and Atlantic, with certainty. Fortunately, thanks to the generous assistance of Richard A. Van Belle, Sint-Niklaas, Belgium, who provided several specimens of L. cancellatus collected at Trébeurden, Bretagne, France, and at Kostenauman, Sweden, it was possible to compare directly eastern Pacific with Atlantic specimens. Side by side, eastern Pacific specimens of L. rugatus ap- peared to be conchologically indistinguishable from the Atlantic L. cancellatus in size, color, shape, tegmental sculp- ture and articulamentum; and although the Atlantic speci- mens seemed to have a slightly higher and rounder back, and a lesser tendency to have spinule-like girdle elements than their Pacific counterparts, the differences were trivial and short of specific significance. There were, however, appreciable differences in the radulae. Comparing the radulae of 10 specimens of Le pto- chiton rugatus from different localities in the eastern Pacific with those of 3 specimens of the Atlantic L. cancellatus it was observed that the radula of the eastern Pacific speci- mens consistently had (1) considerably more rows of Remarks: THE VELIGER Page 149 mature teeth (> 100 rows in L. rugatus vs. 50-70 rows in L. cancellatus), (2) a differently shaped median tooth (anterior end much narrower than posterior end in L. ruga- tus vs. anterior end as wide or wider than posterior end in L. cancellatus), (3) major lateral teeth with a distinctly different head (unicuspid, very long and slender in L. ruga- tus vs. bicuspid, relatively short and thick in L. cancellatus), and (4) outer marginal teeth much shorter than wide (length/width ratio about 0.5) in L. rugatus but longer than wide (length/width ratio 1.2 to 1.6) in L. cancellatus. In view of these marked radular differences, it is con- cluded that the 2 populations, Atlantic and Pacific, de- spite their very great conchological similarities, repre- sent 2 different species, with the eastern Pacific species taking the name proposed by Pivssry (1892), L. rugatus. Leptochiton cancellatus (Sowerby, 1839) is not present in the Pacific collections examined. The synonymies of Leptochiton assimilis Thiele, 1909, and L. alascensis Thiele, 1909 are based on the original de- scription and figuresof thespecies; the examination of sev- eral specimens of L. assimilis collected at the Okhotsk Sea and Sea of Japan, and generously donated by Drs. Golikov and B. Sirenko, Zoological Institute, Academy of Sci- ences, Leningrad, U.S.S.R., demonstrated their con- specificity with L. rugatus. Leptochiton internexus (Carpenter in Pilsbry, 1892) is here regarded as a synonym of L. rugatus, on the basis of the original description as well as on the direct examination of the holotype (USNM 30750), available through the cour- tesy of Dr. Joseph Rosewater. The single type specimen (Figure 7), dry, hasa light brown color; valves i, vii, and viii are disarticulated. In every respect, it corresponds to L. rugatus. Color slides of the holotype are deposited in the California Academy of Sciences Invertebrate Zoology (CASIZ), Color Slides Series. The conspecificity of L. inter- nexus was further confirmed through the study of a lot of 40 specimens so labelled probably on account of its locality and beautiful orange-gold color of the shells (LACM 63-69, 8km SE of Point Conception, Santa Barbara County, Cali- fornia, at 13-17m, under kelp, leg. J. H. McLean, July 18- 19, 1963). There were no significant differences in shell, girdle, and radula, between these specimens and those of other populations of L. rugatus. Leptochiton nexus Carpenter, 1864 (Figures 3 to 6, 8, 35, and 36) Leptochiton nexus CARPENTER, 1864: 612 (nomen nudum), 650 [Reprinted, 1872: 98, 136]. 1866: 212-213. CoopEr, 1867: 23 — DALL, 1879b: 319 — LOWE, 1904: 19 - SMITH, 19474: 6. 1947b: 17 — SMITH & GORDON, 1948: Page 150 THE VELIGER Vol. 22; No. 2 205 — PALMER, 1958: 262; plt. 28, fig. 2 — BURGHARDT & BuRGHARDT, 1969: 25 Lepidopleurus nexus (Carpenter). Pitspry, 1892, 14: 11. 1898: 287 — Oxproyp, 1911: 74 — Dati, 1921: 187 (in section Xiphiozona) — OLDROYD, 1927: 853 (in section Xiphiozona) — WILLETT, 1935: 42 (with syn. L. heathi Berry, 1919, and L. ambustus Dall, 1919 — BURCH, 1942: 7 — BURCH & BURCH, 1943: 5 — ABBOTT, 1974: 393 Chiton (Leptochiton) nexus (Carpenter). ORCUTT, 1885: 544 Lepidopleurus ( Xtphiozona) heathi BERRY, 1919a: 5 — HANSELMAN, 1977: 62 Lepidopleurus heathi (Berry). DALL, 1921: 187 (in section Xiphiozona) — OLpRoYD, 1927: 853 (in section Xiphi- ozona) — WILLETT, 1935: 42 (syn. of L. nexus) Leptochiton (Xtphiozona) heathi (Berry). BERRY, 1919b: 6-8; pit. 2, figs. 1-2; plt. 2 Leptochiton heathi (Berry). SMITH, 19474: 4. 1947b: 17 - SMITH & GORDON, 1948: 205 — BURGHARDT & BURGHARDT, 1969: plt. 2, fig. 33 (with syn. L. ambustus Dall, 1919) Lepidopleurus ambustus BERRY, 1907: 47 (nomen nudum) Lepidopleurus (Leptochiton) ambustus DALL, 1919: 499-500 Lepidopleurus ambustus Dall. DALL, 1921: 187 (in section Leptochiton) — OLpRoYD, 1927: 849-850 (in section Lep- tochiton) — WILLETT, 1935: 42 (syn. of L. nexus) Lepidopleurus (Pilsbryella) ambustus Dall. LELoup, 1940: 4-6, figs. 1-7 Leptochiton ambustus (Dall). SMITH, 19474: 3-4. 1947b: 17 - SMITH & GORDON, 1948: 205 — BURGHARDT & BURG- HARDT, 1969: 24 (syn. of L. heatht). Lepidopleurus (Leptochiton) lycurgus DALL, 1919: 500 Lepidopleurus lycurgus Dall. Dat, 1921: 187 (in section Leptochiton) — OLpRoyp, 1927: 850 (in section Lepto- chiton) — ABBOTT, 1974: 393 Leptochiton lycurgus (Dall). SMITH, 19474: 4. 1947b: 17 - BURGHARDT & BURGHARDT, 1969: 25 Type Material: Leptochiton nexus Carpenter, 1864. Holotype (USNM 16270) Leptochiton (Xiphiozona) heathi Berry, 1919.Holotype (S.S. Berry colln., Cat. No. 3513), “off Monterey, California.” Not examined. Lepidopleurus ambustus Dall, 1919: Lectotype [herein] and 2 paralectotypes (USNM 274120) ; accompanying label gives locality as “Sta Barbara Chan./Sta. Barbara Ids.,” California. Lepidopleurus (Leptochiton) lycurgus Dall, 1919: Holotype (USNM 274119); accompanying label reads “Catalina Id.” Type Locality: Catalina Island, Los Angeles County, California [33°23’N; 118°24’ WI. Description: Carpenter’s original description in the “Guide to the Diagnosis of the Vancouver and California Shells” (1864: 650) is far from adequate: “236. Leptochi- ton nexus, n.s. Like asellus: scarcely sculptured: mantle- margin with striated chaffy scales, like Magdalenensis, in- terspersed with transparent needles. 20-80fm. Cp.” The species is not figured; in a table on the same p.650, it is indi- cated that the specimen(s) was collected in the islands of the Sta. Barbara Group, by Cooper. Seemingly quoting from Carpenter’s unpublished manu- script, Prtspry (1892, 14: 11) enlarged upon this descrip- tion: “Shell small, whitish-ashen, valves gothic-arched; lat- eral areas scarcely defined; entire surface ornamented with series of subquadrate granules, the series longitudinal upon the central, radiating upon the lateral areas and end valves, very close, scarcely interrupted. Jugum elevated, subacute; umbones inconspicuous. Mucro conspicuous, median. In- side with strong sutural lobes and a wide plane sinus to the middle valves; insertion plates obsolete. Girdle having a narrow, Close, striated scales and needle-shaped, crystalline bristles here and there and around the margin. (Cpr.)” To which Pilsbry added, later on the page: “A variety is pret- tily variegated with olive, has stronger sculpture, and the valves slightly beaked. The spicules which appear on the girdle among the striated, elongated scales, are very nu- merous. It is curiously like the young of Ischnochiton radi- ans. (Cpr.).” The species was left unfigured until PALMER (1958, plt. 28, fig. 2). Figure 35 Leptochiton nexus Carpenter, 1864 Girdle scales and spines. Specimen collected at Hyperion stack, Los Angeles, California, at a depth of 62m (LACM-AHF 5714-38) On January 6, 1919, Berry described and figured Lepi- dopleurus (Xiphiozona) heathi [Type locality: Monterey Bay, California] ; five months later, on June 7, 1919, DALL published his description of Lepidopleurus (Leptochiton) ambustus [Type locality: Santa Barbara Channel, Califor- Vol. 22; No. 2 THE VELIGER Page 15! nia]. Although WILLETT (1935) regarded these two nom- inal species as junior synonyms of Leptochiton nexus, the taxonomic position of these species has remained uncertain. The problem was finally solved when, thanks to the un- failing generosity of Dr. Joseph Rosewater, it became pos- sible to examine the holotype of Carpenter’s Lepidopleurus nexus (USNM 16270) and verify its conspecificity with Leptochiton heathi (Berry, 1919) and its already recognized synonym, Leptochiton ambustus (Dall, 1919). The speci- men (Figures 3 & 4) is accompanied by a label which reads, “Type/Catalina Id., Cooper”; a second label reads: “Lep- tochiton nexus Cpr./Type/Cooper.” The specimen, dried and partly disarticulated (loose valve vii, fragment of valve viii, and of valve [?] i), has an estimated length of 7.5mm. Color brownish gray, except for a longitudinal creamy- white stripe along the jugum. Tegmentum paved with granules, mostly rounded, about 40 xm in diameter, which tend to align themselves into longitudinal rows in the cen- tral areas (about 50 rows per half side), and in radial rows in the lateral areas (about 15 rows per lateral area). Lat- eral areas not otherwise defined. Jugum subcarinated, most particularly at the apex of the valves. Mucro central. Girdle shows a few short crystalline spicules, interspersed. Color slides of the holotype specimen at CASIZ, Color Slides Series. The examination of the syntype series of Lepidopleurus ambustus Dall, 1919 (USNM 274120), was possible again through the kindness of Dr. Joseph Rosewater. The accom- panying label reads, “20-40f. Sta. Barb. Chan./Sta. Bar- bara Ids. F. A. Woodworth/Types.” The series consists of 3 dried specimens; the largest, here designated as lectotype (Figure 6), measures 17.5 mm in length, 9.0mm in width, 3.3mm in height. All specimens show the same slight sub- carination, and the same tegmental sculpture seen in the holotype of L. nexus (Figure 5). The tegmental granules are generally round, and measure about 40 um in diameter. Color slides of the lectotype and paralectotypes, at CASIZ Color Slide Series, Nos. 1928-1930. The understanding of the species, Leptochiton nexus, comes clear from the joint descriptions of CARPENTER in Pirssry (1892), Berry (1919), and Dati (1919). However, a few observations must be added: In size, specimens of L. nexus do not usually exceed 20mm; however, an un- usually large specimen collected at 12-18 m, off Santa Cruz Island, California (LACM-AHF 1197-40) has an estimated length of 25mm. Mean width/length ratio, based on 10 specimens from several localities is 0.59. The color of Leptochiton nexus is remarkably constant, a dark tone of brown, often with a lighter creamy-tan stripe along the jugum, and minute specks of a darker brown or lavender, interspersed. Girdle, distinctly spiculose, with many hyaline spines (up to 400-500 um in length) strewn throughout (Figure 35). The radula is figured here for the first time. In a speci- men 7.8mm long (LACM-AHF 5714-58: “34 fathoms [62 m] 6.25 miles from Hyperion stack, Los Angeles, Cali- fornia, April 18, 1958”) measures 5.2mm in length (66% of the specimen’s total length) and has 35 rows of mature teeth. The median tooth (Figure 36) is larger in the front 100 pm Figure 36 Leptochiton nexus Carpenter, 1864 Radula: Median tooth, first lateral teeth and head of second lateral tooth. Specimen collected at Hyperion stack, Los Angeles, Califor- nia at a depth of 62m (LACM-AHF 5714-38) where it bears a small blade 110 wm wide. First lateral teeth have a conspicuous notch in the middle of the anterior bor- der. Major lateral teeth have a bicuspid head, which meas- . ures about 180 um in length, and 90 um in width. Outer marginal teeth are about 125 um long, 55 wm wide, with a length/width ratio of 2.27. Distribution: Leptochiton nexus seems to have a contin- uous distribution from Alaska to Baja California. The veri- fied northernmost record is Cohen Island, Kachemak Bay, Kenai Peninsula, Alaska [59°32.5°N; 151°28.5' W] (LACM 73-20, leg. J. H. McLean & R. Baxter, July 31, 1973, 20+ specimens); the southernmost record, Punta Abreojos, Baja California, Mexico [26°16’N; 113°41’ W] (LACM-AHF 1710-49, coll. March 7, 1949). Judging from the available collections, L. nexus seems to be easier to find between the latitudes of 45° to 28° N. It has been collected at most offshore islands: Catalina (USNM 16270; USNM 274119; LACM-AHF 2132-52; LACM 32-1), Anacapa (LACM-AHF 874-38), Santa Cruz (LACM-AHF 1155; LACM-AHF 1191-40 D1; LACM 1191-40 D2; LACM- AHF 1197-40; LACM-AHF 1303-41; LACM-AHF 1418- 41 D2), Santa Rosa (LACM-AHF 1022; LACM-AHF 995-39) Cedros (LACM-AHF 1253-41; LACM-AHF Page 152 THE VELIGER Vol. 22; No. 2 1247-41; LACM 71-159), and Natividad (LACM-AHF 1706-49). Recently, L. nexus was reported at Bahia de Los Angeles (29°00’ N; 133°35’ W) in the Gulf of California (HANSELMANN, 1977); thanks to the generosity of Col. G. A. Hanselman, I was given the privilege of examining and verifying the identification of the specimens, one collected by D. Mulliner at 17m, in May, 1970, another (11.5mm long) dredged at go m by R. Poorman. Bathymetrically, Leptochiton nexus ranges from the in- tertidal zone to 139-141m (LACM-AHF 1247-41, off Cedros Island, Baja California, Mexico). Median depth, based on the lots examined, 50 m. Remarks: Leptochiton nexus is much less abundant than L. rugatus. It clearly differs from L. rugatus by its darker color, larger size, and wider body. A comparison between 10 adult specimens of each species revealed that the mean length/width ratio of L. nexus specimens (ratio, 0.59) was significantly greater than that of L. rugatus specimens (ratio, 0.53), (Mann-Whitney test, p < 0.05- one-tailed). Differences in the girdle and radular characteristics of the two species were also marked. However, differences in the tegmental sculpture were more difficult to ferret out in view of the considerable intraspecific variation that both species display in the appearance and disposition of the teg- mental granules. A relatively reliable observation in the quick differential diagnosis between the two species is the total absence of carination (i.e., a perfectly round back) in L. rugatus while subcarination, particularly at the posterior end of the intermediate valves is usually discernible in L. nexus. The synonymy of Leptochiton lycurgus Dall, 1919, is based on the examination of the holotype (USNM 274119) available through the courtesy of Dr. Joseph Rosewater. The specimen (Figure 8), dry, curled, is whitish, and slightly carinated; anterior valve is missing; estimated length of the specimen, 7mm. The accompanying label reads: “Type/ Catalina Id., Dall.” In all its characteristics it corresponds to Da.u’s description (1919: 500). Color slides of the specimen, CASIZ Color Slide Series Nos. 1189-1191. Leptochiton alveolus (Lovén, 1846) (Figures 9 to 16, 37 and 38) Chiton alveolus Loven, 1846: 159-160 — JEFFREYS, 1882: 668 Lepidopleurus alveolus (Lovén). Sars, 1878: 110; plt. 7, figs. 3a-3i; pit. I, fig. 7 (radula) — Pivssry, 1892, 14: 6; plt. 2, figs. 23-31 — THIELE, 1902: 282 — NIERSTRASS, 1905: 4-5 — THIELE, 1909: g; plt. 1, figs. 2-4 — JOHNSON, 1915: 7 — JAKOVLEVA, 1952: 51-52; fig. 18, plt. 1, fig. 6 — La- RocQuE, 1953: 7 + ABBOTT, 1974: 393 Leptochiton alveolus (Lovén). DALL, 1879b: 317; 1882: 411 — Happon, 1886: 1, 12-13 — THIELE, 1893: 387; plt. 31, fig. 34 Leptochiton belknapt DALL. 1879: 1; 1879b: 317-318, 1882: 411 — Hapvpon, 1886: 1, 10-13; plt. 1, fig. 2, plt. 2, figs. 2a-2d — SMITH, 19474: 4; 1947b: 17 — BURGHARDT & BURGHARDT, 1969: 23-24 — ABBOTT, 1974: 393 Lepidopleurus belknapi (Dall). Prrspry, 1892, 14: 7-9; plt. 1, figs. 18-22. NIERSTRASS, 1905: 4-5 — DALL, 1921: 187 (in section Leptochiton) — OvpRoyp, 1927: 851 (in section Leptochiton) — LaRocQuE, 1953: 8 Leptochiton benthus Happon, 1886: 1, 10-13; plt. 1, fig. 1; pit. 2, figs. 1a-1m Lepidopleurus benthus (Haddon). Pissry, 1892, 14: 9; plt. 1, figs. 1-13 — NIERSTRASS, 1905: 4-5 Lepidopleurus (Leptochiton) benthus (Haddon). HANSEL- MAN, 1977: 62 Lepidopleurus mesogonus DALL, 1902: 555-556; 1921: 187 (in section Leptochiton) - OLDROYD, 1927: 852-853 - La RocguE, 1953: 8 — BURGHARDT & BURGHARDT, 1969: 25 — ABBOTT, 1974: 393 Leptochiton mesogonus (Dall). SMITH, 19474: 4; 1947b: 17 Lepidopleurus halistreptus DALL, 1902: 556; 1908: 218, 354 — THORPE in KEEN, 1971: 882 Lepidopleurus halistreptus abbreviatus DALL, 1908: 354 Explanation of Figures 9 to 16 Figure 9: Lepidopleurus farallonis Dall, 1902. Holotype (USNM 109025) Figure 10: Leptochiton belknapi Dall, 1879. Holotype (USNM 30972): Close-up to show detail of tegmental sculpture Figure 11: Lepidopleuridus luridus Dall, 1902. Holotype (USNM 109027: close-up of anterior, intermediate, and posterior valves Figure 12: Lepidopleurus mesogonus Dall, 1902. Holotype (US NM rogo1g) : Close-up to show detail of tegmental sculp- ture Figure 13: Lepidopleurus halistreptus Dall, 1902. Holotype (US NM 109032) : Close-up to show detail of sculpture Figure 14: Lepidopleurus simplex NierstraB, 1905. Lectotype (herein designated) (ZMA) Figure 15: Lepidopleurus giganteus NierstraB, 1905. Lectotype (herein designated) (ZMA) Figuré 16: Close-up of the specimen in Figure 15 to show detail of tegmental sculpture [FERREIRA] Figures 9 to 16 Tue VELIGER, Vol. 22, No. 2 Vol. 22; No. 2 THE VELIGER Page 153 Lepidopleurus abbreviatus (Dall). THoRPE in KEEN, 1971: 882 Leptdopleurus luridus DALL, 1902: 556-557; 1908: 218, 355; 1921: 187 (in section Leptochiton) — OLDRoYD, 1924: 184. 1927: 851-852 (in section Leptochiton) - La RocguE, 1953: 8 — BURGHARDT & BURGHARDT, 1969: 24- 25 — THORPE in KEEN, 1971: 882 — ABBOTT, 1974: 393 Leptochiton luridus (Dall). Smrru, 1947a: 4. 1947b: 17 Lepidopleurus farallonis DALL, 1902: 557. 1908: 218-355 — PACKARD, 1918: 291-292 — DALL, i921: 187 (in section Leptochiton) — OLpRoyD, 1927: 852 (in section Lepto- chiton) — BURGHARDT & BURGHARDT, 1969: 24 — THORPE in KEEN, 1971: 882 — ABBOTT, 1974: 393 Leptochiton farallonis (Dall). SMITH, 1947a: 4. 1947b: 17 Lepidopleurus giganteus NiERSTRASS, 1905: 3-4; figs. 1-2, 39- 42 Lepidopleurus simplex NiERSTRASS, 1905: 4-6; figs. 3, 44-47 Lepidopleurus opacus DALL, 1908: 218, 354-355 - THORPE in KEEN, 1971: 882 Leptochiton opacus (Dall). SMITH & FERREIRA, 1977: 84; figs. 34 Lepidopleurus japonicus THIELE, 1909: 11; plt. 1, figs. 21-29 - TAK & TAKI, 1929: 162-163 — JAKOVLEVA, 1952: 50- 51; fig. 17, pit. 1, fig. 5 — TAKt, 1962: 31 — Tart, 1964: 408. Lepidopleurus japonicus aequivaluus BERGENHAYN, 1933: 4-5 figs. 1a-1c — TARI, 1962: 32 — TAkI, 1964: 408 Lepidopleurus aequispinnus BERGENHAYN, 1933: 6-8 text figs. id-ig; plt. 1, fig. 1; plt. 2, fig. 20 - Taxi, 1962: g2 - TARI, 1964: 40 Type Material: Chiton alveolus Lovén, 1846: Unknown, presumably lost or never designated. Leptochiton belknapi Dall, 1879: Holotype (USNM 30972); locality, “Lat. 58°8’ N; Long. 171°19’ W.” Leptochiton benthus Haddon, 1886: Holotype (BMNH 1889.11.9.1); locality, HMS Challenger station 241, ‘Lat. 35°41’ N; long. 157°42’ E.” Not examined. Lepidopleurus mesogonus Dall, 1902: Holotype (USNM 109019); locality, USS Albatross station 3342, “off the Queen Charlotte Islands, British Columbia,” Canada. Lepidopleurus halistreptus Dall, 1902: Holotype (USNM 109032); locality, USS Albatross station 3415, “off Acapulco, Mexico.” Leptdopleurus luridus Dall, 1902: Holotype (USNM 109027); locality, USS Albatross station 3393, at “Panama Bay.” Lepidopleurus farallonis Dall, 1902: Holotype (USNM 109025); locality, USS Albatross station 3104, “‘off the Faral- lones Islands, near San Francisco, California.” Lepidopleurus giganteus NierstraB, 1905: Lectotype [herein designated] at Zoologisches Museum of Amsterdam (ZMA); locality, Siboge station, 221, “6°24’ S, 124°309/ W.” Lepidopleurus simplex NierstraB, 1905: Lectotype [herein designated] at ZMA; locality, Siboga station, 88, “0°34.6’ N, 119°8.5’ W.” Lepidopleurus opacus Dall, 1908: Holotype and paratype (USNM 110664); locality, USS Albatross station 4647, “‘be- tween the Galapagos Islands and the Peruvian coast.” Lepidopleurus japonicus Thiele, 1909: Types in Zoolo- gisches Museum, Humboldt Universitat, Berlin [“1 type spec- imen from Kajiyama (in alcohol) and 1 set of plates from Enoshima (also marked as a type),” Dr. R. Kilias, in litt., 6 September 1978]. Not examined. Leptdopleurus japonicus aequivalvus Bergenhayn, 1933: Type at the Zoologiska Museet, Uppsala, Type collection No. 186: locality, Sagami Bay, Japan. Not examined. Lepidopleurus aequispinnus Bergenhayn, 1933: Type at the Zoologiska Museet, Uppsala, Type collection No. 142; locality, Sagami Bay, Japan. Not examined. Type Locality: Scandinavia. “Boh[us]-Bergen” (Loven, 1846: 159), Original Description: ‘210. C. alveolus Sars ms. — T. oblonga, convexa, alba, squamults limbi inaequalibus, an- gustis;12/5 mm — T. satis convexa, vix culminata; valv. 8,1,7,2,3,0:4,5, arcuatae, longae, long. ad lat. in quinta = 1:3, postico rectae, vix mucronatae, margine antico leviter arcuato, medio emarginato, limbali rotundato. V. secundo antico subangulata, prima semicircularis, ultima subellip- tica. Laminae spatio separatae 1/3 tot. lat. fere superante, rotundatae. Series granulorum in areis lateralibus trans- versae, densae, fere secundum strias incrementi arcuatae. Limbus albus, angustus, tenuis, squamis elongatis, mucro- natis irregulariter imbricatus. Boh.-Bergen.” (Loven, 1846: 158-159). Extended Description: Sars (1878) figured the species (op. cit.: plt. 7, figs. 3, a-i), and added to the description: “Corpus colore nune pallidiore, albido-ctnereo, nunc ob- scurtore, interdum fere omnino nigro, forma quam solito angustiore, elongato-oblongum, plus duplo longius quam latius, zona perangusta, squamulis inaequalibus, nonnulis spinaeformibus. Testa sat convexa, dorso aequaliter arcu- ato ne vestigium quidem carinae vel arearum lateralium praebente, valvulis sat elongatis, postica antica majore, semiorbiculari, antice ad lineam rectum truncata, medits subaequalibus, margine postico recto, antico medio leviter emarginato. Super ficies valuularum tuberculis minutis ova- tis regulariter dispositis ubique sculpta. Long. 16mm” (op. cit.: 110-111). Piispry (1892, 14: 6) translated part of Sars’ description and reprinted his illustrations of the species (of. cit.: plt. 2, figs. 23-31). The radula, first figured by Sars (1878: plt. I, fig. 7) was again described and illustrated by THIELE (1893: 387; plt. 31, fig. 34), and JAKOVLEVA (1956: 51- 52; text fig. 18). THIELE (1909: Qg; plt. 1, figs. 2-4) en- larged upon the description of the girdle scales. From this study of several lots of Leptochiton alveolus (LovEN, 1846) in the eastern Pacific, the following obser- vations must be added: Shell uniformly colored, creamy Page 154 white to a light tan; occasional specimens may show orange or black, bituminous-looking deposits. Length, often over 3 cm; the largest specimen examined (CASIZ 008945, Ore- gon coast, 1829 m), if fully extended, it is estimated to meas- ure over 40mm in length. One of the specimens described by Nrerstrass (1905) as L. giganteus is reported to be 43 mm long. Based on 10 specimens from several localities, the mean width/length ratio is 0.39. Valves high arched, sub- carinated to markedly carinated; jugal angle, also quite variable, often less than go°. In young (small) specimens the valves may be slightly beaked. Mucro central to slightly posterior, often elevated. Tegmental surface covered throughout with granules, round to oval, and disposed in quincunx; the granules are usually well defined and clearly separated by a space about equal to their diameter. In size, shape, and degree of sepa- ration from each other, the granules vary considerably among specimens. As a rule, the tegmental granules tend to be smaller and more crowded together in the lateral than in the central areas. Lateral areas otherwise poorly defined in most specimens; however, they may be slightly elevated, or vaguely demarcated by a shallow sulcus. Con- centric growth lines, in the form of 3 or 4 rugae, are often seen in the lateral areas, quite conspicuously so in some specimens. Anterior valve, and postmucro area of posterior valve may also show similar growth lines, as well as crowded and smaller sized tegmental granules. 100 pm Figure 37 Leptochiton alveolus (Lovén, 1846) Girdle scales and spines. Specimen collected off Catalina Island, California, at a depth of 550m (LACM-AHF 1427-41) THE VELIGER. Vol. 22; No. 2 Girdle relatively narrow, covered by somewhat poly- morphic scales, mostly conical (up to 100 pm in height, 30 um in diameter), displaying a few variably marked lon- gitudinal striations (Figure 37); interspersed, there are a variable number of long, pointed, hyaline spines, up to 300 pm in length. In this respect, Leptochiton alveolus speci- mens seem to be quite variable, even within the same collec- tion site inasmuch as some may have a conspicuously large number of spines (ca. 100/mm/’), while others may show only very few (ca. 10/mm_’) or, perhaps, even none. The radula of a specimen 23mm long (LACM-AHF 1427-41, off Catalina Island, California, at 550m) meas- ures 10.0mm in length (43% of the total specimen’s length), and has 45 rows of mature teeth. Median tooth, rectangular, slightly large in the front where it bears a thin blade, 180 pm wide (Figure 38). First lateral teeth rectangular, with 100 zm See Figure 38 Leptochiton alveolus (Lovén, 1846) Radula: Median tooth, first lateral teeth, and head of second lateral tooth. Specimen collected off Catalina Island, California, at a depth of 550m (LACM-AHF 1427-41) slightly arcuate outer edge. Second (major) lateral teeth with unicuspid head which measures about 300 um in length, 140 um in width; the head’s outer edge is smoothly convex, but its inner edge displays a small bump (not quite a denticle) at its middle point, as illustrated in Sars (1878), THIELE (1893), and JAKOvLEVA (1952). Outer marginal teeth measure about 220 um in length, 150 um in width, with a length/width ratio of 1.46. Distribution: Leptochiton alveolus (Lovén, 1846) has long been considered a North Atlantic species ranging from Scandinavia and the Bay of Biscay in the east, to the Gulf of St. Lawrence and the Gulf of Maine in the west. JaKov- LEVA (1952) acknowledged its presence in the Sea of Ok- hostk, along the eastern coast of U.S.S.R. This study Vol. 22; No. 2 THE VELIGER Page 155 establishes the presence of L. alveolus in the eastern Pacific, (USNM 30972: L. belknapi Dall, 1919, holotype); the as well as in the Sea of Japan and the Bering Sea. southernmost record, 4°33’ N (USNM 110664: L. opacus From the data listed in Table 1, the northernmost record Dall, 1919,holotype and paratype). It has been collected in of Leptochiton alveolus in the eastern Pacific is 58°08’N the Philippine Islands, at 1911 m, by the HMS Challenger Table 1 Lots of Leptochiton alveolus (Lovén, 1846) examined, with latitudes, number of spec- imens per lot, maximum length of specimens in the lot, and collecting depth. B25 tow rs ie) < Lot Latitude o's ie Sa 28 é Sa z ac Depth mm m USNM 30972, L. belknapi, Type 58°08’ N HW i 1 840 USNM 109019, L. mesogonus, Type 53°00’ N 130 2.906 LACM 67-35 49°43/N 1 10 164 Cowan Coll. No. 5374 (Univ. Brit. Columbia) 48°02’N 2tO 977 CASIZ [R/V Ancona] 44°38’-40’N 20 15 800-2000 CASIZ 008945, R/V Ancona 44°-31' N 5 40 1 829 CASIZ [USS Mulberry; R/V Scofield] 37°48’ N 7 20 732-2 104 USNM 109025, L. farallonis, Type 37°45’ N 1 10 715 CASIZ 002893 36°47'N 8 20 567-640 CASIZ 009515 36°45’ N 6 21 954-1044 CASIZ 009524 36°45’ N nh UR 166 LACM 60-21 36°38’ N yO 183 LACM 63-52 36°38’ N tz 183 LACM-AHF 1471-41 33°34 N 1 23 523-549 LACM-AHF 1400-41-D1 33°24’ N 1 24 417 LACM-AHF 1156-40 33°24’N 2 15 421-695 LACM-AHF 1306-41-D1 33°24’ N 1 20 417-488 LACM-AHF 2004-50 33°24’ N 1 13 439-457 LACM-AHF 1425-41 33°13'N v5 B12 LACM-AHF 1019-39 33°04’ N 3 20 457-549 CASIZ 009521 (Gulf of California) 22°48’ N 1 10 366-1 336 CASIZ 009520 21°25’ N 4 15 3 000 USNM 109032, L. halistreptus, Type 16°51’ N 1 30 3 437 SDNH 68405 15°00’ N 1 8 3.962 USNM 109027, L. luridus, Type 8°50’ N » 15 1 866 USNM 110664, L. opacus, Type 4°33 N 2 35 3 667 ZMA, L. simplex, Type 0°35/N 1 7 1301 ZMA, L. giganteus, Type 6°24’S yey 2798 CASIZ 009516 [ex Golikov, USSR] Kunile Is. YG) ? CASIZ 009517 [ex Golikov, USSR] Bering Sea nt 22 ? CASG 52423 [USS Albatross] Hokkaido, 2 20 320 Japan CASIZ 001818 [ex Sirenko, R/V Vityaz] Honshu, 20 2 500 Japan B. Sirenko Colln. Barents Sea n 08 155 P. Kaas Colln. Norway lo 8) 100 R. Van Belle Colin. Gulf of 1 8 1 886 Biscay Page 156 (Haddon, 1886, as L. belknapi). The presence of L. alveolus in the southern hemisphere cannot be considered suffi- ciently documented at this time, except for NrersTRASS’ (1905) report of 3 specimens of what he named Lepido- pleurus giganteus, collected by the Siboga (Station 221: 6°24’ S, 124°39' W, at 2798 m). The bathymetric range of Leptochiton alveolus in the eastern Pacific based upon the specimens examined in this study, extends from 164m to 3962m, with a median of 550m, which places the species favored habitat in the upper bathyal to lower bathyal or abyssal zone (HEDGPETH, 1957). Remarks: Until now, Leptochiton alveolus was known with certainty only in the North Atlantic. Its presence in the Pacific is firmly established here with the examination of Dall’s several eastern Pacific types of Leptochiton species made available for study through the generosity of Dr. Joseph Rosewater. An account of Dall’s type specimens, and their photographs are given here for the first time. Leptochiton belknapi Dall, 1874: Holotype (USNM 30972). Accompanying labels read: “1006 fms. sd. & sh./ Lat. 58°8’ N., Long. 171° 19’ W, Belknap,” and “Type.” A single specimen (Figure 70) dried, curled, carinated, valves slightly beaked, whitish color. Estimated length 15mm, width 6mm, height 3.5. Tegmental granules in quincunx throughout. Lateral areas not raised, defined by faint diag- onal sulcus, with coarse concentric growth lines. Mucro central. Girdle covered with small spicules crowded to- gether. Two discrepancies in previous descriptions must be noticed: Dart (1874) gives the size of the specimen as “Lon. 10, lat. 3mm,” much smaller than my estimate; and Prissry (1892) gives locality longitude as 17°, an obvious typographical error for 171° as stated by Dall, and written on the specimen’s label. Color slides at CASIZ Color Slides Series. THE VELIGER Vol. 22; No. 2 Lepidopleurus mesogonus Dall, 1902: Holotype (USNM 109019). Labels read: “Sta. 3342, 1588 fms. ooze, 53°.3/ British Columbia off Q. Charl. Ids.” and ““Type.” A single specimen (Figure 12), dried, dirty white, curled, strongly carinated. Estimated length 30mm, width romm, height 7mm; jugal angle about 80°. Strong concentric growth lines in lateral areas and continuing into adjacent pleural area. Mucro central. Girdle sandy looking, with many small spicules interspersed. Color slides at CASIZ Color Slides Series. Lepidopleurus halistreptus Dall, 1902: Holotype (USNM 109032). Labels read: “Sta. 3415, 1879 fms., m. 36°/off Acapulco,” and “Type.” A single specimen (Figure 13), dried, curled, whitish in color, strongly carinated. Jugal angle about 90°. Estimated length 30mm, width 10mm, height 7mm. Valves i, vii, and viii disarticulated. Lateral areas faintly defined by a diagonal sulcus. Mucro slightly posterior. Girdle arenaceous looking, with spicules (up to 500 wm long) interspersed. Color slides at CASIZ Color Slides Series. Lepidopleurus luridus Dall, 1902: Holotype (USNM 109027). Labels read: “Sta. 3393, 1020 fms., m. 36°.8,” “Panama Bay,” and “Type.” Single specimen (Figure 11), dried, curled, whitish, carinated (but less so than in the pre- viously mentioned type specimens). Estimated length 15 mm, width 6mm, height 3mm. Valves i, vii, and viii dis- articulated. Tegmental granules relatively larger and more widely separated than in above mentioned type specimens. Lateral areas modestly defined and raised, with a few con- centric growth ridges. Posterior valve distinctly longer than anterior valve, with a truncated looking anterior edge. Girdle sandy looking, with occasional spicules interspersed. Color slides at CASIZ Color Slides Series. Lepidopleurus farallonis Dall, 1902: Holotype (USNM 109025). Labels read: “Sta. 3104, 391 fms, coral, 41° off the Farallones,” and “Type.” Single specimen (Figure 9), Explanation of Figures 17 to 24 Figure 17: Hanleyella oldroydi (Dall, 1919). 6.2mm long. Mon- terey Bay, California (LACM 63-50) Figure 18: Close-up of specimen in Figure 17 to show detail of central area, lateral areas, and girdle Figure 19: Lepidopleurus oldroydi Dall, 1919. Holotype (USNM 218767) Figure 20: Oldroydia percrassa (Dall, 1894). 26mm long (dried). Catalina Island, California (AJF collection) Figure 21: Leptochiton diomedeae Berry, 1917. Holotype (USNM 215625) : Close-up of anterior, posterior, and intermediate valves Figure 22: Close-up of specimen in Figure 21 to show detail of the tegmental sculpture of intermediate valves Figure 23: Lepidopleurus lineatus NierstraB, 1905. Lectotype (herein designated) (ZMA) Figure 24: Close-up of specimen in Figure 23 to show detail of tegmental sculpture THE VELIGER, Vol. 22, No. 2 [Ferreira] Figures 17 to 24 Vol. 22; No. 2 dried, curled, whitish, carinated. Valve i fragmented; valve ii broken but in place; valve viii disarticulated. Tegmental granules about 60-80 wm in diameter; average space be- tween granules in the central areas, 20-30 um. Girdle with numerous spicules (measuring up to 200 pm in length) in- terspersed. Color slides at CASIZ Color Slides Series. Lepidopleurus giganteus Nierstrass, 1905: The specimen examined on a loan arranged through the kindness of Dr. H. E. Coomans, Curator of Malacology, Zoologisch Mu- seum of Amsterdam (ZMA) is well preserved in ethyl alco- hol. The specimen, marked “Type,” conforms well with the description given by NiersTRass (1905). It is accompanied by a museum label which reads “Lepidopleurus giganteus Nierst./Stat. 221/figured specimen by Schepman/ fig. 1 & 2,” a reference to the illustrations in “Die Chitonen der Siboga-Expedition” (1905). The specimen, the smallest of the 3 specimens found at the Station 221 of the Siboga ex- pedition, measures 24.2 mm in length, and 8.2 mm in width, including girdle. It is here designated as lectotype, and illustrated (Figures 15 & 16). The other two specimens of the syntype series of L. giganteus are also in the repository of the Zoologisch Museum of Amsterdam (Dr. H. E. Coo- mans, in litt., June 13, 1978). Color slides of the lectotype at CASIZ, Color Slides Series. Lepidopleurus simplex NierstraB, 1905: A specimen marked “Type” examined also through the generosity of Dr. H. E. Coomans (ZMA). The accompanying label reads “Lepidopleurus simplex Nierstr./Stat. 88/1 specimen.” The specimen, preserved in ethyl alcohol, measures 7.2mm in length, 3.3mm in width; it conforms well to Nier- STRASS’ (1905) original description. Of the 20 specimens of L. simplex mentioned in the Siboga Report (1905), 18 are in the Zoologisches Museum of Amsterdam, “one in the Mus. of Brussels, [and] one probably was kept by Nierstra&. We are not able to recognize the figured [in the Siboga Report] specimen” (Dr. H. E. Coomans, in litt., June 13, 1978). The examined specimen is here illustrated (Figure 14), and designated as lectotype. Color slides at CASIZ Color Slides Series. Lepidopleurus opacus Dall, 1908: Holotype and para- type (USNM 110664) examined through the generosity of Dr. Joseph Rosewater. Labels read, “Sta. 4647, 2005 fms., ooze, 35.4°/near Galapagos Ids./off Peruvian coast/Lat. 4°33’ S/Lon. 87°42’ W,” and “Types.” Specimens dried, curled, sharply carinated. Photograph of holotype in SmirH & FERREIRA (1977, figs. 3 & 4). Color slides, CASIZ Nos. 1581 (holotype), and 1582-1583 (paratype). The placement of Leptochiton benthus Haddon, 1886, Lepidopleurus japonicus Thiele, 1909, and Lepidopleurus aequispinnus Bergenhayn, 1933, in the synonymy of Lepto- chiton alveolus (Lovén, 1846) is based on the published THE VELIGER Page 157 descriptions and illustrations of the respective nominal spe- cies; examination of the respective types was not under- taken. Leptochiton albemarlensis Smith & Ferreira, 1977 Leptochiton albemarlensis SurrH & Ferreira, 1977: 89-84; figs. 1, 2. Type Material: Holotype (CASG, Type Collection No. 58247), and paratype (CASG, Type Collection No. 58248). Type Locality: Tagus Cove, Isla Isabela (Albemarle Island), Galapagos Archipelago, Ecuador [0° 16'S; 91°22’ W], dredged at 20m. Remarks: The species is known only from the type lo- cality and the type material. On account of its spiculose girdle, Leptochiton albemarlensis is similar to L. nexus from which it differs by (1) its white color, (2) the quincunx arrangement of the tegmental granules with only a vague tendency to align themselves in diagonal (not longi- tudinal) rows, and (3) the round back, i.e. with no trace of carination. Compared to Leptochiton alveolus, the Gala- pagan L. albemarlensis differs by (1) its round back, (2) the tendency of the tegmental granules to align in diagonal rows,and (3) itsmuch wider body, as revealed by a width/length ratio (for the holotype) of 0.55. CASIZ Color Slides Nos. 1978-1979. Leptochiton incongruus (Dall, 1908) (Figure 29) Lepidopleurus incongruus DALL, 1908: 218, 355 — THORPE in KEEN, 1971: 882. Type Material: Holotype (USNM 122969) Type Locality: “U.S.S. Albatross, station 3354 [7°09'45” N; 80°50’00”W], Feb. 23, 1891, Gulf of Panama, in 322 fathoms [589 m]” (Daxt, 1908: 355). Original Description: “Animal with the gills forming a very short posterior row; girdle narrow with extremely mi- nute, close-set spines, giving the effect of a naked, velvety surface. Intermediate valves pale brown, the median line slightly obtusely angular; jugum slightly mucronate be- hind, the jugal area sculptured with very fine, close-set, rounded, longitudinal threads; pleural tracts with ten or twelve larger threads with wider, flat interspaces, the threads sometimes breaking off abruptly, leaving double- width interval the rest of the way;lateral areas prominent, Page 158 very finely granulose, conspicuously concentrically rippled; anterior plate with feeble concentric ripples and similar granulation; posterior valve with prominent subcentral mucro, the central area sculpture like the anterior valve; penultimate valve with a length of 2.3, a width of 6.0, and an altitude of 2.5mm. Articulations as usual in the genus.” (Dat, 1908: 355). Extended Description: Thanks to the generosity of Dr. Joseph Rosewater of the National Museum of Natural His- tory, the holotype of Lepidopleurus incongruus Dall, 1908 was studied and photographed. It is figured here for the first time (Figure 29). The specimen, a bit curled, preserved dry, has a uniform light beige color. Estimated length, 12 mm; it measures 5.5mm in width, 3 mm in height. Valves moder- ately carinated, slightly beaked. Lateral areas distinctly defined, moderately raised. Central areas with small, flat- topped granules, arranged in longitudinal rows which con- verge anteriorly; in the pleural areas, these rows become well defined ridges, about 8-10 per side. Lateral areas, valve i, and postmucro area of valve viii display strong concentric growth wrinkles. Mucro central, not prominent; postmucro moderately concave. Girdle has a sandy appearance, with many short spicules, and a few longer spicules interspersed. Color slides of the holotype at CASIZ Color Slides Series. Distribution: In addition to the type locality, Panama Bay, Leptochiton incongruus was also found off Salina Cruz, Gulf of Tehuantepec, Mexico [15°40’N; 95°20’ W] at 3541-3612m of depth (CASIZ 009525, Scripps coll., 18 Nov. 1958, Loc. No. P-127-58, To 58-2, D-4), 2 specimens, preserved in alcohol, a bit curled, both measuring an esti- mated 15mm in length, and beige in color. (CASIZ Color Slides Nos. 724-726). Remarks: There can be no question that Leptochiton incongruus is a valid species, characterized, as Dall (1908) noted, by ‘“‘a unique sculpture.” Hanleyella Sirenko, 1973 Definition: Central areas with rows of flat granules sep- arated from each other by spaces larger than their diam- eters. Girdle with spicules, and by a large number of long spines. Inner edge of sutural laminae slope gently relative to the front border of valve. No insertion plates. Radula with tricuspid major lateral teeth. Type Species: Hanleyella astatica Sirenko, 1973, by OD. THE VELIGER Vol. 22; No. 2 Remarks: Hanleyella asiatica, described from Simusir in the Kurile Islands, and H. oldroydi (Dall, 1919), from the northeastern Pacific, are the only two known representa- tives of the genus which SmrENKo (1973) suggests to be “a link between the genera Lepidopleurus Risso and Hanleya Gray.” Hanleyella oldroydi (Dall, 1919) (Figures 17 to 19, 39 and 40) Lepidopleurus (Leptochiton) oldroydi DALL, 1919: 500-501 Lepidopleurus oldroydi Dall. DAL, 1921: 187 (in section Leptochiton) — OLpRoyD, 1927: 850 (in section Lepto- chiton) — ABBOTT, 1974: 393 Leptochiton oldroydi (Dall). SMITH, 19474: 6. 1947b: 17 — SMITH & GORDON, 1948: 205 — BURGHARDT & BURGHARDT, 1969: 26 Hanleyella oldroydi (Dall). SIRENKO, 1973b: 1569 Type Material: Holotype (USNM 218767) Type Locality: ‘Monterey, California [36°45’N; 121° 55° W]” (DALL, 1919: 501). Original Description: ‘‘Chiton white, with a blackish spot on either side of the jugal area, strongly sculptured; back moderately arcuate, anterior valve semicircular, sculptured with irregularly disposed small prominent round pustules; posterior valve with subcentral prominent mucro, the central area granulose, the periphery pustulose; inter- mediate valves with axially punctostriate jugal area, lateral areas prominent by reason of the conspicuous pustules which are relatively large for the size of the animal; the pleural tracts coarsely axially grooved, the grooves more or less distinctly punctate; girdle with crowded minute spines of equal length, giving an arenaceous effect; interior whit- ish, the jugal sinus wide with a straight edge, the sutural laminae small, subtriangular. Length of dry animal, 5; width, 2; height, 0.7mm.” (DaLL, 1919: 500-501). Extended Description: The examination of the holotype (USNM 218767), available through the kindness of Dr. Joseph Rosewater, disclosed the following points not in- cluded in, or differing from Dall’s original description: The labels accompanying the specimen read “Monterey Har- bour/West Coast, California,” and “Type.” The single specimen (Figure 19) dried, partly disarticulated, has an estimated length of 5.3mm. The five anterior valves are broken but in place, except for large fragments of valves i Vol. 22; No. 2 THE VELIGER Page 159 and iv which are available but detached from the speci- men; valve vii, disarticulated. Valves moderately carinated, slightly beaked. Pleural areas display about 12 ridges per side. Girdle shows a couple of long spines, about 0.2 mm in length, on the right side of the specimen. Valves show no slits or insertion plates. Color slide of the holotype at CASIZ Color Slides Series No. 1926. Specimens of Hanleyella oldroydi are very small; the largest examined measures 8.2mm in length, 4.3mm in width, 1.9mm in height (LACM 63-26). Specimens are usually high arched, moderately carinated (Figures 17 & 18). Valve i, postmucro area of valve viii, and lateral areas of intermediate valves display large round tubercles, ran- domly distributed. Central areas of intermediate valves have granules that align themselves in up to 15 rows or ridges which run parallel to the jugum, or diverge forward slightly. Mucro relatively high, slightly anterior. Girdle (Figure 39) covered with minute scales, about 60 x 30 um in size, which show some irregular striae; in addition there are numerous spinules interspersed, and rather large hya- line spines. These large spines (often 400 wm in length by Figure 39 Hanleyella oldroydi (Dall, 1919) Girdle scales and spines. Specimen collected at Hope Island, British Columbia, at a depth of 39-90m (LACM 63-26) 30 um in width) are mostly interspersed throughout the girdle, although with a definite tendency to bunch up and be larger at the sutures, and close to the margin of the girdle where they may constitute (particularly in young speci- mens) a conspicuous marginal fringe. It is worth noting that, in my experience, these long spines break rather easily in dry specimens, and seem to disappear in OHK solutions. The radula of a specimen 8.2mm long (LACM 63-26, Hope Island, British Columbia, leg. I. McT. Cowan & J. H. McLean, on Ekholi, May 22, 1963, at 39-90 m) measures 3.5 mm in length (42% of the total specimen), and has 45 rows of mature teeth. Median tooth (Figure 40), rectangular, 50um Figure 40 Hanleyella oldroydi (Dall, 1919) Radula: Median tooth, first lateral teeth, and head of second lateral tooth. Specimen collected at Hope Island, British Columbia, at a depth of 39 - 90m (LACM 63-26) slightly wider in front than in back, maximum diameter at anterior blade, 42 um. First lateral uncharacteristic except perhaps by the presence of a notch in the middle of anterior edge. Second (major) lateral with a tricuspid head which measures about 100 wm in length, and 45 wm in width. Outer marginal teeth, 58 um long, 35 um wide; length/ width ratio 1.65. Distribution: BURGHARDT « BURGHARDT (1969), SIRENKO (1973), and Assott (1974) stated that the range of Han- leyella oldroydi extends to Alaska. In this study the north- ernmost verified record is Edna Bay, Kosciusko Island, Alaska [55°57’N, 133°40’ W] (CASG 32430, leg. G. D. Hanna, July, 1947); the southernmost record, Cabo San Quintin, Baja California, Mexico (30°17’40" N; 115°54’ 40” W) (LACM 71-150, leg. J. H. McLean « P. LaFollette, Searcher, Sta. 226-227, October 17, 1971, at 36-55 m). The species has been collected at Monterey Bay (LACM 60-21, LACM 60-22, LACM 60-24, LACM 63-50, LACM 66-56), Catalina Island (LACM A.375, LACM 32-1, LACM 65-6, LACM-AHF 1323-41, LACM-AHF 1350- 41, LACM-AHF 1430-41 D1, LACM-AHF 1523-41), San Page 160 THE VELIGER Vol. 22; No. 2 Clemente Island (LACM-AHF 911-39), and Santa Rosa Island (LACM-AHF 1290-41). Hanleyella oldroydi is a relatively deep water species; the recorded bathymetric range is from 18m (LACM 6o- 22) to 420-455m (LACM-AHF 1992-50); the median depth, based on the lots examined, at about 100m. Remarks: tions of Hanleyella asiatica, and from the examination of specimens generously donated by Dr. B. Sirenko, Zoologi- cal Institute, Academy of Sciences of U.S.S.R., Leningrad, H. oldroydz is found to differ clearly from its western Pacific counterpart by 1) amuch stronger tegmental sculpture, 2) the granulose aspect of the pleural ridges (almost smooth in H. asiatica), 3) the frankly pustulose lateral areas (almost sculptureless in H. asiatica), and 4) the tendency of the long girdle spines to bunch up in tufts (a tendency not noticed in the specimens of H. asiatica exam- ined). In addition, while females of H. asiatica are known to carry young specimens in the mantle cavity (SIRENKo, 1973), the brooding habit has not been observed in H. old- roydi. From the original description and illustra- Oldroydia Dall, 1894 Definition: “Valves heavy, strongly sculptured, with ir- regular transverse ribs that in life are well separated by narrow extensions of girdle reaching to jugum, resulting in coverage that is partial to apical only; articulamentum well developed, unslit; tegmentum with posterior extension be- tween rather large sutural laminae; jugal area prominent, sculptured differently from pleural areas; lateral areas not differentiated.” (SMITH, 1960: 52). First described as a section of Lepidopleurus; elevated to generic rank by THIELE (1910). Type Species: Lepidopleurus (Oldroydia) percrassus Dall, 1894, by OD. Remarks: For its heavy and strongly sculptured valves, the monotypic genus Oldroydia Dall bears close affinity to Lepidopleurus Risso of which it may be considered to be the Pacific counterpart. Oldroydia percrassa (Dall, 1894) (Figure 20) Lepidopleurus percrassus DALL, 1894: go-91 (in new section Oldroydia) Lepidopleurus (Oldroydia) percrassus Dall. BERRY, 1907: 47 Oldroydia percrassa (Dall). THIELE, 1910: 71, 105; plt. 7, figs. 1-8 — DALL, 1921: 187 — OxpRoyD, 1927: 854-855 — STRONG, 1937: 194 — BURCH, 1942: 7 — BuRCH & BURCH, 1943: 5 — SMITH, 19474: 7. 1947b: 18 — SMITH & Gor- DON, 1948: 205 — JAKOVLEVA, 1952: 52-53; text. fig. 19; pit. 19, fig. 1 —- TAKI, 1964: 408 — BURGHARDT & Burc- HARDT, 1969: 33; plt. 4, fig. 68 — ABBOTT, 1974: 393 Hanleya hanleyi (Bean). OLDROYD, 1927: 855 — SMITH, 19474: 7. 1947b: 18 - SMITH & GORDON, 1948: 205 — JAKoy- LEVA, 1952: 54. [not Chiton hanleyi BEAN in THORPE, 1844] Hanleya spicata BERRY, 1919: 8-10; plt. 1, fig. 3; text figs. 5-6 — SMITH, 19474: 7. 1947b: 18 — SMITH & GoRDON, 1948: 205, Hanleya hanleyi spicata Berry. DALL, 1921: 188 — OLpRoyn, 1927: 855-856. Type Material: Lepidopleurus percrassus Dall, 1894: Holotype, “larger, partially disarticulated specimen marked ‘type’ ’, and 2 para- types (USNM 107274) (Dr. J. Rosewater, in lttt., 3 April, and 27 July 1978). Paratypes (CASG-SU Type no. 6239), 5 speci- mens. Hanleya spicata Berry, 1919: Holotype (S. S. Berry private colln., Cat. No. 4102); “off Point Pinos, Monterey Bay, Cali- fornia.” Not examined. Type Locality: “Specimens obtained by Mr. T. S. Old- royd from a stone pulled up from about 75 fathoms [137m] in the Santa Barbara Channel off San Pedro [33°45’N; 118°11’ W], California” (DALL, 1894: 91). Explanation of Figures 25 to 32 Figure 25: Lepidopleurus nicomedes Dall, 1919. Holotype (US NM 96935): anterior valve Figure 26: Lateral view of the specimen in Figure 25 Figure 27: Posterior view of the specimen in Figure 25 Figure 28: Lepidopleurus agesilaus Dall, 1919. Lectotype (herein designated) (USNM 96227) Figure 29: Lepidopleurus incongruus Dall, 1908. Holotype (US NM 122969) Figure jo: Lepidopleurus rissot NierstraB, 1905. Lectotype (here- in designated) (ZMA) Figure 31: Close-up of the specimen in Figure 30 to show detail of lateral and central areas Figure 32: Posterior valve of specimen in Figure jo [FERREIRA] Figures 25 to 32 6 N i.) =] 5} a0 on) . 19 Figure 25 TuHE VELIGER, Vol. 22, No. 2 8 Figure 2 Figure 29 igure 32 FE hs — oe —- 4 () ‘a Aa __CorDovA GULF or ALASKA 60°00’ 147°00° Figure 1 Map of Prince William Sound, Alaska, showing the location of Simpson Bay where specimens of Protothaca staminea were col- lected for this study alveolar lumen. An increasing number of stalked oocytes importance of amphinucleoli in determining ripeness of protrude into the lumen. ova). Late active -— Most ovocytes spherical with slender Ripe — A majority of ova free of the alveolar wall. stalks; many show amphinucleoli (see ALLEN, 1953 for Ripe ova and remaining late ovocytes prominent. Page 184 THE VELIGER Vol. 22; No. 2 Spawning - Follicle cells generally reduced to 1 or 2 rows but seldom absent. Alveoli partially emptied of ripe ova. Post spawning — Follicle cells beginning to fill in the alveoli. Some primary ovocytes appearing between follicle cells. Some free ova still present. Amoebocytes abundant within and in the vicinity of the ovary. Males Early active - Follicular tissue partially or completely filling the alveoli. Characterized by the presence of pri- mary spermatocytes on the germinal epithelium, and the appearance of some spermatids. Middle active - Follicular cells have disappeared. Spermatids arranged in radial rows. Late active — Characterized by a central lumen with- in the alveolus. Spermatids have increased in number. Ap- pearance of some spermatozoa. Ripe — Spermatozoa fill the alveolus in thick radial columns with their tails projecting into the central lumen. Spawning — The appearance of free spermatozoa within the alveolus. Formation of a single row of follicle cells at the alveolar membrane with more appearing as spawning proceeds. Post spawning — Alveoli nearly filled with follicle cells with a few residual spermatozoa present. Primary spermatocytes have begun to appear on the germinal epithelium. The meats of an additional 199 clams were dried to a constant weight, at 80°C, and dry meat-wet meat values calculated. All clams were approximately the same size. Dry Meat Weight Internal Shell Volume calculated for 168 of these clams (see WALNE, 1970; WESTLEY, 1961, for methodology). A condition index was not calculated for January 1973. The technique of Husss & Huss (1953) is used to compare monthly values for dry-wet weight ratios and condition indices. A significant difference (a—0.05) is demonstrated by this technique when the standard errors of the means of the 2 values to be compared fail to overlap. A condition index ( X 100) was RESULTS HISTOLOGICAL STUDY Females The females were inan active stage for the first 4 months of 1973 (Figure 2). In May, 25% of the clams were YZ Yj 80 j; 7 Go Y le 3 60 VE : =|= a 40 G Y a . BIZ | Rea AN Ma Ay MAR MONN Wace AMES Tek y OM NDE (20) (20) (6) (7) (18) (6) (14) (1) (12) (28) (26) (30) (30) (26) | 1973 |-1974 = Earl Middl actin active. dae [iI] Ripe Spawn Es ane Figure 2 The female (above) and male (below) reproductive cycles of Protothaca staminea. The length of each shaded area represents the percent frequency of clams in each reproductive phase. ripe; all others were in one of the 3 active stages. In June most of the individuals were in the spawning phase, some post-spawning individuals were present. All females were in the spawning stage in July. By August over 80% were still in the spawning phase; the rest were in an early active state. An increasing percentage of females in the early active stage was evident in September and October. In October some post-spawning individuals were also pre- sent. A spent phase was not observed. No spawning stages Vol. 22; No. 2 were present in the last 4 months. Clams from November 1973 through January 1974 were in early and middle active stages. THE VELIGER Page 185 spawning was reflected by a significant (a—=0.05) de- crease in DWR during the period of May 18 through June 6. The ratio continued to decline until July 1, when Dry Weight/Wet Weight (DWR) spawning was apparently completed. Thereafter, DWR increased significantly, and then remained relatively stable until January 1974, when an increase occurred. Males All reproductive stages were present for the first 3 months of the year (Figure 2). In April and May all stages, except spawning, were present; early active stages were found only in small males. From June through Jan- uary, the majority of males were in the spawning phase. A spent condition was never observed. CONDITION INDEX The condition index (CI) was lowest in February and March 1973. Significant (@—0.05) increases in CI oc- curred monthly from March 6 to May 18, the months preceding spawning (Figure 4). A significant decrease in CI, representing spawning, was observed between June 6 and July 1. In general, there was little change in CI from July through December. An increase in CI occurred in January 1974. WEIGHT RELATIONSHIPS The period preceding spawning, January 20 to May 18, was characterized by a steady increase in the ratio of dry to wet meat weight (DWR; Figure 3). Initiation of 0.4500 DISCUSSION HISTOLOGICAL STUDY 0.3000 Histological examination of the female reproductive cycle indicates a single annual reproductive period for Protothaca staminea, with ovaries in a spawning phase from early June through September (Figure 2; NickKEr- SON, 1977). However, instead of entering a spent phase, where alveoli were empty of ripe ovocytes, many ripe or residual ova (or both) were present after the major spawning in June (Figures 3, 4), and follicle cell devel- (18) opment was evident in most females during the final months of the spawning phase. Initiation of spawning by males also occurred in June (Figure 2), but a spawning period was less clearly defined. Males were in a spawning phase throughout most of the year. The maintenance of 0.2500 + ° bo ° fe) °o a REET PORES SIEBER SBBES eB (16) (19) (15) (© adjacent column) Figure 3 Dry meat weight/wet meat weight (DWR) indices for a spawning cycle of Protothaca staminea. The single vertical line represerits the range, the white box the standard deviation, the dark box two standard errors of the mean, and the horizontal line the mean. A significant difference (2=0.05) is demonstrated when the standard errors of the means of two values fail to overlap. The total number of clams examined each month is included in parentheses [Fe CMA Me) JON yo AS 8S (20) (20) (6) (7) (18) (6) (1) | ne Oo N D J (12) (28) (26) (30) (g0) (26) 1974 CONDITION INDEX Page 186 (15) of (14) a) (13) (16) (15) Tee rhy Nua ACCENT Ten) IL AGILAS) AO (20) (20) (6) (7) (18) (6) (1) (12) | 1973 NepDE (28) (26) (30) (30) (26) Figure 4 The condition indices for a spawning cycle of Protothaca staminea. The single vertical line represents the range, the white box the standard deviation, the dark box the two standard errors of the mean, and the horizontal line the mean. No condition index was calculated for January, 1973. A significant difference (#=0.05) is demonstrated when the standard errors of the means of two values fail to overlap. The total number of clams examined each month is included in parentheses a minimal level of gonad development throughout the year has been previously reported for other Veneridae (see ANSELL & TREVALLION, 1967, for discussion). THE VELIGER 1974 Vol. 22; No. 2 WEIGHT RELATIONSHIPS anpb CONDITION INDEX The dry weight-wet weight ratio (DWR) decreased significantly from May 18 to June 6 (Figure 3). The de- crease in this ratio and the histological observation of post-spawning females in early June (Figure 2) suggest initiation of spawning occurred prior to June 6. A con- current significant decrease in the condition index (CI) did not occur during the same period. The decrease in DWR was probably the result of water uptake after in- itiation of spawning. Increase in water content of bivalve tissues at the time of spawning has been documented for other bivalves (ANSELL & TREVALLION, 1967). In the cal- culation of CI, only dry tissue weight and shell volume are considered, and water uptake is not observable. Only a decrease in dry meat weight will be detected by the CI method. Thus, only a limited spawning took place before June 6. However, a significant decrease in CI occurred between June and July, indicating a major release of gametes at this time (Figure 4). The high values for DWR and CI in January 1974, as contrasted with January 1973, suggest a better state of health for the clam population or a greater retention of gametes in 1974, or both. The reasons for these differences in DWR and CI at these times are unknown. CONCLUSIONS Calculations of weight ratios and condition indices are necessary to determine the period of peak spawning for Protothaca staminea in Prince William Sound. With his- tological techniques alone, male littleneck clams appear in a spawning condition throughout most of the year and female clams for 4 months of the year. The weight rela- tionship studies show that the major spawning effort of P. staminea is restricted to the month of June. In British Columbia, spawning occurs from April to October (QuayLE, 1943). A shortened spawning period is characteristic of clams at the northern and southern limits of their range, and appears to be primarily a tem- perature-related phenomenon (see discussions by BAYNE, 1976; Orton, 1920; WiLson & Hopcxkin, 1967, on factors controlling reproductive cycles). Vol. 22; No. 2 THE VELIGER Page 187 SUMMARY The annual reproductive cycle is described for the little- neck clam, Protothaca staminea, in Prince William Sound, the northern limit of its range. Reproductive activity was determined by histological examination of gonads and by calculation of monthly dry meat-wet meat weight ra- tios and condition indices. Histological examinations sug- gest that females and males are in a spawning phase from June until September and June to January, respectively. However, dry weight-wet weight ratios indicate a limited spawning in late May with continued spawning during June. Condition indices indicate intensive spawning in June. ACKNOWLEDGMENTS This work is a result of research sponsored by the Alaska Sea Grant Program cooperatively supported by the NOAA office of Sea Grant, U. S. Department of Com- merce, under Grant No. 04-3-158-41, and by the Univer- sity of Alaska with funds appropriated by the State of Alaska. We thank the crew of the R/V Acona and Merle Hanson for aid in collection of specimens. This work is Contribution No. 379 from the Institute of Marine Sci- ence, University of Alaska, Fairbanks. Literature Cited Auten, R. D. 1953. Fertilization and artificial activation of the egg of the surf clam, Spisula solidissima. Biol. Bull 105: 213 - 239 AnseELL, A. D. & A. TREVALLION 1967. Studies on Tellina tenuis Da Costa. I. Seasonal growth and bio- chemical cycle. Journ. Exp. Mar. Biol. Ecol. 1: 220 - 235 Bayne, B. L. 1976. Marine mussels: their ecology and physiology. Progr. 10. Cambridge, 506 pp. Davenport, H. A. 1960. Histological and histochemical technics. Philadelphia. 401 pp. Husss, C. L. « L. Husss 1953. An improved graphical analysis and comparison of series of samples. Syst. Zool. 2: 49-56 Nickerson, R. B. 1977. A study of the littleneck clam (Protothaca staminea Conrad) and the butter clam (Saxidomus giganteus Deshayes) in a habitat permitting coexistence, Prince William Sound, Alaska. Proc. Nat. Shellfish Assoc. 67: 85 - 102 Orton, G. H. 1920. Sea-temperature, breeding and distribution in marine animals Journ. Mar. Biol. Assoc. 12: 339 - 366 Paut, A. J. s Howarp M. Feper 1973- Growth, recruitment and distribution of the littleneck clam, Protothaca staminea, in Galena Bay, Prince William Sound, Alaska. Fish. Bull. 7 (3): 665 - 677 1975- The food of the sea star Pycnopodia helianthoides (Brandt) in Prince William Sound, Alaska. Ophelia 14: 15-22 Paut, A. J., Jupy M. Paut & Howarp M. Fever 1976. Recruitment and growth in the bivalve Protothaca staminea, at Olson Bay, Prince William Sound, ten years after the 1964 earthquake. The Veliger 18 (4): 385-392; 5 text figs. (1 April 1976) Porter, R. G. 1974. | Reproductive cycle of the soft shell clam, Mya arenaria, at Skagit Bay, Washington. Fish Bull. 72 (3): 648-656 Quaytg, P B. 1943. Sex, gonad development and seasonal gonad changes in Paphia staminea (Conrad). Journ. Fish. Res. Brd. Canada 6 (2): 140-151 Ropgs, J. W. 1968. Reproductive cycle of the surf clam, Spisula solidissima, in offshore New Jersey. Biol. Bull. 195 (2): 349-365 Ropgs, J. W.s A. P StickNry 1965. Reproductive cycle of Mya arenaria in New England. Biol. Bull. 128 (2): 315-327 Watng, P. R. 1970. The seasonal variation of meat and glycogen content of seven populations of oysters, Ostrea edulis L. and a review of the literature. Fish. Invest. Ser II 26 (3): 1-95 West ey, R. E. 1961. Selection and evaluation of a method of quantitative measure- ment of oyster condition. Proc. Natl. Shellfish. Assoc. 50: 145 - 149 Witson, Barry R. « E. PR Honexin 1967. A comparative account of the reproductive cycles of five species of marine mussels (Bivalvia: Mytilidae) in the vicinity of Fremantle, Western Australia. Austral. Journ. Mar. Freshwater Res. 18: 175 - 203 Internat. Biol. W. B. Saunders Co., Page 188 THE VELIGER Vol. 22; No. 2 Life-Cycle Completion of the Freshwater Clam Lasmigona compressa (Bivalvia : Unionidae) in an Experimental Host, Lebistes reticulatus ALEX S. TOMPA Division of Molluscs, Museum of Zoology, University of Michigan, Ann Arbor, Michigan 48109 THERE ARE ONLY A FEW WORKS on the known hosts of North American freshwater bivalves of the family Unioni- dae, and in most cases the host is a freshwater fish (the occurrence of Simpsoniconcha ambigua on the mudpup- py, Necturus, is an exception), although in a few cases, such as with Strophitus undulatus, Anodonta zmbecilts and Obliquaria reflexa, there may be no fish host and development proceeds directly (FULLER, 1974). Lists of fish hosts of the various bivalves are scattered widely in the literature, but are summarized in LEFEVRE & Curtis (1912) and especially in FULLER (1974). Hosts of the clam investigated in this study, Lasmigona compressa (Lea, 1829), are not mentioned in these stud- ies, nor does the present work deal with the normal hosts for this species. The purpose of this paper is to call attention to a system of glochidial infection and metamor- phosis which is easy to study in the laboratory, since the clam is abundant in its range. The experimental fish host is the easily obtained pet guppy fish, Lebistes reti- culatus, and the whole life cycle (7.e., maturation of the larval clam) takes less than 2 weeks. Since attachment of larval hooked glochidia occurs on the fins and not on the gills, the infection can be easily observed with a dis- secting microscope. The fins infested with glochidia can be easily removed for more detailed examination at any time during development by simple excision. Adult Lasmigona compressa were collected from the Huron River, in Washtenaw County, Michigan, in Au- gust, 1978. These clams were maintained in a 72 L capa- city aquarium with river vegetation. The clams were given Tetramin flakes of fish food with the expectation that they would feed upon this; previous experience in this laboratory had shown that various clams do well on this diet for 6 months and longer. Female clams of this species were found to pass glochi- dia — as well as orange colored ova — into the water in December, for a period of one week. Free glochidia were taken from the bottom of the aquarium with a long pi- pette rather than being dissected out of the gills as per- formed by other workers (LEFEVRE & CurTIS, 1912), for it was found that glochidia removed by dissection were noninfective. The glochidia are 320 umX260 4m when closed, and each hook (one per valve) is go um long. The hooks are studded with rather sharp little spines, 7 4m in diameter, 3 per row for much of the hook’s length. About 100 glochidia were placed into each shallow Petri plate with water, and male and female guppies were allowed to swim in this for a period of 2-5 minutes, during which they were continuously examined under a dissecting microscope. This interval of time was sufficient to allow an average of 10 glochidia to attach to each fish. A heavier infection was thought to be undesirable because of the small size of these fish. Similarly, goldfish of 5 cm length were also exposed to glochidia in separate containers. Usually, 10 fish of each species were used per experiment. Vol. 22; No. 2 THE VELIGER Page 189 The goldfish were found to attract glochidia, causing the latter to attach to the fins, but the infections were not successful; these larval clams fell off or were shed in 2 days and died. The first infections of guppies were successful in the sense that glochidia attached well to all the fins. But after the infected guppies were isolated and kept in aerated tap-water, every one of 10 fish developed fungus infections at the sites of attachment and the glochidia died and fell off in 4-6 days. Not a single larva meta- morphosed under these conditions. Uninfected fish kept in the same water remained completely healthy and free of fungus. A separate group of guppies was then infected, but place into aerated tap-water into which Furazone (Dy- na-Pet, Campbell, Calif.) was dissolved at the concen- tration recommended for fish by this company to dis- courage bacterial and fungal growth. Under these con- ditions, glochidia remained attached to all the guppies, and the former underwent successful metamorphosis. The criterion for metamorphosis is only generally defined, but involves the young clam falling off or detaching itself from the fish fin after showing some adult features, es- pecially the development of a large, muscular foot with which the young clam now crawls. Gills are also visible at this time. No significant growth was seen during this period. The time for completion of metamorphosis under these laboratory conditions, at room temperature (20° C) was 10-12 days with almost 100% success. When the glochidia finally detached from their hosts, they were placed into aerated tap-water, containing the small freshwater diatom, Navicula pelliculosa. The glo- chidia kept in this water inside a 50mL glass-stoppered flask lived thus and grew for 2 weeks, after which the experiments were terminated. During this time, a new growth-margin developed on each clam shell, amounting to an added new width of 46 um. This new growth area was characterized by a smooth surface, unlike the orig- inal glochidial shell surface, which is coarse, and studded with large calcium carbonate crystals. A few simple experiments were conducted on the newly shed glochidia, which apparently live only one or two days unless they find a suitable fish host. Under normal, quiet conditions, the glochidia lie with the 2 valves spread widely apart, ventral side upward; these animals are nor- mally motionless. Several guppies were ground up in a Waring blender, and the supernatant of this fluid was added dropwise to a dish containing quiescent glochidia. One drop of such fish fluid in 20mL of water was enough to cause most glochidia to snap a few times, after which some stayed closed for several minutes. In 10 minutes all these glochi- dia were open again. The amino acid, L-glutamic acid, was then dropped as crystals into these containers, near the glochidia. Again, a positive response was elicited, as shown by the snapping or snapping-shut of these larvae. Hearp & HENpRIX (1964) found that 20 amino acids, but not the 3 basic amino acids, arginine, histidine and lysine, elicited glochi- dial response in the clams which they tested. Finally, the responsiveness of glochidia to physical stimulation was tested. A very fine, etched piece of tung- sten wire was used to touch various areas of the glochidi- um, to see if pure mechanical stimulation is enough to elicit contraction, 7.é., attachment behavior, and if so, to see which part of the glochidium is most sensitive. It was found that the glochidia are indeed sensitive to mechanical stimulation, without prior chemical sensiti- zation. The most sensitive part of the glochidium was in a swelling near the hinge line, where a larval thread has been shown to originate in other glochidia (see Woop, 1974). It is possible that glochidia of other species could also infect the guppy under similar laboratory conditions. Such a system as this one, with Lasmigona compressa and the guppy, followed by feeding with Navicula pelli- culosa, offers much promise for more detailed studies of fish susceptibility, hardiness to heavy infections, for pos- sible class demonstrations of the life cycle of freshwater clams, and especially in elucidating details of how clams metamorphose on freshwater fish. Such studies are all the more important since it is likely that this early larval life and immediate post-metamorphosis may be the most critical period in the life-cycle of freshwater clams. If such larval and young clams are more sensitive to aquatic pollution of various kinds than are adult clams, then this would explain why so many endangered areas of rivers have adult clams only and lack any (re- cruitment of) young. More detailed studies along these lines could be of great help in re-establishing clams into recently cleaned up rivers and ponds using some method of commercial or laboratory rearing of the young. Voucher specimens of Lasmigona compressa and the in- fected fish have been deposited in the collections of The University of Michigan, Museum of Zoology. Page 190 THE VELIGER Vol. 22; No. 2 Literature Cited Futter, S. 1974. Clams and mussels. In: C. Hart « S. Fuuuer, eds. Pollu- tion ecology of freshwater invertebrates. Acad. Press, N. Y. pp. 215-273 Hearp, W. « S. HENDRIX 1964. Behavior of unionid glochidia. Amer. Malac. Union Bull. 31: 2-4 (abstract) (1 December 1964) Lerevre, G. « W. Curtis 1912. Studies on the reproduction and artificial propagation of fresh- water mussels. Bull. U.S. Bur. Fish. go: 105 - 201 Woop, E. 1974. Development and morphology of the glochidium larva of Ano- donta cygnea. Journ. Zool. London 178: 1-13 Vol. 22; No. 2 THE VELIGER Page 191 Philippia (Psilaxis ) radiata: Another Indo-West-Pacific Architectonicid Newly Found in the Eastern Pacific (Colombia) BY ROBERT ROBERTSON Department of Malacology, Academy of Natural Sciences, Nineteenth and the Parkway, Philadelphia, PA 19103 (1 Plate; 1 Map) INTRODUCTION In 1976 I REPORTED the discovery of the architectonicid Heliacus trochoides (Deshayes, 1830) on the mainland coast of Ecuador (RoBerTson, 1976b). The species had previously been known only between South Africa and the Marquesas. The present paper records a similar ex- tension of known range for a second architectonicid, Philippia (Psilaxis) radiata (Roding, 1798). This species too had been known only between South Africa and the Marquesas (RoBERTSON, 1970, fig. 5). The new record of it is from a single empty shell collected in September 1977 at Gorgonilla, Gorgona Island (2°57’N; 78°12’W), about 30km off the mainland Pacific coast of Colombia, 6500km ENE from the Marquesas. ACKNOWLEDGMENTS I am grateful to the malacological group of the Univer- sidad del Valle, Cali, Colombia (Jaime Ricardo Cantera Kintz, Francisco Borrero, Rafael Contreras, Elizabeth Buttkus and Fernando Zapata) for donating the speci- men to the Academy of Natural Sciences of Philadelphia (ANSP 348879) and for allowing me to publish this paper on their discovery. The work was supported in part by National Science Foundation Grant DEB76- 18835. COLOMBIAN SHELL (Figures 1 to 3) Description: The fairly thick, brown and white shell is 12.2mm high and 16.0mm wide, with 4.9 teleoconch whorls. The outline is trochiform, with the spire fairly high and rounded in profile. The spire angle steadily decreases from about 130° to about 100°, but cannot be measured accurately because of the roundness. The whorls are moderately inflated. The abapertural half of the shell is much more corroded than the apertural half. The protoconch is fairly large (1.2 to 1.3mm in im- mersed width), with a short anal keel. (Both these char- acters distinguish Philippia radiata from the closely sim- ilar species P oxytropis A. Adams, 1855.) The protoconch is not visible through the narrow teleoconch umbilicus. The apical side of the teleoconch is smooth, and mainly brown with numerous radii extending to the periphery (about 28 on the last whorl) (the name radiata came from these radii). At the periphery there is one spiral cord bordered on either side by a fainter cord. The upper suture is attached to the lower side of the major peri- pheral cord. The basal side of the teleoconch is mainly smooth. Outwardly there is a wide brown spiral band. Inwards from this are some brown spiral lines. There are two periumbilical sulci; the cord between them is checkered with about 20 brown marks. There are irregular peri- umbilical nodes (about 12 of them on the last whorl) ; these are white (or tinged with brown near the aper- ture). The umbilicus is narrow (1.5mm wide). (Philippia oxytropis would have a wider umbilicus.) The umbilical wall is recessed, and is smooth except for axial growth lines. The umbilical suture is outside the periumbilical sulci. The outer lip is straight in profile, prosocline, at an angle of 20° to the axis of coiling. The aperture is more or less oval in outline. There is a deep channel opposite the periumbilical nodes, a shallower channel opposite the cord between the periumbilical sulci, and a faint third Page 192 NS , THE VELIGER equator e TORS o) fe °S N° Adee Port Elizabeth South Africa channel the other side of the deep channel. Periostracum thin, pale yellowish brown. Operculum missing. DISCUSSION anp CONCLUSIONS This constitutes the first record of the genus Philippia Gray, 1847 in the Eastern Pacific (RoBERTSON, 1976a). The species is in the subgenus Psilaxis Woodring, 1928 (RoBERTSON, 1970,1973) and is definitely Philippia (Psil- axis) radiata, not the closely similar Pacific Ocean spe- cies P (PR) oxytropis or the Atlantic Ocean species P (P) krebsiu Morch, 1875. Protoconchs in the subgenus Psilaxis are noteworthy for their large size (1.2- 1.8mm in diameter). The veli- ger larvae remain for a long time in the plankton and are transported great distances by near-surface ocean currents (ROBERTSON, SCHELTEMA & ADAMS, 1970). It is likely that the Colombian animal was transported as a veliger from Polynesia — just as was hypothesized for the Ecuadorian Heliacus trochoides (RoBERTSON, 1976b). There is no indication that breeding populations of ei- ther species are established in the Eastern Pacific. Adult Philippia feeds on scleractinian corals (RoBERT- SON, SCHELTEMA & ADAMS, 1970). The empty shell of P Philippia (Psilaxis) radiata g I Vol. 22; No. 2 ~en O © Hawaii @ e @ Line Islands Isla e Galapagos Gorgon; @ e Colombia eis © Marquesas @ @ | % ooo km ZE 2 Figure 4 Geographic records of Philippia (Psilaxis) radiata (Réding) from RoserTson, 1970: fig. 5 and the Colombian record herein. The two literature records from the Gulf of Suez are probably erroneous and have been omitted radiata was collected at Isla Gorgona under a stone on a muddy rock substratum. Some of the corals around the island are species of Pocillopora, Porites and Pavona. EMERSON (1978) has published a useful review of the mollusks with Indo-Pacific faunal affinities in the Eastern Pacific. CosEL (1977) recorded Mitra mitra, another Indo-West-Pacific gastropod from Isla Gorgona. The paucity of marine species common to the Indo- West-Pacific and the Eastern Pacific means either that there is a barrier to larval dispersal east to west and west to east or that larvae are transported the requisite dis- tances but encounter ecological barriers which prevent the successful establishment of breeding populations. The lat- ter explanation seems to hold for Heliacus trochoides, Philippia radiata, and other species known to have long- distance larvae. It is noteworthy that no Eastern Pacific species seems to occur sporadically in Polynesia (the cur- rents flow prevailingly in the westward direction). Tue VELIGER, Vol. 22, No. 1 [RoBERTSON] Figures 1 to 3 Figure 2 Philippia (Psilaxis) radiata (Réding) Isla Gorgona, Colombia. Shell 16.0mm wide Figure 1: apertural view Figure 2: apical view Figure 3: basal view 1 rae Vol. 22; No. 2 THE VELIGER Page 193 Literature Cited CoseL, Rupo von 1977. First record of Mitra mitra (Linnaeus, 1758) (Gastropoda: Prosobranchia) on the Pacific coast of Colombia, South America. The Veliger 19 (4) 422-424; 1 plt.; 3 text figs. (1 April 1977) Emerson, WILLIAM KeziTH 1978. Mollusks with Indo-Pacific faunal affinities in the Eastern Pa- cific Ocean. The Nautilus 92 (2): 91-96 (27 April 1978) RoserTSon, RoBert 1970. Systematics of Indo-Pacific Philippia (Psilaxis), architectonicid gastropods with eggs and young in the umbilicus. Pacif. Sci. 24 (1): 66-83; 17 text figs. (January 1970) 1973. On the fossil history and intrageneric relationships of Philippia (Gastropoda: Architectonicidae). Proc. Acad. Nat. Sci. Philadel- phia 125: 37-46 1976a. Faunal affinities of the Architectonicidae in the Eastern Pacific. Bull. Amer. Malac. Union (1975) 41: 51 1976b. Heliacus trochotdes: an Indo-West-Pacific architectonicid newly found in the Eastern Pacific (mainland Ecuador). The Veliger 19 (1): 13-18; 1 plt.; 1 map (1 July 1976) RosrrTson, Rospert, Ruportr S. SCHELTEMA & FRANK W. ADAMS 1970. The feeding, larval dispersal, and metamorphosis of Philippia (Gastropoda: Architectonicidae). Pacif. Sci. 24 (1): 55-65; 7 text figs. (January 1970) Page 194 THE VELIGER Vol. 22; No. 2 The Packaging of Ova in the Egg Cases of Aplysia californica PAUL KANDEL anp THOMAS R. CAPO Division of Neurobiology and Behavior, Departments of Physiology and Psychiatry, College of Physicians and Surgeons, Columbia University, New York, New York 10032 (5 Text figures) INTRODUCTION THE MARINE MOLLUSK Aplysia is a prolific egg-layer. Large animals are known to lay about 5 X 10° eggs during the several months of their reproductive life (MacGun1- TIE, 1934). Egg-laying behavior has other interesting features. Animals generally do not lay small amounts of eggs continuously, but deposit a large number in all-or- none episodes (KUPFERMANN, 1970; KANDEL, 1976). In addition, egg-laying is one of the rare examples of a be- havior in which the neurobiological substrate underlying it is relatively well understood (for reviews see KUPFER- MANN, 1972; KANDEL, op. cit.). A hormone controlling egg-laying is released by 2 cell clusters called bag cells; each cluster contains about 400 neuroendocrine cells. The active egg-laying hormone released by the bag cells is a polypeptide with a molecular weight of approximately 6000 dalton (Torvs & BRAcKENBURY, 1969; GAINER et al., 1974; ArcH, 1972). A 25000 dalton precursor poly- peptide is produced in the cell bodies and is thought to be converted to a small molecule that is transported along the processes of the bag cells for release (ArcH, 1972). In response to an appropriate stimulus, the bag cells discharge synchronously, causing the hormone to be released into the neurohemal sheath (KuUPFERMANN, 1970; KUPFERMANN & KANDEL, 1970; ARCH, 1972; PINSKER & DUDEK, 1977; DUDEK & BLANKENSHIP, 1977). The hormone then appears to cause muscle fibers in the ovotestis to contract, thereby triggering the first step in the release of eggs (CoGGESHALL, 1967). Once fertilized, the ova are packaged in irregular shaped egg cases (or capsules), each containing several ova. The egg cases are strung together and deposited as an egg ribbon or cordon (Figure 1). We have analyzed certain aspects of egg-laying in wild specimens of Aplysia californica in the laboratory and have examined the fac- tors that determine the varying number of eggs that are Egg Capsule HO rN ry) Mucous Layer —{| ese ae, Figure 1 Egg laying in Aplysia californica. The adult deposits egg masses consisting of a long string of egg capsules, each containing varying amounts of eggs (after KriecsTEIN et al., 1974) packaged in each egg case by different animals of the species. Two aspects of this problem have often been com- mented upon (for review see BRIDGES, 1975; SWITZER- DuN Lop & HapFIELD, 1977). First, what accounts for the considerable observed variability in the number of eggs per capsule in the egg masses of different individuals of the same species? Thus Swirzer-DUNLop & HADFIELD (op. cit.) found 5 to 7 eggs per capsule in A. dactylomela, Vol. 22; No. 2 THE VELIGER Page 195 while OsTERGAARD (1950) found 7 to 15 eggs per capsule. Are these differences related to age, individual variations or nutritional state of the parent? Second, what accounts for the marked differences in the number of eggs per cap- sule between the various species of Aplysia? BripcEs (1975) suggests that among the Aplysiidae an inverse relationship exists between the diameter of the egg and the number of eggs per capsule. In this paper we have used size of the animal as an index of age and examined the first question. We found that the number of eggs per capsule is directly related to the size (and therefore presumably to the age) of the animal. Small and young animals invariably package fewer eggs per capsule, whereas larger and older animals package many. METHODS Wild animals collected off the coast of California were shipped to New York and maintained in 720L capacity, temperature controlled, recirculating aquaria at 16 to 20° C. Animals were fed a daily portion (about 10% wet body weight) of the algae Ulva and Gracilaria. No spe- cial measures were taken to induce spawning. Spawn was collected daily, freed of excess water, and weighed. Three lengths of cordon, a minimum of 1ocm, were then measured. The weight of each section was deter- mined as rapidly as possible and the mean weight per centimeter calculated. The mean number of ova per cap- sule was determined ty triplicate counts on 3 randomly chosen sections (9 observations). The section to be count- ed was placed on a microscope slide, depressed with a cover slip and counted with a binocular microscope at 256 magnification. To determine the number of capsules per centimeter a minimum of 3 0.5cm portions were cut from the cordon and centrifuged at 2000 rpm (1000Xg) for 20 minutes to compact the ova. A squash preparation was made and 0.5m portions counted at 160X magnification. The mean number of capsules per centimeter was calculated. The diameters of cordon and ova were determined with the binocular microscope fitted with an ocular microme- ter; a standard procedure of rounding off partial units was adopted. Three cordon portions were cut at random and kept immersed in sea water. Duplicate determina- tions of the diameter of each portion were made with a binocular microscope at 160 magnification. After the diameters of the cordon were taken, they were each covered with a cover slip, depressed, and the diameters of 3 ova chosen at random and from different capsules were taken at 400 X magnification. The mean number of ova per centimeter was the product of the mean number of ova per capsule and the mean number of capsules per centimeter. Coefficients of correlation (r) and lines of best fit were obtained from a least squares regression using a PDP11 laboratory computer. RESULTS Collections of spawn were made from (wild) animals, ranging in size from small (presumably young), weighing as little as 19g to large (presumably older) animals weighing to 1 300g. In the course of examining the spawn we found that the mean number of eggs per capsule of cordon ranged from 3 to 93. Because of this remarkably wide range we explored a number of variables to deter- mine which might affect how many eggs are packaged per capsule. We first examined the relation of number of eggs per capsule to body weight (a crude index of the age of the animals). Three sections of egg mass were examined and the mean number of eggs per capsule determined. In the 78 animals examined a linear and fairly steep correlation was found between animal weight and number of eggs per capsule (Figure 2). This correlation was highly sig- Mean Number of Ova per Capsule ° 150 300 450 600 750 900 1050 1200 1350 Body Weight (g) Figure 2 Mean number of ova per capsule as a function of body weight. Animals ranged in weight from 20g to 1300g and the ova varied in number from 3 to 93. Each datum point is the mean of g obser- vations on 1 animal (see Methods for details). There is a signifi- cant correlation (n=78; r=0.92; p< 0.01) between the number of ova per capsule and the body weight Page 196 THE VELIGER Vol. 22; No. 2 nificant: r=o0.92, p 0.01; slope =-0.38); beginning at 400g, the decrease is less (n> =43; T=0.77; p< 0.01; slope= —o.06). In this and in all subsequent figures, the dotted lines indi- cate an extrapolation of the plotted functions Part B: Mean number of ova per centimeter of egg string as a function of body weight. The number of ova increases throughout the reproductive life of the animal, but during later stages of the life cycle (> 300g; n=13; r=0.33; p=NS) the increase is less than it is early in the life cycle (20-300g; n=22; r=0.79; p <0.01) Part C: Mean diameter of the egg cordon as a function of the animal’s body weight. From 20g to 600g (n=54; r=0.59; p < 0.01). From 600g to 1300g (n=26; r=0.09; p=NS) Part D: Mean diameter of ova as a function of body weight (n= 40; r=0.38; p < 0.05) Vol. 22; No. 2 up to 400g the animal seems to compensate for the in- crease in number of eggs per capsule by increasing the volume of each capsule. Animals heavier than 400g seem to pack the ova more densely into capsules of similar size. Independent measurements of the number of ova per cm providesome of the support for thisargument (Figure 3B). As a check on this finding we explored 2 other vari- ables: the diameter of the cordon and the diameter of the ova. We found (Figure 3C) that as animals grew up to 500 g the diameter of the cordon increased about 31% presumably indicating an increase in the size of the egg capsules (n=54, r=0.59, p<_o.1). Beyond 600g the diameter did not increase significantly (n= 26, r=o.09, p=ns). This finding suggests that after a certain point the capsules remain about the same size or at least in- crease less. Although this finding could simply indicate that the same sized egg capsules are simply packed dif ferently, we also found that the size of the ova decreased slightly (about 13%) as a function of size of the animal (Figure 3D). Is there a correlation between the weight of the animal and the total number of eggs deposited? To answer this question we examined the weight of the spawn as a func- tion of weight of the animal and found a roughly linear correlation (n= 64, r—o0.70, p< 0.01) between these 2 variables (Figure 4A). Although there was a great deal of variability, small animals tended to lay small amounts of spawn (less than 1g), large animals tended to produce great amounts of spawn (more than 25g). When we next compared the relation of spawn as a percent of body weight to body weight (Figure 4B) we found that inde- pendent of size, animals produce egg masses that were about 2.1% of their body weight (with a range of 0.22 to 5.97%). DISCUSSION We have examined the egg laying capability of Aplysia californica as a function of age; using weight of animal as a crude index of age (see KRIEGSTEIN et al., 1974; CoccESHALL, 1976). Two key findings emerge from this study: 1) animals produce spawn throughout much of their postmetamorphic life. Animals as small as 19g (1.5% of maximum body weight in our samples) already produce viable ova; 2) the number of ova packaged per capsule varies systematically with aging. On the basis of these findings we propose a schema illustrated in Figure 5. As animals grow they package progressively more ova per egg capsule than do smaller animals. At first the increase in ova per capsule is com- THE VELIGER Page 197 n az S tof) ice) 25g as the ani- mals mature (n=64; r=0.70; p < 0.01) Part B: Egg mass as percent of total body weight (n=64; r=o0.11; p=NS) pensated for by an increase in size of the capsule (see middle age animals in Figure 5). However, beyond 600g the size of the capsule may not keep pace with the in- crease in the number of ova per capsule, resulting in a relatively tighter package of ova (old animals; Figure 5). The mechanisms that determine the density of packing and the consequences for the ovum of alterations in density are interesting questions that need now to be ex- plored. Page 198 THE VELIGER Vol. 22; No. 2 ‘S) 5 Ova 600 g Figure 5 Schematic drawing indicating changes in number of ova per capsule and inferred changes in packing density Larger animals also lay larger (more) spawn than do smaller animals. This increase in spawn is a reflection of the fact that animals generally lay egg masses of a rela- tively fixed average percentage —- about 2% of their body weight. ACKNOWLEDGMENTS We thank David Leibowitz, Vincent Castellucci, Carl Berg, and Eric Kandel for their comments on this manu- script, and Susan E. Perritt, James Sliney and Claire Advokat for technical assistance. This research was supported by a grant from the Esther and Joseph Klingenstein Foundation. Literature Cited Arcg, S. 1972. Biosynthesis of the egg-laying hormone (ELH) in the bag cell neurons of Aplysia californica. Journ. Gen. Physiol. 60: 102-119 Brivczs, C. B. 1975- Larval development of Phyllaplysia taylori Dall, with a discus- sion of development in the Anaspidea (Opisthobranchiata: Anaspidea). Ophelia 14: 161 - 184 CoccEsHALL, R. E. 1967. A light and electron microscope study of the abdominal gan- glion of Aplysia californica. Journ. Neurophysiol. go: 1262 - 1287 Dupek, F E. « J. E. BLANKENSHIP 1977. Neuroendocrine cells of Aplysia californica. I. Bag cells action potentials and afterdischarge. Journ. Neurophysiol. 40: 1g01 - 1318 Gainer, H. @ Z. WoLLBERG 1974. Specific protein metabolism in identifiable neurons of Aplysia californica. Journ. Neurobiol. 5: 243 - 261 KanbeL, Eric R. 1976. Cellular basis of behavior. cisco, California; 727 pp.; illust. KrizosTzIn, ARNOLD R., VincenT CastTetitucci e Eric R. KANDEL 1974. Metamorphosis of Aplysia californica in laboratory culture. Proc. Nat. Acad. Sci. 71 (9): 3654 - 3658; g text figs. KuPFERMANN, I. 1970. Stimulation of egg-laying by extracts of neuroendocrine cells (bag cells) of abdominal ganglion of Aplysia. Journ. Neurophysiol. 33: 877 - 881 1972. Studies on the neurosecretory control of egg-laying in Aplysia Amer. Zool. 12: 513 - 519 KuPFERMANN, I. & E. R. KANDEL 1970. Electrophysiology properties and functional interconnections of two symmetrical neurosecretory clusters (bag cells) in abdominal gan- glion of Aplysia. Journ. Neurophysiol. $3: 865 - 876 MacGrniT1z, Grorce EBER 1934. The egg-laying activities of the sea hare, Tethys californicus (Cooper). Biol. Bull. (Woods Hole) 67: 300 - 303 OSTERGAARD, JENS MATHIAS 1950. Spawning and development of some Hawaiian marine gastro- pods. Pacif. Sci. 4: 75-115 Pinsxer, H. e FE E. Dupex 1977. Spontaneous bag cell activity and egg laying in freely behaving Aplysia. Science 197: 490 - 493 Switzer-DunLaP, Marityn & MicHazL G. HapFietp 1977- Observations on development, larval growth and metamor phosis of four species of Aplysiidae (Gastropoda, Opisthobranchia) in laboratory culture. Journ. Exper. Mar. Biol a Ecol. 29: 245 - 261 Torevs, L. A. S. & R. W. BRACKENBURY 1969. Bag cell-specific proteins and the humoral control of egg-lay- ing in Aplysia californica. Comp. Biochem. Physiol. 29: 207 - 216 W. H. Freeman & Co. San Fran- Vol. 22; No. 2 THE VELIGER Page 199 Modiolus aurum Osorio, spec. nov., from Juan Fernandez Archipelago, Chile (Mollusca : Bivalvia : Mytilidae) CECILIA OSORIO R. Laboratorio de Hidrobiologia, Departamento de Biologia, Facultad de Ciencias, Universidad de Chile Casilla 147, Santiago-Chile (9 Text figures) INTRODUCTION In 1970 I RECEIVED 21 specimens of Modiolus from Juan Fernandez Archipelago. Specific identification was not possible, because of encrusting epizoans. In 1973 another sample of 97 specimens came in, not only without epi- zoans, but more representative and in better condition. Some additional specimens then were noted in the collec- tions of the National Museum of Natural History of Santiago. ODHNER (1922) cited “Modiola plumescens Dunker, 1868” as occurring on Juan Fernandez Island, but Soor- RYEN (1959) suggested that until confirmed by further collecting this species should not be included in Chilean faunal lists. The specimen mentioned by Odhner as in the Stockholm Museum seems now to be missing. However, other specimens, from Australia, determined by Odhner as M. plumescens, were available to me on loan. Also I examined material from ile Nou and Broome, Australia, now in the Museum of Paris. The type locality of Dun- ker’s species is Samoa, Viti-Ins, Uvea. Because Odhner had only a single specimen from Juan Fernandez on which to base his determination of “Modi- olus plumescens,” misidentification is understandable. The 2 forms have a similar outline. Table 1 shows the differ- ences between the Australian and the Juan Fernandez forms. As a result of the comparative study I conclude that there are 2 distinct species. For the Juan Fernandez form, which is therefore considered to be undescribed, I propose the name Modiolus aurum, in reference to the golden color of the shell. SYSTEMATIC TREATMENT MytTILDAE Modiolus Lamarck, 1799 (nom. con- serv., ICZN, 1955) Modiolus aurum Osorio, spec. nov. (Figure 1) “Modiola plumescens Dunker” of ODHNER, 1922: 221. Soot- RYEN, 1959: 25- Osorio & BAHAMONDE, 1970: 192. (Non Modiola plumescens DunKgr, 1868: 115; 1875: 175). Description: Shell mytiliform, relatively high; dorsal margin curved, with a pronounced dorso-posterior angle; anterior margin narrow, rounded at the tip; posterior margin broad, evenly rounded; ventral margin straight. Beaks evident, located slightly behind the anterior end, close together, ligaments not evident. External surface of shell with evident periostracum of a golden-yellow color with greenish and brownish hues; periostracum well de- veloped dorsally, becoming weak or wanting anteriorly; periostracal hairs of several patterns (Figure 2), type a-b occurring only near the umbones of larger specimens but over entire surface of small individuals; type c-d oc- curring only on the dorso-posterior end of large speci- mens; all hairs with a smooth axis, lacking denticulations. A heart-shaped lunule present, anterior to the beaks. Hinge area completely smooth within, broad (Figure 4), ligamental area elongate, slightly curved (Figure 4), originating behind beaks. Interior surface of the shell, hinge area and ligament ue; these were especially examined. The exhalant or dorsal branchial opening is oval, with thick, smooth, and Page 200 THE VELIGER Vol. 22; No. 2 Figure 1 Modiolus aurum Osorio, spec. nov. ; Figure 3 Lunule — Se Imm : 1mm left valve right valve a Imm Figure 4 a b e d Figure 2 Different forms of “hairs” of the periostracum of Modiolus aurum Muscle scars well marked, the posterior adductor sub- circular, continuous with retractor muscles, separated into 2 groups; anterior adductor small. According to Soot-RYEN (1955), the anatomical structures of the branchial openings have systematic val- well defined margins; the inhalant or ventral branchial opening is separated from it by a broad septum, the mem- brane prolonged laterally and slightly festooned, forming a fold that is continuous with the mantle lobules. Type Material: Holotype, in Museo Nacional de His- toria Natural, Santiago, Chile, no. 100230. Two para- types, A 100231, B 100232. One hundred additional para- types in the author’s collection. Dimensions: Size of holotype: length, 26.7 mm; width, 15.7mm. Paratype A, length, 26.3 mm, width, 15.6mm; Vol. 22; No. 2 THE VELIGER Page 201 height, mm 16P 14h 5 104 y =1.606 + 0.52 | ne oS at r =0.929 \ 8 12 16 20 24 28 32 3e""" length, mm Figure 7 Figure Seite Proportion of length to height Muscular scars A — retractor muscles Figure 6 Branchial openings in Modiolus aurum Paratype B, length, 21.8mm, width, 13.2 mm. Type Locality: Bahia Cumberland (Lat. 33°37’S; Long. 78°49'W). Date: 18 April 1973. Figure 8 Collectors: C. Moreno and J. Zamorano. Different forms of “hairs” of the periostracum of Substrate: on buoys or adhered to coastal rocks. Modiolus plumescens Dunker, 1868 Page 202 THE VELIGER Vol. 22; No. 2 Table 1 Morphological differences between Modiolus aurum and Modiolus plumescens. Species Characters Modiolus aurum Osorio, spec. nov. Modiolus plumescens Dunker, 1868 Colour Shells golden-yellow, with brown hues. Shells brown-olivaceous, with reddish hues. Lunule Heart-shaped, small, circumscribed by a deep Oval, big, occupying half of the total length, line (Fig. 3). circumscribed by a slight line. Periostracum “Hairs” of different form, axis smooth, without “Hairs” of more or less similar form, axis with denticulations or membranes and its free end with diverse forms (Fig. 2). Branchial openings (Fig. 6). Dorsal relatively large, oval, margins smooth and well defined. Ventral clearly separated from the dorsal opening by a septum. The lateral membrane of the ventral opening terminates between the two openings. denticulations in a row, and its free end more or less pointed (Fig. 8). (Fig. 9). Dorsal relatively small, oval with its inferior margin truncated. Ventral very near the dorsal opening, septum small. The lateral membrane of the ventral opening termi- nates by intruding into the dorsal opening. Depth: jects. o to 45m from littoral pools to submerged ob- Geographical Distribution: Juan Fernandez Archipelago: a) Robinson Crusoe Island in beaches of El Pan- gal, Palillo and Cumberland Bay. b) Alexander Selkirk Island (place of collection was not indicated). DISCUSSION Length-width relationships are shown in Figure 7. The linear regression was estimated, based on analysis of 168 specimens. The formula is: y==1.606 + 0.52x for the Juan Fernandez population. The largest specimen meas- ured 38.2 mm in length, 20.8mm in width. Morphologic differences between Modiolus aurum and M. plumescens may be shown as follows. Soot-RYEN (1955) lists 8 species for the western coast of America, from among which KEEN (1971) indicates only 3 for South America: Modiolus capax (Conrad, 1837) — Santa Cruz, Califor- nia to Payta, Peri Modiolus eiseni Strong « Hertlein, 1937 — Gorda Bank and Guaymas, Gulf of California to Pera Figure 9 Branchial openings in Modiolus plumescens Dunker, 1868 Modiolus pseudotulipus Olsson, 1961 — Bahia Magda- lena, Lower California to Pera. Vol. 22; No. 2 All these are from tropical waters and are distinctly different from Modiolus aurum in form of shell, perios- tracal hairs, lunule, muscle scars, and branchial openings. Only M. pseudotulipus has a similar outline, but the un- branched periostracal hairs, broad base, and the lateral membrane distinguish it clearly from M. aurum. ACKNOWLEDGMENTS The author expresses her thanks to Dr. K. Busse, Ham- burg University and to Professors C. Moreno, J. Zamo- rano, Universidad Austral de Chile, for the material col- lected. To Dr. P Arnaud, Endoume Marine Station, Marseille, who kindly sent bibliography; to Mrs. P. Bouchet of the National Museum of Natural History, Paris and A. An- dersson of the National Museum of Natural History Stockholm, for the loan of collection material. To Dr. W. O. Cermohorsky and Dr. R. B. Dell of the Auckland Institute and National Museum of New Zea- land, respectively; to R. P Kenny, School of Biological Sciences University, S. Stevenson, Natural History Muse- um of Victoria, Australia. THE VELIGER Page 203 To Dr. M. Keen, Stanford University and Professor N. Bahamonde for revision of the work; to Mrs. S. Mann for the translation into English and to Miss M. Cariceo for typing it; Dpto. Biologia, Faculdad de Ciencias, Uni- versidad de Chile. Literature Cited Cvessin, STEPHAN 1889. Mytilidae. In: EF H. W. Martini « J. H. Chemnitz, Systema- tisches Conchylien Cabinet. 20d ed. 8 (3): 1-170; 36 col. plts. Niirn- berg. Dunxer, G. WILHELM 1868. | Museum Godeffroy, Catalog IV: 115, Brit. Mus. (Nat. Hist.), London 1874. Museum Godeffroy, Catalog V: 175. Brit. Mus. (Nat. Hist.) London Ken, A. Myra 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru. 28d ed. Stanford Univ. Press, Stanford, Calif i- xiv+ 1064 pp.; ca. 4000 text figs.; 22 col. plts. (21 September 1971) Opuner, Nits HJALMAR 1921. Mollusca of Juan Fernandez and Easter Islands. In: C, SxorrsBerG, The natural history of Juan Fernandez and Easter Island 3 (2): 219-254; pits. 8, 9 Osorio, Ceciria & N. BAHAMONDE 1970. Lista preliminar de lamelibranquis de Chile. Hist. Nat. 31: 185 - 256, Minist. Educc. Santiago, Chile Soot-Rygn, TRON 1955. A report on the family Mytilidae (Pelecypoda). Allan Han- cock Pacif. Exped. 20 (1): 175 pp.; 10 pits. 78 text figs. Univ. So. Calif. Press, Los Angeles (10 November 1955) 1959. -Pelecypoda. Reprt. Lund. Chile Exped. (1948-49). Arssk. N. FE (2) 55, nr. 635: 1 - 86; 4 plts.; § text figs. Lund, Sweden Bol, Mus, Nac. Page 204 THE VELIGER Vol. 22; No. 2 Range Extensions of Mollusk Species Found on the Tropical Coast of El Salvador MARCO ANTONIO HERNANDEZ BY Curador de la Seccién de Malacologia y de Invertebrados Marinos del Museo de Historia Natural de El Salvador Tue coasts oF Ex SALvapor form part of the tropical Pacific coastline of Central America, making it part of the Panamic Province. This province extends from Cabo San Lucas (Gulf of California, approximately 30°30’N) to Cabo Blanco, north of Pera (4°15/S). It is possible that the coasts of this region are some of the most representative of the malacological fauna in the American Pacific. Stuarpo (1964) estimated the diversi- ty of this area to be 2200 species, including the gastro- pods, pelecypods, scaphopods and _ polyplacophorans. KEEN (1971), studying the same tropical zone, found that the diversity of mollusks is much higher and reports ap- proximately 3 317 species. Early in 1977, the Museo de Historia Natural de El Salvador started in a systematic manner a study aimed at an inventory of the malacological fauna of the country. To date, this work has identified close to 250 species, 26 of which are not previously reported in El Salvador (Tables 1 and 2). Generally, the species of mollusks that inhabit the coasts of El Salvador live in environments that are relatively uniform, with gradual climatological fluctuations. Sruar- Do (1964) suggests that among the environmental fac- tors having the greatest influence on these invertebrates is temperature. OLSSON (1961) reports the surface temperature of the waters of the Panamic Province as varying between 26.6° and 29.4°C, with the exception of temporary cold water upwellings. The coasts of El Salvador are characterized by their volcanic origin; some parts demonstrate the geological history of the area (GmERLOFF-EMDEN, 1971). The sandy beaches present a topography in which strong waves remove great masses of sand, and set up surface currents; for example, Costa del Sol, El Pimien- tal and Los Blancos. Of the rocky beaches, most representative are Los Cébanos and Maculis, Table 1 Extensions of range northward to the coast of El Salvador Species New northern record Tegula pellisserpentis (Wood) Turbo saxosus (Wood) Astraea buschitt (Philippi) Littorina varia Sowerby Cerithidea pulchra (C. B. Adams) Anachis rugosa (Sowerby) Microcithara cithara (Reeve) Cancellaria bulbulus Sowerby Conus patricius (Hinds) Anadara similis (C. B. Adams) Chama buddiana C. B. Adams Protothaca beili (Olsson) Tellina ecuadoriana Pilsbry & Olsson Strigilla disjuncta (Carpenter) Donax dentifer Hanley Corbula tumaca (Olsson) Periploma pentadactylus Pilsbry & Olsson Gastropoda Los Cébanos, Sonsonate E] Tamarindo, estuary E] Tamarindo, estuary Barra de Santiago and El Tamarindo E] Tamarindo, estuary Solimar; also E] Tamarindo Los Cébanos, Sonsonate Costa del Sol; also, El] Tamarindo Costa del Sol; also, El] Tamarindo Pelecypoda Los Cébanos; also, El Tamarindo Maculis, La Union E] Tamarindo, estuary E] Tamarindo, estuary E] Tamarindo, estuary Barra de Santiago; also, El Tamarindo E] Tamarindo, estuary Costa del Sol; La Paz Added in Proof Gastropoda Cyclothyca corrugata Stearns Playa El Zonte, La Libertad Vol. 22; No. 2 THE VELIGER Page 205 Table 2 Extensions of range southward to the coast of El Salvador Species New southern record Gastropoda Astraea olivacea Los Cébanos; Costa del Sol; El (Wood) Tamarindo Cerithium maculosum Los Cébanos, Sonsonate (Kiener) Planaxis obsoletus Los Cébanos and Maculis, La Menke Union Acanthina tyrianthina Los Cébanos, Sonsonate Berry Barra de Santiago and E] Tamarindo Costa del Sol and E] Tamarindo Solenostetra gatesi Berry Northia northiae (Griffith & Pidgeon) Fusinus ambustus Julupita, La Libertad, estuary (Gould) Pelecypoda Cardita affinis El Tamarindo and Maculis, La Sowerby Union Amphichaena kindermanni Metalio and El Tamarindo Philippi Added in Proof Pelecypoda Plicatula anomioides Keen __ E] Pital, La Libertad Los Cébanos is characterized by having the formation of a true reef. The beach is strewn with large quantities of organic material (shell fragments mixed with sand). In the Golfo de Fonseca (El Tamarindo, Playitas and Maculis) relatively tranquil waters provide favorable conditions for an abundant invertebrate fauna and some marine mammals. Literature Cited GrerLorr-Empen, H. G. 1971. La Costa de El Salvador. Prim. Edic. Direcc. Public. del Ministerio de Educc. El Salvador: 273 pp. Kazgn, A. Myra 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru, 2d ed. Stanford Univ. Press, Stanford, Calif. i-xiv+ 1064 pp.; ca. 4000 text figs.; 22 col. plts. (21 September 1971) Oxsson, AxEL ADOLF 1961. Mollusks of the tropical eastern Pacific, particularly from the southern part of the Panamic-Pacific faunal province (Panama to Peru). Panamic-Pacific Pelecypoda. Paleont. Res. Inst. Ithaca, N. Y, 574 pp., 86 pits. Stuarpo, Josz 1964. Distribucién de Ios moluscos marinos litorales en Latinoamérica. Bol. Inst. Biol. Mar. del Mar del Plata no. 7: 79-91 Page 206 THE VELIGER Vol. 22; No. 2 A Re-Evaluation of the Northwestern Range of the Melongena corona Complex ISABEL C. LEVY (3 Text figures) INTRODUCTION Melongena corena Gmelin, 1791 is a common species along the coasts of Florida and eastern Alabama (CLENCH & TURNER, 1956: 171-172; plts. 100-105). This mollusk inhabits oyster barsand low energy intertidal zones. HatH- AWAY (1957) completed the most thorough biological study of M. corona. A systematic review of the Melon- genidae of the Western Atlantic was published by CLENcH & TURNER (op. cit.), who described the M. corona com- plex. The Melongena corona complex consists of 3 morpho- logically different and geographically isolated subspecies: M. c. altispira, M. c. corona, and M. c. johnstonei. The genus as a whole is a highly variable group especially in terms of shell coloration and spination (CLENcH & TurR- NER, 1956: 171-172) (Figure 1). Melongena corona altispira is distributed along the east coast of Florida, from St. Augustine to Miami. This 30° Melongena corona altispira '—— Melongena corona — johnstont Melongena : aroha Florida corona 24> Figure 1 Map of occurrence of subspecies of the Melongena corona complex, according to CLENCH & TURNER, 1956 subspecies is characterized by a high ratio of shell length to shell width (CLENcH « TURNER, 1956: 176-178; plt. 104). The range of Melongena corona corona extends from Cape Sable, Monroe County, to Keatons Beach, Taylor County, Florida; M. c. johnstonei is distributed from Panacea, Wakulla County, Florida to Gulf Shores, Bald- win County, Alabama. Apalachee Bay was described as an area of overlap where a clinal relationship between the 2 subspecies occurs (CLENCH & TURNER, 1956: 172). (Figure 1) Spination is the primary morphological character used to differentiate between Melongena corona corona and M. c. johnstone: (Figure 2). The former subspecies has 2 rows of spines on the shell, a shoulder row and a basal row. The latter has only a shoulder row of spines. Coloration was used as a secondary character for subspeci- shoulder spines Figure 2 Subspecies of Melongena corona of the Gulf of Mexico a — Melongena corona corona b — Melongena corona johnstonei Vol. 22; No. 2 fic classification (CLENCH & TURNER, 1956: 173-174, 178; plts. 94-97, 100-103, 105). In my preliminary studies shell spination and colora- tion were shown to be variable characters throughout the range of the 2 subspecies. The objective of this study was to determine the actual value of spination and coloration as taxonomic characters for the 2 subspecies: Melongena corona corona and M. c. johnstonez. METHODS Melongena corona were collected at 5 stations along the west coast of Florida: Pensacola (Escambia County), Port St. Joe (Gulf County), Shell Point (Wakulla Coun- ty), and Seahorse Key (Levy County) (Figure 3). Samples from Pensacola and Wakulla Beach were from Thalassia and Spartina habitats, respectively. Sandflats were sampled at Port St. Joe and Shell Point; at Seahorse Key collections were made on oyster bars. Sampling was done by collecting all Melongena corona found in randomly selected areas at each station. All col- lections were done during low tide when the majority of snails were active (not buried in the sand). Sex, shell length, shell width, shoulder width, shell color, spire length, and spination were recorded for each snail. Spination data consisted of the presence or absence of the basal row of spines and number and size of spines. Table 1 shows the location of the stations sampled, sample size, as well as the percentage at each station of individ- uals with basal spines. THE VELIGER Page 207 Sex of the snail was determined by secondary sexual characters. A prominent yellow area around the entrance to the pedal gland distinguishes mature females. The penis of males is located on the right side of the head. Im- mature females lack both characters mentioned. Non-parametric tests for equality of proportions were carried out on the binomial data obtained. Relation- ships between the treatments (geographical location, snail size) and the effect (spination) were thus investigated. Coloration could not be quantified due to the extreme variability. Post-hoc tests were completed on all significant chi- square values. This type of test involved pairwise compari- sons between all treatment levels. In this manner the levels that caused significant frequency differences were identified. RESULTS anp DISCUSSION Table 2 summarizes the criteria used by CLENcH & Tur- NER (1956) for the subspecific classification of Melon- gena corona corona and M. c. johnstonet. The stations sampled in this study included the range of both sub- species and the cline between them. The cline theory by CLENCH & TURNER (op. cit.) was not supported by my data. Shell color (banding) was found to be extremely vari- able at all locations sampled. The variability found among populations was similar to the variability represented Florida Panhandle Pensacola, _ t N e Collection site Figure 3 Florida Panhandle. Collection sites of this study Page 208 THE VELIGER Vol. 22; No. 2 within each station. Consequently, coloration was con- sidered an unreliable character for this species. No difference was found in the proportion of shell shoulder width to spire length among stations sampled. Shell spination, although variable, was quantified at all locations studied. The basal row of spines (character- istic of Melongena corona corona) was found through- out the ranges of both subspecies and their cline (Table 1). Table 1 Sample % with Station Location size basal spines A Pensacola 53 26% Escambia County B Port St. Joe 74 10% Gulf County C Shell Point 52 79% Wakulla County D Wakulla Beach 61 40% Wakulla County E Seahorse Key 51 55% Levy County The chi-square values of non-parametric tests for equal- ity of proportions for spination as related to snail size were significant (0.01 level). In other words, the percentage of snails with basal spines is different according to size. Post-hoc tests of these data demonstrated that snails longer than 6.1cm exhibit higher frequencies of basal spination than snails below 6.0 cm. Similar tests were run on shell spination with respect to geographic location. The analysis of these data showed that basal spine frequencies are different due to geo- graphic location, the chi-square value was significant to the o.o1 level (Table 2). No trends were apparent. The sample from Pensacola, Florida, was analyzed for sex-linked spined morphology. No statistically significant differences were found between the sexes (26.9% of males and 23.8% of females had basal spines). The results draw attention to the fact that individuals with basal spines are not confined to the range of Melon- gena corona corona, as defined by CLENCH & TURNER (1956), but are also found throughout the range of A. c. johnstonei. Spination is not sex-linked, but is related to snail size and geographic location. Field data obtained on mating pairs included all possible combinations be- tween spined and non-spined individuals. Characters previously used for subspecific classification, other than spination, display high degrees of intrasub- specific variability and lack of intersubspecific character exclusion. Therefore, they can not be used to define the subspecies. CONCLUSIONS Because of their sedentary habit, encapsulated develop- ment, and absence of planktotrophic larvae, Melongena Table 2 Melongena corona corona Melongena corona johnstonet 1. Shoulder is usually horizontal and narrow 2. Shell reaching 205 mm 3. Color. Ivory with spiral orange- brown bands, very variable 4. Spire subdepressed to extended 4. Spire extended 5. Usually there is a single row of rather large spines at the base of the whorl. In other colonies the spines may be reduced to a single row at the shoulder whorl, the basal row being absent 6. May have two smaller rows of 6. Not mentioned spines between the ones mentioned above 1. Whorls shouldered 2. Shell reaching 171 mm Stations A-E 1. Within a given population both types were found 2. Continually growing species 3. Color. Ivory with two or three 3. Within a given population all color bands of dark brown morphs for both subspecies were found 4. The spire was subdepressed to extended 5. Sculpture consisting usually of a 5. All areas studied showed significant single row of strong, erect or recurved spines on the margin of the shoulder whorl. Rarely there exists a rather weak series of spines near the base of the shell percentages of individuals with basal spines 6. Not found in areas studied Vol. 22; No. 2 THE VELIGER Page 209 corona populations have a tendency to remain within the same area. Ege capsules that can be moved by mechanical means or tides are the most probable mechanisms of dis- persal. Prior to construction of the intercoastal waterway, the only potential barrier to tidal dispersal in northwest Florida was the area between West Bay and Destin, where for approximately 80km there are high energy beaches and no interconnecting marshes. Low energy beaches are encountered again in the Destin area, Oka- loosa County, and continue along Santa Rosa Sound to Gulf Shores, Baldwin County, Alabama. Melongena corona thrives in all appropriate habitats throughout Florida and Alabama. Due to the snail’s in- herent variability, small semi-isolated populations may be slightly different from each other. The Melongena co- rona complex possibly represents a polymorphic species, as opposed to a polytypic one. Selection for certain physiological adaptations, which may or may not be apparent, are possibly due to specific environmental conditions. If the adaptations are pheno- typically apparent, such morphs, as well as their delimita- tions are then as difficult to describe as the delimitations of their causative factors (Mayr, 1969). Melongena corona populations from southern Florida may display a third row of spines, between the shoulder and basal rows. This character was not found in this study. The southern range of the species should be re- evaluated. Data from the present study indicate that CLENCH & TuRNER’s (1956) descriptions of subspecies of Melongena corona from the northeastern Gulf of Mexico are not valid. My data show that subspecific classification is not warranted by the characters previously used. ACKNOWLEDGMENTS I express my appreciation to Mr. Dan O'Leary, Jr., of Crawfordville, Florida, for allowing me to study his Mel- ongena corona collection. Many thanks to Ms. Karen Swift and Ms. Sandra Mossburg for their help in the col- lection of specimens. I am also grateful to Dr. Jerry Oglesby for his advice on statistical procedures and Mr. Ray Johnson for his aid in the analysis of the data. This research was partially supported by an Environmental Protection Agency grant to Dr. Paul V. Hamilton. I am most grateful to Dr. Charles N. D’Asaro for encouraging me to write this paper and for the long hours he spent editing it. Literature Cited Cuzncw, Wiruam James « Ruts Dizon Turner 1956. The family Melongenidae in the western Atlantic. Johnsonia 3 (35): 161 - 188 Hatuaway, R. R. 1957. Studies on the crown conch, Melongena corona. Master’s thesis, Florida State Univ., Tallahassee, Florida, 95 pp. Mayr, ERNST 1969. Principles of systematic zoology. xi+42B pp.; iltust. New York, McGraw-Hill, Page 210 THE VELIGER Vol. 22; No. 2 Rediscovery of the Holotype of Cymatium (Cymatium) ranzanu (Bianconi, 1850) (Gastropoda: Prosobranchia ) BY BRUNO SABELLI', MARCO TAVIANI* anp STEFANO TOMMASINI' (1 Plate) DuRING A RECENT REARRANGEMENT Of the malacological collections in the Zoological Museum of the University of Bologna, Italy, we found the mollusks collected by G. Fornasini from Mozambique. This material was studied by the Bolognese naturalist Giovanni Giuseppe Bianconi (1809 - 1878), who published on it during the period 1847 - 1869 in “Specimina Zoologica Mosambicana.” He described some new species among which was a rare gast- ropod, a Cymatium. This was described as a new species by Bianconi February 21, 1850 during a meeting of the Accademia delle Scienze dell’Istituto e della Societa di Bologna (Branconi, 1850) in the following diagnosis: “Triton Ranzanu, Bianc. “— T, Testa fusiformi subtrigona transversim obscure sulcata, et striata, flavo rufescente, anfractibus superne angulatis, ultimo subtriangulari, ad angulum tuberculo instructum; latere sinistro bituberculato, dextro incavato; fauce postice apecta; columella nigro maculata, cauda recta longiuscula.” The following year he published the 4™ part of Specimina Zoologica Mosambicana (Bian- cont, 1851), figuring the specimen and adding to the original diagnosis this description: “Descriptio. Conchylium fusiforme, oblongum, subtrigon- um. In spira ejus octo circumvolutiones ad mediam cari- natas, declives, superne striatae, subtus vix concavae nu- merantur. Tubercula depressa inaequalia, et inaequaliter disposita super spiram, quaemadmodum super volutam exteriorem. Haec voluta, spiram excedens, retro valde carinata est; carina vero rotundata, quae in triangulum transversim evolvitur. Tria tubercula, unum in unoquo- que trianguli angulo, duo in latere sinistro. Latus dexter- um caret tuberculis, immo profunde concavum est. Super- fictes ultimae volutae tuberculis caret; tantummodo strus ornatur et sulculis, inter costulas sitis ut in Trit. femorale, at vix elevatis. Faux triangularis, oblonga, in latere dex- tero sulcata, postice valde angulata. Ejus margo dexter aliquantulum dilatatus, et undulatus; canalis omnino rec- tus; columella recta inferne, incavata postice, et inflexa. Color in profunda fauce violaceus: maculae brunneo- nigrescentes marginem inspergunt; quae majores in colu- mella, quam usquam alibi. Facies externa, si amoveas epi- dermidem sive indumentum marinum filamentosum, fla- veola est, infults brunneis distincta.” Although the original figures of Bianconi are very fine, we think it useful to give a photograph of the holotype. In fact by comparing the 2 figures (Figures 1, 2), some differences are evident. The siphonal canal is longer and more curved than in Bianconi’s figure; the aperture is apparently narrower and the upper part of the outer lip Explanation of Figures 1 to 5 Figure 1: Cymatium ranzanit, holotype, Museum of Zoology of Figure 3: Original label accompanying the holotype, slightly en- the University of Bologna XI larged Figure 2: Original drawing of Cymatium ranzani from Biancont Figure 4: Cymatium ranzanii, holotype, apical view, Museum of (1851) XI Zoology of the University of Bologna x1 Figure 5: Original drawing of the apical view of Cymatium ran- zanit from Bianconi (1851) X11 ‘Istituto e Museo di Zoologia dell’Universita di Bologna §_ * Laboratorio di Geologia Marina del C.N.R. di Bologna Tue VELIGER, Vol. 22, No. 2 [SaBELLI & Taviani] Figures 1 to 5 Figure 2 Figure 3 Figure 4 Vol. 22; No. 2 THE VELIGER Page 211 is less angular. Furthermore, the specimen was drawn slightly bent, the anterior varix thus appearing upward directed. Bianconi’s figure looks to be more compact than the specimen. The so well developed periostracum is now almost completely lost. The height of the holotype is 137 mm. Cymatium (Cymatium) ranzanii is one of the rarest Cymatiidae, with a geographic range from the eastern part of the Indian Ocean to the East African coast and the Red Sea (Mienis, 1976). Recently it was well fig- ured by some authors (EMERSON & D’ATTILIO, 1962; ANONYMOUS, 1971; DANCE, 1972; Kriss, 1973; ANON- YMOUS, 1974; MIENIs, op. cit.). Very close but different species, according to the authors, are C. (C.) tigrinum (Broderip, 1833) of the Panamic province, and the Caribbean C.(C.) femorale (Linnaeus, 1758). Literature Cited ANONYMOUS 1971. Cymatium ranzanit. text fig. Brancont, GIovANNI GIUSEPPE 1850. Rendicont dell’adunanza del 21 Febbraio. Nuovi Ann. Sci. Natur. e Rendicont. Lav. Accad. Sci. Istituto Societ. agrar. Bologna (III) 11: 39-42 1851. Specimina zoologica mosambicana. Sci. Bologna 3: 1-18; 2 text figs. Dance, STANLEY PETER 1969. Rare shells. Faber & Faber, London; 128 pp.; 24 pits. Emerson, WILLIAM KgiTH & ANTHONY D’ATTILIO 1962. Remarks on Triton ranzantt Bianconi (Mollusca, Gastropoda). Amer. Mus. Novit., New York, 2108: 1-8; 4 text figs. Kitiras, RuDoiF 1973- Cymatiidae. Mients, Henx K. 1976. Notes on the distribution of Cymatium ranzonit (Bianconi), a rare species from the Indian Ocean. Malacol. Opstellen 29-33; 2 pits. Nicotay, Kety 1974. Conchiglie di tutti i mari: Gen.: Cymatium, Charonia. La Conchiglia 60: 9-10; 4 figs. Hawaiian Shell News 19 (2): 73; 1 Fasc. IV Mem. Accad. Das Tierreich 92: vilit235 pp.; 149 figs. Page 212 THE VELIGER Vol. 22; No. 2 BOOKS, PERIODICALS, PAMPHLETS The Family Cerithiidae in the Indo-Pacific. Part I: The Genera Rhinoclavis, Pseudovertagus and Clavocerithium by Ricuarp S. Housrick. Monographs of Marine Mol- lusca, No. 1: 1 - 130; 98 plts. $17.50 (15 December 1978) Detailed taxonomic monographs seldom make good bedtime reading and are generally consulted only for making identifications or in subsequent revisionary tax- onomy. Richard S. Houbrick’s recent monograph of the cer- ithiid genera Rhinoclavis, Pseudovertagus and Clavo- cerithium contains much more than the average taxo- nomic monograph, and I recommend it both as a model of the diversity of information that can and should be incorporated in taxonomy and as a readable and interest- ing source of information and insight into a major group of tropical reef-associated gastropods that cannot be as- sessed taxonomically or placed in an evolutionary con- text working from populations of highly polymorphic shells alone. Treatment includes 47 living and fossil spe- cies. Those involved in taxonomic research and in making phylogenetic inferences will appreciate the care with which the author has considered characters and their weighting and his explicit discussion of the data base for his concept of species. In particular, it is good to see life history and ecological data incorporated in taxonomic de- cision making. In addition to shell characters, morpho- logical analysis includes soft anatomy and radular char- acters. Scanning electron micrographs of radulae provide a helpful adjunct to verbal descriptions, although they have contrast problems that obscure detail and fail to resolve important features. On the basis of this work we may justifiably look forward to the promised monographic treatments of additional cerithiid genera. The format of the new journal, Monographs of Marine Mollusca, will be immediately familiar to those who know Indo-Pacific Mollusca. It is published by American Malacologists, Inc., Greenville, Delaware, under the editorship of R. Tucker Abbott and serves as a continu- ation of Indo-Pacific Mollusca, currently an inactive pub- lication series of the Delaware Museum of Natural History. Carole S. Hickman Department of Paleontology University of California Berkeley, CA 94720 A New Monoplacophoran Limpet from the Continental Shelf off Southern California by James H. McLean. Contributions in Science, Natural History Museum of Los Angeles County, No. 307; pp. I - 19; 25 figures in text. 16 March 1979 Although discovery of living coelocanths in the 1950s made larger headlines, the collecting of a living mono- placophoran — Neopilina — was a more signal event for paleontologists and malacologists. The deep-water habi- tat of that and several subsequent finds left questions about mode of life unanswered. Now a new find may bring some answers, for specimens of another new neo- pilinid have been taken in relatively shallow water in an area at present easy of access. The new form is surprisingly small — less than 3 mm in length — yet, with the aid of modern techniques of study and photography, McLean and the colleagues who assisted him have been able to make detailed exami- nation of internal and external anatomy and to obtain good radular mounts. The holotype, which is 1.94mm in length, is figured at a magnification of X100 on a full- page plate. The photograph shows not only shell structure but also the form of the soft parts — the mouth, the foot, the 6 pairs of gills, and the mantle margin. A new subgenus is proposed for the reception of the new species — Neopilina (Laevipilina) hyalina McLean, spec. nov. The type locality is on Santa Rosa-Cortes Ridge, depth 373-384m. Other records of the species from the general region range as low as 174m depth. A photograph of the bottom at the type locality shows small rocks thinly covered with sediment, and an early sample was on a rock fragment that was snagged up by a fishing line. We may hope that more material will now become available. A. Myra Keen Vol. 22; No. 2 THE VELIGER Page 213 “Evolutionary Systematics of Bivalve Mollusca ”: A discussion organized for the Royal Society and the Malacological Society of London by Sir Maurice Yonge, E R. S., and T: E. Thompson. Philosophical Transactions of the Royal Society of London, Series B, Biological Sciences (ISSN 0080-4622), Vol. 284 (1001): pp. 199- 436, (paper). £17.00, overseas, postpaid. 16 Nov. 1978 This impressive issue of “Phil. Trans.” is the product of a symposium held in London in May, 1977, devoted to the evolution and systematics of bivalve molluscs. Noting that 2 of the contributors were paleontologists, and the rest distinguished neontologists and malacologists, your reviewer entered upon his task with determination, but also with some trepidation. The terminological jungle in which I found myself was at first formidable but, by dint of persistence, I came to perceive some fascinating areas new to me. I must confess I found difficult the papers by Scarlato « Starobogatov, and by Pojeta, on the early evo- lution of bivalves. The former authors outlined 14 orders and 25 suborders of bivalves; the latter, despite 15 crowded plates of fossil specimens, left me figuratively not seeing the forest for the trees (my own deficiency, I think, rather than that of the paper). Stanley, in a lucid paper on the Trigoniidae, invokes both biology and paleontology to argue that the living genus Neotrigonia is not a “living fossil,” even though it contains the survivors of a flourishing Mesozoic group, but a genus well-adapted to modern conditions. Incidentally, it is regrettable that this volume contains no paper com- parable to Stanley’s illuminating account of the great Mesozoic radiation of bivalves (Journ. Paleontol. 42: 214- 229, 1968), a paper the general reader might do well to read before tackling some of the more specialized papers of this volume. Bivalves are exceptionally favorable material for the study of evolutionary rates, by virtue of the abundance of fossil material in numerous lineages, found in exten- sive stratigraphic sequences for, or from which, eco- logical and radiometric data are available. Kauffman, in a closely-reasoned paper based on studies of many Cre- taceous lineages, argues that both modes and rates of evolution (in new taxa per unit time or duration of taxa) can vary widely even within lineages. Rapid rates of species formation may occur at times of colonization of unoccupied ecological space, or during crises of eco- logical stress. Although the “simple” bivalves have com- monly been considered to have evolved slowly and steadi- ly, the actual rates are as high as those claimed for late Cenozoic mammals! Yet one may wonder if the small differences which seem to separate fossil bivalve species may not magnify the apparent rate of speciation or, as a later contributor, Boss, puts it, whether the tendencies “to proliferate nomina and to elevate taxa partially ob- fuscate reality.” One should read the original and judge for oneself. To me, one of the most illuminating papers was that of Runnegar, on the pseudo-bivalved Rostroconchia, recog- nized as a new class as recently as 1972. Here is the group transitional between the univalved Monoplaco- phora and the bivalves: Paleozoic forms with a bilobed but unhinged shell, precisely fitting the “hypothetical filter-feeding monoplacophoroid with undivided shell” postulated by Stasek as ancestral to the true Bivalvia, the first of which, Fordilla, appeared in the Cambrian. Waller’s paper on the classification of the Pteriomor- phia includes some difficult material on hinge ligaments; in this connection it is regrettable that the contribution of C. M. Yonge on the ligament of the Bivalvia is repre- sented only in abstract. One may need to read Yonge’s paper, published elsewhere (Proc. Roy. Soc. London, B, 202: 231-248, 1978), before Waller’s. Contributions on bivalve soft parts are fewer. Owen clarifies Atkin’s groupings of Macrociliobranchia and Microciliobranchia by establishing through scanning elec- tron microscopy that the Ostraeidae belong to the former group, despite features other than gill ciliation that seem to contradict this placement. Allen’s paper on deep-sea Protobranchia reveals that this order is of more impor- tance in the deep sea than most of us shallow-water bio- logists have been assuming, and he also shows how little their shells may reveal of variation in their internal soft anatomy. Levinton and Lassen discuss enzyme polymor- phism in Mytilus and make a case for selection of popula- tion isolates in spatially close but ecologically distinct areas, as in estuarine conditions in Long Island Sound. Boss cautions against the danger inherent in the “super- fluity” of bivalve nomenclature and argues for a simplified but phyletically meaningful classification. The volume closes with a thoughtful analysis by Purchon of an at- tempt to base a bivalve classification on 9 “useful” cate- gories of morphological data, on both soft parts and shell characteristics. This seems a wholesome trend away from Page 214 THE VELIGER Vol. 22; No. 2 the over-reliance of paleontologists on dentition. One can return to Newell’s introductory paper, “A paleontolo- gist’s view of bivalve phylogeny,” with a little more understanding and a great deal more sympathy for the problems faced by those who would unravel the evolution of bivalves, Ralph I. Smith Department of Zoology University of California Berkeley, California 94720 The Genus Arcinella (Mollusca: Bivalvia) in Venezuela and Some Associated Faunas by J. Gisson-SmMitH and W. Gisson-SmirH, GEOS, Escuela de Geologia y Minas, Universidad Central de Venezuela, No. 24; pp. 11 - 32; 3 plts. January 1979 This paper describes and illustrates Caribbean forms of the chamid genus Arcinella, from Miocene to Holocene, and plots their biostratigraphic distribution and phylo- geny. Iwo new early Miocene species are described and a new subgenus (Nicolia, type-species Chama draconis Dall, 1903, from the Chipola Formation of Florida) is proposed. The authors’ extensive collections of Venezu- elan Tertiary mollusks form the basis for many additional faunal notes, bearing on the development of Neotropical faunas and Neogene chronostratigraphy of the Caribbean area. A number of synonymies in other molluscan groups are introduced, and earlier authors’ accounts reappraised. The new gastropod species Purpura weisbordi is described from the Mare Formation of middle or late Pliocene. Barry Roth THE VELIGER is open to original papers pertaining to any problem concerned with mollusks. This is meant to make facilities available for publication of original articles from a wide field of endeavor. Papers dealing with anatomical, cytological, distributional, ecological, histological, morphological, phys- iological, taxonomic, etc., aspects of marine, freshwater or terrestrial mollusks from any region, will be considered. Even topics only indi- rectly concerned with mollusks may be acceptable. In the unlikely event that space considerations make limitations necessary, papers dealing with mollusks from the Pacific region will be given priority. However, in this case the term “Pacific region” is to be most liberally interpreted. It is the editorial policy to preserve the individualistic writing style of the author; therefore any editorial changes in a manuscript will be sub- mitted to the author for his approval, before going to press. Short articles containing descriptions of new species or lesser taxa will be given preferential treatment in the speed of publication provided that arrangements have been made by the author for depositing the holotype with a recognized public Museum. Museum numbers of the type specimens must be included in the manuscript. Type localities must be defined as accurately as possible, with geographical longitudes and latitudes added. Short original papers, not exceeding 500 words, will be published in the column “NOTES & NEWS”; in this column will also appear notices of meetings of the American Malacological Union, as well as news items which are deemed of interest to our subscribers in general. Articles on “METHODS & TECHNIQUES” will be considered for publication in another column, provided that the information is complete and tech- niques and methods are capable of duplication by anyone carefully fol- lowing the description given. Such articles should be mainly original and deal with collecting, preparing, maintaining, studying, photo- graphing, etc., of mollusks or other invertebrates. A third column, en- titled “INFORMATION DESK,” will contain articles dealing with any problem pertaining to collecting, identifying, etc., in short, problems encountered by our readers. In contrast to other contributions, articles in this column do not necessarily contain new and original materials. Questions to the editor, which can be answered in this column, are in- vited. The column “BOOKS, PERIODICALS, PAMPHLETS?” will attempt to bring reviews of new publications to the attention of our readers. Also, new timely articles may be listed by title only, if this is deemed expedient. Manuscripts should be typed in final form on a high grade white paper, 81/2” by 11”, double spaced and accompanied by a carbon copy. A pamphlet with detailed suggestions for preparing manuscripts intended for publication in THE VELIGER is available to authors upon request. A self-addressed envelope, sufficiently large to accom- modate the pamphlet (which measures 52” by 81/2’), with double first class postage, should be sent with the request to the Editor. EDITORIAL BOARD Dr. Donan P. Assorrt, Professor of Biology Hopkins Marine Station of Stanford University Dr. WarrEN O. AppicoTT, Research Geologist, U. S. Geological Survey, Menlo Park, California, and Consulting Professor of Paleontology, Stanford University Dr. Hans Bertscu, Curator of Marine Invertebrates San Diego Museum of Natural History Dr. Jerry DononvE, Professor of Chemistry University of Pennsylvania, Philadelphia, and Research Associate in the Allan Hancock Foundation University of Southern California, Los Angeles Dr. J. Wyatr Duruam, Professor of Paleontology Emeritus University of California, Berkeley, California Dr. Cavet Hann, Professor of Zoology and Director, Bodega Marine Laboratory University of California, Berkeley, California Dr. Carore S. Hickman, Assistant Professor of Paleontology University of California, Berkeley, California Dr. A. Myra KEEN, Professor of Paleontology and Curator of Malacology, Emeritus Stanford University, Stanford, California Dr. Victor Loosanorr, Senior Biologist, Emeritus U.S. National Marine Fisheries Service EDITOR-IN-CHIEF Dr. Rupoir STOHLER, Research Zoologist, Emeritus University of California, Berkeley, California Dr. Joun McGowan, Professor of Oceanography Scripps Institution of Oceanography, La Jolla University of California at San Diego Dr. Franx A. Pite.xa, Professor of Zoology University of California, Berkeley, California Dr. RoBert Rosertson, Pilsbry Chair of Malacology Department of Malacology Academy of Natural Sciences of Philadelphia Dr. PETER U. Roppa, Chairman and Curator, Department of Geology California Academy of Sciences, San Francisco Dr. Ciype FE. E. Roper, Curator Department of Invertebrate Zoology (Mollusca) National Museum of Natural History Washington, D. C. Dr. JupirH Terry Smiru, Visiting Scholar Department of Geology, Stanford University Stanford, California Dr. Ratpu I. Smiru, Professor of Zoology University of California, Berkeley, California Dr. Cuartes R. STASEK, Bodega Bay Institute Bodega Bay, California Dr. T. E. Tuompson, Reader in Zoology University of Bristol, England ASSOCIATE EDITOR Mrs. JEAN M. Care Rancho Santa Fe, California GE VELIGER A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California VOLUME 22 JANUARY 1, 1980 NUMBER 3 CONTENTS Acochlidium bayerfehlmanni spec. nov. (Gastropoda : Opisthobranchia : Acochlidi- acea) from Palau Islands. (2 Plates) ERHARD WaAwRA Variations in the Shell-Flesh Relationships of Mytilus: The Value of Sea Mussels as Items of Prey. (1 Text figure) R. SEED The Nudibranch Aegires albopunctatus (Polyceratacea : Aegiretidae) Preys on Leucilla nuttingi (Porifera : Calcarea) . (1 Plate; 2 Text figures) Hans BertscH Gea) fh PRO Ty ss OA el ah AUPE | Eat Sa Karyotypes of Six Eastern Pacific Acmaeid Gastropods. (1 Plate;~4 Text figures ) Davi M. Cuapin « PAuL A. RosBerts eats Cepek nals Mee ey ceumnE EES The Distribution of Shallow-Water Marine Prosobranch Gastropod Molluscs Along the Coastline of Western Australia. (5 Text figures) Frep E. WELLS Predator Boreholes in Periploma margaritaceum With a Brief Survey of Other Periplomatidae (Bivalvia : Anomalodesmata). (1 Plate; 1 Text figure) JosePpH RoSEWATER , Se ilie Sri Si aMoreet ty cube Habitat, Food and Reproductive Activity of the Nudibranch Hexabranchus san- guineus on Tongatapu Island, Tonga. (4 Text figures) Ma tco.io P. FRANCIS [Continued on Inside Front Cover] 5 DUG - 222 . 225 . 232 - 252 Distributed free to Members of the California Malacozoological Society, Inc. Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc. Volume 22: $30.- plus $1.50 for postage (U.S. A. only) For ALL foreign countries: Swiss Francs 60.- plus SF 7.- for postage Single copy this issue $14.00; postage extra Send subscription orders to California Malacozoological Society, Inc. 1584 Milvia Street, Berkeley, CA 94709, U.S.A. Address all other correspondence to Dr. R. SrouuEr, Editor, Department of Zoology University of California, Berkeley, CA 94720 Second Class Postage Paid at Berkeley, California ConTENTs — Continued Depth Distributions of Nautilus pompilius in Fiji and Nautilus macromphalus in New Caledonia. (1 Plate; 6 Text figures) Peter D. Warp « ArTHUR W. MarTIN as Notoacmea gabatella (Berry), an Outer Coast Form of Notoacmea depicta (Hinds) (Gastropoda : Acmaeidae). (1 Plate) Davi R. LINDBERG . Reinstatement of Two Species of Murexiella (Gastropoda : Muricidae) from the Tropical Eastern Pacific. (1 Plate; 1 Text figure) Leroy H. PoorMan . MMe re AU aa eee hed Tyas tia stna ag hl AM Larval and Early Benthic Stages of Brachidontes granulata (Bivalvia : Mytilidae). (2 Plates) Bernarpita Campos M. « Luis Ramorino M. AU ser ar Deep Water Collections of Opisthobranchs in Central California. (2 Text figures ) RoBERT CHAMBERLAIN & Davip W. BEHRENS isa Some Aspects of Food Intake in Octopus joubini Robson. (3 Text figures) JENNIFER A. MaTHER Predation by a Rockfish, Sebastes chrysomelas, on Lamellaria diegoensis Dall, 1885. Doue.as E. Hunt ; 4 Observations of Spawning in Calliostoma ligatum (Gould, 1849). Douctias E. Hunt NOTES & NEWS Trematodes in Chilean Fissurellid Molluscs. Marta BRETOS & CLAUDINA JIRON BOOKS, PERIODICALS & PAMPHLETS . - 259 . 265 - 273 - 277 . 282 . 286 . 291 5 DOW - 203 - 207 Note: The various taxa above species are indicated by the use of different type styles as shown by the following examples, and by increasing indentation. ORDER, Suborder, DIVISION, Subdivision, SECTION, SUPERFAMILY, Famiy, Subfamily, Genus, (Subgenus) New Taxa Vol. 22; No. 3 THE VELIGER Page 215 Acochlidium bayerfeldmanni spec. nov., (Gastropoda : Opisthobranchia : Acochlidiacea ) from Palau Islands BY ERHARD WAWRA 3. Zoologische Abteilung, Naturhistorisches Museum Wien, A-ro1o Vienna, Austria (2 Plates) INTRODUCTION AFTER I HAD SUBMITTED a manuscript on material of Acochlidium collected by Sutter in 1949 on the island of Sumba (BUHLER & SUTTER, 1951), some specimens from the locality of Acochlidium amboinense (Strubell, 1892) in the Palau Islands, as described by BAYER & FEHLMANN (1960), were obtained. The specimens from Sumba identified by van BEent- HEM JUTTING (1955) as Acochlidium weber (Bergh, 1896) proved to represent a new species, A. sutter1s WAw- RA (in press). Because of several discrepancies between the description of BercH (1896) of the radula and penial armature and the results of my dissections, the Sumba specimens could not be identified as A. weberi. The specimens from Sumba differ also from the descrip- tion of A. amboinense in which the penis opens to the right behind the buccal mass (BicKiNc, 1933). In A. sutteri a duct is formed by the penial sheath leading from the penis to the base of the right rhinophore where it opens to the outside (Waw,ra, in press). Regrettably, BAYER & FEHLMANN (1960) did not in- dicate where the opening for the penis in their specimens might be, whether in agreement with the description of BickINc (1933), whose work was based on the original specimens of STRUBELL (1892), or otherwise. Because the type material of A. amboinense is destroyed (Rudo von Cosel, Universitat GieBen, FRG, personal communica- tion), attempts were made to examine specimens of A. amboinense from Palau. Acochlidium bayerfehlmanni Wawra, spec. nov. 1960. Acochlidium amboinense, F. M. Bayer « H. A. FEHL- MANN, Proc. Biol. Soc. Wash. 73: 185 Diagnosis: Length of living animal 25mm, preserved 11mm; foot well developed, 2 pairs of tentacles. Radula: Radular formula: 1-1: 2Xn, n approximately 56; left marginal plate lacking, rhachidian tooth finely serrated. Hermaphroditic, the vas deferens leading from visceral sac to base of right rhinophore, then to penial glands; penial armature consisting of a double row of hooks and a ‘cluster of slender thorns; penis opening at the base of right rhinophore. Type Locality: Arakitaoch River, Island Babelthuap, Palau Islands (BAYER & FEHLMANN, 1969: 185) Type Specimens: The holotype has been deposited in the Molluskensammlung, Naturhistorisches Museum, Wien, Austria (Inv. Nr. 81.232), along with 2 paratypes (Inv. Nr. 81.233: partially dissected, radula mounted in polyvinyllactophenol on 2 slides with subnumbers 161, 162, and a sample of the gonad sectioned at 5 um and stained in Heidenhain — Azan on a third slide with the subnumber 163; Inv. Nr. 81.234: partially dissected, a sample of the gonad with the subnumber 164). One para- type has been deposited in the Naturhistorisches Museum, Basel, Switzerland (Nr. 11.1174). The specimens were collected around the end of January and beginning of February 1978 by G. Bright. Page 216 THE VELIGER Vol. 22; No. 3 Named in honour of FE M. Bayer and H. A. Fehlmann who were the first to work on Acochlidium from the Palau Islands. Unfortunately, the specimens studied by Bayer « Fehlmann have been misplaced (J. Rosewater, U. S. National Museum, Washington, D. C., personal com- munication). DESCRIPTION Preliminary Remarks: Although I have not seen the material described by Bayer & Fehlmann, I assume that it is identical, on the specific level, with the specimens received for this paper. Bayer « Fehlmann gave a detailed description of their specimens, and now only a correction and an addition to their results are necessary, the latter, however, with systematic consequences. General Appearance: The general appearance of these Palauan freshwater opisthobranchs is similar to that of Acochlidium amboinense and A. sutteri. The only differ- ence is that the visceral sac is smooth as pointed out by BAYER & FEHLMANN (1960: 185). This difference may not necessarily be of diagnostic value, rather it could be the result of more careful fixation in the 2 recent samples. Radula: The radula of several species of the Acochlidi- acea has proved to be asymmetrical, e. g., Hedylopsis suecica Odhner, 1937 (Marcus, 1953); H. spiculifera (Kowalevsky, 1901) (Poizat, 1978); H. cf. H. spicult- fera (Wawra, unpublished record); Microhedyle crypt- ophthalma Westheide « Wawra (WESTHEDE & WAwRA, 1974); Strubellia paradoxa (STRUBELL, 1892) and A- cochlidium sutteri (Wawra, in press). Therefore, the observations of BicKING (1933) and BAYER & FEHLMANN (1960) needed reconsidering. In the specimens examined, the radular formula is asymmetrical (1:1°2 Xn, n=53, with a few of the younger rows lost during preparation) because the left marginal plate is lacking (Figures 4 and 5). The form and size of the radular plates otherwise correspond with the drawings and measurements of BAYER & FEHLMANN (1960) and the number of’ crossrows, very unlikely con- stant, is approximately the same. Table 1 compares the measurements of the radular elements found by BAYER & FEHLMANN (1960) and those of my own observations. Attention should be paid to the difference in size of the 2 lateral plates, although the sum of the widths on the right side plus the space between the plates equals the width of the left plate. Table 1 Measurements of radular elements (in mm), those taken from Bayer & FEHLMANN (1960) in parentheses. Height of rhachidian tooth 0.21-0.23 (0.24-0.27) Width of right lateral 0.12 (0.12) Width of right marginal 0.02 (0.03) Width of left lateral 0.15 (— ) Height of right marginal 0.04 (0.055) Male Genital System: The dissection of the specimens did not at first reveal any differences from the animals of Acochlidium sutteri until the penial sheath was opened. As in A. sutteri, the vas deferens, which is embedded in the right lateral body wall of the anterior body, runs parallel to the foot from the visceral sac to the base of the right rhinophore. With close inspection, this part of the vas deferens is visible from the outside as a thin cord (Figure 1). After a U-turn to the left, the vas defer- ens leaves the body wall and continues as a small duct running underneath another, larger duct formed by the penial sheath across the cerebral commissure to the penial glands (Figure 2). The only difference from the animals of Acochlidium suttert appears in the construction of the armature of Explanation of Figures 1 to 3 Acochlidium bayerfehlmanni Wawra, spec. nov. Figure 1: Holotype, (NHMW 81.232): Female genital opening (go), vas deferens (vd) X 20 Figure 2: Paratype (NHMW 81.233): Penial duct formed by the penial sheath (pd), vas deferens (vd) Figure 3: Paratype (NHMW 81.233) after dissection of the penial sheath: Penis armature X 50 J Tue VELIGER, Vol. 22, No. 3 Figure 2 Figure 7 [Wawra] Figures 7 to 3 Vol. 22; No. 3 the glans penis. Whereas the penial armature in A. ambo- inense, with one longer and one shorter tip or thorn in addition to a double corona of hooks as drawn by Biickinc (1933: figs. 6 & 7), corresponds to the armature in A. suttert (WAwRA, in press, fig. 4), in the animals of A. bayerfehlmanni the glans penis lacks the large thorn- like ending of the vas deferens and the smaller thorn is replaced with a bundle of thin, slender thorns (Figure 3). These observations are in good agreement with the findings of Bayer « FEHLMANN (1960) with the excep- tion that there is no free space between the smaller hooks and the thin thorns as illustrated by BAYER « FEHLMANN (1960: fig. 2b). The histological sections of a sample of the gonad taken from a specimen with a penis show a true hermaphroditic condition, spermatids and mature sperm occurring at the same time as oocytes with yolk- material (Figures 6, 7). The gonad of a specimen lacking the penis and its glands was still undifferentiated. DISCUSSION Bayer & FEHLMANN (1960) considered their specimens of Acochlidium as representing A. amboinense despite differences between the drawings of BicK1Nc (1933) and their own observations, differences which they thought were due to different “artistic representation” or “defi- ciencies in observations.” Insofar as the radula is con- cerned I would possibly agree, because my observations differ from those of Bayer & Fehlmann as well as those of Biicking. But by employing phase contrast microscopy and using a suitable mounting medium the asymmetry of the radula of A. bayerfehlmanni can easily be demonstra- ted. However, figure 4 of BiicKx1ne (1933) leaves the pos- sibility for a symmetrical radula (2:1-2) in A. amboin- ense, unless there is proof to the contrary. I cannot agree with BaYer & FEHLMANN (1960) re- garding the penial armature. The same armature as illus- trated by BUckince (1933) for Acochlidium amboinense has been found in A. sutteri and therefore it is hard to believe that Biicx1ne’s (1933) drawings were not precisely done. However, the description of the glans penis and its armature by Bayer & FEHLMANN (1960) is confirmed. THE VELIGER Page 217 The slight differences in the placement of the hooks be- tween the figures of these authors and my specimens may perhaps be caused by different muscle contraction during fixation. One might consider the lack of the longer thorns in Acochlidium bayerfehlmanni as an earlier developmental stage of specimens of A. sutteri, but the specimens re- ported by Bayer & FEHLMANN (1960) were collected in September 1957 and October 1958, while my specimens were collected in January/February 1978, and none of the Palauan specimens examined had a penial armature as described for A. sutteri. CONCLUSION The gross anatomy of Acochlidium bayerfehlmanni in- dicates a close relationship to A. suttert which is morpho- logically separated only by a different penial armature. The isolated localities of the freshwater streams of Palau and of Sumba, respectively, speak rather more for specia- tion than against. Differences from A. amboinense are more distinctive and include the location of the male genital opening, the course of the vas deferens, the penial armature, and differences in the radula, assuming a symmetric radula for A. amboinense. ACKNOWLEDGMENTS I am very grateful to Dr. E. Alison Kay, University of Hawaii, Honolulu, to her student J. Heslinga, and to Dr. Greg Bright, who together made it possible for me to obtain 4 specimens from Palau. I am also indebted to Dr. Rainer Hacker, Naturhistorisches Museum Wien, and again to Dr. Kay for critical reading of the manuscript. Literature Cited Bayer, FREDERICK MERKLE « H. A. FEHLMANN 1960. The discovery of a freshwater opisthobranchiate mollusk, A- cochlidium amboinense Strubell, in the Palau Islands. Proc. Biol. Soc. Wash. 73: 183 - 194 BENTHEM JUTTING, WouTERa S. S. vAN 1955: Siisswassermollusken von Sumba. Basel 66: 49 - 60 Verh. naturf Gesell. Page 218 THE VELIGER Vol. 22; No. 3 BercH, Lupwic SopHus Rupo.ir 1896. Die Hedyliden, eine Familie der kladohepatischen Nudibran- chien. Verh. zool. bot. Ges. Wien 45: 1-12; plts. I, IT Bicxine, G. 1933. Hedyle amboinensis (Strubell). 549 - 582; plt. 2 Buuer, A. & E. SutTer 1951. Wissenschaftliche Ergebnisse der Sumba-Expedition des Museums fiir Vélkerkunde und des Naturhistorischen Museums in Basel, 1949: Reisebericht und allgemeine Einfiihrung. Verh. naturf. Gesell. Basel 62: 181 - 217; 6 plts. Marcus, E. 1953. Three Brazilian sand opisthobranchs. Letr., Zool. 18: 165 - 203; plts. 1-9 Zool. Jahrb., Abt. Syst. 64: Bol. Fac, Fil.Ciéne. WesTHEIDE, W. & ERHARD WAWRA Porzat, C. 1978. Gastéropodes mésopsammiques de fonds sableux du Golfe de Marseille: Ecologie et reproduction. Thése, Univ. d’Aix-Marseille, Fac. Sci. Tech. Saint-Jerome: Fasc. I: 1-301,; Fase. II: figs. 1 - 84; pits. I- XVII STRUBELL, A. 1892. [no title] Verhandl. naturhist. Verein preuss. Rheinlande 4gster Jahrg., Sitzung niederrhein. Gesell. 13 Juni 1892: 62 Wawra, ErRHarD 1974. The rediscovery of Strubellia paradoxa (Strubell) (Gastropoda: Euthyneura : Acochlidiacea) on the Solomon Islands. The Veliger 17 (1): 8-10; 1 plt.; 3 text figs. (1 July 1974) in press Acochlidium sutteri nov. spec. (Gastropoda, Opisthobranchia, Acochlidiacea) von Sumba. Indonesien. Ann. Naturhist. Mus. Wien 82 1974. Organisation, Systematik und Biologie von Microhedyle crypt- ophthalma nov. spec. (Gastropoda, Opisthobranchia) aus dem Bran- dungsstrand des Mittelmeeres. 27-41 Helgol. wiss. Meeresunters. 26: Explanation of Figures 4 to 7 Acochlidium bayerfehlmanni Wawra, spec. nov. Figure 4: Paratype (NHMW 81.233/162): Radula, right marginal plate and distal part of right lateral plate. Phasecontrast X goo Figure 5: Same as Figure 4, but left lateral plate Figure 6: Paratype (NHMW 81.233/163): Ovotestis, yolkmate- rial (yo) Figure 7: Same as Figure 6: Sperm and spermatids XK 155 X 800 [Wawra] Figures 4 to 7 THE VELIGER, Vol. 22, No. 3 - * Ee eat y > " ee us * a Vol. 22; No. 3 THE VELIGER Page 219 Variations in the Shell-Flesh Relationships of Mytilus: The Value of Sea Mussels as Items of Prey BY R. SEED Department of Zoology, University College of North Wales, Bangor LL57 2UW, Wales, United Kingdom (1 Text figure) OVER THE PAST DECADE or so there has been a growing interest in the different strategies adopted by predators in their search for food. An optimal foraging strategy repre- sents the combination of those factors associated with maximising an organism’s energy intake relative to the time and effort involved in acquiring food (e. g., EMLEN, 1968; SCHOENER, 1971; CHARNOV, 1976; PyKE, PULLIAM & CHARNOV, 1977). A predator expends energy searching for and handling suitable items of prey and this must therefore be set against the amount of energy eventually yielded by the prey item. Predators feeding mainly on relatively sedentary species such as bivalves will generally select their prey on the basis of its size and availability (e. g., ELNER & HuGHES, 1978; O’CoNNorR & BRown, 1977). Mytilus, however, which constitutes a major food re- source for several contrasted predators (e. g., sea-stars, gastropods, crabs, birds) possesses a strong protective shell the size and strength of which will be a major factor in determining a predator’s handling time. For practical reasons it is assumed that shell strength is proportional to shell weight (at least for mussels of similar size and shape) and the latter is known to vary not only from one locality to another but also with respect to tidal level (Table 1). Furthermore, the size and strength of the adductor muscle is also known to vary significantly even amongst mussels of comparable shell length (Hancock, 1965). This short communication considers the relationships between shell and flesh weights in Mytilus populations experiencing dif- ferent degrees of aerial exposure and briefly comments on the possible importance of such considerations to studies of foraging behaviour in those predators that are known to feed extensively on sea mussels. Table 1 Variations in Shell Weight in Mytilus edulis Shell weight (g) Shell iHigh shore iLow shore iffigh shore IV Low shore YHigh shore viLow shore Length (cm) Filey Brigg Filey Brigg Filey Bay Filey Bay Robin Hood’s Bay Whitby Harbour 1.50 0.20 0.16 0.25 0.21 0.21 0.19 2.50 0.98 0.83 1.30 0.85 1.09 0.85 3.50 3.45 2.57 4.49 2.31 — 2.73 4.50 — — 7.93 5.01 — 5.81 5.50 — = 11.45 7.87 — 10.71 6.50 _ _ = = = 18.80 n 432 373 490 360 71 102 Filey Brigg and Robin Hood’s Bay are high energy, waveswept shores. Filey Bay and Whitby Harbour are low energy, protected shores. All sites situated on the North Sea cost of N.E. England. Page 220 Rao (1953)demonstrated that shell weight in both Mytilus edulis Linnaeus, 1758 and M. californianus Con- rad, 1837 decreased as exposure to air increased, the heaviest shells for any given flesh weight occurring a- mongst permanently submerged mussels. He attributed this relationship to the propensity of mussels for extracting calcium from seawater -— thus the longer a mussel re- mained exposed to seawater the heavier its shell became. Rao (1953a) also noted an apparently similar phenom- enon (though with somewhat different causes) in relation to high and low latitudinal mussel populations. Fox « CoE (1943), however, found that M. californianus from the higher intertidal zone had thicker (and therefore presumably heavier) shells than conspecifics of similar length further down the shore, an observation which was subsequently corroborated by Kopp (1979) for M. californianus and by Bairp & DRINNAN (1957) for M. edulis. Fast growing subtidal populations, by contrast, tend to have relatively thin rather brittle shells. Barp « DRINNAN (op. cit.) explained their findings in terms of requirements for basal metabolism. When mussels are exposed to air, basal metabolism reduces flesh weight at a greater rate than chemical erosion reduces the shell. Higher intertidal mussels will therefore have higher shell- flesh ratios than those from the lower intertidal zone. More recently, SEED (1973) obtained similar results for M. edulis from several contrasted localities. When mussels of similar shell length (though of differing age since growth rates also vary widely with intertidal height) were compared, those from the high shore had consistently heavier shell weights than those from the low shore (see also Kopp (1979) for M. californianus). Moreover, shell- flesh ratios generally increased with increasing shell length suggesting that as mussels age, their shells account for a progressively greater proportion of their total body weight. Figure 1 illustrates the relationship between shell and tissue weights as functions of intertidal height in mussels from 2 contrasted rocky shores on the NE coast of England. Mussels from the high shore consistently had heavier (—stronger) shells for any given weight of tissue than those from the low shore. Although previous explanations for the observed dif- ferences in shell-flesh relationships amongst vertically separated mussel populations appear to have been sought in physiological terms, other possibly equally important factors such as longevity and population stability * ought perhaps also be considered. Stability and longevity in rocky shore mussel populations are known to increase ™ T use the term ‘stability’ to denote the absence of any marked temporal change in population structure THE VELIGER Vol. 22; No. 3 Tissue Weight*'° g 8 / A fe 1.5 6 vy 78 1.0 I 4 ee eo” 13 wll 0.5 fom 2 Oo “79 eo Vi 4 sf ‘4 2 4 6 8 10 12 I 2 3 Shell Weight - ¢ Figure 1 The relationship between shell and tissue weights as functions of intertidal height in mussels of known age and size from (A) Filey Bay — a comparatively low energy shore and (B) Filey Brigg — a severely wave exposed high energy shore. Solid symbols repre- sent high shore populations; open symbols, low shore populations. Numbers indicate the estimated age of mussels of known size progressively with increasing intertidal height (SEED, 1976; see also LEwis, 1972). In the low shore, heavy settlement and rapid growth are generally accompanied by exceedingly intense predation resulting in a rapid turn- over of mussels and wide oscillations in population density. By contrast, the higher intertidal area provides a spatial refuge largely free of mussel predators and although recruitment and growth are here substantially reduced, enhanced longevity confers considerable long term popula- tion stability. High shore populations are therefore fre- quently characterised by relatively high proportions of old, slow-growing mussels with typically high shell-flesh ratios. It is by no means unusual for many of these upper shore populations to consist of 20 or more year classes even though the largest mussels in such populations may measure only 2-3 cm in shell length (SEED, 1969). On certain shores, however, mussels can also attain substantial immunity from predation by way of large body size, thereby surpassing the ability of any single predator to consume them (PAINE, 1976). This will be partly de- termined by local predation pressure but it will also be considerably enhanced in habitats which are optimal for Vol. 22; No. 3 THE VELIGER Page 221 mussel growth, e. g., the low shore and subtidal regions. Size-limited predation may conceivably be more charac- teristic of Mytilus californianus populations since this species appears to be rather less attractive than M. edulis to certain predators, probably by virtue of its strong, heavily ribbed shell and its ability to grow to a much larger body size (HARGER, 1972). However, low shore populations of unusually large and sometimes apparently very old M. edulis are occasionally encountered, more especially in habitats sheltered from wave action where local growth rates may be substantially improved. Al- though these large bodied, apparently predator-free mus- sels will increase the stability and structural diversity of such populations thereby enabling both predator and prey to coexist in close proximity, their presence will also substantially alter the microenvironment available to newly recruited mussels. Small mussels which settle in amongst these larger individuals will experience severe intraspecific competition for food and their growth rates can therefore be greatly reduced (e. g., SEED, 1969). At the same time, however, the matrix of large mussels will offer substantial protection from predators and many of these populations could therefore exhibit many of the characteristic features more generally associated with high shore mussel populations (7.¢., greater stability, enhanced longevity, and high shell-flesh ratios). Con- sideration of the structure and relative stability of mussel populations can therefore go some way towards explaining the apparently contradictory literature regarding shell- flesh relationships in mussels collected from different local habitats. It may therefore be relevant to note that the sub- tidal mussels (M. edulis) studied by Rao (1953) came from the underside of floats, a habitat in which mussels may be substantially protected against predation. More recently, size-limited predation has been demonstrated in subtidal Modiolus modiolus (Linnaeus) (SEED & Brown, 1978) and these populations too exhibit considerable long term stability. From this brief account it will perhaps be appreciated that sea mussels vary considerably in their shell-flesh characteristics according to the very local nature of both the physical and biological environment. Accordingly, the amount of time a predator might spend handling mussels of comparable size but different shell strength could vary within quite broad limits as indeed could the eventual food value yielded by such mussels. Furthermore, many of the common predators of sea mussels forage over rela- tively wide areas whilst some also exhibit regular tidal and seasonal migrations. Since an extremely wide range of mussel types could therefore be encountered by these foraging predators, rigorous investigations of the size structure and shell-flesh characteristics of locally occurring mussel populations will clearly be fundamental to any proper understanding of the foraging behaviour of those predators known to utilise mussels as a major food resource. Literature Cited Bairp, R. H. « R. E. Drinnan 1957- The ratio of shell to meat in Mytilus as a function of tidal exposure to air. Journ. du Conseil 22: 329 - 336 Cuarnoy, E. L. 1976. Optimal foraging: attack strategy of a mantid. Amer. Natural. 110: 141-151 ELNerR, R. W.a R. N. HucHes 1978. Energy maximization in the diet of the shore crab, Carcinus maenas. Journ. Anim. Ecol. 47: 103 - 116 EMLEN, JoHN MERRITT 1968. Optimal choice in animals. Fox, Denis L. & WEsLEY ROSWELL CoE 1943. Biology of the Californian sea mussel Mytilus californianus. II. Nutrition, metabolism, growth and calcium deposition. Journ. Exp. Zool. 93: 205 - 249 Hancock, Donatp A. 1965. Adductor muscle size in Danish and British mussels in relation to starfish predation. Ophelia 2: 253 - 267 Harcer, JoHN Rosin 1972. Competitive co-existence: maintenance of interacting associations of the sea mussels Mytilus edulis and Mytilus californianus. The Veliger 14 (4): 387-410; 8 text figs. (1 April 1972) Kopp, James C. 1979. Growth and the intertidal gradient in the sea mussel Mytilus californtanus Conrad, 1837 (Mollusca : Bivalvia : Mytilidae). The Veliger 22 (1): 51-56; 3 text figs, (1 July 1979) Lewis, J. R. 1972. Problems and approaches to baseline studies in coastal com- munities. In: Marine pollution and sea life (M. Rurva, ed.): 401 - 403 Fishing News Books (Ltd,), London O’Connor, R.J. « R. A. Brown 1977. Prey depletion and foraging strategy in the oyster-catcher Haematopus ostralegus. Oecologia (Berlin) 27: 75-92 PAINE, ROBERT TREAT 1976. Size-limited predation: an observational and experimental ap- proach with the Mytilus-Pisaster interaction. Ecology 57: 858 - 873 Rao, K. PAMPAPATHI 1953. Shell weight as a function of intertidal height in a littoral popu- lation of pelecypods. Experientia 9: 465 - 466 19532. Rate of water propulsion in Mytilus californianus as a function of latitude. Biol. Bull. of MBL, Woods Hole, Mass. 104: 171 - 181 ScHoener, T. W. 1971. Theory of feeding strategies. 2: 369 - 404 SzED, RAYMOND 1969. The ecology of Myt#lus edulis L. (Lamellibranchiata) on ex- posed rocky shores. II Growth and mortality. Oecologia (Berlin) 8? 317 - 350 1973. | Absolute and allometric growth in the mussel Mytilus edulis L. (Mollusca: Bivalvia). Proc. Malacol. Soc. London 40: 343 - 357 1976. Ecology. In: Marine mussels: their ecology and physiology. B. L Bayne (ed.), I. B. P 10: 13 - 65; Cambridge Univ. Press Szep, RayMonpD & R. A. Brown 1978. Growth as a strategy for survival in two marine bivalves, Ceras- toderma edule (L.) and Modiolus modiolus (L.). Journ. Anim. Ecol. 47: 283 - 292 Amer. Natural. 102: 385 - 389 Ann. Rev. Ecology System. Page 222 THE VELIGER Vol. 22; No. 3 The Nudibranch Aegires albopunctatus (Polyceratacea : Aegiretidae) Preys on Leucilla nuttingt (Porifera : Calcarea) HANS BERTSCH Department of Marine Invertebrates, Natural History Museum, Balboa Park P.O. Box 1390, San Diego, California g2112 (1 Plate; 2 Text figures) IT HAS BEEN CONSDERED axiomatic that cryptobranchiate dorid nudibranchs (Doridacea or Eudoridacea) with simple or denticled hamate teeth are primarily sponge feeders (YOUNG, 1970) and that phanerobranchs (Poly- ceratacea) usually feed on bryozoans (e. g. MILLER, 1961). A recent literature compilation for Californian nudi- branchs (McDonaLp « NyBAKKEN, 1978) has shown the known diet of numerous species; all 6 species of Poly- ceratacea (= Nonsuctoria) feed on bryozoans; 13 of the 14 cryptobranch species listed feed on various demospon- giae (the exception, Hallaxa chani Gosliner & Williams, 1975, feeds on tunicates, and has a distinctly different radular morphology) . Radular morphology indicates prey specificity (BeERTSCH, 1974): thus it is curious to find a sponge- feeding radula type in a superfamily characteristically reported to feed on bryozoans. Within the Polyceratacea, species of Aegires Lovén, 1844, and Notodoris Bergh, 1875, have typical sponge-feeding radulae with simple, hamate teeth. Pruvot-Fot (1954: 245) commented on the similarity between radulae of the European Aegires punctilucens (Orbigny, 1837) and those possessed by other rasping sponge feeders: “La radula est banale, rapprochant cette famille des Archidoridinae . . . Tout semble indiquer que, malgré des divergences, les Aegirétidés doivent étre rap- prochés des Doridiens vrais, mangeurs d’éponges.”’ Para- doxically, MILLER (1961: 106) suggests that A. puncti- lucens feeds on bryozoans (Membranipora and Electra) and THOMPSON & Brown (1976: 108) state its colora- Explanation of Figures 7 to 4 Scanning Electron Micrographs of the Radula of Aegires albopunctatus SEMs by Robert Pettyjohn and Hans Bertsch Figure '1: View of the entire radula. X 6o Figure 2: Outermost teeth of the anterior 6 rows. A basal flange is clearly visible on the inner side of the teeth (providing within-row structural support). The overlapping rows pro- vide between-row strengthening. BertscH et al. (1973: 292) comment on similar aspects of radular functional mor- phology in the Chromodorididae (cryptobranchiate sponge- feeding dorids) . X 300 Figure 3: Central part of the radula showing naked rachidian re- gion and the smaller size of the innermost 4 or 5 teeth. X 420 Figure 4: Posterior region of the radula; note the thin, developing innermost teeth. X 300 [BerTscH] Figures z to 4 THE VELIGER, Vol. 22, No. 3 ~ Vol. 22; No. 3 THE VELIGER Page 223 tion camouflages A. punctilucens to its prey item, “en- crusting polyzoans.” Radulae of the California Aegires albopunctatus Mac- Farland, 1905, bear 15-22 rows of simple, hamate teeth (Figure 1). The morphology of these teeth (Fig- ures 2 and 3) is similar to the radular teeth shape of known cryptobranch sponge feeders (compare with illus- trations in FERREIRA & BERTSCH, 1975; BLoom, 1976; BERTSCH, 1976; and throughout the opisthobranch liter- ature). Newly forming teeth (Figures 7 and 4) are weak and thin. The teeth grow and thicken first on the outer portion of the posterior rows. The most recently formed teeth occur in the center of the developing rows (Figure 4): MacFarianp (1906: 133) reports that Aegires albo- punctatus was “especially common upon sponges in a tunnel-like grotto formed by the waves near Pebble Beach, on Carmelo Bay” (Monterey, California), and RIcKETTS & CALVIN (1968: 107) state that A. albopunc- tatus can be found on a “harmless variety” of white sponge. Except for these 2 obscure statements, there is no other published mention of a food source for A. albo- punctatus. On 21 April 1979, while leading several teams of scuba divers underwater, I found 9 specimens of Aegires albopunctatus (8 - 19mm in total length, x = 12.3mm). Some were preying on the sponge Leucilla nuttingi (Urban, 1902). Five of the nudibranchs were in the ie — Woe Figure 5 The nudibranch Aegires albopunctatus, crawling in the midst of a colony of Leucilla nuttingi. The nudibranch is 16mm long Drawing by Anthony D’Attilio, based on underwater photographs taken in situ, in the Point Loma kelp beds off San Diego, California midst of different colonies of L. nuttingi (Figure 5). Clear- ly visible were sponges with pieces chewed out by the nudibranchs (Figure 6). These preliminary observations were made during a 25-minute timed search, on a 50 foot (15m) dive, in the kelp beds off Point Loma (San Diego, California), northwest of the Cabrillo Lighthouse. ANY WAM Hy WS Say y PN a = Figure 6 Aegires and Leucilla; the upper sponge has had % of its width eaten away by Aegires, leaving only a thin piece of the normally inflated-tube-shaped sponge. Drawing by Anthony D’Attilio, based on underwater photographs taken in situ, in the Point Loma kelp beds off San Diego, California To survey the association of Aegires and Leucilla, I returned to the Point Loma kelp beds with 5 pairs of scuba diving teams, on the morning of 11 May 1979. We measured every A. albopunctatus found, counted all colonies of L. nutting:, and measured the proximity of the 2 organisms. Diving depths varied from 40 - 55 feet (12-17m) ; total underwater search time of the teams was 3 hours and 16 minutes. This area was within 2km of the original location. We found 25 Aegires albopunctatus (Table 1), of which 76% were either touching or less than 30cm away from a Leucilla colony. The nudibranchs varied in total length from 4mm to 20mm (x=10.4mm). Of the L. nuttingi colonies counted, 26% were associated with the nudibranch predator (on the colony or less than 30cm distant). The habitat studied was a smooth rock substrate, with periodic sandy areas, in the midst of the kelp beds. Other Page 224 Table 1 Subtidal survey of Aegires albopunctatus and Leucilla nuttingr Total Leucilla nuttingi colonies 74 Total Aegires albopunctatus 25 Number of Aegires directly touching Leucilla 10 Number of Aegires < 30 cm from Leucilla Aegires with no proximate sponge molluscan species common in the area included the proso- branchs Haliotis spp., Mitra idae Melvill, 1893, Kelletia kelletu (Forbes, 1852), and the nudibranchs Cadlina flavomaculata MacFarland, 1905, Doriopsilla albopunc- tata (Cooper, 1863), Tritonia festiva (Stearns, 1873), Anisodoris nobilis (MacFarland, 1905), and Discodoris sandiegensis (Cooper, 1863), and, more rarely, Laila cockerelli MacFarland, 1905, and Archidoris odhneri (MacFarland, 1966). The sea urchin, Strongylocentrotus franciscanus (Agassiz, 1863) and its commensal alpheid shrimp Betaeus macginitieae Hart, 1964, were also ex- tremely common. CONCLUSION Aegires albopunctatus feeds on the calcareous sponge Leucilla nuttingi. Scuba divers have found 76% of the A. albopunctatus associated with (orin theclose proximity of) this sponge subtidally in May. Further studies are required to determine whether there is any seasonal variation in the predator-prey relationship between A. albopunctatus and L. nuttingi, or whether Aegires is monophagous. The phylogenetic relationships and superfamilial placement of the Aegiretidae need additional investigation. ACKNOWLEDGMENTS This is a contribution from the Marine Awareness Pro- gram of the PADI International College in California. I thank Commander Jim Williams for providing use of the boats Kwajalein and Kona Princess; David K. Mul- liner who teaches the PADI marine biology course with me; my fellow divers: Mark Ashbrook, Buck Buchanan, John Cuccia, Scott Haeker, Marc Harris, Jeff Knutson, Chris Miles, Mike Ortega, and Ira Stein; Robert Petty- john for assistance with the scanning electron micro- THE VELIGER Vol. 22; No. 3 scopy; Frank Tose for photographic darkroom work; Judith Bertsch for comments and help during the prepa- ration of this note; Anthony D’Attilio for his drawings; and James R. Lance for comments on the manuscript. Literature Cited BertscH, Hans 1974. Nudibranch radular morphology and prey specificity. West. Soc. Malacol. Ann. Rprt. 7: 33 (12 November 1974) 1976. Intraspecific and ontogenetic radular variation in opisthobranch systematics (Mollusca: Gastropoda). Syst. Zool. 25 (2): 117-122; 7 text figs. (9 July 1976) BzatscH, Hans, Antonio J. Frrrema, Westey M. Farmer & THOMAS L. Hayes 1973. | The genera Chromodoris and Felimida (Nudibranchia : Chrom- odorididae) in tropical west America: Distributional data, description of a new species, and scanning electron microscopic studies of radulae. The Veliger 15 (4): 287-294; 3 plts.; 3 text figs. (1 April 1973) BercH, Lunwic SopHus Rupo.F 1875. Neue Nacktschnecken der Siidsee, malacologische Untersuchun- gen. III. Journ. Mus. Godef. 3(8): 53-100; pits. 7-11 Bioom, STEPHEN A. 1976. Morphological correlations between dorid nudibranch predators and sponge prey. The Veliger 18 (3): 289-301; 1 text fig. 1 Janu 1976 Ferreira, ANTONIO J. e Hans BerTscH oases 3) 1975. Anatomical and distributional observations of some opistho- branchs from the Panamic faunal province. The Veliger 17 (4): 323 - 330; 3 plts.; 1 text fig. (1 April 1975) Loven, Sven Lupvic 1844. Om nordiska hafs-mollusker. Ofvers. K. Vet. Akad. Forh. Stockholm 1 (3): 48-53 MacFarvanp, Frank Mace 1905. A preliminary account of the Dorididae of Monterey Bay, Cali- fornia. Proc. Biol. Soc. Wash. 18: 35 - 54 (2 February 1905) 1906. Opisthobranchiate Mollusca from Monterey Bay, California, and vicinity. Bull. Bur. Fish. (1905) 25: 109-151; plts, 18-31 (24 May 1906) 1966. Studies of opisthobranchiate mollusks of the Pacific coast of North America. Mem. Calif. Acad. Sci. 6: xvit+546 pp.; 72 plts. (8 April 1966) McDonatp, Gary R. & James WILLARD NYBAKKEN 1978. Additional notes on the food of some California nudibranchs with a summary of known food habits of California species. The Veliger 21 (1): 110-119 (1 July 1978) Mitier, MicHazt CHares 1961. Distribution and food of the nudibranchiate Mollusca of the south of the Isle of Man. Journ. Anim. Ecol. 30 (1): 95-116 (May 1961) Orsieny, Ace CHarzes Victor DESSALINES D’ 1837. Mémoire sur les espéces et sur des genres nouveaux de l’ordre des nudibranches observés sur les cotes de France. Mag. de Zool. 7 Class 5: 1-16; plts. 102-109 [not seen; fide RussELL, 1971: 15] Pruvot-Fot, AuicE 1954. Mollusques opisthobranches. 1 plit.; 173 text figs. Ricketts, Epwarp FE « Jack Ca.vin 1968. Between Pacific tides. 4th ed. rev. by Joel W. Hedgpeth. xiv+ 614 pp.; illus. Stanford Univ. Press, Stanford, Calif. Russe.__, Henry DrummMoNnD 1971. Index Nudibranchia. Faune de France 58: 1 - 460; Delaware Mus. Nat. Hist. iv+141 pp. (1 July 1971) THompson, THomAs Everett & Grecory H. Brown 1976. British opisthobranch molluscs. Synopses of the British Fauna (N.S.) no. 8. London, Acad. Press, 203 pp.; 1 color plt.; 105 text figs. (1 September 1976) Ursan, F 1902. Rhabdodermella nuttingi nov. gen. et nov. spec. Zeitschr. wissensch. Zool. 71 (2): 268-275; plt. 14; 1 text fig. Younc, Davw KENNETH 1970. The functional morphology of the feeding apparatus of some Indo-West Pacific dorid nudibranchs. Malacologia 9 (2): 421-446; 17 text figs. (20 July 1970) Vol. 22; No. 3 THE VELIGER Page 225 Karyotypes of Six Eastern Pacific Acmaeid Gastropods DAVID M. CHAPIN anp PAUL A. ROBERTS Department of Zoology, Oregon State University, Corvallis, Oregon 97331 (1 Plate; 4 Text figures) INTRODUCTION INTERTIDAL LIMPETS OF THE FAMILY Acmaeidae are globally widespread and particularly well represented in the Eastern Pacific, where at least 24 species can be found from Baja California to Alaska. Among these many have widely overlapping ranges and sympatric associations of five or more are not uncommon. The systematics of the Acmaeidae have been extensively studied by Test (1938) and McLean (1966). Test based her arrangements mainly on radular ribbon characteristics, McLean on both radular teeth characteristics and shell morphology. The two studies are in close agreement as to their designations of species but differ substantially in their generic and subgeneric groupings. Test included all but one species (Lottia gigantea Sowerby, 1834) in the one genus Acmaea. But McLean (1966), whose arrangement is currently accepted, divides the Eastern Pacific species into three genera: Acmaea, Collisella, and Lottia. Mc- Lean now (1969) regards his subgenera of Collisella, C. (Collisella) and C. (Notoacmea), as full genera. Karyotypes provide another taxonomic character to consider in the systematics of any species group, and it is the intention of this study to contribute cytotaxonomic evi- dence to Acmaeid systematics. Here, the karyotypes of 3 members each from Notoacmea and Collisella are pre- sented: N. scutum (Rathke, 1833), N. fenestrata (Reeve, 1855), N. persona (Rathke, 1833), C. digitalis (Rathke, 1833), C. pelta (Rathke, 1833), and C. strigatella (Car- penter, 1864). Chromosome number, size, and centromere position are the karyotypic characteristics examined. The only other cytotaxonomic study of the Acmaeidae (see PATTERSON, 1969, for the most recent review of mol- luscan chromosomes) was that of NisHikawa (1962) on 6 Japanese limpets. Three of these he classified in the genus Notoacmea: N. schrenckii (Lishke, 1868), N. concinna (Lishke, 1870), and N. fuscoviridis Teramachi (1949); and 3 in the genus Patelloida: P. saccharina lanx (Reeve, 1855), P. pygmaea (Dunker, 1860), and P. lampanicola Habe (1944). In all of these species, and in 3 species of the closely related family Patellidae (both are in the superfamily Pa- tellacea), he found a consistent haploid number of g. Al- though details of chromosome morphology are lacking from Nishikawa’s study, his findings provide an opportu- nity to compare chromosome counts of Western Pacific Acmaeids to those of the Eastern Pacific. MATERIALS anp METHODS Limpets of the 6 species of Acmaeidae examined in this study were collected from rocky intertidal habitats along the central Oregon coast, between Yachats on the south and Neskowin on the north. Only those individuals that could be clearly classified as to species (according to the keys of Licut et al. (1975), KozLorr (1974) and McLean (1969)) were used. The animals were kept in a running saltwater facility at the Oregon State University campus until they were ready to be examined for karyotypes. Two methods were used to obtain karyotypes of these 6 species: the first employed colchicine arrested metaphase I spermatocyte nuclei; the second, cultured embryos, which yielded mitotic cleavage nuclei. Spermatocytes were obtained by first injecting 0.2mL of 0.5% colchicine into the body cavity (to enrich the re- covery of metaphase nuclei), by dissecting out the testis two hours later, and then submitting the testis for about 15 min. to a hypotonic 1% Na,citrate solution to swell the nuclei. Testes were subsequently macerated, fixed in freshly made Carnoy’s solution (3 prts. 100% ETOH: 1 prt. glacial acetic acid) for 2 h, and stained for 5 min. on a siliconized glass slide with 1% lactic-acetic orcein. Stain- Page 226 THE VELIGER Vol. 22; No. 3 ing was followed by rather vigorous squashing under a cover slip. With this technique, only highly condensed metaphase I nuclei could be observed (Figure 1), useful for chromosome counts but unsuitable for any characterization of chromosome morphology. With cultured embryos, however, large numbers of mi- totic metaphases could be obtained, showing both chromo- some number and karyotypic details of chromosome size and centromere position (Figure 3). Fertilization was car- ried out following the procedure reported by STRATHMAN (1975). Embryos were subjected to a colchicine treatment during the last five hours of culture (4mL of 0.5% colchi- cine per 200 mL sea water), which markedly increased the number of metaphase nuclei in embryos beyond the 32 cell stage. Unfertilized embryos were usually arrested at late diplotene, and thus were useful for confirming chro- mosome counts (Figure 2). At the end of the culture period the embryos were put in a hypotonic solution (1 prt. sea water: 2 prts. distilled water) to swell the nuclei and then fixed for several hours in fresh Carnoy’s. The fixed embryos were treated over- night in a 1 part chloroform: 1 part ethanol/methanol solution to extract the yolk and then stored in 70% eth- anol. Mitotic nuclei from these embryos were obtained by staining a few embryos on a siliconized glass slide with 1% lactic-acetic orcein for 5 to 10 min. and then squashing them under a siliconized cover slip with light to moderate pressure. Broken nuclei were often mixed making chromo- some counts difficult, but this was less of a problem with more mature embryos. Only discrete, well-separated nu- clei were used to establish karyotypes. Photographs (such as Figure 3) proved inadequate for a clear characterization of karyotype; consequently, cam- era lucida drawings were made with a Zeiss phase contrast microscope using a 25X ocular and a 100X objective (Figures 5 and 6). To obtain the karyotypes, drawings of individual chro- mosomes were measured to the nearest half millimeter, the lengths of the 2 short arms (S) and 2 long arms (L) were each averaged, and with these two values the arm ratio (R=L/S) and % of total genome (% = = a (100)) were calculated. From these values L. + S: individual comeare were plotted (R vs. % of total genome) to distinguish pairing relationships (Figure 4a). From 3 such karyotypes (one from each of 3 separately fertilized cultures) a pattern of matched chromosome pairs was determined (Figure 4b). Although there was some scatter within a chromosome pair of a sample karyotype and for each chromosome pair of a species karyotype, a definite clustering of pairs from the sample karyotypes permitted determination of the species karyotype. In some cases (e.g. chromosome pairs 6 and 8, Figure 4a) where alternative pairing relationships were reasonably possible for a sample karyotype, the pairing which was used was the one most consistent with the other sample karyotypes for that species. Distortion during squashing most likely accounts for the scatter within a chromosome pair, but possible intraspecific chromosomal variation within the limits of the scatter was not resolvable. To summarize, from three separately fertilized embryos of each species the average size (% of total genome) and centromere position( R value) for particular chromosome pairs were determined. The karyotype of each species is the series of chromosomes, in order of ascending size, char- acterized by these values. RESULTS anp OBSERVATIONS For all six species the modal number of chromosomes in mitotic cleavage nuclei was 2n=20. This number was found in virtually all discrete yet well spread nuclei exam- ined, and was substantiated by a haploid count of 10 made in several metaphase I spermatocytes (Figure 1) or unfer- tilized oocytes (Figure 2) for each species. Although these animals have separate sexes, no typical sex chromosome characteristics (such as heteromorphic pairs or differential staining of any elements) were observed. Quantified karyotypic data (% of total genome and R values) of both the sample and species karyotypes are pre- sented in Table 1. Comparative karyotypes of camera lu- Explanation of Figures 1 to 3 Figure 1: Photographs of 2 metaphase I spermatocyte nuclei (squash preparations) from Collisella digitalis. Ten bivalents are clearly visible in each photograph X 625 Figure 2: Photograph of late diplotene oocyte nucleus (squash preparation) from Notoacmea fenestrata. Ten bivalents vis- ible X 250 Figure 3: Photograph of colchicine metaphase nucleus from cleav- ing embryo of Notoacmea scutum. 2n=20 X 250 THe VELIGER, Vol. 22, No. 3 [CHapin & Roperts] Figures 7 to 3 Figure rb Vol. 22; No. 3 THE VELIGER Page 227 E (sj 6 ia ~~ 5 ij 0 f=] 2 Il a4 2.0 3.0 4.0 5.0 6.0 7.0 8.0 4.0 % of total genome 7 B 3 Eo S 5 a 6 7 E 7 5 g 3 be SOS, ‘ {e) 4 5 7 =! G2{0) I 6 I 6 6 9 fas 5 10 7 7 “aoe qos oe a no Oa 7 8 7 5 4.0 6.0 8.0 10.0 12.0 14.0 16.0 % of total genome Figure 4 a) Individual mitotic chromosomes from one metaphase cleavage nucleus of Collisella digitalis plotted as R vs. % of total genome. Numbers designate chromosome pairs. b) Graph of chromosome pair values (small circles) for R and % of total genome from 3 nuclei of Collisella digitalis and of their cida drawn chromosomes can be seen in Figure 6, and an idiogram for the karyotype of Collisella digitalis, con- structed from the data in Table 1, is shown in Figure 7. Certain aspects of these data need comment. Firstly, chromosome pairs 3 and 4 were not consistently distin- guishable in Notoacmea persona and N. fenestrata; hence, they were averaged. Secondly, in the karyotypes of both Collisella strigatella and N. scutum pair 6 is out of size order. In each case 2 of the 3 samples warranted the size order designated in the karyotypes, and the centromere mean values (large circles). Pair values for each sample determined from mean of individual homolog values. Mean values for R and % of total genome from 3 samples characterize species karyotype. Small numbers refer to samples 5, 6, and 7, and large numbers designate chromosome pair number positions of that chromosome for all 3 samples of the 2 species were close to those of chromosome 6 from the other 4 species. Putting these 2 chromosomes of anomalous size in any other order would indicate karyotypic differences that may not be real. But because of these and other varia- tions among the samples of a species karyotype, the resolu- tion of this method of determining karyotypic differences between species is somewhat reduced. If we err, we err on the side of a conservative estimate of the amount of chro- mosomal rearrangement in the Acmaeidae. No. 3 b) Vol. 22 THE VELIGER Page 228 Size and arm ratios of chromosome pairs from sample and species karyotypes of six Eastern Pacific Acmaeidae. Table 1 Chromosome number Sample Species teases 1 2 3 4 5 6 7 8 9 10 mean of cammygles J I ©)P % IR Cle % I Cle % ik Ce % IX ©P FH we Ce Gir ©P GY Ir CGP BIR Ge % IR Gp Notoacmea 3 4.4 2.38 8.3 3.17 8.3 2.03 9.1 2.75 9.6 2.94 10.0 1.26 10.7 1.48 12.6 1.66 12.8 1.47 14.0 1.55 scutum 7 5.0 1.41 6.4 4.59 8.5 2.79 9.8 2.73 10.2 2.84 7.5 1.37 10.1 1.27 12.9 2.41 12.9 1.53 14.2 1.24 8 4.9 1.63 6.2 3.50 8.4 2.62 9.0 4.50 9.2 2.45 9.9 1.68 11.3 1.65 12.8 2.82 13.6 1.48 14.5 1.95 mean 4.8 1.81 sm_ 7.0 3.75 st 8.4 2.48 sm 9.4 3.33 st 9.7 2.74 sm 9.1 1.44 m 10.7 1.47 m 12.8 2.30 sm 13.1 1.49 m 14.2 1.58 m Notoacmea 5 4.5 1.54 6.8 3.64 8.9 2.22 9.5 1.25 10.6 1.52 12.3 1.29 12.4 2.00 13.1 1.13 13.3 1.65 fenestrata 6 4.8 1.93 7.2 4.14 9.2 2.11 nos3&4 10.1 1.16 11.8 1.82 11.5 1.25 12.3 1.75 12.3 1.16 13.4 1.89 7 4.5 2.43 6.3 3.38 8.2 2.03 averaged 8.7 1.20 11.2 1.74 12.0 1.20 12.1 1.48 13.8 1.13 15.3 1.99 mean 4.6 1.97 sm_ 6.8 3.72 st 8.8 2.12 sm 9.4 1.20 m 11.2 1.69 m 11.9 1.25 m 12.3 1.74 sm 13.1 1.14 m 14.0 1.84 sm Notoacmea 7 3.4 1.76 6.4 2.90 8.8 1.93 10.6 1.71 10.3 1.19 11.2 1.15 12.8 2.71 12.6 1.39 14.4 1.57 persona 9 4.4 1.74 6.4 2.33 8.4 2.04 nos. 3 & 4 9.4 2.03 11.2 1.49 12.0 1.39 11.4 2.84 12R ely, 15.6 1.57 10 4.4 1.78 6.6 2.61 8.2 2.36 averaged 9.4 2.24 10.4 1.09 11.5 1.23 12.8 2.95 13.6 1.36 15.6 1.24 mean 4.1 1.76 sm 6.5 2.61 sm 8.5 2.11 sm 9.8 1.99 sm 10.6 1.26 m 11.6 1.26 m 12.3 2.83 sm 13.0 1.31 m 15.2 1.36 m Collisella 8 5.0 2.23 7.9 2.08 9.2 3.63 8.3 2.65 9.6 2.06 10.8 1.13 10.3 1.51 12.1 1.09 11.7 1.88 16.1 1.56 pelta 9 3.5 2.07 8.1 2.25 8.3 3.09 9.2 2.42 9.5 2.16 11.0 1.20 11.3 2.03 12.7 1.30 12.8 1.88 14.6 1.26 10 3.4 1.77 7.7 2.68 8.2 3.40 8.7 2.90 9.7 1.98 10.8 1.16 10.7 2.03 12.3 1.12 13.0 1.56 15.0 1.77 mean 4.0 2.02 sm 7.9 2.34 sm 8.6 3.37 st 8.7 2.66 sm 9.6 2.07 sm 10.9 1.16 m 10.8 1.86 sm 12.4 1.17 m 12.5 1.77 sm 15.2 1.53 m Collisella 5 4.5 1.66 7.6 1.14 7.6 2.78 8.8 2.25 10.5 2.38 10.7 1.29 11.5 1.86 11.6 1.23 13.2 2.04 14.1 1.51 digitalis 6 3.6 1.82 7.9 1.12 8.8 2.60 9.0 1.91 9.5 2.33 11.3 1.31 11.4 2.56 12.2 1.36 12.3 1.96 14.9 1.60 7 48 1.91 8.4 1.26 8.8 3.85 8.4 2.26 10.6 2.22 10.8 1.53 11.4 1.99 11.6 1.30 12.8 1.57 13.4 1.35 mean 4.3 1.80 sm 8.0 1.17 m_ 8.4 3.08 st 8.7 2.14 sm 10.2 2.31 sm 10.9 1.38 m 11.4 2.14 sm 11.8 1.30 m 12.8 1.86 sm 14.1 1.51 m Collisella 5 4.8 2.21 7.5 2.24 8.1 2.67 9.1 2.96 10.0 2.64 9.9 1.22 11.5 1.12 11.0 1.55 13.2 1.64 14.8 1.70 strigatella 6 4.0 2.21 7.7 1.92 8.7 3.20 9.3 3.19 10.3 2.11 7.8 1.12 11.0 1.17 11.9 1.45 13.1 1.27 16.2 1.71 8 4.4 1.98 8.0 3.11 9.2 3.58 10.3 3.95 9.8 2.81 9.6 1.27 10.2 1.28 11.6 1.70 12.1 1.38 14.8 1.31 mean 4.4 2.13 sm 7.7 2.42 sm_ 8.7 3.15 st 9.6 3.37 st 10.0 2.52 sm 9.1 1.20 m 10.9 1.19 m 11.5 1.57 m 12.8 1.43 m 15.3 1.57 m % = % of total genome m = median region (1.0 < R < 1.7) R = long arm length/short arm length CP = centromere position mean sample values = species karyotype values sm = submedian region (1.7 < R < 3.0) st = subterminal region (3.0 < R <7) Vol. 22; No. 3 THE VELIGER Page 229 The karyotypes of these six species show several general similarities. The range in size is fairly consistent among all the karyotypes: one very small chromosome comprising 4 to 5% of the total genome length (about 24m to 3 wm; see scale in Figures 5 and 6) with the other 9 chro- mosomes in a more or less continuously ascending series of size up to 14 to 16% of the total genome length(about9 um to 10 um). Centromere position is generally median to sub- median, with no pairs having terminal centromeres. Over- all, the larger chromosomes have more median centro- meres than the smaller ones. ARR RB Ah AY PYSK sap WSANP YE Figure 5 Camera lucida drawings of chromosomes paired from graph in Figure 4a (Collisella digitalis, sample 5) In a comparison of the chromosomes of equivalent size among the 6 species, few marked changes in centromere position are observed. In Notoacmea fenestrata the centro- mere of chromosome 5 has a more median position than those of the equivalent chromosomes from the other spe- cies. Notoacmea persona has a noticeably more terminal centromere position for chromosome 8 than do the other species. Collisella digitalis shows two differences in com- parison to the other species, one of these shared by C. pelta. Chromosome 2 of C. digitalis has a distinctly median cen- tromere in contrast to the sub-median to sub-terminal centromere of no. 2 in the other 5 species. Chromosome 7 of both C. digitalis and C. pelta is more sub-median in centromere position than the median no. 7 of the other species. Centromere shifts may occur by pericentric inversion (by 2 breaks) or by the rarer transposition of the centro- mere containing segment into another part of the chromo- some (3 breaks required). The few centromere shifts re- I 2 3 4 5 eR YY Notoacmea fenestrata sO HSK AK Notoacmea persona HHP Uf 4 Collisella pelta APAPA SUNY gitalis RAAPIK ABR TAT Figure 6 Karyotypes of 6 acmaeid species examined in this study. Only 1 chromosome per pair is presented (n= 10). All chromosomes in each karyotype are camera lucida drawings from 1 sample nucleus. Arrows indicate karyotypic differences discussed in text corded may be due to a modest number of pericentric inversions fixed in the course of speciation of these limpets. One might make the criticism that due to size variation among the samples comprising each species karyotype, designated chromosome numbers may not reflect the same homologous pair throughout the 6 species; consequently, the differences just pointed out may be accounted for by changes in the designation of chromosome number. But after making any reasonable rearrangements in size order within a karyotype, these reported differences represent a minimum estimate of karyotypic change among the 6 species. Page 230 THE VELIGER Vol. 22; No. 3 8 chromosome number between these 6 Eastern Pacific spe- 2 i4] cies (n= 10) and the 6 Western Pacific Acmaeidae (n=9) e 12 hi examined by NisHikawa (1962) is striking. to 10 A a Onl if i a no A change of one in chromosome number can be brought 2 6 mill q about by either a dissociation or fusion. Although the pre- 3 Stef il | wil ili wil | cise mechanisms are not well understood, a dissociation g Chol can be thought of as the division of a metacentric chromo- 15679 3 4 5 6 7 8 9 Figure 7 Idiograms of species karyotype (thick chromosomes) and sample karyotypes (thin chromosomes) for Collisella digitalis. Small num- bers below chromosomes (5, 6, and 7) indicate order of sample karyotypes in idiogram. Idiograms were constructed from R (long arm length/short arm length) and % of total genome values for Collisella digitalis presented in Table 1 and Figure 4b DISCUSSION It appears from these data that speciation of Acmaeid gastropods has not been accompanied by extensive chro- mosomal rearrangement. Nevertheless, it must be kept in mind that rearrangements that do not visibly alter chro- mosome length or centromere position (e.g. symmetric pericentric inversions or paracentric inversions) may be present in these species. Although genomic reorganization at the level of detect- able karyotypic differences can sometimes act as a genetic isolating mechanism through the promotion of hybrid sterility or inviability (WHITE, 1973), the present results indicate that chromosomal rearrangements are not likely to play an important role in the reproductive isolation of sympatric Acmaeids. The question of the nature of isolat- ing mechanisms remains open, although Fritchman’s study of Acmaeid reproductive periods suggests that non-coin- cident spawning stimuli could in some cases result in repro- ductive isolation of sympatric limpets (FrRITCHMAN, 1961a, 1961b, 1961c, 1962). There is a paucity of comparative karyotypic data re- ported so far on gastropod species (see PATTERSON, 1969, for review), but a study by BurcH (1967) on the Japanese melaniid genus Semisulcospira shows that the Gastropoda are not all as karyotypically conservative as the Acmae- idae. Burch reported variations in chromosome number from n=7 to n=20 among the Semisulcospira species ex- amined and numerous differences in centromere position in the karyotypes of those species with the same number of chromosomes. Although few differences in karyotype among the East- ern Pacific Acmaeidae were found here, the difference in some into two acrocentrics and a fusion as the joining of two acrocentrics into a metacentric, a process sometimes called Robertsonian translocation. It is not clear which of these two events is more likely to occur; therefore, one can not say whether n=9 or n= 10 is the more primitive con- dition in the Acmaeidae. The difference in chromosome number between the Eastern and Western Pacific Acmaeidae does, however, indicate that the species of Nishikawa’s study and those analyzed here may represent separate phylogenetic groups: one n=Q group, including 3 species of Patelloida (P. sac- charina lanx, P. pygmaea, P. lampanicola) and 3 species of Notoacmea (N. schrenckit, N. concinna, N. fuscoviridis) ; and a n=1I0 group, including 3 species of Collisella (C. digitalis, C. pelta, C. strigatella) and 3 species of Notoac- mea (N. scutum, N. persona, N. fenestrata). But this pre- sents a perplexing taxonomic situation if members of the same genus, Notoacmea, have members in each cytotaxo- nomically distinguishable group. The present generic groupings of Patelloida, Collisella, and Notoacmea are based primarily on radular teeth char- acteristics: Collisella has uncini (rudimentary marginal teeth), Notoacmea lacks them, and Patelloida has two pairs of elongate marginal teeth (McLgEan, 1966, 1969). No distinct shell differences among these 3 genera are evident, with the exception that no species of Notoacmea possesses heavy radial ribbing, whereas some some species of Col- lisella and Patelloida are heavily ribbed. (The genus Acmaea is clearly distinguishable from the other Acmaeid genera by both shell and radular teeth characters (Mc- LEAN, 1966).) Based mainly on the one morphological criterion of radular teeth characteristics, these classifica- tions would seem to be unambiguous. But reliance on pri- marily one characteristic (radular teeth) for classification may be less than adequate if the taxonomy of a species group is to reflect its phyletic relationships. Because a mis- representation of the phylogenetic history of the Acmae- idae can only confuse further evolutionary studies, it is important to determine whether the cytotaxonomic evi- dence warrants a reexamination of Acmaeid systematics. It is possible that NisHikawa’s cytological technique (1962) failed to reveal a small chromosome (for example chromosome 1) common to the karyotypes of all 6 species in this study. Although Nishikawa’s data are presented Vol. 22; No. 3 THE VELIGER Page 231 LLL en RES only as camera lucida drawings of metaphase I spermato- cyte nuclei, the clarity of his drawings and the fact that all of the metaphase I spermatocyte chromosomes of the spe- cies examined by us are darkly stained and distinct (Figure 1) make it unlikely that Nishikawa miscounted in all 6 spe- cies from the Western Pacific. Since the chromosome counts of the Eastern Pacific Acmaeidae reported here are well substantiated, it appears that there is a fundamental discrepancy between taxonomic classifications based on morphological criteria and the chromosomal evidence. Although Test’s groupings (subgeneric) based on radular ribbon criteria differ substantially from McLean’s (ge- neric), her arrangement of the species included in Nishi- kawa’s and the present study is also found to be contradic- tory to the cytotaxonomic data. In light of the inconsistencies between the classification based primarily on the single morphological criterion of radular teeth and the karyotypic data, it would seem de- sirable to confirm Nishikawa’s results and to examine more anatomical and physiological characteristics of this group. Differences in enzyme electrophoretic patterns or hybrid viability studies could reveal phyletic groupings; and kary- optic studies (at least of chromosome number) need to be extended to more species for a more complete picture of cytotaxonomic groupings in the Acmaeidae. Perhaps, too, the phylogenetic significance of radular characteristics needs to be reevaluated. With these and other approaches possible, taxonomic work on the Acmaeidae certainly de- serves more attention, for an adequate understanding of the interesting and complex evolutionary history of this family has yet to be achieved. SUMMARY 1. Chromosome numbers in 3 species of Notoacmea (N. scutum, N. persona, N. fenestrata) and 3 species of Collisella (C. pelta, C. digitalis, C. strigatella), all found along the Pacific coast of temperate North America, were determined to be n=10 in each case. 2. Few differences in chromosome size or centromere position were observed among any of the 6 species karyotypes determined in this study. Collisella pelta, Notoacmea persona, N. fenestrata all show one clear change in centromere position and C. digitalis shows 2. 3. The count of n= 10 differs from that of n=9 reported by Nishikawa (1962) for 3 species of Notoacmea and 3 species of Patelloida (also in the Acmaeidae) from the Western Pacific. In light of these differences, cur- rent taxonomic groupings of the Western and Eastern Pacific Notoacmea should be reevaluated. ACKNOWLEDGMENTS We would like to thank Christopher J. Bayne and James R. Allen for their helpful comments and suggestions. Literature Cited Burcu, JoHNn Bayragp 1967. Cytotaxonomy of some Japanese Semisulcospira ( Streptoneura: Pleuroceridae). Journ. Conchol. 107: 3-51 FritcuMan, Harry Kr, II 1961a. A study of the reproductive cycle in the California Acmaeidae (Gastropoda), Part I. The Veliger 3 (3) :57-63; plt. 10 (1 January 1961) 1961b. A study of the reproductive cycle in the California Acmaeidae (Gastropoda), Part II. The Veliger 3 (4): 95-101; plts. 15-47 (1 April 1961) 1g961c. A study of the reproductive cycle in the California Acmaeidae (Gastropoda), Part III. The Veliger 4 (1): 41-47; plts. 9-14 (1 July 1961) 1962. A study of the reproductive cycle in the California Acmaeidae (Gastropoda), Part IV. The Veliger 4 (3): 134-140; plts. 30-32 (1 January 1962) Koz.orr, EucENE 1974. Keys to the intertidal invertebrates of Puget Sound, the San Juan Archipelago, and adjacent regions. Univ. Wash. Press, Seattle, Wash. x+226 pp.; illust. McLean, James Hami_Ton 1966. West American prosobranch gastropods: superfamilies Patella- cea, Pleurotomariacea, and Fissurellacea. Ph. D. thesis, Stanford Univ., Stanford, Calif. x+225 pp.; 7 plts. 1969. Marine shells of southern California. Los Angeles County Mus. Nat. Hist. Sci. Ser. 24, Zool. 11: 104 pp.; 54 figs. NIsHIKAWA, §S. 1962. A comparative study of chromosomes in marine gastropods with some remarks on cytotaxonomy and phylogeny. Journ. Shimono- seki Coll. Fish. 11 (3): 149-186 PATTERSON, CHARLOTTE MorGAN 1969. Chromosomes of molluscs. Symp. on Mollusca II: 635 - 686 Suiru, Ravpo INcraM & James T. CariTon (eds.) 1975. Light’s Manual. Intertidal invertebrates of the central California coast. grd ed., xviilit 716 pp.; 156 ples. (8 May 1975) STRATHMAN, MEGUMI 1975. Methods in developmental biology. Friday Harbor Labor- atories, Friday Harbor, Washington Test, Avery Ransome GRANT 1938. A systematic revision of the genus Acmaea Eschscholtz, includ- ing consideration of ecology and speciation. Ph. D. thesis. Univ. Calif., Berkeley, California, 432 pp.; 35 plts. 1945. Ecology of California Acmaea. Ecology 26 (4): 395-405 (1 October 1945) Univ. Michig. Mar. Biol. Assoc. India, Proc. 1946. Speciation in limpets of the genus Acmaea. Contr. Lab. Vert. Biol. g1: 1-24 Wuite, Micuagt JAMES DENHAM 1973- Animal cytology and evolution. Press, Cambridge viii+961 pp.; illust. 3m ed. Cambridge Univ. Page 232 THE VELIGER Vol. 22; No. 3 The Distribution of Shallow-Water Marine Prosobranch Gastropod Molluscs Along the Coastline of Western Australia BY FRED E. WELLS Western Australian Museum, Francis Street, Perth, Western Australia 6000 (5, Text figures) INTRODUCTION THE STATE OF WESTERN AUSTRALIA is the largest of the Australian States, occupying fully a third of the Austral- ian continent. The coastline of the State is vast, about 7000 km long, and has a wide variety of habitats. The coast spans some 21° of latitude (14° to 35°S) and 15° of longitude (114° to 129° E). This western portion of Australia was regarded as a wasteland by the early settlers who bypassed it on their way to the east coast. As the process of exploring and settling the eastern states con- tinued, scientific knowledge of the fauna there surged ahead. With no population and little exploration in the west, the study of the fauna of Western Australia lagged behind. The first settlement in Western Australia was founded in Albany in 1827. By that time Sydney was al- ready a thriving town and the Australian Museum was being established. The early history of the scientific exploration of the Western Australian coastline was summarized by Hedley (1916), who compiled the published descriptions of Western Australian molluscs and found that about 800 species had been recorded in the State to that time. Of these about 550 were marine gastropods. While many of the species have since been synonymized or transferred to (adjacent column —) Figure 1 Map of Western Australia showing the locations of points men- tioned in the text Dampier Barrow Archip. @ Grediand ieee Feds WESTERN AUSTRALIA Esperance Naturaliste Cape Leeuwin Vol. 22; No. 3 THE VELIGER Page 233 other genera, Hedley’s compilation is still the most exten- sive listing of Western Australian molluscs. HEDLEY (1926) later published an analysis of the zoogeography of Austral- ian marine molluscs, but the species ranges in Western Australia were poorly known at that time. Many collec- tions have been made in the intervening 74 years and the details of the distributional patterns of marine gastropods along the Western Australian coastlines can now be eluci- dated. MATERIALS anp METHODS The distributions in Western Australia of 440 species of prosobranch gastropods were established by analyzing the collections of the Western Australian Museum (WAM). Cate (1964; 1968) and Witson & GILLETT (197!) also used the WAM collections but had access to private col- lections which extended the known ranges of some species Table 1 Prosobranch families investigated in this paper. Number of Family species investigated Archeogastropoda Haliotidae 10 Trochidae 37 Turbinidae U7 Neritidae 8 Mesogastropoda Littorinidae 9 Strombidae 15 Cypraeidae 61 Naticidae 14 Cassidae 14 Tonnidae 6 Neogastropoda Muricidae 26 Thaididae 22 Columbellidae 22 Fasciolariidae 13 Nassariidae 19 Olividae 9 Mitridae 51 Volutidae 23 Conidae 48 Terebridae 16 beyond that recorded in the Museum holdings; the ex- tended ranges have been used in this analysis. The 20 families selected for analysis are those that are best repre- sented in the collection and are well known taxonom- ically. Table I lists the 20 families and the number of species examined in each; Appendix I lists the species examined and their ranges. The analysis provides a good coverage of the three prosobranch orders: 72 archeo- gastropods, 119 mesogastropods, and 249 neogastropods are discussed. This encompasses most of the prosobranchs whose range can be established with the WAM collections and the results can be considered to be representative. The WAM collections are the result of an intensive field collecting programme conducted throughout the State during the last 14 years. Only material from the inter- tidal or the shallow depths that can be reached by SCUBA techniques, is analyzed in this paper. The overall outlines of the distributions of shallow-water prosobranchs can be established, but not every species has been collected at all of the major localities. In the case of widespread tropical Indo-Pacific or warm temperate southern Australian spe- cies the lack of a specimen from a particular locality in the middle of its range is considered to be an artifact and the ranges discussed assume the species will be recorded later at intermediate localities. This generalization of species ranges might obscure areas along the coast where groups of species are absent. RESULTS Overall Patterns of Distribution Three general distributional patterns emerge from the overall analysis of species distributions: the majority (308) of the species are widely distributed tropical species found in the Indo-Pacific region and along the tropical coasts of northern Australia, extending into Western Australia; a smaller group of 94 species are warm temperate forms distributed along the southern coast of Australia, includ- ing the south coast of W.A.; the smallest segment of the fauna is a group of 38 species that are endemic to West- ern Australia. There are no major distributional features exhibited by the tropical species along the north coast (Figure 2). Of the 308 tropical species, 230 extend as far west as the North-West Cape-Barrow Island area. The 78 species that do not reach the North-West Cape area drop out grad- ually along the north coast. The North-West Cape area is the major geographical limit for the tropical fauna with Page 234 go species having their range limits in the area. Two smaller areas of substantial range limits occur along the west coast at Shark Bay, where 43 species terminate, and the Houtman Abrolhos (47). Only 47 of the 308 tropical species extend south of the Houtman Abrolhos. These decrease gradually along the west coast until only 9 have been recorded at Cape Leeuwin at the southwestern cor- ner of the State. Four tropical species (Clanculus consor- binus, Cypraea caputserpentis, C. helvola, and Natica qualtieriana) are found along the south coast to Albany. One species, Natica sagittata, is recorded as far as Esperance. A similar, but reverse pattern, is exhibited by the warm temperate species (Figure 2). Eighty-eight of the 94 spe- cies are distributed along the entire south coast to Cape Leeuwin. Seventeen of these extend only to the Cape Leeuwin - Cape Naturaliste area. The major distribution limit for the warm temperate group is in the Perth area, which is the extreme range for 31 species. Forty of the southern species occur north of the Perth area, but 24 of these do not extend north of the Houtman Abrolhos- Geraldton area. The remaining 16 exhibit a gradual de- cline further north on the west coast, and only three extend to the north coast: Nassarius particeps, which ex- tends to the Dampier Archipelago, and Eunaticina din- geldi and Oliva australis, both of which are found as far as Broome. The distributions of the 38 species endemic to Western Australia are shown on Figure 2, and the species are listed on Table 2. Most occur only along the west coast though some extend to the north or south coast. The percentages of endemic species are about the same in the 3 proso- branch orders: 7.0% in the Archeogastropoda, 8.4% in the Mesogastropoda, and 9.2% among the Neogastropoda. None of the orders has a rate of endemicity substantially different from the average rate of 8.6% endemic. The presence of two overlapping faunas on the west coast provides an interesting area for the examination of the changes which occur in species populations near the extreme limit of their ranges. Figure 3 shows the relative percentages of northern and southern species along the west coast. There is a clear decline in the percentage of tropical species southward along the coast. The molluscan fauna of both North-West Cape and Shark Bay is almost completely tropical. The Abrolhos is 72% tropical. South of the Abrolhos the percentage of tropical species declines sharply, decreasing to 39% in the Perth area. The fauna of the southwest corner of the State is predominately warm temperate. The major overlap between the tropical and warm temperate faunas occurs on the stretch of coastline between Perth and the Houtman Abrolhos. THE VELIGER Vol. 22; No. 3 Table 2 Species investigated that are endemic to Western Australia. Archeogastropoda Haliotidae Haliotis elegans H. semiplicata Trochidae Calliostoma ciliaris C. lepidus Clanculus occidus Mesogastropoda Littorinidae Nodilittorina australis N. nodosa Tectarius rusticus Cypraeidae Cvpraea armeniaca C. brevidentata C. decipiens C. pulicaria C. rosselli C. venusta Cassidae Phalium whitworthi Neogastropoda Muricidae Haustellum wilsont Pterynotus westralis Dermomurex antontus Thaididae Cronta avellana Columbellidae Zafra mitriformis Fasciolariidae Latirus walkeri Mitridae Mitra backae M. chalybeia M. gilbertsont M. hansenae M. marrow Volutidae Amonia ellioti A. grayi A. irvinae A. macandrewt A. nivosa Volutoconus coniformis V. hargreavesi Conidae Conus clarus C. dorreensts C. kenyonae C. nodulosus Terebridae Duplicaria crakei Vol. 22; No. 3 THE VELIGER Page 235 Temperate Tropical Kimberleys Derby Broome Pt. Hedland Dampier North Coast Onslow North-West Coast Endemic Cypraea Conus Trochidae Carnarvon Shark Bay Abrolhos Perth Naturaliste West Coast Leeuwin Albany Esperance Figure 2 Distribution of marine prosobranch gastropods in Western Australia There is a tendency for the Houtman Abrolhos and Rott- nest Island to have a more tropical fauna than the adja- cent inshore areas; this is particularly true of the Abrolhos. At Geraldton, 60 km east of the Abrolhos, the coastal fauna is much more temperate in nature, though the percentages of tropical species used here do not indicate this. The dif- ference between the molluscan faunas of the Abrolhos and Geraldton could be better shown by a detailed analysis which included relative abundances instead of just pres- ence and absence. The ocean waters at Geraldton reach a minimum of 18° C during the winter (HopcKIN & PuiL- LIPS, 1969), while the minimum at the Abrolhos is 19° C (Mars, 1976). The difference, though small, is crucial since a minimum of 20°C is generally accepted as the limit of a tropical fauna (Briccs, 1975). Distribution of the Family Cypraeidae The genus Cypraea has been extensively studied in West- ern Australia. Cate (1964; 1968) and WILson & GILLETT (1971) provide data on the distributions of almost all of the cowries occurring in the State. In addition, WiLson & SuMMe_Rs (1966) and Witson « McComs (1967) discuss the systematics and distributions of the members of the subgenus Zozla in detail. The genus Cypraea is characteristic of shallow tropical marine environments. The distribution of the 61 species recorded in Western Australia is shown on Figure 2, which clearly demonstrates the tropical nature of the group. Fifty of the species are tropical, 5 are temperate and 6 are en- demic. The tropical C'ypraea follow the generalized dis- Page 236 THE VELIGER Vol. 22; No. 3 eS TROPICAL - TEMPERATE 7 8 North-West Cape Carnarvon Shark Bay @ Geraldton Naturaliste Leeuwin Figure 3 Percentages of tropical (left) and temperate (right) marine proso- branch species along the west coast of Western Australia. These figures exclude endemic species tribution pattern discussed above. Nine species have been collected only along the north coast east of North-West Cape and an additional 10 have been reported at but not south of North-West Cape. The number of tropical Cyypraea declines sharply along the west coast, and only 12 pyisnuaa (2107) pavighy qassoL (DjL0Z) vav.dh pjoulsivu (D]107) vavighy upuauf (poz) vaviddy suatqisap (ojt07Z) vaviddy popiuausp (vYyiquiyy) vavigay pupvsyng (vavighr0j0N) vaviddg pyisagig (vav1g4I0j0N)) pavigay 103g wWo2 (vav1ghI0j0N) pavigay tangas (vanigas0usnp ) pavigay Kimberleys Derby Broome Pt. Hedland a) n SS ° 16) G za i] iC} a Dampier Onslow North-West Coast Carnarvon Shark Bay Abrolhos Perth West Coast Cape Naturaliste Cape Leeuwin Albany Esperance Eucla South Coast Figure 4 Distribution of cowries of the subgenera Austrocypraea, Notocyp- raea, Umbilia, and Zoila in Western Australia have been reported south of the Houtman Abrolhos. South of the Abrolhos there is a gradual decline in the number of tropical species. The extreme ranges are exhibited by C. helvola and C. caputserpentis, which range along the south coast to Albany. Four subgenera of Cypraea, with a total of 14 species, are restricted to Australia: Austrocypraea, Notocypraea, Umbilia, and Zoila. Nine species of these subgenera occur in Western Australia, and 5 are endemic to the State (Fig- ure 4). Eight of the 9 species are found predominately in the coldwater areas of the lower west coast and the south Vol. 22; No. 3 THE VELIGER Page 237 coast. Only one species, Cypraea (Zoila) decipiens, is found on the north coast, where it is distributed between North- West Cape and the Buccaneer Archipelago. Aside from the members of the Australian subgenera only a single species of Cypraea, C’. brevidentata, is en- demic to Western Australia. This species is restricted to the north coast between North-West Cape and Broome. Distribution of the Family Conidae Another tropical genus, Conus, has 49 representatives in Western Australia. Five of these are endemic and 5 are temperate species (Figure 2). The North-West Cape area is again the major distributional limit of the tropical spe- cies, with 15 species not being recorded farther to the south. Shark Bay and the Houtman Abrolhos are the other major limits for cones, and only one tropical species, C. liv- zdus, reaches as far south as Rottnest Island. Distribution of the Family Trochidae In contrast to the 2 families just discussed the trochids have a fairly even split of tropical (14 species) and tem- perate (20) forms. Three species: Calliostoma ciliaris, C. lepidus and Clanculus occidus are limited to Western Australia. The number of species of tropical trochids de- clines progressively southward along the coast (Figure 2) with no major distributional limits. The southern species show a similar pattern, with a gradual decrease in the number of species proceeding northward. Distribution of the Family Volutidae The highest rate of endemicity occurs in the neogastropod family Volutidae, where 7 of the 23 species, or 30%, are endemic. In addition,Amoria dampieria is distributed pri- marily in Western Australia but extends into the Northern Territory. Volutes reproduce by depositing benthic egg capsules from which the young emerge as crawling juve- niles (WiLson & GILLETT, 1971). The lack of a pelagic distributional stage contributes substantially to the high rate of endemicity in the group. The distributions of the endemic volutes are shown on Figure 5. The figure shows that not only are these species endemic to Western Australia, but two have restricted dis- tributions within the State. Amoria macandrewi is found only off Barrow Island and Monte Bello Island, a distance of only 50km. Amoria ellioti has a range of 200km from Dampier to Port Hedland. Both species are found in shal- DsonUu DIOP aDUuInd DUO idpid DoW HOUj2IDIAOULY, wsaaDalsADY Snuoz0jNjJOA SImMAOfIuo? snuoz0INjJOA IMaLPUDIDUL DILOULP Kimberleys Derby Broome Point Hedland Dampier ~ n 3 ° Oo S r=] ~ iS) Zz Onslow North-West C. Carnarvon Shark Bay Abrolhos Perth West Coast Cape Naturaliste Cape Leeuwin Albany Esperance South Coast Figure 5 Distribution of volute species endemic to Western Australia low waters and their known ranges are not likely to be extended substantially by continued collecting. In contrast to the species with restricted ranges are sev- eral endemics found over large portions of the Western Australian coastline. Amoria gray: extends along the entire north coast westward from the Kimberleys and southward on the west coast to Cape Naturaliste. Amoria irvinae has a southern distribution from Albany on the south coast to Jurien Bay on the west coast. Three other species (Amoria nivosa, Volutoconus coniformis and V. hargreavesi) have distributions over reasonably wide areas of the coastline. Thus there is substantial variation in the ranges occupied by the volutes endemic to Western Australia. It would be interesting to elucidate the mechanisms which account for these variations. Page 238 THE VELIGER Vol. 22; No. 3 DISCUSSION The detailed analysis of prosobranch distributions along the coast of Western Australia presented here elaborates the general outlines presented by HEDLEY (1926), WILSON & GILLETT (1971), and WILSON & STEVENSON (1977). There are two distinct faunas in the State: a northern tropical fauna which extends into eastern Australia and the Indo-Pacific region and a warm temperate fauna which is continuous with the remainder of the coastline of southern Australia. Most of the tropical species reach the North-West Cape area and almost all of the temper- ate forms are distributed along the south coast to Cape Leeuwin. The two faunas overlap on the west coast be- tween Cape Leeuwin and North-West Cape, with the major overlap area located between Perth and the Hout- man Abrolhos. The division of the Western Australian coastline into faunal regions has received attention from a number of workers investigating different phyla. The tropical coast of Australia was divided into a Solanderian fauna east of Cape York and a Damperian fauna extending from the west of Cape York to the Houtman Abrolhos and Gerald- ton (HEDLEY, 1926). This pattern was followed for echi- noderms by CiarK (1946) and Marsu (1976) but ENDEAN (1957) suggested merging the Damperian and Solanderian faunas into a single tropical Australian prov- ince. Marsu (op. cit.) rejected this proposal because 20% of the shallow-water asteroids of north Western Australia are endemic to the waters west of Torres Strait. Eighty- eight percent of the prosobranch gastropods investigated here which occur on the north coast of Western Australia also occur in eastern Queensland or the Indo-Pacific re- gion. This suggests that in the case of prosobranchs there is no need to subdivide the tropical waters of Australia into two faunal regions, and to do so would obscure the exten- sive similarities between the two areas. A similar conclu- sion was reached by WILSON & STEVENSON (1977) in their discussion of the distributions of cardiid bivalves. Hep ey (1926) described an Adelaidean region, which has since been renamed the Flindersian region; extending along the south coast of Australia westward from Bass Strait, and including all of the south coast of Western Australia and the west coast to Geraldton. The Flindersian region was subsequently restricted to the south coast east of Albany by Kort (1952) and a Baudinian region estab- lished between Perth and Albany. The proposal of a sep- arate Baudinian region was based on four species of ascid- ians endemic to the area. Figure 2 shows that while a number of endemic prosobranchs occur between Albany and Perth, most extend further to the north or are found entirely outside Kott’s Baudinian region. The analysis of prosobranch distributions indicates that there is no neces- sity for the separation of the lower west coast into a distinct faunal area. As has been indicated,the major overlap zone between the tropical and temperate prosobranchs occurs in the region of Perth to the Houtman Abrolhos and Geraldton. Since the marine fauna at Geraldton has a distinctly more temperate character than the offshore Abrolhos the place- ment of the northern limit of the Flindersian region at Geraldton by HEDLEY (1926) is substantiated. Similarly ENDEAN (1957) divided the fauna of the Queensland coast into two groups: an inshore component influenced by freshwater runoff from adjacent land masses and an offshore reef area. A similar inshore and offshore division could be made on the Western Australian coast, but there would be a substantial overlap between the two groups. The overall rate of endemism, 8.6%, is low. Among the species presently regarded as endemic are 9 described since 1965. The ranges of these species are poorly known and some of them may well be eventually found outside of Western Australia. Undoubtedly other new forms may be described as the molluscan fauna of Western Australia receives more attention. Some of the endemic species whose distributions are well established are clearly valid species. Included in this category are species such as Conus dorreensis which cannot be confused with any other species. Many of the shallow-water forms occurring along the Western Australian tropical coastline are closely related to, or conspecific with, populations in the Indo-Pacific re- gion or eastern Australia. The populations in Western Australia have diverged from the main body of the species because of reproductive isolation, but whether they have diverged sufficiently to achieve species status is a matter of taxonomic judgement. One example of this is Conus nodu- losus which is regarded as a species in this paper and by Witson & GILLETT (1971). However, Wilson and Gillett state that the status of the species is uncertain. It has a close affinity to C. victoriae and may prove on detailed investigation to be a subspecies of C’. victoriae. The differ- ent judgements on such questions that could be made by taxonomists affect the rate of endemism presented here. The rate of 8.6% endemics is not an unchallenged figure, but it does indicate the presence in Western Australia of a low proportion of prosobranch species unique to the area. Two factors appear to be important in limiting species in Western Australia: temperature and the availability of suitable habitats. The limiting role of temperature is well known (see Briccs, 1975). Temperate species en- Vol. 22; No. 3 THE VELIGER Page 239 counter progressively warmer waters further north on the west coast until the upper tolerance limit is met; tropical species encounter colder waters to the south. The tempera- ture tolerance range of each species is modified by a suite of physical and biological environmental factors, but at some point the limit is encountered and the species can go no further. The Perth area is the northern limit for one-third of the temperate species. Most of these prosobranchs live on rocky substrates that are largely lacking in the area be- tween Perth and Geraldton. It seems likely that some of these species would be able to colonize the shorelines fur- ther north if suitable substrates were available. Many of the tropical species live on coral reefs, and the absence of some of the reef habitats south of North-West Cape may limit tropical prosobranch species. This evades the question of why coral development is restricted south of North-West Cape, and the factor(s) limiting coral development may also be acting on the molluscs. The pattern of oceanic surface currents along the faunal overlap zone of the west coast obviously has an important bearing on the distributions of marine prosobranch species, most of which have planktonic larval stages. The current structure is poorly known. A Western Australian Current flows northward offshore of the Continental Shelf during the summer months of November to March (Wyrrtk1, 1973). A southerly flowing countercurrent has recently been found over the shelf (STEEDMAN et al., 1977). This brings warm surface waters southward along the west coast, with the 26°C isotherm reaching as far south as 31°S, just north of Perth. During the winter months the countercurrent dissipates and a series of semi-permanent vortices is established on the west coast. Surface tempera- tures over the Continental Shelf are 2° to 7° C higher than in the waters west of the Continental Shelf (STEEDMAN et al., 1977). The relatively warm surface waters moved southward by the currents allow tropical prosobranch spe- cies to extend further to the south than would otherwise be the case. ACKNOWLEDGMENTS This paper is based on Western Australian Museum collec- tions developed by two curators, Dr. B. R. Wilson and Mrs. S. M. Slack-Smith. Without their work and that of a number of specialists who have identified particular groups over the last 14 years this paper could not have been written. Drs. R. W. George and B. R. Wilson crit- ically read the manuscript and made a number of very helpful suggestions. Mrs. G. Hansen examined the species distributions. Mrs. M. Wallis typed this and a number of my other manuscripts. APPENDIX HALIOTDAE Haliotis asinina Linnaeus, 1758. North coast eastwards from North-West Cape. H. conicopora Péron, 1816. South coast and northwards along the west coast to Fremantle. H. cyclobates Péron, 1816. South coast westwards to Albany. H. elegans Philippi, 1899. South coast westwards from Albany and northwards along the west coast to the Houtman Abrolhos. H. ovina Gmelin, 1791. North coast eastwards from Barrow Island. H. roei Gray, 1826. South coast and northwards along the west coast to Shark Bay. H. scalaris (Leach, 1814). South coast and northwards along the west coast to Jurien Bay. e H. semiplicata Menke, 1843. South coast westwards from Esperance and northwards along the west coast to Perth. H. squamata Reeve, 1846. North coast and southwards along the west coast to Shark Bay. H. varia Linnaeus, 1758. North coast and southwards along the west coast to the Houtman Abrolhos. TROCHIDAE Angaria tyria (Reeve, 1843). North coast and southward along the west coast to Cockburn Sound. Austrocochlea constricta (Lamarck, 1822). South coast and north- ward along the west coast to the Houtman Abrolhos. A. rudis (Gray, 1826). South coast and northward along the west coast to the Murchison River. Calliostoma australe (Broderip, 1835). South coast and northward along the west coast to Fremantle. C. ciliaris (Menke, 1843). South coast westward from Esperance and northwards along the west coast to Perth. C. interruptum (Wood, 1828). South coast and northward along the west coast to Rottnest. C. lepidus (Philippi, 1846). West coast from Jurien Bay to the Houtman Abrolhos. C. monile (Reeve, 1843). North coast eastwards from Monte Bello Island. C. spinulosum Tate, 1893. North coast and southward along the west coast to Rottnest Island. Cantharidella beachportensis (Cotton and Godfrey, 1934). South coast and northward along the west coast to Cape Naturaliste. C. ocellina (Hedley, 1911). South coast and northward along the west coast to Rottnest Island. Clanculus atropurpureus (Gould, 1846). North coast and south- ward along west coast to Kalbarri. Page 240 THE VELIGER Vol. 22; No. 3 C. consorbrinus Tate, 1893. North coast, west coast, and eastward on south coast to Albany. C. denticulatus (Gray, 1827). South coast and northward along the west coast to the Houtman Abrolhos. C. dunkeri (Koch, 1843). South coast as far west as Albany. C. limbatus (Quoy and Gaimard, 1834). South coast and northward along the west coast to Bunbury. C. maxillatus (Menke, 1843). South coast and northward along the west coast to Kalbarri. C. occiduus Cotten « Godfrey, 1934. South coast from Hopetoun west and northward along the west coast to Rottnest Island. C. personatus (Philippi, 1846). South coast and northward along the west coast to Rottnest Island. C. plebejus (Philippi, 1851). South coast and northward along the west coast to Geraldton. C. ringens (Menke, 1843). South coast and northward along the west coast to Perth. Gibbula macculochit Hedley, 1907. North coast and southward along the west coast to Perth. G. preissiana (Philippi, 1848). South coast and northward along the west coast to Perth. Monilea callifera (Lamarck, 1827). North coast and southward along the west coast to Cockburn Sound. Monodonta labio (Linnaeus, 1758). North coast and southward along the west coast to Shark Bay. Odontotrochus cf. O. baudini (Fischer, 1878). North coast and southward along the west coast to Fremantle. O. chlorostoma (Menke, 1843). South coast and northward along the west coast to North-West Cape. Phasianotrochus eximus (Perry, 1811). South coast and northward along the west coast to Fremantle. Prothalotia lehmanni (Menke, 1843). South coast and northward along the west coast to Kalbarni. P. pulcherrima (Wood, 1828). South coast and northward along the west coast to the Houtman Abrolhos. P. ramburi (Crosse, 1864). South coast and northward along the west coast to Cape Naturaliste. Tectus pyramis Born, 1778. North coast and southward along the west coast to Rottnest Island. Thalotia conica (Gray, 1827). South coast and northward along the west coast to Rottnest Island. Trochus fenestratus Tate, 1893. North coast and southward along the west coast to Shark Bay. T. hanleyanus Reeve, 1842. North coast and southward along the west coast to the Houtman Abrolhos. T. lineatus Lamarck, 1822. North coast eastward from North-West Cape. T. maculatus Linnaeus, 1758. North coast and southward along the west coast to the Houtman Abrolhos. TURBINIDAE Astraea pileola Reeve, 1842. North coast and southward along the west coast to Carnarvon. A. rotularia (Lamarck, 1822). North coast eastward from Port Hedland. A. squamifera (Koch, 1844). South coast and northward along the west coast to Jurien Bay. A. stellare (Gmelin, 1791). North coast eastward from North-West Cape. A. tentorium (Thiele, 1931). Endemic from Cockburn Sound to the Houtman Abrolhos. Lunella cinereus Born, 1798. North coast eastward from North- West Cape. Marmarostoma pulcher (Reeve, 1842). South coast and northward along the west coast to Shark Bay. Ninella haynesi Preston, 1914. North coast and southward along the west coast to Rottnest Island. N. torquata Gmelin, 1791. South coast and northward along the west coast to Geraldton. Phasianella australis (Gmelin, 1791). South coast and northward along the west coast to Geraldton. P. ventricosa (Swainson, 1822). South coast and northward along the west coast to Geraldton. Subninella undulata (Gmelin, 1791). South coast and northward along the west coast to Geraldton. Turbo cf. T. argyrostomus Linnaeus, 1758. North coast and south- ward along the west coast to Carnarvon. T. chrysostoma Linnaeus, 1758. North coast and southward along the west coast to Carnarvon. T. foliaceus Philippi, 1846. North coast eastward from North- West Cape. T. jourdani Kiener, 1839. South coast and northward along the west coast to the Houtman Abrolhos. T. petholatus Linnaeus, 1758. North coast and southward along the west coast to Shark Bay. NERITIDAE Nerita albicilla Linnaeus, 1758. North coast and southward along the west coast to the Abrolhos. N. atramentosa Reeve, 1855. South coast and northwards along the west coast to North-West Cape. N. chamaeleon Linnaeus, 1758. North coast eastwards from North- West Cape. N. lineata Gmelin, 1791. North coast and southward along the west coast to Shark Bay. N. plicata Linnaeus, 1758. North coast and southwards along the west coast to the Houtman Abrolhos. N. reticulata Karsten, 1789. Kimberley region of the north coast. N. undata Linnaeus, 1758. North coast and southwards along the west coast to Shark Bay. LrrrorRINDAE Bembicium auratum (Quoy and Gaimard, 1834). South coast and northward along the west coast to the Houtman Abrolhos. B. melanostoma (Gmelin, 1791). South coast and northward along the west coast to Perth. Littorina scabra (Linnaeus, 1758). North coast and southward along the west coast to Shark Bay. L. undulata Gray, 1939. North coast eastward from North-West Cape. L. unifasciata Gray, 1826. South coast and northward along the west coast to North-West Cape. Vol. 22; No. 3 Nodilittorina australis (Gray, 1826). Endemic from Esperance to North-West Cape. N. nodosa (Gray, 1839). Endemic from Geraldton to the north coast. N. pyramidalis (Quoy and Gaimard, 1833). North coast and south- ward along the west coast to Fremantle. Tectartus rusticus (Philippi, 1846). Endemic to the Kimberley re- gion. STROMBIDAE Lambis chiragra (Linnaeus, 1758). North coast eastwards from North-West Cape. L. lambis (Linnaeus, 1758). North coast eastwards from North- West Cape. Strombus campbell: Griffith and Pidgeon, 1834. North coast and southwards along the west coast to Fremantle. S. dilatatus Swainson, 1821. North coast eastwards from North- West Cape. S. epidromus Linnaeus, 1758. North coast eastwards from North- West Cape. S. gibberulus Linnaeus, 1758. North coast eastwards from North- West Cape. S. lentiginosus Linnaeus, 1758. North coast eastwards from the Dampier Archipelago. S. marginatus Duclos, 1844. North coast eastwards from North- West Cape. S. mutabilis Swainson, 1821. North coast and southwards along the west coast to Cape Leeuwin. S. plicatus (R6ding, 1798). North coast eastwards from Onslow. S. urceus Linnaeus, 1758. North coast eastwards from North-West Cape. S. vomer (RGding, 1798). North coast and southwards along the west coast to Shark Bay. S. wilsoni Abbott, 1967. North coast east from North-West Cape. Rimella cancellata Lamarck, 1816. North coast eastwards from the North-West Cape. Terebellum terebellum (Linnaeus, 1758). North coast eastwards from North-West Cape. CyPprRAEDDAE Cypraea annulus Linnaeus, 1758. North coast southwards along the west coast to Rottnest Island. C. arabica Linnaeus, 1758. North coast southwards along the west coast to Shark Bay. C. argus Linnaeus, 1758. Along the north coast eastwards from North-West Cape. C. armeniaca Verco, 1912. South coast as far westwards as Eucla. C. asellus Linnaeus, 1758. Along the north coast eastwards from North-West Cape. C. brevidentata Sowerby, 1870. North coast endemic between North-West Cape and Broome. C. caputserpentis Linnaeus, 1758. North coast, southwards on the west coast, continuing on to Albany on the south coast. €. carneola Linnaeus, 1758. North coast and southwards along the west coast to the Houtman Abrolhos. C. caurica Linnaeus, 1758. North coast southwards along the west coast to Shark Bay. THE VELIGER Page 241 C. cernica Sowerby, 1870. North coast and southwards along the west coast to Bunbury. C. chinensis Lamarck, 1822. Along the north coast eastwards from North-West Cape, and southwards on the west coast to Cape Naturaliste. C. cicercula Linnaeus, 1758. North coast eastwards from North- West Cape. C. clandestina Linnaeus, 1767. North coast and southwards along the west coast to Cape Naturaliste. C. comptoni Gray, 1847. South coast to Cape Naturaliste. C. crtbraria Linnaeus, 1758. North coast southwards on the west coast to Cape Naturaliste. C. cylindrica Born, 1778. North coast southwards on the west coast to Shark Bay. C. decipiens Smith, 1880. Along the north coast from North-West Cape to the Buccaneer Archipelago. C. eglantina Duclos, 1833. North coast and southwards along the west coast to the Houtman Abrolhos. C. erosa Linnaeus, 1758. North coast and southwards along the west coast to the Houtman Abrolhos. C. errones Linnaeus, 1758. North coast and southwards along the west coast to Cape Naturaliste. C. fimbriata Gmelin, 1791. North coast southwards along the west coast to Cape Naturaliste. C. friendti Gray, 1831. South coast and northwards along the west coast to North-West Cape. C. gracilis Broderip and Sowerby, 1829. North coast southwards along the west coast to Cape Naturaliste. C. hammondae (Iredale, 1939). North coast southwards along the west coast to the Houtman Abrolhos. C. helvola Linnaeus, 1758. North coast southwards along the west coast and along the south coast to Albany. C. hirundo Linnaeus, 1758. North coast and southwards along the west coast to Shark Bay. C. histrio Gmelin, 1791. Along the north coast eastwards from North-West Cape. C. isabella Linnaeus, 1758. North coast and southwards along the west coast to the Houtman Abrolhos. C. kieneri Hidalgo, 1906. North coast eastwards from North-West Cape. C. labrolineata Gaskoin, 1848. North coast eastwards from North- west Cape. C. limacina Lamarck, 1810. North coast and southwards along the west coast to Cape Naturaliste. C. lutea Gmelin, 1791. North coast eastwards. from North-West Cape. C. lynx Linnaeus, 1758. North coast and southwards along the west coast to the Houtman Abrolhos. C. marginata Lamarck, 1811. South coast and northwards along the west coast to Geraldton. C. mauritania Linnaeus, 1758. North coast in the Kimberley area. C. miliaris Gmelin, 1791. North coast and southwards along the west coast to Shark Bay. C. moneta Linnaeus, 1758. North coast and southwards along the west coast to the Houtman Abrolhos. C. nucleus Lamarck, 1811. Along the north coast eastwards from North-West Cape. C. ovum Gmelin, 1791. North coast and southwards along the west coast to the Houtman Abrolhos. Page 242 THE VELIGER Vol. 22; No. 3 C. pallidula Gaskoin, 1849. North coast and southwards along the west coast to Cape Naturaliste. C. piperita Gray, 1825. South coast and northwards along the west coast to Cape Naturaliste. C. poraria Linnaeus, 1758. Along the north coast and southward on the west coast to Cape Naturaliste. C. pyriformis Gray, 1824. Along the Kimberley region of the north coast. C. pulicaria Reeve, 1846. West coast from Cape Leeuwin to Rott- nest Island. C. punctata Linnaeus, 1771. North coast eastwards from North- West Cape. C. quadrimaculata Gray, 1824. Along the north coast in the Kim- berley region. C. reevei Sowerby, 1832. South coast northwards along the west coast to the Houtman Abrolhos. C. rosselli (Cotton, 1948). West coast from Cape Leeuwin to Shark Bay. C. saulae Gaskoin, 1843. Along the north coast eastwards from the Dampier Archipelago. C. staphylaea Linnaeus, 1758. North coast and southwards along the west coast to Rottnest Island. C. stolida Linnaeus, 1758. North coast and southwards along the west coast to the Houtman Abrolhos. C. subviridis Reeve, 1835. North coast and southwards along the west coast to Rottnest Island. C. talpa Linnaeus, 1758. Along the north coast eastwards from North-West Cape. C. teres Gmelin, 1791. North coast and southwards along the west coast to Cape Naturaliste. C. testudinaria Linnaeus, 1758. Along the Kimberley region of the north coast. C. tigris Linnaeus, 1758. North coast and southwards along the west coast to the Houtman Abrolhos. C. ursellus Gmelin, 1791. Along the north coast eastwards from Broome. C. venusta Sowerby, 1847. South coast northwards along the west coast to Shark Bay. C. vitellus Linnaeus, 1758. North coast and southwards along the west coast to Cape Naturaliste. C. walkeri Sowerby, 1832. Along the north coast eastwards from Broome. C. ziczac Linnaeus, 1758. North coast and southwards along the west coast to Shark Bay. NATICIDAE Ectosinum zonale (Quoy and Gaimard, 1833). South coast and northward along the west coast to Rottnest Island. Eunaticina dingeldi (Iredale, 1931). South coast, west along the coast, and eastward along the north coast to Broome. Mamamilla opaca (Récluz, 1851). North coast and southward along the west coast to Shark Bay. M. simiae (Deshayes, 1838). North coast eastward from the Dam- pier Archipelago. Mazmillaria powisiana (Récluz, 1844). North coast and southward along the west coast to Geraldton. Natica fasciata (Réding, 1798). North coast eastward from the Dampier Archipelago. N. gualtieriana Récluz, 1844. North coast, west coast, and eastward along the south coast to Albany. N. sagitatta Menke, 1843. North coast, west coast, and eastward along the south coast to Esperance. N. seychellium Watson, 1886. North coast and southward along the west coast to Cape Naturaliste. Sigaretotrema umbilicatum (Quay and Gaimard, 1833). South coast and northward along the west coast to Perth. CAssDDAE Casmaria ertnacea (Linnaeus, 1758). North coast eastward from North-West Cape. C. ponderosa (Gmelin, 1791). North coast eastward from Barrow Island. Cassts cornuta (Linnaeus, 1758). North coast eastward from Ons- low. C. fimbriata Quoy and Gaimard, 1833. South coast and northward along the west coast to the Houtman Abrolhos. Phalium adcocki (Sowerby, 1896). South coast as far westward as Eucla. P. areola (Linnaeus, 1758). North coast and southward along the west coast to Carnarvon. P. bandatum (Perry, 1811). North coast and southward along the west coast to the Houtman Abrolhos. P. bisulcatum (Schubert and Wagner, 1829). North coast and south- ward along the west coast to Carnarvon. P. glabratum (Iredale, 1927). Kimberley region of the north coast. P. pauctruge (Menke, 1843). South coast as far northward as Shark Bay. P. pyrum (Lamarck, 1822). South coast as far westward as Bremer Bay. P. semigranosum (Lamarck, 1822). South coast and northward along the west coast to Fremantle. P. sinuosum (Verco, 1904). South coast and northward along the west coast to Bunbury. P. whitwortht Abbott, 1968. West coast from Rottnest to Gerald- ton. TOoNNDAE Tonna allium (Dillwyn, 1817). North coast eastward from North- West Cape. T. canaliculata (Linnaeus, 1758). North coast and southward along the west coast to Shark Bay. T. chinensts (Dillwyn, 1817). North coast and southward along the west coast to Cape Leeuwin. T. perdix (Linnaeus, 1758). North coast and southward along the west coast to the Houtman Abrolhos. T. tessellata (Linnaeus, 1758). North coast eastward from the Dam- pier Archipelago. T. variegata (Lamarck, 1822). South coast and northward along the west coast to North-West Cape. Vol. 22; No. 3 THE VELIGER Page 243 Muricwar Bedeva hanleyi (Angas, 1867). South coast and northward along the west coast to Shark Bay. B. paivae (Crosse, 1864). South coast and northward along the west coast to Pt. Quobba. Chicoreus banksii (Sowerby, 1840). North coast eastward from the Dampier Archipelago. C. cervicornis (Lamarck, 1822). North coast and southward along the west coast to Rottnest Island. C. cornucervi (Roding, 1798). North coast and southward along the west coast to the Houtman Abrolhos. C. lacintatus (Sowerby, 1841). North coast eastward from Onslow. C. ramosus (Linnaeus, 1758). Kimberley region of the north coast. C. rubiginosus (Reeve, 1845). North coast and southward along the west coast to the Houtman Abrolhos. C. torrefactus (Sowerby, 1841). North coast eastward from North- West Cape. Dermomurex antonius Vokes, 1974. Dampier Archipelago area of the north coast. Haustellum macgillivrayi (Dohrn, 1862). North coast eastward from North-West Cape and southwards on the west coast to Jurien Bay. H. multiplicatum (Sowerby, 1895). North coast and southward along the west coast to Rottnest Island. H. wilsoni Old and d’Attilio, 1971. West coast in the Jurien Bay area. Hexaplex stainforthi (Reeve, 1842). North coast and southward along the west coast to the Houtman Abrolhos. Homalocantha secunda (Lamarck, 1822). North coast eastward from North-West Cape. Murex acanthostephes Watson, 1883. North coast eastward from Broome. M. brunneus (Link, 1807). Kimberley region of the north coast. M. coppingert E. A. Smith, 1884. North coast eastward from North-West Cape. M. triremus (Perry, 1811). North coast and southward along the west coast to Shark Bay. Muricopsis planilirata (Reeve, 1845). South coast and northward along the west coast to Perth. Pterotyphis angasi (Crosse, 1863). South coast and northward along the west coast to Fremantle. Pterynotus acanthopterus (Lamarck, 1816). North coast and southward along the west coast to Cape Leeuwin. P. bednalli (Brazier, 1877). Kimberley region of the north coast. P. lowei (Pilsbry, 1931). North coast eastward from the Dampier Archipelago. P. westralis Ponder and Wilson, 1973. West coast from Bunbury to North-West Cape. Typhis yatesi Crosse and Fischer, 1865. South coast and northward along the west coast to Rottnest Island. THADDAE Cronia avellana (Reeve, 1846). North coast, west coast, and east- wards along the south coast to Albany. Dicathais aegrota (Reeve, 1846). South coast and northward along the west coast to the Houtman Abrolhos. Drupa morum Réding, 1798. North coast eastwards from North- West Cape. D. ricinus (Linnaeus, 1758). North coast and southward along the west coast to the Houtman Abrolhos. Drupella cornus (Réding, 1798). North coast and southwards along the west coast to the Houtman Abrolhos. Drupina grossularia Réding, 1798. North coast eastwards from Barrow Island. D. lobata (Blainville, 1832). North coast eastwards from North- West Cape. Lepsiella flindersi Adams and Angas, 1863. South coast and north- wards along the west coast to Cockburn Sound. L. vinosa (Lamarck, 1822). South coast and northwards along the west coast to Cockburn Sound. Manicella manicella (Linnaeus, 1758). North coast and southwards along the west coast to Shark Bay. M. tuberosa (Réding, 1798). Kimberley region of the north coast. Morula granulata (Duclos, 1832). North coast and southward along the west coast to Pt. Quobba. M. margariticola (Broderip, 1832). North coast eastwards from North-West Cape. M. spinosa (H. and A. Adams, 1853). North coast and southwards along the west coast to the Houtman Abrolhos. M. uva (Roding, 1798). North coast and southward along the west coast to the Houtman Abrolhos. Nassa francolina (Bruguiére, 1789). North coast and southwards along the west coast to Rottnest Island. Thais aculeata Deshayes, 1844. North coast eastwards from North- West Cape. T. alvina (R6ding, 1798). North coast eastward from the Dampier Archipelago. T. bufo (Lamarck, 1822). North coast eastwards from North-West Cape. T. echinata (Blainville, 1832). North coast and southward along the west coast to Shark Bay. T. kieneri (Deshayes, 1844). North coast eastwards from North- West Cape. Vexilla vexillum (Gmelin, 1791). North coast eastwards from North-West Cape. CoLUMBELLIDAE Dentimitrella austrina (Gaskoin, 1852). South coast and north- ward along the west coast to Kalbarri. D. lincolnensis (Reeve, 1859). South coast and northward along the west coast to Rottnest Island. D. pulla (Gaskoin, 1852). South coast as far westward as Pt. d’Entre- casteaux. D. semiconvexa (Lamarck, 1822). South coast and northward along the west coast to Cape Naturaliste. Lavesopus essingtonensis (Reeve, 1859). North coast eastward from Pt. Hedland. Macrozafra angasi Brazier, 1871. South coast as far westward as Hopetoun. M. speciosa (Angas, 1877). South coast and northward on the west coast to Fremantle. Page 244 Mitrella albina (Kiener, 1841). North coast and west coast as far southward as Perth. M. ligula (Duclos, 1840). North coast and west coast as far south- ward as Shark Bay. M. marquesa (Gaskoin, 1852). North coast eastward from North- West Cape. M. miser (Sowerby, 1844). North coast and southward along the west coast to Pt. Quobba. M. puella (Sowerby, 1844). North coast eastward from North-West Cape. Pseudamyoia dermestoidea (Lamarck, 1822). South coast and northward along the west coast to Cape Naturaliste. Pyrene bidentata (Dall, 1889). South coast and northward along the west coast to North-West Cape. P. flava (Bruguiére, 1789). North coast eastwards from North-West Cape. P. punctata (Bruguiére, 1789). North coast and southward along the west coast to the Houtman Abrolhos. P. testudinaria (Link, 1807). North coast eastward from the North- West Cape. P. tuturina (Lamarck, 1822). North coast and southward along the west coast to the Houtman Abrolhos. P. varians (Sowerby, 1832). North coast and southward along the west coast to Rottnest Island. Rettzafra calva (Verco, 1910). South coast and northward along the west coast to Rottnest Island. Zafra mitriformis A. Adams, 1860. Swan estuary only. Z. troglodytes (Souverbie, 1866). Dampier Archipelago area of the north coast. FASCIOLARIDAE Fusinus australis (Quoy and Gaimard, 1833). South coast and northwards along the west coast to the Houtman Abrolhos. F. tessellatus (Sowerby, 1880). South coast and northwards along the west coast to Geraldton. Fusus colus (Linnaeus, 1758). North coast and southwards along the west coast to Shark Bay. Lattrus belcheri (Reeve, 1847). North coast eastwards from North- West Cape. L. paetelianus Kobelt, 1876. North coast eastwards from Onslow. L. recurvirostris Schubert and Wagner, 1829. North coast eastwards from North-West Cape. L. walkeri Melvill, 1895. North coast eastwards from North-West Latirolagena smaragdula (Linnaeus, 1758). North coast eastwards from Rowley Shoals. Microcolus dunkevi (Jonas, 1844). South coast and northwards along the west coast to the Houtman Abrolhos. Peristernia incarnata (Deshayes, 1830). North coast and southwards along the west coast to Geraldton. Pleuroploca australasta (Perry, 1811). South coast. P. filamentosa (Réding, 1798). Kimberley region of the north coast. P. trapezium (Linnaeus, 1758). North coast eastwards from the Montebello Islands. THE VELIGER Vol. 22; No. 3 NASSARIDAE Nassarius albescens (Dunker, 1846). North coast eastwards from North-West Cape. N. albina (Thiele, 1930). North coast and southward along the west coast to the Houtman Abrolhos. N. bicallosus (E. A. Smith, 1876). North coast and southwards along the west coast to Shark Bay. . burchardt (Dunker in Philippi, 1849). South coast and north- wards along the west coast to Perth. N. concinnus (Powys, 1835). North coast eastwards from North- West Cape. N. coronatus (Bruguiére, 1789). North coast eastwards from North- West Cape. N. crematus (Hinds, 1844). Kimberley region of the north coast. N. dorsatus (Réding, 1798). North coast eastwards from North- West Cape. N. gaudiosus (Hinds, 1844). North coast and southwards along the west coast to Perth. N. glans (Linnaeus, 1758). North coast and southwards along the west coast to Geraldton. N. livescens (Philippi, 1849). North coast and southwards along the west coast to Shark Bay. N. nigellus (Reeve, 1864). South coast and northwards along the west coast to Cockburn Sound. N. particeps (Hedley, 1915). South coast, west coast, and north coast east to Dampier. N. pauperatus (Lamarck, 1822). South coast and northwards along the west coast to Geraldton. N. pyrrhus (Menke, 1843). South coast and northwards along the west coast to Fremantle. N. rufula (Kiener, 1834). South coast westwards from Albany and northwards along the west coast to Geraldton. N. stigmarius (A. Adams, 1852). North coast eastwards from North- z West Cape. N. variegatus (A. Adams, 1852). North coast eastwards from Ons- low. OLIVIDAE Alocospira monilifera (Reeve, 1864). South coast and northward along the west coast to Cape Leeuwin. Amalda elongata (Gray, 1847). North coast and southward along the west coast to Shark Bay. Ancillista cingulata (Sowerby, 1830). North coast and southward along the west coast to Shark Bay. Oliva australis Duclos, 1835. South coast, west coast, and eastward along the north coast to Broome. O. caerulea (Réding, 1798). North coast and southward along the west coast to the Houtman Abrolhos. O. caldania Duclos, 183%. North coast eastward from North-West Cape. O. lignara Marratt, 1868. North coast and southward along the west coast to Cockburn Sound. Vol. 22; No. 3 O. oliva (Linnaeus, 1758). North coast eastward from North-West Cape. O. sidelia Duclos, 1835. North coast eastward from Onslow. MITRIDAE Canctlla circula (Kiener, 1838). North coast eastward from Onslow. Imbricaria cf. I. conovula Quoy & Gaimard, 1833. North coast east- ward from Dampier. Mitra ambigua Swainson, 1829. North coast eastward from North- West Cape. M. atjehensis Oostingh, 1939. North coast and southward along the west coast to Dongara. M. aurantia (Gmelin, 1791). North coast eastward from Barrow Island. carbonaria Swainson, 1822. South coast and northward along the west coast to the Houtman Abrolhos. chalybeia Reeve, 1844. West coast endemic from Pt. d’Entre- casteaux to Port Gregory. coffea Schubert and Wagner, 1829. North coast eastward from the Dampier Archipelago. cucumerina Lamarck, 1811. North coast eastward from North- West Cape. ferruginea Lamarck, 1811. North coast eastward from North- West Cape. fraga Quoy and Gaimard, 1833. North coast and southward along the west coast to Geraldton. gilbertsoni (J. Cate, 1968). Endemic from the Houtman Abrol- hos to Port Hedland. glabra Swainson, 1821. South coast and northward along the west coast to Perth. litterata Lamarck, 1811. North coast eastward from North- west Cape. M. luctuosa A. Adams, 1853. North coast and southward along the west coast to Perth. M. puncticulata Lamarck, 1811. North coast eastward from North- west Cape. M. rosacea Reeve, 1845. North coast and southward along the west coast to Rottnest Island. M. rubiginosa Reeve, 1844. North coast eastward from North-West Cape. M. scutulata (Gmelin, 1791). North coast and southward along the west coast to Cape Naturaliste. M. sowerbyi kingae Cernohorsky, 1972. North coast and southward along the west coast to the Houtman Abrolhos. M. stictica (Link, 1807). Kimberley region of the north coast. M. ticaontca Reeve, 1844. North coast and southward along the west coast to Geraldton. M. variabilis Reeve, 1844. Eastward along the north coast from North-West Cape. Scabricola backae Cernohorsky, 1973. Entire west coast, eastward along the north coast to Onslow. S. desetangsii (Kiener, 1838). North coast eastward from the Dam- pier Archipelago. S. flammigera (Reeve, 1844). North coast eastward from Onslow. S. incarnata (Reeve, 1845). North coast eastward from Onslow. S. interlirata (Reeve, 1844). North coast eastward from North-West Cape. = 5 § § &§ &§ &§ § & THE VELIGER Page 245 S. lacunosa (Reeve, 1844). North coast eastward from the North- West Cape. S. ocellata ekerae Cernohorsky, 1973. North coast eastward from North-West Cape. S. praestantissima Réding, 1798. North coast eastward from North- West Cape. Vexillum amanda (Reeve, 1845). North coast and southward along the west coast to Rottnest Island. V. angustissimum (E. A. Smith, 1903). North coast eastward from the Dampier Archipelago. V. australe Swainson, 1820. South coast and northward along the west coast to Shark Bay. V. cadaverosum (Reeve, 1844). North coast and southward along the west coast to Shark Bay. V. crocatum (Lamarck, 1811). North coast eastward from the Dam- pier Archipelago. V. hansenae Cernohorsky, 1973. Endemic from Cape Leeuwin to Fremantle. V. lincolnensis Angas, 1878. South coast as far west as Hopetoun. V. marrowt Cernohorsky, 1973. Endemic from Hopetoun to Kal- barri. V. microzonias (Lamarck, 1811). North coast eastward from Bar- row Island. V. modestum (Reeve, 1845). North coast eastward from Onslow. V. obeliscus (Reeve, 1844). North coast and southward along the west coast to Rottnest Island. V. pacificum (Reeve, 1845). North coast and southward along the west coast to Shark Bay. V. percnodictya (Melvill, 1888). North coast eastward from North- West Cape. V. plicarium (Linnaeus, 1758). North coast eastward from North- West Cape. V. radius (Reeve, 1845). North coast eastward from North-West Cape. V. rugosum (Gmelin, 1791). North coast eastward from North-West Cape. V. suluense (Adams and Reeve, 1850). North coast eastward from North-West Cape. V. unifasciatum (Wood, 1828). North coast and southward along the west coast to Shark Bay. V. vulpecula (Linnaeus, 1758). North coast eastward from North- West Cape. V. zebuense (Reeve, 1844). North coast and southward along the west coast to the Houtman Abrolhos. VOLUTIDAE Amoria damoni Gray, 1864. North coast and southward along the west coast to Rottnest Island. A. dampiera Weaver, 1960. North coast eastwards from the Monte Bello Islands. A. elliotti (Sowerby, 1864). North coast eastwards from Port Hed- land. A. exoptanda (Reeve, 1849). South coast. A. grayi Ludbrook, 1953. North coast from the Kimberleys west- ward and southward along the west coast to Geographe Bay. Page 246 THE VELIGER Vol. 22; No. 3 a A. irvinae Smith, 1909. South coast westward from Albany and northwards along the west coast to the Houtman Abrolhos. A, jamrachi Gray, 1864. North coast eastward from North-West Cape to the Kimberleys, and also occurs in Indonesia. A. macandrewi (Sowerby, 1887). Region of Barrow Island. A. nivosa (Lamarck, 1804). North coast and southward along the west Coast to Cockburn Sound. A. praetexta (Reeve, 1849). North coast between North-West Cape and Broome. A. turnert (Gray in Griffith and Pidgeon, 1834). Kimberley region of the north coast. Cottonia nodiplicata (Cox, 1916). South coast and northward along the west coast to Jurien Bay. Ericusa fulgetrum (Sowerby, 1825). South coast to Albany. E. papillosa (Swainson, 1822). South coast and northward along the west coast to Bunbury. Livonia roadnightae (McCoy, 1881). South coast and northward along the west coast to Rottnest Island. Lyria mitraeformis (Lamarck, 1811). South coast westward to Hopetoun. Melo amphora (Solander, 1786). North coast and southward along the west coast to Shark Bay. M. miltonis (Gray in Griffith and Pidgeon, 1834). South coast and northward along the west coast to the Houtman Abrolhos. Notopeplum kreuslerae (Angas, 1865). South coast. Volutoconus bednalli (Brazier, 1879). Kimberley region of the north coast. V. coniformis (Cox, 1871). North coast between the Dampier Arch- ipelago and Broome. V. hargreavesi (Angas, 1872). Between Shark Bay and the Dampier Archipelago. ConmAE Conus achatinus Gmelin, 1791. North coast and southward along the west coast to Jurien Bay. C. anemone Lamarck, 1810. South coast and northward along the west coast to the Abrolhos Islands. C. arenatus Hwass, 1792. North coast eastward from North-West Cape. C. capitaneus Linnaeus, 1758. North coast eastward from North- West Cape. C. catus Hwass, 1792. Kimberley region of the north coast. C. ceylanensis Bruguiére, 1792. North coast and southward along the west coast to Shark Bay. C. chaldeus (Réding, 1798). North coast and southward along the west coast to the Houtman Abrolhos. C. cholmondelyi Melvill, 1900. Kimberley region of the north coast. C. clarus Smith, 1881. North coast eastward from North-West Cape. C. cocceus Reeve, 1844. South coast and northward on the west coast to Rottnest Island. C. coronatus Gmelin, 1791. Eastward along the north coast from North-West Cape. C. dorreensis Péron, 1807. South coast westward from Albany and northward on the west coast to the North-West Cape. C. ebraeus Linnaeus, 1758. North coast and southward along the west coast to the Houtman Abrolhos. C. eburneus Hwass, 1792. North coast eastward from Rowley Shoals, C. frigidus Reeve, 1848. North coast eastwards from Barrow Island. C. generalis Linnaeus, 1767. North coast eastward from North- West Cape. C. geographus Linnaeus, 1758. North coast and southward along the west coast to the Houtman Abrolhos. C. giluus Reeve, 1849. North coast eastward from the Dampier Archipelago. C. glans Hwass, 1792. North coast eastward from N.W. Cape. C. infrenatus Reeve, 1848. West coast from Cape Naturaliste to the Houtman Abrolhos. C. kenyonae Brazier, 1896. North coast and southward along the west coast to the Houtman Abrolhos. C. klemae (Cotton, 1953). South coast and northwards along the west coast to the Houtman Abrolhos. C. lemniscatus Reeve, 1849. North coast eastward from Adele Is- land. C. lividus Hwass, 1792. North coast and southward along the west coast to Rottnest Island. C. macarae Bernardi, 1857. North coast and southward along the west coast to Shark Bay. C. marmoreus Linnaeus, 1758. North coast eastward from Rowley Shoals. C. miliaris Bruguiére, 1792. North coast eastward from North- West Cape. C. monachus Linnaeus, 1758. North coast and southward along the west coast to Jurien Bay. C. mustelinus Bruguiére, 1792. Kimberley region of the north coast. C. nanus Sowerby, 1833. North coast and southward along the west coast to Shark Bay. C. nodulosus Sowerby, 1864. West coast from Fremantle to Shark Bay. C. nussatella Linnaeus, 1758. Kimberley region of the north coast. C. obscurus Sowerby, 1833. North coast eastwards from N.W. Cape. C. planorbis Born, 1778. North coast eastwards from North-West Cape. C. pulicarius Hwass, 1792. North coast eastward from North-West Cape. C. rattus Hwass, 1792. North coast eastward from North-West Cape. C. rutilus Menke, 1843. South coast and along the west coast to Rottnest Island. C. segravei Gatliff, 1891. South coast westward to Cape Leeuwin. C. spectrum Linnaeus, 1758. North coast and southward along the west coast to Shark Bay. C. sponsalis Hwass, 1792. North coast eastward from Onslow. C. striatus Linnaeus, 1758. North coast eastward from North-West Cape. C. suturalis Reeve, 1844. North coast and southwards along the west coast to Shark Bay. C. terebra Born, 1780. North coast and southward along the west coast to Carnarvon. C. tessellata Born, 1778. Kimberley region of the north coast. Vol. 22; No. 3 THE VELIGER Page 247 C, textile Linnaeus, 1758. North coast eastward from North-West Cape. C. trigonis Reeve, 1848. North coast eastward from North-West Cape. C. vexillum Gmelin, 1791. North coast eastward from North-West Cape. C. victoriae Reeve, 1843. North coast eastward from North-West Cape. TEREBRIDAE Duplicaria addita Deshayes, 1859. North coast and southward along the west coast to Cape Naturaliste. D. bernardi (Deshayes, 1857). North eastward from Broome. D. crakei Burch, 1965. Broome area on north coast. D. duplicata (Linnaeus, 1758). North coast and southwards along the west coast to Shark Bay. D. evoluta (Deshayes, 1859). North coast and southward along the west coast to Shark Bay. Hastula nitida (Hinds, 1844). North coast and southwards along the west coast to Rottnest Island. H. rufopunctata (E. A. Smith, 1877). North coast and southward along the west coast to Shark Bay. H. strigilata (Linnaeus, 1758). North coast eastward from Pt. Samson. Terebra affinis Gray, 1834. North coast and southward along the west coast to the Houtman Abrolhos. T. areolata (Link, 1807). North coast eastward from North-West Cape. T. commaculata (Gmelin, 1791). North coast and southward along the west coast to Shark Bay. T. crenulata (Linnaeus, 1758). North coast eastward from North- West Cape. T. dimidiata (Linnaeus, 1758). North coast eastward from North- West Cape. T. felina (Dillwyn, 1817). North coast eastwards from North-West Cape. T. nebulosa Sowerby, 1825. North coast and southward along the west coast to Shark Bay. T. triseriata Gray, 1834. North coast eastwards from Onslow. Literature Cited Brices, Joun CaRMoNn 1975. Marine Zoogeography. 65 figs. Cats, CrRawForpD NEILL 1964. Western Australian cowries (Mollusca : Gastropoda). The Veliger 7 (1): 7-29; pit. 5 ; 1 map (1 July 1964) 1968. Western Australian cowries, a second, revised, and expanded report. The Veliger 10 (3): 212-232; plts. 21-34; 5 maps (1 January 1968) Publ. Carnegie Inst. McGraw-Hill, New York, 475 pp.; Crarx, H. L. 1946. The echinoderm fauna of Australia. 566: 1 - 567 ENDEAN, RoBEeRT 1957- The biogeography of Queensland’s shallow water echinoderm fauna (excluding Crinoidea) with a rearrangement of the faunistic provinces of tropical Australia. Austral. Journ. mar. freshwat. Res. 8: 233 - 273 Hepiey, CHARLES 1916. A preliminary index of the Mollusca of Western Australia. Journ. Roy. Soc. West. Austral. 1: 1-77 1926. Zoogeography. Austral. Encyclop. 2: 743 - 744 Hopcxin, Ernest P « Bruce F Puiuips 1969. Sea temperatures on the coast of southwestern Australia. Journ. Proc. R. Soc. West Austral. 53: 59 - 62 Kort, Patricia 1952. Ascidians of Australia. $: 205 - 333 MarsH, LotseTTe M. 1976. Western Australian Asteroidea since H. L. Clark. Jugoslav. 12: 213 - 225 STEEDMAN, R. K. e¢ al. 1977. Preliminary study of oceanographic and meteorologic condi- tions as affecting offshore drilling on WA-59-P, Abrolhos Island area, Western Australia. MS Rprt. submitted to ESSO Australia. 128 pp. Witson, Barry R. «& Keir GILLETT 1971. Australian shells. Sydney, A. H «& A. W. Reed; 168 pp., ca. half of them color plates Witson, Barry R. & JENNIFER A. McComs 1967. The genus Cypraea (subgenus Zoila Jousseaume). Indo- Pacif. Moll. 1: 457 - 484 Witson, Barry R. « SUZANNE STEVENSON 1977. Cardiidae (Mollusca, Bivalvia) of Western Australia. West. Austral. Mus. Spec. Publ. 9: 1-114 Witson, Barry R. & Ray SUMMERS 1966. Variation in the Zoila friendi (Gray) species complex (Gastro- poda, Cypraeidae) in south-western Australia. Journ. Malacol. Soc. Austral. 1 (9): 3-24 Wyrtx!, Kraus 1973. Physical oceanography of the Indian Ocean. In: Zzitz- ‘SCHEL & SAGERLAGH (eds.) The biology of the Indian Ocean. Berlin, Springer Verlag: 18 - 36 Austral. Journ. Mar. freshwat. Res. Thalassia Page 248 THE VELIGER Vol. 22; No. 3 Predator Boreholes in Periploma margaritaceum, With a Brief Survey of Other Periplomatidae (Bivalvia : Anomalodesmata ) BY JOSEPH ROSEWATER Department of Invertebrate Zoology (Mollusks) National Museum of Natural History, Smithsonian Institution, Washington, D. C. 20560 (1 Plate; 1 Text figure) Stupy oF THE Nationa, Museum or Natura History collection of Periplomatidae revealed a population sample of Periploma margaritaceum (Lamarck, 1801) in which approximately 42% of the 443 valves were bored by a gastropod predator. The distribution of the boreholes ap- pears to be fairly consistent within P. margaritaceum. A survey of boreholes in seven different species in the family reveals a fairly similar distribution in all specimens ex- amined. Predation, while long recognized as an important factor in guiding evolution, has recently received some intensive study as it affects marine animals and especially mollusks (VERMEIJ, 1978). Of the several methods of predation reviewed by Vermeij, including crushing and prying-apart of valves, swallowing whole and partial mutilation by fishes, birds, and other invertebrates, one of the most im- portant causes of mortality is through drilling. Bivalves offer few defenses to predation other than thickened shells and some sculptural adaptations. In groups where these structural safeguards are absent, Vermeij suggests that some bivalves survive as species only by high reproduc- tive rates. The Periplomatidae may be classed with the latter group which also includes some Tellinacea, Mactracea, other Pandoracea and Poromyacea, among others. Little is known of the natural history of Periplomatidae other than notes supplied by Morse (1919) for Anatina papyra- cea Say [sic, probably = Periploma fragile (Totten, 1835)], and the detailed study by ALLEN (1958) on Cochlo- desma praetenue (Pulteney, 1799). Allen’s work gives the only insight we have into the habit of the family. He found C. praetenue to live infaunally more often on its left side in fine gravelly to muddy sand below low water mark. Allen placed Cochlodesma in Laternulidae, al- though it usually is considered to belong in Periplomatidae. The two families are fairly similar in many features, especially in the possession of thin shells which are split near the umbos by “transverse external slits’ (MorTON, 1976). Morton suggested that some Laternulidae may pro- tect themselves from predation with the aid of pallial eyes enabling the bivalves to initiate camouflage activities to evade predators. It is improbable that shell boring gastro- pods would find such defences a deterrent to predation. Nevertheless, I was unable to find a single Laternulid shell in the collection of the U.S. National Museum of Natural History that showed evidence of having been bored. The resulting supposition could be that their defenses are most effective, but the possibility is that their partially gaping valves allow easy access to predators without the need for boring. STANLEY (1970: 77-80) pointed out that a number of Pandoracea, the superfamily containing Periplomatidae, and members of the Tellinacea having inequivalve shells often lie on their sides within the substrate, or at least often exhibit non-vertical orientations. As Periplomatidae are uniformly inequivalve it should probably be expected that Periploma margaritaceum would orient itself on one side with either the right or left valve uppermost (see ALLEN, 1958). Periplomatidae probably can close their valves quite tightly utilizing the peculiar structural slit arrangement described by Morton (1976) which allows the shells of Laternulids to function as if they consisted of four valves instead of two. It is therefore apparently nec- essary for a predator either to pull the valves apart, crush them or penetrate them by boring to obtain a meal. The subjects of the present study succumbed to boring. Vol. 22; No. 3 The lot of P. margaritaceum alluded to above (USNM 465183/465234) contains 443 separated, recently dead valves from Matagorda Bay, Indianola, Calhoun County, Texas, collected during the early years of this century. The numbers of specimens bored and unbored are given in Table 1 and the locations of the boreholes as found in right or left valves are indicated in Table 1 and Figure 1. The boreholes appear to be the result of predation by a gastropod, probably Polinices duplicatus (Say, 1822), the main naticid predator in the vicinity of Matagorda Bay (Dr. H. Harry, personal communication). They resemble those illustrated by Carriker and Yochelson (1968, pl. 2, figs. 6-9; pl. 3, figs. 2-8) which they termed “parabolic countersunk holes” (see Figure 2, herein). B Periploma margaritaceum from Matagorda Bay, Texas Composites of 185 valves showing locations of boreholes. A ~ right valve; B — left valve; dashed lines indicate areas of shells (as - anterior slope; d - disc; ps — posterior slope) Note: due to crowding in some areas not all boreholes are indicated (see Table 1) THE VELIGER- Page 249 Table 1 Predation in Periploma margaritaceum from Matagorda Bay, Texas (see Figure 1) Position of Bore Hole Number Number unbored bored Disc Anterior slope Right valve 137 83 38 45 Left valve 121 102 26 76 Some attempts have been made toward numerical lo- calization of boreholes in gastropods (BERG & NisHENKOo, 1975). In both Gastropoda and Bivalvia diagrammatic localization of holes may also be effective (ANSELL, 1960; MENGE, 1974) and has been adopted here (Figure 1). The terms anterior slope, posterior slope and disc, although not consistently applicable to all bivalve shells, are used here as follows: anterior slope = area anterodorsal to a line drawn from umbo to junction of anterior and ventral margins; posterior slope = area posterodorsal to a line drawn from umbo to junction of posterior and ventral margins; disc = the remaining central area of the shell between those lines (see Figure 1). It should be reemphasized that right and left valves had been separated in this lot so there is little possibility for identifying complete individuals. It is assumed that the presence of a borehole in one valve of a bivalve indicates the individual was killed by this attack, and that some- where there exists (or existed) an opposite unbored valve (see THomaSs, 1976). If we assume that this sample repre- -sents a natural assemblage, there is a slight preference for the predator to drill the left valve, 55% of those bored being the left ones. There is a more obvious preference in the location on the shells where the borings are made, 75% of the holes drilled in the left valves and 54% of holes in the right valves are located on the anterior slope; the remaining 25% and 46%, respectively, of the borings are located on the disc (see fig. 1). No holes are located on the narrow posterior slope. Most holes are located nearer the umbos dorsally than they are to the ventral margin. The fact that most boreholes are nearer the dorsal edge of the valve than in any other position indicates feeding behavior of the predator similar to that described by FRETTER & GRAHAM (1962, pp. 572-574). The latter de- scribed predation by Natica which manipulates the bivalve prey with its foot, turning the bivalve so that it can bore into it in a precise manner. This complexity of behavior Page 250 seems to characterize some gastropod predation and espe- cially that entered into by Naticidae (CarRRIKER & YOCHEL- Son, 1968; Epwarps, 1975; BERG & NISHENKO, 1975). Location of the boreholes mostly in the dorsal area of valves of the population of Periploma margaritaceum may indicate predator preference for boring in that site. Since shells of P margaritaceum are rather uniformly thin, it seems probable that boreholes are made in this area of the shell because it provides the most efficient access to the predator’s meal. As pointed out by CarRIKER & YOCHEL- SON (1968) location of Naticid borings may be limited by the grasping ability of the predator. While that interpre- tation would place matters of preferences into the cate- gory of genetically programmed behavior, it is extremely likely that the grasping ability of the predator has been subject to intense selective pressure favoring most efficient feeding on available prey species. ANSELL (1960) found Natica alderi (Forbes, 1838) to bore preferentially in the ventral regions of the shells of Venus striatula (da Costa, 1778). He suggested that one reason for this was that the relatively thick shell of Venus is thinnest near the shell margins, outside the pallial line. It appears that in N. aldert, also, borehole location is dic- tated by greatest feeding efficiency. Franz (1977) dem- onstrated that it was the smaller valves of Spisula solidis- sima (Dillwyn, 1817) that were bored by Lunatia heros (Say, 1822). Presumably, then, either due to choice or evolutionary pressure, ease of valve penetration may be an important factor in determining where boreholes are made (also see CARTER, 1968: 41). To detect valve preference and compare locations of predator boreholes in other members of the family Peri- plomatidae a survey was made of Peritploma margarita- ceum from other localities and seven other species having different shell shapes and sizes. The results are shown in Table 2 and Figure 2. Specimens of P margaritaceum in- THE VELIGER Vol. 22; No. 3 cluded in Table 2 are from lots other than those tabulated in Table 1. Certain tendencies may be noted. In all species most boreholes are located near the dorsal valve margins. Again, in P margaritaceum most holes are in the anterior slope. However, in contrast to the initial lot of RP margan- taceum, in this sample, the right valve was bored consider- ably more often than the left indicating there is variation in the valve chosen by the predator for boring or perhaps variation in which valve is uppermost. Table 2 Survey of Borehole Locations in Periplomatidae (see Figure 2) Valve Position of Bored Borehole Anterior , Left Right] Disc Slope Periploma margaritaceum Lamarck 19 30 3 46 Periploma ovatum Kuroda and Habe 1 1 Cochlodesma leanum Conrad 25 19 40 4 Periploma fragile Totten 13 30 34 9 Periploma stearnsii Dall 1 1 2 Periploma fracturum Boshoff 1 1 Periploma discus Stearns 1 1 1 1 Periploma planiusculum Sowerby 2 3 3 2 Of the other species surveyed, specimens of Cochlo- desma leanum and Periploma fragile were numerous enough to give an indication of predator selectivity in bor- ing sites (see Table 2 and Figure 2, E-H). In C. leanum there appears to be a slight preference for left valves (579%) versus right (43%) as a site for boring. The area of Explanation of Figure 2 A, B. Periploma margaritaceum Lamarck, Matagorda Bay, Indi- anola, Calhoun County, Texas, USNM 465183. A. left valve, length 18 mm; B. borehole, largest diameter (including bevel) 2.5 mm C, D. Periploma planiusculum Sowerby, Dominical, Costa Rica, USNM 597917. C. left valve, length 37.1mm; D. borehole, diameter 4mm E, E Cochlodesma leanum Conrad, south of Gay Head, Martha’s Vineyard, Dukes County, Massachusetts, USNM 95546. E. left valve, length 33.7mm; F borehole, diameter 3.8mm G, H. Periploma fragile Totten, Sand Island, New Brunswick, Cana- da, USNM 444773. G. left valve, length 14.1 mm; H. bore- hole, diameter 1.9mm I, J. Periploma stearnsit Dall, off Point Fermin, Gulf of California, USNM 110548, Holotype. I. right valve, length 44.8mm; J. borehole, diameter 4.5 mm K, L. Periploma fracturum Boshoff, Anton Bruun Station gg1-F, northeast of Durban, South Africa, 77m, USNM 718117. K. right valve, length 17.4mm; L. borehole, diameter 1.8mm; M, N. Periploma discus Stearns, Long Beach, California, USNM 126931, Holotype. M. right valve, length 43mm; N. borehole, diameter 3.5 mm ci THe VELIGcER, Vol. 22, No. 3 [RosEWaTER] Figure 2 Vol. 22; No. 3 the disc (90%) is clearly more often penetrated than the anterior slope (9%). In P. fragile right valves (70%) are more often penetrated than left (30%). The disc (79%) again appears to be the boring site chosen over the anterior slope (21%). It is not known what predator species were involved in attacking the bivalves surveyed in Table 2, except that the boreholes indicate Naticid predation. No boreholes were found in posterior slopes of any specimens examined in this survey. That area of the shells of periplomatids tends to be truncated, generally reduced in size, and may be in an inconvenient area for penetration by Naticids. SUMMARY 1. Of 443 valves of Per'ploma margaritaceum from Ma- tagorda Bay, Texas, 185 or 42% were bored by a naticid gastropod, probably Polinices duplicatus. 2. In this sample a slightly larger percentage of left valves of Periploma margaritaceum is bored than right valves. In a smaller sample of the same species it is the right valve that is more frequently bored. 3. While Peritploma margaritaceum and other Periplo- matids may lie horizontally on one side within the substratum, there is little evidence from location of boreholes as to which side is uppermost. 4 In almost all instances boreholes are made in the dor- sal part of the valves, either in the anterior slope or disc. These data are consistent with evidence from other workers indicating that naticid predators ma- nipulate prey and select areas for boring that allow most efficient penetration and feeding. 5: Whereas in Periploma margaritaceum the anterior slope is the preferred site of boring, in Cochlodesma leanum and Periploma fragile it is the disc. This dif- ference may be due to the fact that the former is strongly inequilateral while both C’. leanum and P. - fragile are more equilateral than P. margaritaceum. 6. No boreholes were noted in the posterior slope, prob- ably due to its small size. ACKNOWLEDGMENTS I am grateful to R. S. Houbrick, E. L. Yochelson and T. R. Waller for valuable comments and criticism, and to Luana Read and Helen Kafka for tabulating data and illustrating borehole locations in Periploma margarita- THE VELIGER Page 251 ceum; H. Harry suggested the identity of the naticid predator. Literature Cited Auten, J. H. 1958. Observations on Cochlodesma praetenue (Pulteney) [Eulamelli- branchial]. Journ. Mar. Biol. Assoc. U. K. 37: 97-112 ALLER, Rosert C. 1974. Prefabrication of shell ornamentation in the bivalve Laternula. Lethaia 7 (1): 43-56 ANSELL, ALAN Davip 1960. Observation on predation of Venus striatula (DaCosta) by Natica alderi (Forbes). Proc. Malacol. Soc. London 34 (3): 157-164 Bere, Cart J. & SruarT NISHENKO 1975- Stereotypy of predatory boring behavior of Pleistocene naticid gastropods. Paleobiol. 1 (3): 258-260 CarrikeR, MELBOURNE RoMAINE & ELLIs LEON YOCHELSON 1968. Recent gastropod boreholes and Ordovician cylindrical borings. U. S. Geol. Surv. Prof. Paper 593-B: 26 pp.; 5 plts.; figs. Carter, R. M. 1968. On the biology and paleontology of some predators of bivalved Mollusca. Palaeogeogr., Palaeoclimatol., Palaeoecol. 4 (1): 29-65 Epwarps, D. Craic 1975. Preferred prey of Polinices duplicatus in Cape Cod inlets. Bull. Amer. Malacol. Union for 1974: 17 - 20 [published in 1975] Franz, Davp R. 1977- Size and age-specific predation by Lunatia heros (Say, 1822) on the surf clam Spisula solidissima (Dillwyn, 1817) off western Long Island, New York. The Veliger 20 (2): 144-150; 5 text figs. (1 October 1977) Fretrer, VERA & ALASTAIR GRAHAM 1962. British prosobranch molluscs, their functional anatomy and eco- logy. London, Ray Soc. xvi+755 pp.; 316 figs. MENGE, JANE LuBCHENCO 1974. Prey selection and foraging period of the predaceous rocky intertidal snail, Acanthina punctulata. Oecologia (Berlin) 17: 293 - 311 Morse, Epwarp SYLVESTER 1919. Observations of living lamellibranchs of New England. Proc. Boston Soc. Nat. Hist. 35 (5): 139-196 Morton, Brian S. 1976. The structure, mode of operation and variation in form of shell of the Laternulidae (Bivalvia: Anomalodesmata: Pandoracea) Journ. Moll. Stud. 42 (2): 261-278 ROSEWATER, JOSEPH 1968. Notes on Periplomatidae (Pelecypoda: Anomalodesmata) with a geographical checklist. Ann. Rprt. Amer. Malacol. Union for 1968: 37 - 39 STANLEY, STEVEN M. 1970. Relation of shell form to life habits of the Bivalvia (Mollusca). Geol. Soc. Amer. Mem. 125: i-xiiit+296; plts. 1-40; figs. 1-48 4 (December 1970) Tuomas, R. D. K. 1976. Gastropod predation on sympatric Neogene species of Glycy- meris (Bivalvia) from the eastern United States. Journ. Paleont. 50 (3): 488-499 VERMEIJ, GEERAT J. 1978. Biogeography and adaptation patterns of marine life. Har- vard Univ. Press, Cambridge, Mass. xi+332 pp.; illust. Added reference: : Several weeks after returning the galley proofs the following paper ap- peared: WILLIAMS, LORALYNN ) : { 1979. Boring and feeding behaviors (sic) of the California marine gastrapod Ceratostoma nuitallt. Of Sea and Shore 10 (1): 18-203 3 text figs. Page 252 THE VELIGER Vol. 22; No. 3 Habitat, Food and Reproductive Activity of the Nudibranch Hexabranchus sanguineus on Tongatapu Island MALCOLM P. FRANCIS" ’Atenisi Institute, Nuku’alofa, Tonga (4 Text figures) INTRODUCTION THE TAXONOMY OF THE GENUS Hexabranchus Ehrenberg, 1831 is confused and requires revision. More than 20 species have been described (see THompson, 1972), many of them based primarily on differences in colour pattern. Some authors (BERGH, 1900; ELIoT, 1904; THoMPsoN, 1972) consider the many named species to be merely colour varieties of Hexabranchus sanguineus (Rippell & Leuckart, 1828). The validity of the use of colour pat- tern as a taxonomic character remains to be demonstrated in this genus. Nevertheless, a full description of the col- our pattern of Tongan Hexabranchus is given below to facilitate comparative studies in the event that colour does prove to be distinctive. External and internal mor- phology appear to vary little among the described varie- ties (ELioT, 1904) but these features were not investi- gated in Tongan animals. In the absence of any distinctive taxonomic characters, Tongan Hexabranchus are here referred to the type spe- cies of the genus, H. sanguineus (Ruppell « Leuckart). Other described “species” will here be considered simply as colour variations of H. sanguineus. Little work has been done on the biology or ecology of Hexabranchus. The only comprehensive study is that of GouHar & SOLIMAN (1963) on Red Sea specimens. Several authors have commented on feeding and stomach con- tents, and a review of their results is presented below. OSTERGAARD (1950) has described spawning and develop- ment in 2 Hawaiian varieties. The only Hexabranchus previously recorded from Ton- ga is H. flammulatus (Quoy & Gaimard, 1832). Speci- mens 10 to 13cm long were collected during the voyage t Present address: 44 Belmont Terrace, Auckland 10, New Zealand of the Astrolabe from Tongatapu Island (21°10’S; 175° 10’W; see Figure 1). DESCRIPTION The colour of the dorsum and foot of Hexabranchus san- guineus varied from pale pink to dark red. The general appearance was, however, influenced by the colour, size Hoositea ° - Beach Figure 1 Tongatapu Island, Tonga, showing localities mentioned in the text. Dotted line indicates the edge of the fringing reef. Scale=5 km Vol. 22; No. 3 and density of spots covering the dorsum; these ranged from large dense yellow spots (producing an overall orange appearance) to small white spots (which allowed the background colour to dominate the appearance). Most of the animals seen were dark red with these dense yellow spots on the dorsum (Figure 2, zone A). Figure 2 Close-up photograph of a large Hexabranchus sanguineus showing colour zones. See text for explanation of colour pattern of the various zones. Scale=1cm In animals longer than about 6 cm, 5 colour bands were distinguishable on the notum (see Figure 2). Band B was comprised of a translucent layer of white pigment over- lying dark red tissue, producing an overall mottled pink band punctuated with red spots. Band C was a dark red colour. Occasionally, band B was interrupted by a dark red intrusion from band C which linked with the dorsum. These intrusions probably represent areas from which the overlying pigment of band B is absent. Band D was white with dark red lines radiating out from band C. These lines merged to produce a mauve coloured band E. The edge of the notum, band F, was always white. Animals shorter than 6cm were very similar to larger animals, except that bands D and E were frequently merged to form a single light purple band. One small animal (1.8cm long) had a different colour pattern. The notum had only 3 colour bands: a white margin tinged with blue, a central orange band with violet notches along its distal edge, and an inner white band. This could represent the juvenile colour pattern, THE VELIGER Page 253 with the number of bands increasing with animal size as the violet notches coalesce to produce the characteristic red and purple bands of all larger animals. The underside of the notum and the foot were dark red with yellow or white spots. The rhinophores were red, and spotted with yellow or white, and the lamellae were red with yellow edges. The rhinophore collars were densely spotted with yellow or white and had bright red rings around their distal edges. The gills were yellow and veined with red. The only significant difference in colour pattern be- tween Tongan Hexabranchus examined in this study and H. flammulatus described by Quoy & GammarD (1832: plt. 17, figs. 6 & 7) is the presence in the latter of distinct white pennants punctuated with red adjoining the dor- sum. These pennants clearly represent the remains of colour band B after it has been split into segments by dark red intrusions from band C. Distinct segments were not observed in the present study and band B was usually continuous around the whole notum. Distinct segments appear to be a feature of Australian Hexabranchus and are clearly seen in illustrations by GmLetr « McNEILL (1959: plt. 84) and THompson (1972: fig. 2B). Other- wise, Tongan and Australian animals differ little in colour pattern. Hexabranchus marginatus (Quoy & Gaimard, 1832) as illustrated by BERGH (1905; plt. 1, fig. 2) has an almost continuous band B and is very similar to Tongan animals. The only visible difference is that band D in H. marginatus is yellow in Bergh’s figure, although Quoy « Gaimard’s original description (1832: 255) gives it as white. METHODS During a preliminary survey of the coastline of Tongatapu Island, Hexabranchus sanguineus were found to occur predominantly on the shallow fringing reef platforms all around the island except on the sheltered north coast. The animals were most common on the south and south- west coasts where the fringing reef is approximately 1oom wide. Two sections of reef, at Hoositea Beach and Mono- tapu Beach (Figure 1), were chosen for further study and were visited regularly between March and November 1976. HOOSITEA BEACH A section of reef 100m long was searched from the beach to the reef rampart (a distance of 125m) at ap- Page 254 THE VELIGER Vol. 22; No. 3 proximately monthly intervals between March and Octo- ber 1976. Searches were conducted at low tide and cov- ered an area of 1.25 hectares. All Hexabranchus sangut- neus found within the area were counted and measured to the nearest centimeter below extended crawling length (excluding the foot). This measurement corresponds with the standard mantle length, Am, of Risso-DomMINcUEz (1963). Channels in the reef were examined by snorkel diving. Sea surface temperatures were measured at mid-morn- ing over the low tide period in surge channels just inside the edge of the reef. Specimens of Hexabranchus sanguineus were collected, cleaned of adhering debris and placed individually in seawater-filled jars which were then submerged in reef channels. After 1 to 5 hours the animals were released and their faeces collected and preserved in neutral forma- lin. MONOTAPU BEACH The inner 85m of a section of reef 60m long was searched at least once a month from April to November 1976. Searches were conducted at low tide and covered an area of 0.5 hectare. The outer edge of the reef was not searched because preliminary surveys had shown Hexa- branchus sanguineus to be absent from this part of the reef, where a strong current flowed across the reef from the south. All Hexabranchus sanguineus egg coils found within the area were counted and removed (to ensure that no egg coil was counted in a subsequent search). Faecal samples were collected from live animals as for Hoositea Beach. RESULTS HABITAT Hexabranchus sanguineus were abundant on shallow reef platforms on exposed coasts. In one 3-hour period, 45 animals were found in the 1.25 hectare area searched at Hoositea. Hexabranchus sanguineus could only be found readily over the low tide period. At other times they disappeared from view, probably into coral crevices in the reef. Reef platforms on exposed Tongan coasts are protect- ed from the prevailing south-easterly swells by raised ramparts at their seaward edges. The extent of protection afforded to reef platform organisms depends upon the state of the tide: during the low tide the water remaining on the reef is calm, whereas at high tide waves break over the rampart producing turbulent conditions. The rampart at Hoositea shelters the reef for about half of the 12-hour tidal cycle. The reef is almost hori- zontal in profile and is covered by about 1.5m of water at high tide. At low tide the reef dries almost completely, except for the numerous channels in the reef. These channels are up to 2m deep and run at right angles to the beach. Monotapu has a similar reef structure to Hoositea, but the reef is subject to turbulent wave action for about 2 of the tidal cycle, and does not drain completely at low tide. A layer of water 10cm to 30cm deep remains at low tide. The reef platform around most of Tongatapu Island is composed of compacted fused coral, and covered by a turf of coralline algae. Live coral (mainly Porites and Acropora species) is rare and confined to the channels which are water-filled at all times. Patches of dead Acro- pora project up to 30cm above the level of the reef flat. At low tide, Hexabranchus sanguineus were usually found crawling over the substrate, either among dead Acropora branches or on the algal turf. The animals ac- tively avoided live coral, but were frequently seen moving among the algae-encrusted bases of live coral. They were rarely found in the reef channels, but sometimes occurred on the vertical sides of the channels. Hexabranchus sanguineus probably shelter in coral crevices over high tide and thus reduce the risk of being swept from the reef or being damaged by the abrasive action of suspended sediment. The distribution of Hexabranchus sanguineus across the reef was found to be non-random with respect to the size of the animal. The distribution by size was analysed dur- ing one low tide sampling period (1 July 1976), and the results are presented in Figure 3. Animals less than 3 cm long occurred only on the outer edge of the reef near the rampart. Hexabranchus sanguineus between 3cm and 7 cm were found throughout the outer half of the reef, and large animals (exceeding 7cm) occurred over the whole reef, but were most common on the inner half. WATER TEMPERATURE At Hoositea, water covering the outer 4 of the reef platform is continually renewed at all tides by the in- flux of ocean water through the surge channels in the reef rampart. The outer section of the reef platform is there- fore buffered against large diurnal fluctuations in temp- erature and salinity. The inner # of the reef is more sub- ject to temperature changes due to solar heating and Vol. 22; No. 3 120 & ° 99 (2) 3 —Q E 60 & fe) 8 =| = go A fo} i 8 9a) BO. 4 Beg ao oe Length (cm) Figure 3 Distribution of Hexabranchus sanguineus across Hoositea reef by size (n=45). Open circles indicate mean distance of each size class from the beach, and vertical lines indicate the observed ranges. Size-classes with no marked range were represented in the sample by only one specimen each night-time cooling, and salinity changes due to evapora- tion and freshwater seepage from the beach. Temperature and salinity conditions at Monotapu are more constant than at Hoositea. The water level over the reef does not drop as low at Monotapu, and ocean water circulates continuously over most of the reef. Sea surface temperatures measured at Hoositea are plotted in Figure 4, and indicate the trends during the period April to October. Temperatures ranged from 22.5° C to 27.6°C. The highest temperature recorded in the shallow water overlying the inner sections of Hoositea reef was 29.4°C; higher temperatures than this would be expected from January to March. REPRODUCTIVE ACTIVITY The egg coils of Hexabranchus sanguineus found in Tonga were consistent with the description of GoHaR & SoLIMAN (1963: 230-231). Freshly-laid coils were bright red, fading to a dull red-brown colour as the embryos developed. The number of egg coils found at Monotapu fluctuated randomly from April to November (Figure 4). Egg coils were found in varying numbers during the coldest months, but apparently disappeared as the water temperature started increasing again. The absence of samples from the hottest part of the year precludes a full comparison of reproductive activity with seawater temperature. THE VELIGER Page 255 Water Temperature (°C) Oo 28 MT HM ST o Number of Egg Coils Months in 1976 Figure 4 Number of egg coils of Hexabranchus sanguineus found at Mono- tapu (open circles) and sea surface temperature measured at Hoositea (closed circles) Hoositea was less preferred as a breeding site than Monotapu. Only 2 egg coils were found at Hoositea despite the much larger area searched throughout the study period. SIZE STRUCTURE or tHe HOOSITEA POPULATION The size-frequency distributions of the Hoositea Hexa- branchus sanguineus samples are shown in Table 1. Very small individuals were probably present throughout the study period, despite their absence from the first 2 samples. An animal 1.8cm long was found at ’Atele Beach (Figure 1) on 19 April 1976, and a 1.5cm animal (the smallest found in this study) at Hoositea on 24 April 1976. Even the 1.5cm long animal was able to flare its notum when disturbed, and was able to swim. Large adults were present throughout the study, although none larger than 14cm was found. Most of the large samples had unimodal size-frequency distributions with modes between 5cm and 7cm. Small size classes have probably been under-represented because of their cryptic coloration. Since animals less than 4cm were present from April to October, it appears that larval settlement on the reef occurs throughout this period. Lar- val settlement was especially heavy some time prior to the sample of 29 May 1976, as this sample contained Page 256 THE VELIGER Vol. 22; No. 3 Table 1 Size-frequency distribution of samples of Hexabranchus sanguineus taken at Hoositea Beach. Length Date of (nearest cm below) Number sample in in 1976 1 2 3 4 5 6 7 8 9 10 11 12 13. sample 14 March 1 3 2 1 1 2 10 18 April 1 1 2 3 1 1 1 1 2 16 29 May 2 3 1 1 4 5 2 4 4 2 1 1 30 1 July 1 5 5 5 7 9 7 4 1 1 45 30 July 3 6 8 5 6 1 1 1 31 30 August 2 2 4 8 9 7 6 3 1 43 26 September 2 2 4 2 1 3 2 3 2 1 23 22 October 1 2 3 4 7 5 1 2 1 1 27 many small animals. The strong size class could also be detected in subsequent samples. Few animals longer than 10cm were found. FEEDING An examination of faecal material showed that Hexa- branchus sanguineus feeds on sponges. All of the 13 Table 2 Sponge species identified from faeces of Hexabranchus sanguineus and their frequency of occurrence. Number of animals in Sponge species which sponge was found (n = 6) CALCAREA Calcareous triradiates 2 DEMOSPONGIAE CHorIsTIDA Stelletta sp. 1 Ancorina acervus (Bowerbank) 1 Pachastrella sp. 1 HADROMERIDA Chiona sp. 3 HAPLOSCLERIDA Haliclona sp. or Adocia sp. 2 Callyspongia sp. 3 POECILOSCLERIDA Xestospongia exigua (Kirkpatrick) 1 Petrosia sp. 5 Paraesperella sp. 1 Mycale sp. or Zygomycale sp. 4 faecal samples collected contained a large proportion by volume of sponge spicules. 'The species eaten, based upon spicule identification of 6 faecal samples, are shown in Table 2. Hexabranchus sanguineus feeds on at least 11 genera of sponges from 5 orders and 2 classes, and is clearly a non-selective browser. A wide variety of non-sponge material was also identi- fied from the faeces. Non-sponge components of the gut contents found in this study and by other authors are listed in Table 3. This material never formed more than a small proportion of the faecal material of Tongan animals. Table 3 Non-sponge material identified from faeces of Hexabranchus sanguineus. Material Source Algae ABsout-E1a (1959); GoHar & SOLIMAN (1963); present study Foraminifera EALEs (1938); GoHar & SoLiman (1963); present study Hydroids BeErGH (1900); Error (1906) Alcyonarians Gouar & Sottman (1963) Coral fragments Worm tubes Gastropod shells Amphipods Crab chela Polyzoans Echinoderm shells Ascidian spicules Gouar & SOLIMAN (1963); present study Bercu (1900); Eares (1938); present study Fates (1938); present study Present study Present study Exior (1906) EALEs (1938) THompSsON (1972) Vol. 22; No. 3 THE VELIGER Page 257 DISCUSSION mals ingest much extraneous material while feeding on the encrusting sponges in the crevices where other or- REPRODUCTIVE ACTIVITY ganisms, animal skeletons and detritus also collect. Red Sea populations of Hexabranchus continued to breed throughout the year but activity declined during the cold months (GoHar & SoLIMAN, 1963). The num- ber of egg coils at Monotapu showed no relationship to seawater surface temperature during the 8 months of observations (Figure 4). GoHAR & SOLIMAN (1963: 242) believe that spawning in H. sanguineus is induced by in- creasing temperatures following winter low temperatures. The minimum temperatures in the Red Sea (16°C to 19° C [Gouar & SoLIMAN, loc. cit.]) were considerably lower than the lowest temperature measured during the Tongan winter (22.5°C), and it is possible that the temperature of Tongan waters is suitable for uninterrupted year-round breeding. The scarcity of egg coils at Hoositea was probably due to unfavourable environmental conditions. The tempera- ture of water trapped in channels at low tide during the day increases and the dissolved oxygen concentration probably decreases. GoHaR & SOLIMAN (1963) observed retardation of embryo development in stagnant water. Egg coils kept in a non-circulating aquarium in this study disintegrated before larvae were ready to hatch. The continual renewal of seawater at Monotapu produces a more constant environment for larval development than exists at Hoositea. FEEDING Hexabranchus sanguineus is a sponge feeder, consum- ing a wide variety of species. The sponges taken are main- ly inconspicuous encrusting species, probably occurring in crevices and holes in the reef matrix (P. Bergquist, per- sonal communication). Most other workers have also recorded sponges in the diet of Hexabranchus. BERcH (1900: 231) found “fragments of transparent spicules” and “masses of simple and three-rayed silica needles” in one specimen; these were obviously sponge remains. ELI- oT (1906) and Gowar & SOLIMAN (1963) noted that sponges form at least part of the diet of Hexabranchus, and Younc (1966) reported the calcareous sponge Leu- cetta solida from the gut of a Hawaiian specimen. Kay & YouNc (1969) concluded that 3 Hawaiian varieties of Hexabranchus were “rasping sponge feeders.” The wide variety of non-sponge material in the gut contents of the different varieties of Hexabranchus is indicative of an indiscriminate mode of feeding. The ani- The contention that Hexabranchus is entirely herbi- vorous (ABouL-ELA, 1959) is not supported. It is likely that algae are ingested accidentally during feeding, and they appear to pass through the gut undigested. Broom (1976) has demonstrated a correlation between the radula and digestive morphology of dorid nudi- branchs, and the type of sponge preferred as prey. Bloom’s major thesis is that the presence of a gut caecum (a spicule-compacting organ) enables nudibranchs to handle large quantities of large sharp spicules, and they are therefore adapted to feeding on sponges with unorganised or non-reticulated skeletons. Conversely, nudibranchs lacking a gut caecum, but possessing a robust radula, are able to feed on, and digest, sponges with organised, re- ticulated skeletons. The only sponge prey of Hexabran- chus marginatus mentioned by BLoom (1976: table 3) is Leucetta solida, originally reported by Younc (1966). Thus, using limited data, Bloom finds that Hexabranchus, which possesses a caecum and radula teeth with a low degree of hook (Bloom’s measure of tooth robustness) feeds on non-reticulated sponges; this supports his con- tention that a correlation exists between nudibranch morphology and sponge prey. However, the present study has shown that H. sanguineus consumes a wide variety of sponge types, ranging from unorganised non-reticulated sponges (calcareous species) to highly organised reticu- lated speces (e. g., Mycale sp., Petrosia sp., Callyspongia sp.) (see Broom, 1976: table 1, for a description of skeletal structure in each of these groups). The presence of a gut caecum may indeed allow Hexabranchus to feed on non-reticulated sponges, but the absence of robust radula teeth does not prevent it from also eating reticu- lated sponges. Generalist sponge feeders are exceptions to Bloom’s correlations, and other species may also prove to be generalists on closer study. ACKNOWLEDGMENTS I would like to thank Shirley Cummins for her consider- able assistance with field work in Tonga, despite often adverse conditions. I also thank Professor P. R. Bergquist (University of Auckland, N. Z.) for identifying the spon- ges consumed by Hexabranchus. I am very grateful for the criticisms and helpful comments given by Professor P. S. Corbet (University of Canterbury, N. Z.) and Dr. G. N. Soliman (University of Sulaimaniyah, Iraq) while this research was at manuscript stage. Page 258 THE VELIGER Vol. 22; No. 3 Literature Cited Asout-E1a, IpRAHIM AHMED 1959. On the food of nudibranchs. Biol. Bull. 117: 439-442 Bzrox, Rupotr Lupwic SopHus 1900. Die Opisthobranchier. Pacific (Schauinsland 1896-1897). plts. 1-21 1905. Die Opisthobranchiata der Siboga Expedition. prt. 50: 1 - 248; plts. 1-20 Bioom, STEPHEN A. 1976. Morphological correlations between dorid nudibranch predators and sponge prey. The Veliger 18 (3): 289-301; 1 text fig. (1 January 1976) Ergebnisse einer Reise nach dem Zool. Jahrb. Syst. 13: 207 - 246; Siboga Exp. (October 1905) Eaves, Nevur B. 1938. A systematic and anatomical account of the Opisthobranchia. John Murray Expedition, 1933-34. Sci. Reprt. Brit. Mus. (Nat. Hist.) 5: 77-122; 1 plt.; 28 text figs. EHRENBERG, CHRISTIAN GOTTFRIED 1831. Animalia evertebrata, exclusive insectis. In: C. G. Ehren- berg « E G. Hemprich (eds.), Symbolae physicae. Berlin Exiot, CHARLES Norton EDGECOMBE 1904. On some nudibranchs from East Africa and Zanzibar. Part 6. Proc. Zool. Soc. London 2 (I): 268 - 298; plts. 16, 17. (9g June 1904) 1906. Nudibranchiata, with some remarks on the families and genera, and description of a new genus, Doridomorpha. In: J. S. Gar- diner (ed.), The fauna and geography of the Maldive and Laccadive Archipelagoes. 2: 540-573. Cambridge Univ. Press, London GittettT, KerirH & FranK McNEILL 1959. The Great Barrier Reef and adjacent isles. Sydney. 194 pp.; 161 plts.; 3 text figs. Gouar, H. A. E & Gamit N. Souiman 1963. The biology and development of Hexabranchus sanguineus (Riip- pell « Leuckart) (Gastropoda, Nudibranchiata). Publ. Mar. Biol. Sta. Al-Ghardaga 12: 219-247; 37 text figs. Kay, ExvizaBpeTtH ALison & Davin KENNETH YOUNG 1969. The Doridacea (Opisthobranchia; Mollusca) of the Hawaiian Coral Press, Islands. Pacif: Sci. 23 (2): 172 - 231; 82 text figs. (30 April 1969) OsTERGAARD, JENS MaTHIAS 1950. Spawning and development of some Hawaiian marine gastro- pods. Pacif. Sci. 4: 75-115; 42 text figs. (28 April 1950) Quoy, JzEAN RENE CoNSTANT & JosePpH PAuL GalmarD 1832. Voyage de découvertes de l’Astrolabe. Zoologie, Mollusca 2. Tastu, Paris Risso-DomINGuEzZ, Cartos J. 1963. | Measuring nudibranchs: a standardization for descriptive pur- poses. Proc. Malacol. Soc. London 35: 193 - 202; 2 text figs. Ruppert, WILHELM Peter Epuarp S. & FRIEDRICH SIGISMUND LEUCKART 1828. | Neue wirbellose Thiere des Rothen Meers. In: Eduard Riip- pell, Atlas zu der Reise im nérdlichen Afrika 5. pp. 1-47; plts. 1-12 Tuompson, THomaAS EvERETT 1972. Observations on Hexabranchus from the Australian Great Bar- rier Reef (Gastropoda : Opisthobranchia). The Veliger 15 (1): 1-5; 2 text figs. (1 July 1972) Younc, Daviw KENNETH 1966. Systematics, food and functional morphology of the feeding apparatus of some dorid nudibranchs. Ph. D. dissertation, Univ. Hawaii, Honolulu Vol. 22; No. 3 THE VELIGER Page 259 Depth Distribution of Nautilus pompilius in Fiji and Nautilus macromphalus in New Caledonia PETER D. WARD Department of Geology, University of California, Davis; Davis,California 95616 ARTHUR W. MARTIN Department of Zoology, University of Washington, Seattle, Washington 98195 (1 Plate; 6 Text figures) INTRODUCTION AMMONITE woRKERS have long noticed that adults and juveniles of the same species are often segregated by facies. This observation indicates either selective preservation, selective post-mortal drift, or that different growth stages were separated geographically or by depth. KENNEDY & CopBan (1976) have discussed this problem in their review of ammonite paleobiology, and have concluded that at least in some instances the latter hypothesis has been valid. It was therefore suggested that significant migration must have occurred during the lifecycle of some species. Migration for feeding or breeding is well-known in many coleoid cephalopods (PackarD, 1972), but has been little examined in Recent Nautilus, which, because of its greater morphologic similarity to the ammonites than the coleoids, is probably a more useful analogue. We have examined the depth and geographic distributions of 2 species of Nautilus, N. pompilius in the Fiji Islands, and N. macromphalus in New Caledonia. The results of these observations, reported in this paper, indicate that separation of adults and juveniles occurs within these 2 populations. Our interpretation of reasons for this segre- gation in Nautilus may have some applicability in inter- preting distribution patterns of ammonites. METHODS anp CAPTURE SITES The captures of Nautilus were accomplished using meth- ods previously described by Warp et al., 1976. The use of hydraulic winches and sophisticated bottom sounding gear in Fiji, donated by the department of Fisheries, Fiji, enabled us to set traps to depths of 750m. In New Cale- donia, trapping efforts were conducted on small private boats without the benefit of winches, and hence were se- verely limited as far as depth capabilities. No trapping from depths greater than 100m was attempted. The reef environments and trapping areas in Fiji have been previously described (Warp et al., 1977). In New Caledonia, all captures have been made at one area on the barrier reef fore-reef slope near Noumea (Figure 1). Seaward of the tom wide algal ridge the reef flat ex- tends into a zone of actively growing coral heads which are dissected by spur and grooves perpendicular to the reef crest. This zone is several hundred meters wide and descends to a depth of 10 to 15m, where a pronounced break in slope occurs. A steep scarp, descending to depths varying between 40 and 50m, is covered with growing coral in the upper half, but shows mainly algae in the lower portions. At the base of this reef precipice is a talus slope of reef rubble and sand, with little actively growing coral. The talus slope extends downward to a flatter THE VELIGER Vol. 22; No. 3 Figure 1 Trapping area of Nautilus macromphalus, New Caledonia. Depths given in meters terrace of coralline sand and low, hummocky ridges of rubble and algae which commence at depths between 80 and room. All of our trapping in New Caledonia has been from the reef talus and the deeper coralline sand terrace. RESULTS In Fiji we captured 57 Nautilus from known depths and positions during 1976 and 1977. These specimens have been separated into groupings by depths and compared (Figure 2). Twenty-five specimens from depths between 100 and 300m had an average weight (in air) of slightly less than 500g. No specimens of less than 300¢ total weight were captured. Thirty-two specimens from depths greater than 300m had a mean weight of 413g, and 9 animals of less than 300g were captured. If these assem- blages were to be preserved in the fossil record, we would see a shallower facies with no juveniles, and a deeper facies with both juveniles and adults. In Fiji, the deepest captures were from depths of 600 to 650m. Traps were placed at 700m on 5 occasions, with no Nautilus captures resulting. The maximum habi- tat depth of Nautilus is dictated at least in part by the strength of the shell against implosion, and to study this depth limit a number of implosion experiments has been made in pressure devices. DENTON & GILPIN-BROWN (1966) imploded shells at pressures of 53-73 atms., equivalent to depths between approximately 500 and V/A Males Nautilus pompilius 300-600 m Nn = 32, X =413.2g N Females 2 = 0.26%x, = 360.78 [__ ]Unsexed 10 S = 0.74%X,, = 478.38 Number of Specimens on 100 300 500 700 Nautilus pompilius 100-300m n= 25, X = 4908.2¢ 3b =0.71%, xa = 539.028 2 = 0.29%, X, = 400.58 Number of Specimens oO 100 300 500 700 Total Weight (g) Figure 2 Size-sex distributions of Nautilus pompilius, Fiji Islands (Suva Harbor region). The upper group was captured in depths between 300 and 600 m;; the lower from depths of 100 to 300m 7oom. Raup & TAKAHASHI (1966) and SAUNDERS & WeEHMAN (1977) arrived at similar figures. In Fiji we inadvertently imploded the shells of 6 living animals, but at a depth greater than any of those produced experi- mentally in pressure chambers. The newly captured speci- mens were lowered in a trap and buoy system to 700m. One day later, during recovery, we noted that the trap had moved downslope into deeper water. The new resting depth of the trap was deeper than the maximum working depth of the echo sounder on board the recovery vessel, so no measurement of the actual depth was obtained. It Vol. 22; No. 3 THE VELIGER Page 261 is estimated that the trap was in a depth between 800 and goom, however, because only 1000m of line was at- tached, and between 100 and 200m of this was slack on the surface. Retrieval showed that the shells of all animals had imploded (Figures 7A to 7D). In a second experiment, 2 animalswere lowered to the maximum depth of the echo sounder, 750m, and left on the bottom in a closed trap for one hour. Both animals were intact and healthy on recovery, with no apparent excess of cameral liquid. These observations, coupled with our distributional data, indicate that the maximum depth of habitat is probably between 600 and 700m, and that depths much in excess of 800m will implode the shells. The patterns of shell breaks (Figure 7) in the imploded shells are remarkably similar. In each shell both umbili- cal calyces are preserved. The body chambers were broken into large, curved portions. The external shell surrounding the final phragmocone was also preserved, but in smaller pieces than the body chamber fragments. In each shell _ the septa were completely fragmented, and sheared to the edge (suture) of the external shell. No pieces of septa were recovered from the trap (mesh size approximately 5mm), indicating that the force of implosion was violent, breaking all septa and fragmenting the external shell. There is some evidence that Nautilus macromphalus in the New Caledonian region are also size segregated according to depth. Most specimens captured at depths of 100m or less are mature (2. ¢., approximated septa, black band around the apertural margin), and have a mean weight of about 650g. Size and sex distributions of N. macromphalus captured during the periods of Decem- ber-January, 1970-71 and June-July, 1972 are shown in Figure 3. All of these captures were made at depths be- tween 50 and 100m. There appear to be no significant size or sex ratio differences between the summer and winter captures. As in all other collections of Nautilus known to us, males far outnumber females, and no juveniles of less than 100g total weight were captured. Catches from the deep water trapping show a much smaller average size (P. Rancurel, personal communication, 1977) ; of a dozen specimens captured to date by ORSTOM deep water trapping (in excess of 300m) none was mature. The largest was 500 g, and most were between 200 and 400g. WILEY (1902) first indicated that Nautilus macrom- phalus in the region of New Caledonia come into shallow water at night, presumably to feed. This early, and previ- ously unsubstantiated observation has been widely quoted, and misquoted, in subsequent discussions of Recent and fossil cephalopod behavior, and has been influential in interpretation of mode of life of fossil forms (see HEPTON- STALL, 1970). Nautilus macromphalus July-August 1972, 50-100m 15 Males TECH ESE Females So = 0.61%, X,, = 7938 2 = 0.39%, X, = 673g (ia) Unsexed ™ } Number of Specimens Oo 100 300 500 700 900 Number of Specimens 100 300 500 700 goo Total Weight (g) Figure 3 Size and sex distribution of Nautilus macromphalus from Récif To, New Caledonia, during the periods July and August, 1972, and December and January, 1970-1971. All captures from 50 to 100m depth We have captured 7 specimens at night while SCUBA diving outside the barrier reef over 2 years’ time, and none has weighed less than 500 g. Two of these specimens were discovered in water between 5 and 10m deep in the shallow reef platform environments, several hundred meters offshore of the algal ridge. The other 5 Nautilus (September 1977) were captured in depths between 20 and 50m on the talus slope below the reef scarp. All were found swimming singly. According to the chief diving officer in New Caledonia, nocturnal sightings of N. macromphalus by divers are common during the Page 262 THE VELIGER Vol. 22; No. 3 austral winter months, but are never made during the austral summer months. This lack of shallow water sight- ings during the summer months may be due to elevated surface temperatures at this time. We have kept N. macromphalus in the Aquarium de Nouméa at various times of year, in varying temperatures, and have noted that seawater temperatures above 25° C adversely affect the animals. When the seawater in the aquarium reaches 27° C, the animals die. Surface temperatures outside the barrier reef at our trapping areas usually exceed 26°C during the interval of December to March, whereas depths greater than 100m have similar water tempera- tures during all times of the year (Figures 4, 5). Month Jp F M A M J e J A S O N D 27 26 25 24 23 22 21 Temperature (°C), New Caledonia Figure 4 Monthly surface temperatures from Récif To, 1973 The distribution of Nautilus pompilius in the Fiji Is- lands is somewhat different from that of N. macromphalus in New Caledonia, in that no fore-reef observations or captures of Nautilus are known in Fiji from depths less than 100m. Shallow feeding migration to near surface waters have never been observed. M AJJA o Ei ony fe 100 | 4 200 = < 300 o Q 400 March 30 -——— April 25 mers a eceeee June 28 Me 5 —— July 25 aa Ecol ian August 31 F 26 24 22 20 «18 16 14 12 10 Temperature (°C), New Caledonia Figure 5 Temperatures at various depths at Boulari Pass, New Caledonia. These temperatures were taken during 1977 outside the barrier reef system near Recif To Warp et al. (1977) trapped systematically from shal- low depths of the fore-reef slope environments to depths of 600 m in Fiji, and found that the largest concentrations of Nautilus can be captured in excess of 300m. No speci- mens were captured in less than 100m depths, which, in New Caledonia, have proved to maintain signifcant num- bers of N. macromphalus. This difference between the 2 areas can probably be attributed to temperature differ- ences. Although little temperature data are available for our sampling site in Fiji, we do have a recent depth/tem- perature survey for the month of May (Figure 6) which shows significantly higher water temperatures in the shal- lower depths of Fiji as compared to New Caledonia during the same month. These high temperatures prob- ably account for the lack of sightings of Nauéilus in shallow water in Fiji. Explanation of Figures 7A to 7D Body chamber fragments y A> Freshly imploded Nautilus pompilius Xt Phragmocone fragments. Note that septa have been evenly removed to suture by force of implosion xX4 Umbilical calyces. Septa evenly removed as in C X11 [Warp] Figures 7A to 7D THe VE LIcER, Vol. 22, No. 3 Figure 7 oryyerseth Fm ie Vol. 22; No. 3 THE VELIGER Page 263 New Caledonia ole) 200 300 Depth (m) 400 500 600 700 28 24 20 16 12 8 4 Temperature (°C) Figure 6 Temperatures at various depths during May, 1977, in New Cale- donia (Boulari Pass) and Suva Harbor, Fiji DISCUSSION The data presented above, although few in number, strongly suggest that some type of size-depth segregation takes place in these 2 Nautilus populations. In Fiji, adult animals are seemingly distributed at all depths. Smaller animals, however, are more common in depths in excess of 300m. In both New Caledonia and Fiji, the smallest individuals yet captured (less than 200g) have come from depths in excess of 400m. Several lines of evidence have recently suggested to some workers that egg-laying and early development by Nautilus takes place in near-surface waters, and possibly even within the lagoon environment. For instance, Ha- MADA & Mikami (1977) have postulated that N. macrom- phalus migrates into shallow water for egg-laying pur- poses. Eggs are thought to be laid and hatch in depths as shallow as 5 m. Evidence for this hypothesis comes from oxygen isotopic measurements (EICHLER & RISTED, 1966), indicating that early shell secretion takes place in temper- atures between 25 and 30° C; and from the only known observations of apparently newly hatched Nautilus, seen in intertidal depths in Fiji within the lagoon (Davis & Mouorter, 1972). In spite of the arguments above, we favor an alter- native hypothesis of a deeper water reproductive site (at least 100m), for the following reasons. First, it is diffi- cult to believe that juveniles hatch and spend the early parts of their lives in a reef environment at near-surface depths, in temperatures which would be lethal at least to adults. In addition to high temperature, these shallow water sites are also subjected to occasional drops of salinity, due to heavy rain, and to heavy surf and surge during most times of the year. Secondly, we, and a number of other workers, have maintained Nautilus eggs in aquaria at surface tempera- ture and pressure during several years’ time, with never a successful hatching. Even though copulation is common, and the eggs are commonly laid in the aquarium situa- tion, they are never fertile, suggesting either sperm or egg inactivation due to adverse environmental conditions, which are most probably due to unfavorable temperatures or pressure. Finally, if the Nautilus were breeding and hatching in shallow reef environments we should expect some sighting of them. In New Caledonia and Fiji numerous divers and shell collectors are perpetually searching the shallow reef for shells, yet other than the observation of Davis and Mohorter, there is no record of any juveniles from these areas, even though large populations of adults can be trapped immediately offshore in deeper water. In our collections, the smallest individuals are invariably in the_ deepest traps, suggesting to us a deeper water develop- ment and early habitat. The maximum depth from which Nautilus has been captured in both New Caledonia and Fiji is 600m. DENTON & GILPIN-BROWN (1966) raised questions about the accuracy of earlier reports of Nautilus from these depths, and suggested that the nature of the cameral liquid removal system may exclude animals in the process of forming new chambers from depths greater than 250m, since the osmotic gradients maintained between blood and fresh water could only balance a hydrostatic pressure encountered at that depth. CHAMBERLAIN (1978) postu- lated that immature Nautilus would also be found in shal- lower depths with warm water, since he concluded that cameral liquid could be removed more efficiently in these environments due to permeability properties of the si- phuncle. Based on our experience, the observational data are insupportive of these hypotheses. In both Fiji and New Caledonia a number of Nautilus has been captured showing evidence of very recent septal formation (thin septa, final chamber filled with cameral fluid of seawater Page 264 THE VELIGER Vol. 22; No. 3 osmolarity) from depths as deep as 500m, indicating that cameral fluid formation can and does occur near the maximum habitat depth of the animal. Osmotic gradients sufficient to balance hydrostatic pressure depths greater than 250m are therefore accomplished in Nautilus, indi- cating that supersaturation of salt in the siphuncular epi- thelium is occurring. In conclusion, it appears that in at least 2 populations of Nautilus, some size segregation is occurring, with the immature animals in deeper water, and further from shore. Although these deep-water (>300m) samples also contain some mature adults, they differ markedly from shallower samples which have yielded mainly ma- ture adults. This situation somewhat resembles that de- scribed by KENNEDY & CosBAN (1976) for the Western Interior Cretaceous, where young ammonites have been found in deeper, more offshore facies than adults. The reasons for these segregations may be similar in both cases. CHAMBERLAIN (1978) suggests that juvenile Nau- tilus would have a difficult time swimming in the high- turbulence of shallow reef areas. Deeper depths could be expected to be much less turbulent. Secondly, predation pressure by large carnivores, notably reef-fish, moray eels, sharks, and sea-snakes may be more pronounced in shal- low reef areas. It could be that juvenile ammonites also sought deeper, offshore areas of lower turbulence and predation pressure. ACKNOWLEDGMENTS Our research would not have been possible without the support of a large number of people. Dr. and Mrs. Catala, and later Dr. ¥. Magnier of the Aquarium de Nouméa provided aquarium facilities in New Caledonia. Temper- ature data for New Caledonia were provided by Drs. P. Rougerie and Y. Magnier of O. R. S. T.O. M. Tempera- ture data for the Fiji Islands were provided by Mr. Robert Stone. Diving assistance in New Caledonia was provided by Pierre Laboute and Pierre Djemaoun. Literature Cited CHAMBERLAIN, J. A. 1978. Permeability of the siphuncular tube of Nautilus, its ecologic and paleoecologic implications. Neus. Jahr. fiir Geol. und Paleont., Monogr. g: 129-142 Davis, R. A. « W. MoHorTER 1973- Juvenile Nautilus from the Fiji Islands. 925 - 928 Denton, E. J. «& J. B. Girpin-Brown 1966. On the buoyancy of the pearly Nautilus. Assoc. U. K. 46: 723 - 759 Eicuuer, R. & H. Ristept 1966. Isotopic evidence on the early life history of Nautilus pompilius (Linné). Science 153: 734-736 Hamapa, T. « S. MIKAMI 1977. A fundamental assumption on the habitat condition of Nautilis and its application to the rearing of N. macromphalus. Sci. papers Univ. Tokyo 27: 31 - 39 HeEpPToNSTALL, WILu1AM B. 1970. Buoyancy control in Ammonoids. Kennepy, W. « W. Cossan 1976. Aspects of ammonite biology, biogeography, and biostratigraphy. Spec. pap. in Paleontology no. 17: 93 pp. PacKarp, ANDREW Journ. Paleont. 47: Journ. mar. biol. Lethaia 3: 317 - 328 1972. | Cephalopods and fish; the limits of convergence. Biol. Rev. 47: 241 - 307 Raup, D. M. « T. TAKAHASHI 1966. Experiments on strength of cephalopod shells. Bull. Geol. Soc. Amer. (Program for 1966 Ann. Meetg.): 172-173 Saunpers, W. B. « D. A. WEHMAN 1977. Shell strength of Nautilus as a depth limiting factor. Paleo- biol. 3: 83 - 89 Warp, P, R. Stonz, G. WESTERMANN & A. MARTIN 1977- Notes on animal weight, cameral fluids, swimming speed, and color polymorphism of the cephalopod Nautilus pompilius in the Fiji Islands. Paleobiol. 3: 377 - 388 Wittey, A. 1902. Zoological results, part VI: 691 - 830. Press, London Cambridge Univ. Vol. 22; No. 3 THE VELIGER Page 265 Notoacmea gabatella (Berry), An Outer Coast Form of Notoacmea depicta (Hinds) (Mollusca : Acmaeidae) DAVID R. LINDBERG Center for Coastal Marine Studies, University of California, Santa Cruz, California 95064 and Department of Invertebrate Zoology, California Academy of Sciences, San Francisco, California 94118 (1 Plate) INTRODUCTION THIS PAPER WAS ORIGINALLY intended as a short note redescribing, illustrating, and reporting the occurrences of Recent specimens of Notoacmea gabatella (Berry, 1960), with the hope that workers, made aware of its rarity, might determine if it is extant, for the last re- ported specimens were collected in the 1930’s or 1940's. However, my plan was changed when I found a small piece of body tissue adhering to the shell of a museum specimen of N. gabatella (SDMNH 48479). The tissue contained a portion of the radula. This proved to be virtually identical to that of Notoacmea depicta (Hinds, 1842). The specimen was part of the original lot and even had been considered a paratype by some authors. There- fore, I consider N. gabatella to be an oval form of the otherwise laterally compressed N. depicta. PrtsBry in 1891 had identified a USNM specimen of this oval form as N. depicta, but most other workers have not accepted his identification. Acmaeid limpets that have several sztus forms are com- mon (cf. McLEAN, 1966; JoBE, 1968; LINDBERG, 1979). However, those occurring on marine plants and algae tend to have highly specialized shell and radular morpho- logies that limit their success on non-host substrata. Pre- viously only one species occurring on a marine angio- sperm was known to have both oval and compressed forms — Patelloida pustulata (Helbling, 1779), widely distributed in the Caribbean and in the Gulf of Mexico. The compressed form occurs on the turtle grass Thalassia testudinum Banks, 1805, while the oval form occurs on calcium carbonate substrata (e. g., coral heads, encrusting coralline algae). Several species occurring on algae also have both compressed and oval forms that correspond respectively to branching and encrusting coralline species. These species include Collisella triangularis (Carpenter, 1864) and Acmaea rosacea Carpenter, 1864. At one time Notoacmea testudinalis (Miiller, 1776) was thought to have both oval and compressed forms, the former occurring on rocks, the latter on Zostera marina Linnae- us, 1753. However, JAcKSON (1907) studied the radulae of both forms and demonstrated that the Zostera form is distinct. Although this species, Collisella alueus (Con- rad, 1831), is now separated generically as well (Mc- LEAN, 1966), some workers continue erroneously to list it as a form of N. testudinalis (e. g., ABBOTT, 1974). Dati (1871) was one of the first workers to recognize and illustrate the species-specific nature of the acmaeid radula. GRANT (1937) furthered this work when she studied and illustrated the radular ribbons from over 50 acmaeid species. FRIrcHMAN (1960) used radular char- acters to distinguish his Acmaea paradigitalis [= Colli- sella strigatella (Carpenter, 1857) ] from both C. digitalis (Rathke, 1833) and C. pelta (Rathke, 1833). McLean (1966; zn KEEN, 1971; 7n MarINcovicH, 1973) studied the radulae of almost all of the eastern Pacific species and found no species that shared identical radular mor- phologies or had variations that overlapped or inter- graded with another species. LinDBERG (1979) has used radular characters to identify 5 distinct sztus forms of C. ochracea (Dall, 1871). Thus, the acmaeid radula is a powerful tool for resolving taxonomic difficulties en- countered in the convergent and highly variable shell morphologies found in acmaeid limpets. Page 266 ACKNOWLEDGMENTS I am indebted to James T. Carlton, Woods Hole Ocean- ographic Institution, Woods Hole, Massachusetts, who suggested the original topic and provided suggestions and criticisms as the topic evolved. George L. Kennedy, U.S. Geological Survey, Menlo Park, California, sup- plied a list of the invertebrate fauna of LACMIP locality 332, and provided substantial input in this paper, especi- ally the paleoecology section, for which I am most grate- ful. Michael G. Kellogg, Moss Landing Marine Labora- tories, Moss Landing, California and James H. McLean, Natural History Museum of Los Angeles County, Los Angeles, California, also provided criticisms of the manu- script. Hans Bertsch, San Diego Muesum of Natural His- tory, San Diego, California, Barry Roth, California Acad- emy of Sciences, San Francisco, and Edward C. Wilson, also of the Natural History Museum of Los Angeles County, loaned specimens from their respective institu- tions. ABBREVIATIONS USED The following abbreviations are used herein: ANSP — Department of Mollusks, Academy of Natural Sciences, Philadelphia, Pennsylvania CASG — Department of Geology, California Academy of Sciences, San Francisco, California LACMIP - Section of Invertebrate Paleontology, Nat- ural History Museum of Los Angeles County, Los Angeles, California MCZ — Museum of Comparative Zoology, Harvard Uni- versity, Cambridge, Massachusetts SU ~ Department of Geology, Stanford University, Stanford, California (now on permanent loan to CASG) SDMNH - Department of Marine Invertebrates, San Diego Museum of Natural. History, San Diego, California UCMP - Department of Paleontology, University of California, Berkeley, California USNM -— Division of Mollusks, U.S. National Museum of Natural History, Washington, D. C. HISTORICAL REVIEW The first known oval specimens of Notoacmea depicta (“Acmaea” gabatella) were both collected live at San THE VELIGER Vol. 22; No. 3 Diego, California and were catalogued into the USNM collection in 1885 (ex R.E.C. Stearns Collection). In the first report on the specimens Prspry (1891) illus- trated the larger specimen, identifying it as an oval form of N. depicta. Pilsbry’s treatment of the specimen may have been influenced by remarks concerning N. depicta made by Dat (1871) who predicted, “It is quite likely that thorough dredging would result in procuring non- compressed specimens, which might have grown on peb- bles, & c., ...” Pilsbry’s conclusion clearly confirms this prediction. Two more specimens collected from “kelp leaves, stems” at San Pedro, California, were originally identified as “young Acmaea scabra.” This is undoubtedly intended as Acmaea scabra (Reeve, 1855), now considered a syno- nym of Collisella limatula (Carpenter, 1864) ; the name was rejected by Dati (1914) as a secondary homonym. The specimens were probably collected in the early 1900’s and identified prior to Dall’s publication. Avery Ransome Grant, working on her doctoral disser- tation, considered these oval specimens in the USNM col- lection distinct from Notoacmea depicta and not forms of it as suggested by Pmsspry (1891). However, this is evident only by a note left with one of the specimens, “not Acmaea depicta probably not an Acmaea (teste A. R. Grant)”; no mention of the specimens appeared in GrantT’s thesis (1937) or subsequent publications (TeEsrT, 1945; 1946). Another (undated) note left with the speci- men reads, “this may be a Helcion E.PC.” (E.PC. = Emery Perkins Chace of Lomita, California) . The reason- ing for this is not clear to me. Helcion deMontfort, 1810, is a patellid genus characterized by a submarginal apex, a character which the specimen clearly lacks. A series of specimens (at least 7) was dredged by A. M. Strong off Reef Point and Laguna Beach, California. The exact date(s) is(are) unknown, but was(were) probably between 1930 and 1945. After Strong’s death his collection went to SDMNH where the specimens were examined and subsequently described by S. Still- man Berry as “Acmaea” gabatella. In 1963, Roger D. Reimer and George P. Kanakoff col- lected unconsolidated fossiliferous sediments from a cut bank in the 600 block of Miraflores Street, San Pedro, California (LACMIP loc. 332). From this material over 100 specimens of the oval form were recovered making it one of the more abundant “species” in the Pleistocene fauna. McLEAN (1966) considered “Notoacmea gabatella” as part of his doctoral dissertation. McLean was the first worker to recognize the Recent specimens from San Pedro Vol. 22; No. 3 THE VELIGER Page 267 as “N. gabatella” and illustrated Berry’s type material for the first time. He was also the first worker to recognize the Pleistocene specimens as “N. gabatella.” Witson « KENNEDY (1967)listed all of the Strong specimens of “Notoacmea gabatella” at SDMNH as type material and gave the same locality, Reef Point, for all 7 specimens. However, only 2 specimens are from Reef Point and only these were designated as type material by Berry (1960). The remaining 5 specimens from Laguna Beach are not types. AsgotTt (1974) suggested that “Acmaea gabatella”’ may be an aberrant form of Notoacmea depicta. CARLTON (1976) and LinpBerc (1976) both considered “Notoac- mea gabatella” in papers treating the marine plant lim- pets of the northeastern Pacific and it was this work that sparked our interest in this species. SYSTEMATICS Notoacmea depicta (Hinds, 1842) Patelloida depicta Hinps, 1842: 82; plt. 6, fig. 4; - Hips, 1845: 53 — Keen, 1966: 268 Nacella depicta (Hinds). CARPENTER, 1857: 318 — CarPENn- TER, 1864: 650 Acmaea (Collisella) depicta (Hinds). Datu, 1871: 254 — Datu, 1921: 170 — OtpRoyD, 1927: 759 — ABBOTT, 1954: 104 — ABBOTT, 1974: 32; fig. 162 Acmaea depicta (Hinds). Pmssry, 1891: 19; plt. 6, fig. 40 — ARNOLD, 1903: 317 — GRANT & GALE, 1931: 812 —- GRANT, 1933: 59 — KEEN, 1937: 28 - SmitH « Gor- DON, 1948: 199 Acmaea (Acmaea) depicta (Hinds). Grant, 1937: 753 figs. 293, 294, 316, 317, 332,333 — BurcH, 1946: 7 - FritcH- MAN, 1961: 58 Collisella (Notoacmea) depicta (Hinds). McLean, 1966: 112; pit. 3, figs. 20, 21 Notoacmea depicta (Hinds). McLzan, 1969: 19; fig. 6.4 — McLean, 1978: 19; fig. 6.4 - CARLTON, 1976: 22 — LINDBERG, 1976: 26 Acmaea depicta var. Pitspry, 1891: 20, 174; plt. 6, fig. 41 “Acmaea” gabatella Berry, 1960: 118 — Witson & KENNEDY, 1967: 245 Collisella (Notoacmea) gabatella (Berry). McLean, 1966: 124; plt. 3, figs. 4, 5 Notoacmea gabatella (Berry). CARLTON, 1976: 22 - Linp- BERG, 1976: 26 Diagnosis: There are 2 situs forms of this species, a Zostera form and an oval form. A diagnosis for each fol- lows: Zostera form (Figure 1) — Shell thin, small (less than 15 mm in length) ; profile medium; apex posi- tioned in anterior third of shell, anteriorly directed. Ante- rior slope straight toslightly concave; posterior and lateral slopes convex. Aperture laterally compressed, elongate. Sculpture of concentric growth lines. Color yellow-white to yellow-brown with darker red-brown chevron mark- ings that radiate outward from the apex and midline of the shell; interior margin slightly darkened, intermediate area lighter. Myostracum vague; central area yellow; anterior slope darkly stained with brown. Shell diapha- nous; exterior markings clearly visible through shell. Oval form (Figure 2) — Shell thin, small (less than 1omm in length) ; profile low; apex positioned in ante- rior third of shell, anteriorly directed, strongly protrud- ing from anterior slope. Anterior slope straight to slight- ly concave; posterior and lateral slopes distinctly con- vex. Aperture ovoid, slightly narrowed anteriorly. Sculp- ture of concentric growth lines. Color yellow-brown with darker red-brown chevron markings that radiate out- ward from the apex and midline of the shell; interior margin slightly darkened, intermediate area lighter. My- ostracum vague; central area with yellow cast. Shell dia- phanous; exterior markings clearly visible through shell. Radula (Figures 3 and 4): First lateral teeth closely set on anterior edge of ventral plates, slightly overlapping anterior ventral plates. Medial edges of first lateral teeth convex, lateral edges sigmoidal; cusps pointed. Second and third lateral teeth positioned obliquely across ventral plates. Second and third lateral teeth twice as broad as first lateral teeth. Medial edges of second lateral teeth slightly convex; cutting edges straight; lateral edge con- cave, forming small laterally directed cusp. Medial edges of third lateral teeth strongly convex, lateral edge also convex; cutting edge straight; cusp gently rounded. Mar- ginal teeth lacking. Ventral plates squarish with strong anterior process. Ventral plates strongly sutured in vicin- ity of anterior process and along lateral edge. Small posterior process at medial edge of ribbon units. Remarks: Although both forms are similar in size, sculp- ture, and color patterns, the Zostera form is laterally com- pressed, having an elongate aperture and parallel lateral margins that conform in width to its host plant, the eelgrass Zostera marina, whereas the oval form has a broadly oval aperture and shows no trace of lateral com- pression. The oval form is proportionately lower than the Zostera form and lacks the dark brown anterior slope stain found in the Zostera form. In some fossil specimens of the oval form a dark interior stain is present, but it is posterior to the apex rather than anterior, as it is in the Zostera form. McLean (1966) pointed out that larger fossil oval forms show a tendency to have raised lateral margins, a character also present in some modern oval specimens, but not found in the Zostera form. The radulae of oval and Zostera forms show no significant differences Page 268 THE VELIGER Vol. 22; No. 3 and are well within the range of intraspecific variation seen in other species that have several setus forms (e. g., Collisella ochracea; C. pelta; C. triangularis; Patelloida pustulata).'The only radular differences are in the cusps of the second lateral teeth and the lengths of the ribbon segments. In the Zostera form the second lateral tooth cusp is distinctly pointed; in the oval form it is rounded. Also, in the Zostera form the ribbon segments are com- pressed, in the oval form they appear typical of the genus. The remaining radular teeth profiles, arrangement of the teeth, and basal plate morphologies are identical in both forms. Because of the rarity of modern specimens of the oval form, the entire radula was not prepared. Instead, the radula was cut in half and one section preserved with the rehydrated tissues. The shell, tissue, and radula are deposited in the SDMNH alcoholic collection No. 48479. Notoacmea lepisma (Berry, 1940), an extinct middle Pleistocene species from San Pedro, California, has the same shell markings as N. depicta, first noted by Mc- Lean (1966). Notoacmea lepisma, like N. depicta is laterally compressed, with parallel lateral margins. How- ever, the profile is distinctly lower and the apex more anterior than in N. depicta. According to McLEAN (op. cit.) N. lepisma is intermediate in shell morphology be- tween N. depicta and “Notoacmea gabatella.” 'The range of variability of each “species” does not overlap, nor are there any intermediates. My own observations confirm this. If N. depicta and “N. gabatella” represent extremes of the same species, as I suspect, then N. lepisma is also probably a form of N. depicta. However, synonymizing N. lepisma with N. depicta will depend on the results of a morphometric study using coordinate transformation (LINDBERG, in preparation). Recent Distribution: Zostera form — California: Mon- terey County, Monterey (36°36’N) (ANSP 39142; MCZ unnumbered lot; UCMP 2395); and Santa Barbara County, Mugu Lagoon (34°06’N) (SDMNH 41434) to Mexico: Baja California, Cabo San Lucas (23°N) (CA SG 17663A). Oval form — California: Los Angeles Coun- ty, San Pedro (33°45’N) (SU 5478) to Orange County, Laguna Beach (33°33’N) (SDMNH 48479); and San Diego County, San Diego (32°40’N) (USNM 326554). The specimen lots of the Zostera form from Monterey, California, were all collected in the nineteenth century, and represent 3 different collectors and dates, evidence that suggests that these are valid records and not mixed specimen lots nor have erroneous locality data. Warm wa- ter years in northern California (1853 to 1860) (Husss, 1948) may have allowed for the establishment of Notoac- mea depicta in Monterey Bay in the second half of the nineteenth century. It has not been reported since (cf. SMITH & GorDON, 1948). Fossil Distribution: (Pleistocene) Zostera form — Cali- fornia: Los Angeles County, San Pedro, Lomita Marl, (Burcu, 1946), San Pedro Sand (LACMIP Loc. 332) ; Palos Verdes, Palos Verdes Sand (ARNOLD, 1903); Orange County, Costa Mesa, Terrace cover and Palos Verdes Sand (2?) (KaNAKoFF & EMERSON, 1959) ; San Diego County, La Jolla (BurcH, 1946); Mexico: Baja California, La Playa (BurcH, 1946); Ensenada (Burcu, 1946). — O- val form — California: Los Angeles County, San Pedro, San Pedro Sand (LACMIP Loc. 332). Ecology: The Zostera form occurs on the blades of the eelgrass Zostera marina (CARLTON, 1976), not on the surfgrass Phyllospadix species as reported by McLEan (1966). Zostera marina occurs discontinuously through- out the boreal northern hemisphere from the Seas of Okhotsk and Japan to the Baltic and Mediterranean, and is restricted to shallow water, soft bottom habitats (2. é., lagoons and estuaries) (SETCHELL, 1935). The oval form of Notoacmea depicta occurs on the outer coast (Table 1). Living specimens have been dredged at a depth of 18m off Reef Point, and at Laguna Beach. The San Pedro specimens were collected from “kelp leaves, stems,” possibly Macrocystis pyrifera (Linnaeus) Explanation of Figures z and 2 Notoacmea depicta (Hinds, 1842) Figure 1: Zostera form (CASG No. 32762) Shell length - 10.8 mm Figure 3 Notoacmea depicta (Hinds, 1842) Radular ribbon segment of Zostera form (CASG No. 49207) Figure 2: Oval form (SU No. 5478) Shell length - 7.65 mm Figure 4 Notoacmea depicta (Hinds, 1842) Radular ribbon segment of oval form (SDMNH No. 48479) THE VELIcER, Vol. 22, No. 3 [LinpBERG] Figures z and 2 Zostera form ( 7 7 Se a Figure 7 Oval form Figure 2 Lateral Teeth Profiles Zostera form Figure 3 Ae & Oval form Figure 4 = Vol. 22; No. 3 THE VELIGER Page 269 Table 1 Records of Oval forms of Notoacmea depicta Locality (Collector) Habitat Authority. Depository (No. of specimens) CALIFORNIA: Orange County, off Reef Point (A.M. Strong) CALIFORNIA: Orange County, off Laguna Beach (A.M. Strong) CALIFORNIA: Los Angeles County, San Pedro (Unknown) CALIFORNIA: Los Angeles County, San Pedro (Reimer and Kanakoff) CALIFORNIA: San Diego County, San Diego, Ballast Point [east side of Point Loma] (Brannan) CALIFORNIA: San Diego County, San Diego (H. Hemphill) Dredged kelp leaves, stems Pleistocene Dredged 18m Berry, 1960. SDMNH 50945. Holotype (Type No. 1301) and Paratype (Type No. 1302) of “Acmaea” gabatella SDMNH 48479 (5 specimens); (Type Nos. 1303-1307) by Wirson & KEennepy (1967) McLean, 1966. SU 5478 (2 specimens) McLean, 1966. LACMIP loc. 332 (approximately 100 specimens) USNM 32547 (1 specimen) Pirspry, 1891. USNM 32654 (1 specimen) Agardh, 1830, or any of several other large brown algae. In view of the vast amount of research on M. pyrifera that has taken place in southern California, it seems un- likely that the limpet would have been overlooked. No habitat data are available for the San Diego specimens. In view of the cool water fauna of the San Pedro Sand in San Pedro (LACMIP Loc. 332; see below) it is possible that the oval form could occur north of San Pedro today. However, kelp forest studies by McLEan (1962) and Pearse & Lowry (1974) have not reported the form in the Monterey Bay area. Paleoecology: Although modern specimens of the oval form of Notoacmea depicta are few, over 100 specimens are known from a locality (LACMIP Loc. 332) in the San Pedro Sand of San Pedro, Los Angeles County, Cali- fornia. The exposure was a cut bank in the 600 block of Miraflores Street which has subsequently been covered by the southern extension of the Harbor Freeway into San Pedro. Unfortunately, no field notes are available on this locality that might describe the stratigraphic section or indicate whether one or several shell horizons were collec- ted. It is not now possible to tell if the bathymetric dis- crepancies indicated by the molluscan fauna at this local- ity are a product of downslope reworking of detrital shells or a result of collecting through a stratigraphic section in which shoaling might have been involved. The only published discussion on the paleoecology of this locality is that by FrrcH (1967). He reported 38 spe- cies of fish, mainly as otoliths. Several additional fish taxa are also listed by LANGENWATER (in KENNEDY, 1975). Fitch’s study indicated a “cold water” fauna similar to one that would occur off central California today. Analysis of the bathymetric ranges for each species (bathymetric data from Miner & Lea, 1972) gives the following results. Seventy-nine % (30 species) occur in depths less than 15 m, 87% (33 species) occur between 15m and 50m depths, and 90% (35 species) occur between 50m and 70 m depths. Below 70m only 72% (27 species) remain. This suggests a depositional depth between 30m and 70 m, the lesser depth suggested by the upper limits of several deep water species. The habitats represented by the fish are diverse and include epipelagic (sharks), sand or mud bottom (flounders and rays), off shore (herring and an- chovy), and rocky bottom (rock fish, wrasse, and scul- pins). A preliminary list of mollusks from LACMIP Loc. 332 (G. L. Kennedy, in litt., 1978) was used to infer the bathymetric, habitat, and thermal conditions of the mol- luscan fauna. The fauna, taken in aggregate, indicates depth ranges from intertidal to over 500m. However, both intertidal and deep water species are few in number and are represented by few specimens. The major part of the mollusks occur today at depths between 15m and 70m. Woonrinc et al. (1946), using molluscan data from other localities, suggested that the San Pedro Sand repre- sented shallow water deposits at some localities and moderate depths (50m to 90m) elsewhere. VALENTINE (1961) considered the San Pedro Sand to represent depths between 18m and 90m. LACMIP Loc. 332 ap- pears to contain both shallow water and moderate depth species. Page 270 THE VELIGER Vol. 22; No. 3 The molluscan component at LACMIP Loc. 332 is not distinctly cold or warm water. Instead, it includes several cryophilic (7. e., northern) and thermophilic (7. e., south- ern) members. The majority of the species present have modern distributions that include San Pedro. Fircu (1967), like Wooprine et al. (1946), felt the fauna in- dicated a cooler water environment. VALENTINE (1961) stated that the faunas representing depths greater than 27m suggested cooler waters than those in the area today. Habitat data are by far the most confusing. Several bio- topes are represented in the molluscan component in- cluding shallow water embayment, protected rocky coast, kelp forest, and both sand and mud bottoms. Of the 179 species (Kennedy, zn litt., 1978) from LACMIP Loc. 332, the oval form of Notoacmea depicta ranks 23" in abundance (approximately 100 specimens). The Zostera form of N. depicta is represented by only 2 specimens. Other abundant species and their habitats are given in Table 2. The paleoecologicai setting of LACMIP Loc. 332 suggests several possible habitats for the oval form of N. depicta, ranging from shallow embayment to rocky sub- tidal. However, the modern occurrences clearly support the latter, particularly a kelp forest habitat. WEHMILLER et al. (1977) have estimated the age of the San Pedro Sand, based on racemization of leucine in Protothaca shells from LACMIP Loc. 332, at 320000 to 410000 + 80000 BP, thus placing it within the middle Pleistocene as currently defined. SUMMARY Notoacmea gabatella (Berry) is synonymized with N. depicta (Hinds). This action is based on the identical radular morphologies found in both “species.” Notoacmea depicta is considered to be a polytypic species with 2 forms — a laterally compressed form on the eel grass Zostera marina, and an oval form thought to occur on or near large brown algae. The oval form is extremely rare in the Recent fauna, being represented by fewer than ro specimens. In the middle Pleistocene of San Pedro, California, it is represented by more than 100 specimens at a single locality (LACMIP Loc. 332). The paleoecology of LACMIP Loc. 332 is consistent with the Recent habitat and suggests depositional depths between 30m and 7om. Table 2 Abundant (500 specimens) Mollusks from LACMIP loc. 332 Species No. of specimens Depth Gastropoda Lirularia optabilis (Carpenter, 1864) 968 Turritella cooperi Carpenter, 1864 1500 >20 m Bittium armillatum Carpenter, 1864 1309== Bittitum munitum (Carpenter, 1864) 1850+ 24m Bitttum rugatum Carpenter, 1864 > 1000 60-80 m Mitrella tuberosa (Carpenter, 1864) 2070 subtidal Alia carinata (Hinds, 1844) = 500 low intertidal to subtidal Bivalvia Acila castrensis (Hinds, 1843) 600 16-500 m Mysella aleutica (Dall, 1899) 1500 20-130 m Lucinisca nuttalli (Conrad, 1837) Transennella tantilla (Gould, 1853) = 2000 to 36 m Known only from Pleistocene assemblages Probably restricted to Pleistocene many hundreds low intertidal to subtidal low intertidal Habitat Authority McLean, 1964 McLean, 1978 Woonrinc et al, 1946; teste McLean, 1979 fine sand, shale fragments SmitH & Gorpon, 1948 2 CARPENTER, 1866 not uncommon in gravel McLean, 1978 under kelp near surf grass and algae sand, mud bottoms McLEAn, 1978 ScHENCK, 1936; Asport, 1974 Assortt, 1974; SmityH & Gorpon, 1948 McLean, 1978 sandy mud fine sand, shale fragments sand flats—off shore sand, sandy mud semi- HERTLEIN & GRANT, 1972; protected bays and off shore Coan & Cartron, 1974 Vol. 22; No. 3 Literature Cited Assotr, Ropert Tucxz 1954. American seashells. 40 plts.; 100 text figs. 1974. American seashells. Nostrand Reinhold, New York Arno.p, RarPH 1903. The paleontology and stratigraphy of the marine Pliocene and Pleistocene of San Pedro, California. Calif. Acad. Sci. Mem. g: 420 pp.; 37 pits. (27 June 1903) [reprinted as: Stanford Univ., Contributions to Biology from the Hopkins Seaside Laboratory no. 31, same date and pagination] Berry, SAMUEL STILLMAN 1960. Notices of new eastern Pacific Mollusca. — IV. in Malacol. 1 (19): 115-122 Burcu, Jonn Quincy (ed.) 1944-1946. Distributional list of west American marine mollusks from San Diego, California to the Polar Sea. Minutes Conch. Club So. Calif. nos. 33-63 (pagination by issue); 3 plts. (mimeographed) Carxton, James THEODORE 1976. Marine plant limpets of the northeastern Pacific: Patterns of host. utilization and comparative plant-limpet distributions. Ann. Reprt. West. Soc. Malacol. 9: 22 - 25 (12 October 1976) CarRPENTER, PHILIP PEARSALL 1857. Report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Brit. Assoc. Adv. Sci., Reprt. 26 (for 1856): 159-368+4 pp.; pits. 6-9 (pre-22 April 1857) 1864. Supplementary report on the present state of our knowledge with regard to the Mollusca of the west coast of North America. Brit. Assoc. Adv. Sci., Rprt. 33 (for 1863): 517-686 (post 1 Aug.) [reprinted in CarPeNTER, 1872 (A): 1 - 172] [dating, CARPENTER, 1872] 1866. On the Pleistocene fossils collected by Col. E. Jewett, at Sta. Barbara, California; with descriptions of new species. Ann. Mag. Nat. Hist. (3) 17: 274-278 (April 1866) Coan, Eucene Vicror & James THEOpoRE CARLTON 1975- Phylum Mollusca: Bivalvia. In: Ralph I. Smith « James T. Carlton (eds.), Light’s Manual: Intertidal invertebrates of the central California coast. pp. 543-578; Univ. Calif. Press, Berkeley, California DALL, WILLIAM HEALEY 1871. On the limpets; with special reference to the species of the west coast of America, and to a more natural classification of the group. Amer. Journ. Conchol. 6 (3): 227-282; plts. 14-17 (4 April 1871) 1914. Notes on some northwest coast acmaeas. The Nautilus 28 (2): 13-15 (13 June 1914) 1921. Summary of the marine shellbearing mollusks of the northwest coast of America, from San Diego, California, to the Polar Sea, mostly contained in the collection of the United States National Museum, with illustrations of hitherto unfigured species. U.S. Natl. Mus. Bull. 112: 1 - 217; 22 pits. (24 February 1921) Frreu, Joun E. 1967. The marine fish fauna, based primarily on otoliths of a lower Pleistocene deposit at San Pedro, California (LACMIP 332, San Pedro Sand). Los Angeles County Mus. Nat. Hist. Contrib. Sci. 128: 23 Pp.; 31 text figs. (31 May 1967) Frircuman, Harry Kr, III 1960. Acmaea paradigitalis sp. nov. (Acmaeidae, Gastropoda). The Veliger 2 (3): 53-573 plts. 9-12 (1 January 1960) 1961. _—A study of the reproductive cycle in the California Acmaeidae (Gastropoda). Part I. The Veliger 3 (3): 57-63; plt. 10 (1 January 1961) Van Nostrand Co. New York, 541 pp. aed ed. 663 pp.; 24 col. pits. Van Leaflets (31 December 1960) Grant, Avery RANSOME 1933. A revision of the California limpets of the genus Acmaea Esch- scholtz. M. A. Thesis, Univ. Calif, Berkeley, Calif. 142 pp.; 7 plts. 1937. A systematic revision of the genus Acmaea Eschscholtz, in- cluding consideration of ecology and speciation. Ph. D. Thesis, Univ. Calif, Berkeley, Calif 432 pp.; 35 pits. Grant, Utysszs Simpson, IV « Hoyt Ropnry GALE 1931. Catalogue of the marine Pliocene and Pleistocene Mollusca of California and adjacent regions ... . San Diego Soc. Nat. Hist., Mem. I: 1-1036; 32 plits.; 15 text figs. (3 November 1931) Hartizin, Lzo Gzorcz # Utysses Simpson Grant, IV 1972. The geology and paleontology of the marine Pliocene of San Diego, California. Part 2B: Paleontology: Pelecypoda. San Diego Soc. Nat. Hist., Mem:..2: 135 - 409; frontispiece; plts. 27-57; text figs. 7-13 (21 July 1972) THE VELIGER Page 271 Hinps, RicHarp Brins.ey 1842. Descriptions of new shells. 1 pit. Husss, Cart L. 1948. Changes in the fish fauna of western North America correlated with changes in ocean temperature. Journ. Mar. Res. 7 (3): 459-482; 4 text figs. Jackson, Henry, Jr. 1907. The differences between the two New England species of Ac- maea. The Nautilus 21 (1): 1-5; plt. 2; 2 text figs. (13 May ’o7) Jose, ALAN 1968. A study of morphological variation in the limpet Acmaea pelta. The Veliger 11 (Supplement): 69 - 72; plt. 6; 1 text fig. (15 July 1968) Kanaxorr, Gzoror P. & Wituiam Keira EMERSON 1959. Late Pleistocene invertebrates of the Newport Bay area, Cali- fornia. Los Angeles County Mus. Contrib. Sci. no. $1: 1-47; 5 text figs. (14 October 1959) Keen, A. Myra 1937. An abridged check list and bibliography of west North Ameri- can marine Mollusca. Stanford Univ. Press, Stanford, Calif., 84 pp.; g text figs. (29 September1937) 1966. West American mollusk types in the British Museum (Natural History). II. Species described by R. B. Hinds. The Veliger 8 (4): 265 - 275; plts. 46, 47; 6 text figs. (1 April 1966) LANGENWALTER, Paut E., II 1975. The fossil vertebrates of the Los Angeles-Long Beach Harbors region. In: George L. Kennedy, Paleontological record of areas adjacent to the Los Angeles and Long Beach Harbors, Los Angeles, California. Marine studies of San Pedro Bay, California, Part 9, Paleontology. 36-54. Allan Hancock Foundation, Harbors Environ- mental projects and the Office of Sea Grant Programs, Univ. So. Calif, USC-SG-4-75 (10 June 1975) LinpzBerc, Davp R. 1976. Marine plant limpets of the northern Pacific: Neogene phylo- geny and zoogeography. Ann. Reprt. West. Soc. Malacol. 9: 26 (12 October 1976) 1979. Variation of Collisella ochracea (Dall) and northeastern Pa- cific distribution of Notoacmea testudinalis (Miller) (Gastropoda). The Nautilus 93: (2): 50-56; 10 text figs. (23 April 1979) McLzgan, James HamitTon 1962. Sublittoral ecology of kelp beds of the open coast areas near Carmel, California. Biol. Bull. 122: 95-114; 2 text figs. 1964. | West American species of Lirularia. Ann. Reprt. Amer. Malac. Union 1964: 40-41 1966. | West American prosobranch Gastropoda: superfamilies Patella- cea, Pleurotomariacea, and Fissurellacea. Ph. D. dissert., Stan- ford Univ.; x+262 pp.; 7 plts. 1969. Marine shells of southern California. Los Angeles Cty. Mus. Nat. Hist. Sci. Ser. 24, Zool., no. 11: 104 pp.; 54 text figs. (Oct. ’69) 1971. Family Acmaeidae. In: A. Myra Keen, Sea shells of tropical West America: 322-329. Stanford Univ. Press 1973. Family Acmaeidae. In: L. Marincovich, Jr. Intertidal mol- lusks of Iquique, Chile. Nat. Hist. Mus. Los Angeles County, Sci. Bull. 16: 18-23 1978. Marine shells of southern California. Nat. Hist. Mus. Los Angeles County, Sci. Ser. 24 (Rev. Edit.): 1-104; 54 plts. (20 March 1978) Ann. Mag. Nat. Hist. ro: 81 - 84; Oxproyp, Ipa SHEPARD 1927. The Marine Shells of the West Coast of North America. Stanford Univ. Press, Stanford, Calif. 2 (3): 1-339; [605 - 941] pits. 73 - 108 PgarsE, JonN S. & Ltoyp F Lowry 1974. An annotated species list of the benthic algae and invertebrates in the kelp forest community at Point Cabrillo, Pacific Grove, Califor- nia. Coast. Mar. Lab., Univ. Calif, Santa Cruz Tech. Reprt.. I: 1-73; 3 text figs. Pitsspry, HENry AuGusTus 1891. Acmaeidae, Lepetidae, Patellidae, Titisoaniidae. Man. Conch. 13: 1-195; 74 plts. Scuencx, Husert G. 1936. Nuculid bivalves of the genus Acila. Geol. Soc. Amer. Spec. Paper 4: xiv+149 pp.; 18 plts.; 15 text figs. (18 July 1936) SeTcHELL, WILLIAM A. : : 1935. Geographic elements of the marine flora of the North Pacific Ocean. Amer. Natural. 69 (725): 560-577 Test, Avery Ransome GRANT 1945: Ecology of California Acmaea. Ecology 26: 395-405 1946. Speciation in limpets of the genus Acmaea. Univ. Michig. Contr. Lab. Vert. Biol. $1: 1-24 Page 272 THE VELIGER Vol. 22; No. 3 VALENTINE, JAMEs WILLIAM 1961. Paleoecologic molluscan geography of the Californian Pleisto- cene. Univ. Calif: Publ. Geol. Sci. 94 (7): 309-442; 16 text figs. (17 May 1961) WEHRMILLER, JOHN F, Kennetu R. Lajor, Keirn A. KvENvOLDEN, Etta Peterson, Daniet F Berxnap, Grorce L. KENNEDY, WARREN O. AppicoTT, Joun G. VeEppER « RoBert W. WRIGHT 1977. Correlation and chronology of Pacific coast marine terrace deposits of continental United States by fossil amino-acid stereochem- istry-technique evaluation, relative ages, and geological implications. U. S. Geol. Surv. Open-File Reprt. 77-680: 1-191; 18 text figs. Wruson, Epwarp CG. « Grorcz L. KENNEDY 1967. Type specimens of Recent invertebrates (except Arachnida and Insecta) in the San Diego Natural History Museum. Trans. San Diego Soc. Nat. Hist. 14 (19): 237-279 (17 November 1967) Wooprine, WENDELL PHILiirs, MILTON NUNN BRAMLETTE & Wiiuiam STEPHEN WEBSTER KEw 1946. Geology and paleontology of Palos Verdes Hills, California. U. S. Geol. Surv., Prof. Paper 907: 145 pp.; 37 plts.; 16 text figs. _Vol..22; No. 3 THE VELIGER Page 273 Reinstatement of Two Species of Murexiella (Gastropoda : Muricidae) from the Tropical Eastern Pacific BY LEROY H. POORMAN'! Museum Associate, Los Angeles County Museum of Natural History (1 Plate; 1 Text figure) IN THEIR RECENTLY PUBLISHED book on the Muricidae, _ Rapwin « D’Atriuio (1976) synonymized Murexiella keenae and M. laurae, both described by VoxKEs, 1970, with M. humilis (Broderip, 1833). Specimens collected by dredging along the west Mexican coast over the past 24 years suggested that the 3 closely related species recog- nized by Vokes are valid. In order to base my conclusions on as many specimens as possible, material from 7 major collections was exam- ined. The material was identified as follows: Murexiella humilis, 23 lots, 60 specimens; M. keenae, 15 lots, 52 specimens; M. laurae, 8 lots, 45 specimens. The following characteristics were considered signifi- cant: general outline of the shell, form of the nucleus, num- ber of varices, form of the spiral cords, microscopic spiral threads, shape of spines on the shoulder and canal, num- ber of spiral cords and spines on the canal. No difficulty was encountered in separating the mate- rial, including several mixed lots, into 3 distinct and homogeneous groups. Immature shells (under 4mm) of the 3 species, which more nearly resemble each other, were also easily separated. The 3 groups were compared with the holotypes for Murexiella keenae and M. laurae and with the original description and figure for M. humi- lis. The geographic distribution of each species is shown in Figure 7. Specimens from Manzanillo, Colima, Mexico, were se- lected for comparison. They included a significant number of each species and came from the same area of several square kilometers. All 3 species were dredged from the same substrate of coarse gravel and small rocks in 20 - 30 meters off a rocky headland. "Mailing address: 15300 Magnolia Street, Space 55, Westminster, California 92683 Murexiella humilis (Broderip, 1833) (Figures 1, 4) Murex humilis Broderip, 1833; Sowersy, 1834: plt. 65, figs. 46, 47 Murex norristi Reeve, August 1845: plt. 28, sp. 129A-B Murexiella humilis. VoxEs, 1970: plt. 50, figs. 1-3; KEEN, 1971: fig. 988; Rapwiw « D’AtTILI0, 1976: plt. 25, fig. 4 Murex humilis Broderip, 1833, was described with Santa Elena, Ecuador, as type locality. The species was first figured by SowrErBy (1834). The description and figure are of a shell with a globose body whorl, a moder- ately elevated spire of about 5 whorls, and a long, nearly straight canal. Varices are fimbriated along the leading edges and are intersected by spiral cords. The intersections give rise to spines that terminate sharply and are spirally recurved. Below the body whorl, on the upper part of the canal, is a wide band where the axial and spiral sculpture is greatly subdued. Anteriorly on the canal are 3 rows of sharp recurved spines that rise from the terminations of spiral cords at the leading edges of obsolete canals. Based on the large number of specimens examined, further details may be added. The nucleus is conical, of 3 globose whorls, the first small, the other 2 increasing regularly. The tip is immersed. The entire nucleus is covered with microscopic granules. The first postnuclear whorl has 8 low axial ribs that extend over the third nuclear whorl and are attached to it for the lower 4 of its height. The varices are formed by the terminations of successive growth stages. In mature specimens, the thin outer layer of each growth stage is reinforced with successive layers of shell material deposited on the inner surface. The layers are slightly separated along the terminal edge. Page 274 Head of Gulf of California Consag Rock, Gulf of California Cabo Tepoca, Sonora Bahia de Los Angeles, Baja California Norte Guaymas, Sonora Rio Sinaloa, Sinaloa Mazatlan, Sinaloa Teacapan, Sinaloa Cuastecomate, Jalisco Manzanillo, Colima Zihuantenejo, Guerrero White Friars, Mexico Salina Cruz, Oaxaca Puntarenas, Costa Rica Venado Island, Panama Galapagos Islands, Ecuador Santa Elena, Ecuador TL - Type Locality Figure 7 * * ~—— THE VELIGER Vol. 22; No. 3 Each growth stage is begun below the last layer of the preceding one. Thus, the leading edge of each varix is exposed as an expanded frilly cross section of the shell wall. Although all specimens examined began postnucle- ar growth with 8 varices per whorl, some increased the number with growth and some decreased. The range for the number of varices on the adult body whorl was 6 to 9, with 56% of the specimens having 8. There are 5 major cords on the body whorl, 2 on the spire, and 3 anteriorly on the canal. Each major cord is marked by 9 to 14 small grooves and resembles a bundle of fine threads. At the termination, these smaller threads broaden axially and end successively from the outside in toward the heaviest central thread. The entire spine is recurved spirally as much as 180° and terminates sharply. In some cases, asecondary thread separatesand recurves to form a second smaller spine. Between the heaviest cord at the shoulder and the suture is a minor cord with minor spines. The 3 rows of spines on the anterior one-half of the siphonal canal occurred on 81% of the specimens. The other 19% had only 2 rows. In this case, a third obsolete row was apparent on some specimens. On others, a third row was present on earlier growth stages but became obsolete before the final varix. Three rows of spines on the anterior part of the canal seems to be normal for this species. Murex norris Reeve, 1845, was compared in the orig- ginal description with M. humilis. The only significant difference noted was several rows of very small, sharp spines in the nearly smooth region on the canal anterior to the aperture. Reeve’s illustration shows these. All specimens of M. humilis showed 3 to 5 small cords in this region terminating as small straight spines. Most of these were broken off; but some of them were intact and were as much as 2mm in length. Comparable spines were also Explanation of Figures 7 to 6 Figure 1: Murexiella humilis (Broderip, 1833) Figure 2: Murexiella humilis (Broderip, 1833) Figure 3: Murexiella keenae Vokes, 1970 Figure 4: Murexiella keenae Vokes, 1970 Figure 5: Murexiella laurae Vokes, 1970 Figure 6: Murexiella laurae Vokes, 1970 All 3 specimens were dredged in 20 to 30m off Punta Juluapan, Colima, Mexico all figures 5 THE VELIGER, Vol. 22, No. 3 [Poorman] Figures 7 to 6 Figure 1 Figure 9 Figure 5 Figure 2 Vol. 22; No. 3 noted on the other 2 species. This is probably a charac- teristic of the “humilis” group in the sense it is used by VoKES (1970). Murexiella keenae Vokes, 1970 (Figures 2, 5) Maxwellia (?) humilis. KEEn, 1958: 354; fig. 342 Murexiella keenae VoxEs, 1970: 328; plt. 50, figs. 8-10; KEEN, 1971: 519; fig. 989 Murexiella humilis. Rapwin & D’ATTILIO, 1976: 157; plt. 25, fig. 5 The species was very adequately described and figured. The holotype (LACMNH 1259) ismore globose thanother specimens collected recently from the same area. Shells from the northern part of the range tend to be still more slender and less fimbriated; but the distinguishing char- acteristics are strongly apparent throughout the range. The nucleus resembles that of Murexiella humilis, ex- cept that the third whorl does not increase regularly but is much more inflated. Also, the axial ribs of the first post- nuclear whorl extend completely across the third nuclear whorl. Postnuclear growth begins with 8 varices per whorl but decreases to 6 in the fully adult stage. The leading edge of each varix is heavier, more fimbriated, and more re- curved than those of the preceding species. The spines at the terminations of the spiral cords are short and axially triangular, recurved, and flexing toward the apex. The spines on the shoulder are subdued and may be only a slight prominence on the thin lamella-like vari- ces. The rows of spines anteriorly on the canal are reduced to 2. The minor cords, which make up the major cords, fan out axially at their terminations to form spatulate recurved spines that do not terminate sharply. The major and minor cords are somewhat like those of Murexiella humilis; but the entire surface is also covered with microscopically fine spiral striations and, in some cases, growth lamellae. Murexiella laurae Vokes, 1970 (Figures 3, 6) Murexiella laurae VoxEs, 1970: 328; plt. 50, figs. 4, 5; KEEN, 1971: 519; fig. 991 THE VELIGER Page 275 Murexiella humilis. Rapwin & D’AtTiL10, 1976: 157; plt. 25, fig. 3 This species was also well described. It is much different from Murexiella humilis. The nucleus is lower and more turbinate. The axial varices of the first postnuclear whorl extend completely across the third nuclear whorl. Adult growth begins with 8 varices per whorl but de- creases to 6 by the fourth whorl and to 4 on the fully adult shell (only 3 on several specimens). The varices are progressively thicker and more reflected with growth. On the shoulder, they appear as reflected lamellae, fimbriated on the leading edge and with a spine in the middle. The spiral cords on the body whorl of the juvenile shell are very broad, nearly flat surfaced, and with narrow interspaces. The cords are high and irregularly nodose from incremental growth. On the adult whorl, these cords are reduced to low, smoothly rounded ridges especially prominent toward the terminal edge of the varix. The cords become stronger here and, as they join the reflected edge, give rise to low, broadly triangular spines, which are sharply recurved. The spine at the shoulder is the heaviest, recurved and flexed toward the apex. The canal is nearly closed. For the first one-half of its length, it is at an angle of 30° to the left of the axis. Then, it. abruptly flexes to the right and is parallel to the axis for the second one-half. Only the tip is reflected dorsally. The first one-half of the dorsum of the canal is nearly smooth, showing several weak cords between low varices. Anteriorly are 2 rows of spines formed by weak © spiral cords. The spines are spatulate and end in 3 or more of the lesser threads that make up each of the major cords. The entire shell is covered with microscopically fine spiral striations. The holotype of Murexiella laurae (LACMNH 1260) is not fully adult. Only the last 2 growth stages, ending in the typically heavy varices, are adult (“growth stage” is defined here as the growth of the shell from one varix to the next succeeding one). Unfortunately, this does not show in the original illustration (VoKEs, 1970), in which the sculpture at the left of the aperture is typical of the juvenile form. DISCUSSION Murexiella humilis occurs from Santa Elena, Ecuador, northward nearly to the head of the Gulf of California. It has not been reported in the Gulf of Panama. The Page 276 THE VELIGER Vol. 22; No. 3 members of this species prefer silty bottom among rocks in 20 - 30m. Murexiella keenae is uncommon intertidally among rocks at Venado Island, Panama. On the west Mexican coast, it has been dredged as far north as Mazatlan, Sina- loa, in 20- 30m on small rock and gravel bottom off rocky headlands. Murexiella laurae has been reported from White Friars, Mexico, to Guaymas, Sonora. It has been dredged in 20 - 30m on small rock and gravel bottom off rocky head- lands. It is to be expected that more intensive searching will extend the ranges of the last 2 species northward and southward, perhaps to more nearly correspond to that of Murexiella humilis. ACKNOWLEDGMENTS Dr. James H. McLean (LACMNH) was instrumental in getting me started on this project. He also made many valuable suggestions concerning the manuscript. Dis- cussions with Carol Skoglund, Phoenix, Arizona, were also valuable in structuring the comparative aspects of the paper. I am also indebted to the following institutions and individuals for making material available for study: Los Angeles County Museum of Natural History; Allan Han- cock Foundation (gastropod collection on loan to LACM NH); S. Stillman Berry, Redlands, California; Ann Marti, Trinity, Alabama (former resident of the Canal Zone) ; Donald Shasky, Redlands, California; Carl and Laura Shy, Westminster, California. Literature Cited BRopDERIP, WILLIAM JOHN 1833. Characters of new species of Mollusca and Conchifera collect- ed by Mr. Cuming. Proc. Zool. Soc. London, Part II (for 1832): 173-179 (14 January 1833) Keen, A. Myra 1958. Sea shells of tropical west America: marine mollusks from Lower California to Colombia. 1%t ed. Stanford Univ. Press, Stanford, California. viii+624 pp.; ca. 1700 figs.; 10 color pits. (5 Dec. 1958) 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru, 2nd ed. Stanford Univ. Press, Stanford, Calif. i-xiv+ 1064 pp.; ca. 4000 text figs.; 22 col. plts. (21 September 1971) Rapwin, Gzorce Epwarp & ANTHONY D’AtTILIO 1976. Murex shells of the world. Stanford Univ. Press, Stanford, Calif. 1-238; 32 colored plts.; 192 text figs. Reeve, Lovett Aucustus 1845-46. Conchologia Iconica: or illustrations of the shells of mollus- cous animals. Monograph of the genus Murex. London, plts. 1-36 (April 1845 to April 1846) Sowerrsy, Gzorcg BRETTINGHAM 2nd 1834-41. Conchological illustrations. Murex. London, plts. 58 - 67 (30 May to 1 October 1834) VoxeEs, Emity Hoskins 1970. The west American species of Murexiella (Gastropoda : Murici- dae), including two new species. The Veliger 12 (3): 325-329; pit. 50 (1 January 1970) Vol. 22; No. 3 THE VELIGER Page 277 Larval and Early Benthic Stages of Brachidontes granulata (Bivalvia : Mytilidae) BY BERNARDITA CAMPOS M. ann LUIS RAMORINO M. Departamento de Oceanologia, Universidad de Chile, Valparaiso, Casilla 13-D, Vifia del Mar, Chile (2 Plates) INTRODUCTION Brachidontes granulata (Hanley, 1843) is a mytilid dis- tributed from Lobos, Peri (6°S) to Isla Chiloé and Seno Reloncavi, Chile (43°S) (Soort-RYEN, 1959). It may be found attached to undersides of rocks in the lower inter- tidal zone (MarINcovicH, 1973), and forming small groups under the mats of various algae in intertidal pools in the rocky areas of Central Chile (Stuarpo, 1960). In Montemar, Valparaiso (33°S), B. granulata is more abun- dant in the semisheltered zone. Other mytilid species that occur in the same area are: Perumytilus purpuratus (Lamarck, 1819), Semimytilus algosus (Gould, 1850) and Aulacomya ater (Molina, 1782). Brachidontes granulata has been included in the sub- genus or genus Hormomya together with Mytilus adamsi- anus Dunker, 1857 (Soot-RYEN, 1955). However, KEEN (1971), adopting a conservative classification recognizes only the genus Brachidontes. This matter needs clarifica- tion in order to properly evaluate the differences found in the larval stages of the species included in Brachidontes sensu lato. The following contributions have been made in rela- tion to the larvae of bivalve species from the Chilean coast: Ranson (1960), Hotiis & Miiiar (1963), Pa- DILLA & ORREGO (1967), and Sotfs (1967) on Ostrea chilensis; WALNE (1974) on Choromytilus chorus (mis- spelled as C. choro) ; Papitta (1973) on Mytilus edulis chilensis and SoLis, SANCHEZ & NAVARETTE (1976) on bivalve larvae from the Estero de Castro. Only RANSON (op. cit.) and Soris, SANCHEZ & NAVARETTE (0p. cit.) described the morphology of the larval shell. No descriptions of larvae of either Hormomya or Bra- chidontes are known to us, except the paper of CHANLEY (1970) on B. recurvus from the western North Atlantic Ocean. YosHma (1937) described the larvae and early juveniles of B. senhausi (Reeve), but according to Kuro- pa et al. (1971) this species belongs to the genus Musculus. The purpose of this paper is to describe the larvae and early benthic stages of Brachidontes granulata so that they may be identified in planktonic and benthic samples. This information should prove especially useful in areas where other species of Mytilidae are cultivated for com- mercial purposes. MATERIALS anp METHODS Sexually mature adult mussels were collected at the intertidal zone near Montemar. In the laboratory all specimens were washed and placed in plastic dishes with filtered sea water (0.8 um membrane filter) and held in an incubator at 6°C for 12 hours. Thereafter, they were changed to filtered sea water at 16°C where they usually began to spawn about 5 hours later. Unfertilized eggs were poured through a set of nylon screens with mesh openings from 150 to 55 um to remove debris. They were immediately placed in a finger bowl with filtered sea water, where some drops of sperm sus- pension were added. Fertilized eggs were kept without movement for 30 minutes, being transferred to a culture vessel with 2L of filtered sea water at laboratory temper- ature (12 - 16°C). Three days later, when the end of the prodissoconch I was reached, air and a food mixture of Tetraselmis and Nannochloris were added; the concentra- tion of algal food was not controlled. During the experi- ment the larvae were washed, sea water was changed and new food was added every 2 days. As the larvae of Brachi- dontes granulata were only cultured up to the early disso- conch stage, the description of the early benthic stages was made from recently settled individuals collected in the same area where adults came from. Page 278 THE VELIGER Vol. 22; No. 3 Cleaned valves were examined, measured and photo- graphed, using a Leitz Ortholux microscope with an Or- thomat camera. The terms prodissoconch I (= prod. I), prodissoconch II (= prod. II) and dissoconch follow WERNER (1939) and REES (1950). The terms veliger and pediveliger are defined in CHANLEY & ANDREWS (1971). RESULTS Brachidontes granulata is a dioecious species. Spawned eggs were brown in color with a mean diameter of 66.3 pum and a size range of 63 to 73 um; sample size: n= 100. The prod. I was colorless, transparent, and the meas- urements in um were: mean length 117, s.d. 5.3, range 105 - 126; mean height 69.7, s.d. 3.9, range 63 - 73.5; n= 100. The evidence of the prod. II development was a yellow edge in the shell material surrounding the prod. I. Simul- taneously with the initial growth of the prod. II appeared a taxodont dentition, bearing an indeterminate number of small central teeth and 2 more prominent ones at the ends of the hinge line. As the larval shell was growing, the 2 largest teeth increased in size, and in number to 6 or 7, forming the curved end of the provinculum in the pedi- veliger stage. The umbo appeared at about 150m larval length as the result of the growing towards the central side of the largest provincular teeth. The dorsal side of the hinge line remained relatively flat, forming a broadly rounded umbo. The eyespots appeared at 160m larval length. A notorious ligament slightly displaced towards the posteri- or side could be seen in larvae 180 um in length. As larvae attained a length of 190 um, the foot was evident between the valves and became functional at 200um length. The velum disappeared at 215 um larval length. At the pediveliger stage the larval shell of Brachidontes granulata could be characterized as follows: yellow pro- dissoconch, anterior margin curved, posterior end almost straight, ventral margin nearly circular, shoulders almost straight, the anterior being longer than the posterior one and not sloping as steeply, umbo broadly rounded and more prominent in width than in height. The mean length in um of the prod. II was 215, s.d. 5.3, range 200 - 231; n= 100. The correlation of the major features of larval develop- ment and sizes of Brachidontes granulata reared in the laboratory, is summarized in Table 1, and the main char- acteristics of the larval shell are shown in Figures 7 to 6. Table 1 Summary of the major features of larval development in Brachidontes granulata. Temp. 12-16°C; §.34-35%o Stage or distinctive feature Mean size Mean age microns Unfertilized egg 66.3 Trochophore 73.0 20 hours Prodissoconch I 117.0 3 days Early umbo 150.0 12 days Appearance of eyespots 160.0 22 days Ligament evident 180.0 28 days Foot 190.0 30 days Functional foot (pediveliger) 200.0 40 days Loss of velum 215.0 55 days The shell of the early juveniles collected from the intertidal zone shows the following characteristics (Fig- ures 7 to 13) : The dissoconch is colorless and transparent; this feature serves to easily distinguish the limit of the yellow prod. II. About 290 4m in length (maximal dis- tance parallel to the hinge), the shell shows a conspicuous growth towards the ventero-posterior end. At a length of about 400 um, the anterior ventral margin of the shell is straight and the umbo is prominent; also the first disso- Explanation of Figures 1 to 7 Larval and early benthic stages of Brachidontes granulata Figure z: Internal view of the larval shell, right valve; length 126 ym Figure 2: Internal view of the larval shell, right valve; length 190 zm Figure 3: Internal view of the larval shell at the pediveliger stage, right valve; length 210m Figure 4: Hinge structure of the larval shell at the pediveliger stage, dorsal view. Shell length 210 um Figure 5: Hinge structure of the larval shell at the pediveliger stage, internal view. Shell length 210m Figure 6: Larva at late pediveliger stage; length 220um Figure 7: Internal view of the right valve at early benthic stage showing the initial dissoconch; length 240 4m Tue VELIGER, Vol. 22, No. 3 [Campos & RaMmorINo] Figures 7 to —— Vol. 22; No. 3 conch teeth (Cox, 1969) develop along the posterodorsal margin some distance beyond the larval hinge. At shell lengths ranging from 450 to 550 um, a new zone grows in the dissoconch; this new zone is characterized by the presence of radial ribs deeply impressed, a slight sinus in the ventral margin and a white coloration contrasting with the first colorless zone of the dissoconch. DISCUSSION Although the main purpose of this work was the descrip- tion of larval and post-larval stages of Brachidontes gran- ulata, the method used allowed us to obtain additional information on the development of the larvae under laboratory conditions. The common methods to induce spawning of sexually mature bivalves in the laboratory are thermal, chemical and electrical shock, sex products stimulation and stretch- ing the posterior adductor muscle (LoosANorF & Davis, 1963). The review of the literature reveals the difficulties of inducing bivalves to spawn, and that there is no agree- ment as to which stimulus is best for each species. The insertion of a small wooden wedge between the shells and stretching of the adductor muscle was unsuccessful in Brachidontes granulata. However, rapid cooling and warming of the water always stimulated spawning of abundant eggs and sperm, giving rise to healthy larvae. From the investigations carried out to date (CARRIKER, 1961; LoosaNorFF & Davis, 1963; BAYNE, 1965; LoosAN- OFF, Davis & CHANLEY, 1966; DEScHWEINITzZ & LuTz, 1976), it is reasonable to infer that in bivalves many features, such as larval shells. eyespots, foot, etc., appear- ing during the larval stages up to metamorphosis, seem to be primarily dependent on size rather than age, temperature, food, or other environmental factors. This conclusion suggests the validity of correlating the larval dimensions with the shape of the larval shells, as well as the other structures already mentioned, even if the individuals may show small variations in the size at which a given feature appears. This generalization has led us to rear Brachidontes granulata with minimal laboratory requirements, regardless of the duration of the larval development. Cultured larvae reached the pediveliger stage by the 40" day at a mean size of 200m in length. Later, the velum gradually degenerated, disappearing by the 55% day. These larvae were kept alive until the end of the experiment for another 18 days, and no attachment was observed during this time; this fact suggests a delay of metamorphosis in Brachidontes granulata larvae. Our THE VELIGER Page 279 results agree with those of BAYNE (1965) on Mytilus edulis, mainly in relation to the events that occur during the delay of metamorphosis. This same author emphasizes that the major factor influencing the delay of metamor- phosis is the availability of a suitable substrate capable of stimulating the secretion of the byssus gland. Although no experiments on the effects of environmental factors in the metamorphosis were carried out by us, the fact that suitable substrates were not offered to pediveligers, might have been the cause of non-attachment in B. granu- lata larvae. From the studies of LoosaNorF (1961), CARRIKER (1961), LoosaNorF & Davis (1963) and Bayne (1965), we infer that it is difficult to ascertain the precise moment of metamorphosis. It is rather a gradual process, preceded by the pediveliger stage, and ending with the develop- ment of a functional byssus gland and the appearance of the dissoconch shell. Neither functional byssal gland nor growth of the shell were observed during the 33 days after the larvae of Brachidontes granulata reached the pediveliger stage, except for a narrow white band of shell that marked the beginning of the dissoconch. While culturing Anomia simplex, LoosaANoFF (1961) observed the disappearance of the velum, retention of a functional foot, and the beginning of the dissoconch without attachment to the substratum. This experience is very similar to ours, except that the shell of Brachidontes granulata did not grow appreciably as in A. simplex. LoosaNnorF (op. cit.) named this phenomenon ‘partial metamorphosis without attachment to the substratum.’ We do not know if the byssal gland in B. granulata could become functional in case that a suitable substrate had been added after the beginning of the dissoconch growth. These “abnormalities” in the metamorphosis process should be further investigated before any feature delimit- ing the end of metamorphosis can be properly ascer- tained. Larvae of Brachidontes granulata are typical pelagic mytilid larvae. Compared with those of B. recurvus from the coast of Virginia in the North Atlantic (CHANLEy, 1970), the former have a more rounded shell, the large teeth of the provinculum appear at a bigger larval size (126 against 105 wm in B. recurvus) and have a distinct ligament. Egg diameter, umbo shape and size at which the eyespots appear are about the same in both species. A shell area located immediately beyond the prodisso- conch and differing conspicuously from the final disso- conch in the early bottom stages of some mytilids, has been described as an “interdissoconch” (JORGENSEN, 1946; REES, 1950). Page 280 THE VELIGER Vol. 22; No. 3 OcKELMANN (1965) has pointed out that such inter- dissoconchs also occur in some species of other families besides Mytilidae and Pectinidae, so there may be as many as 4 different stages of shell formation in marine bivalves. On the other hand, CarrikeR (1961) described 2 different zones in the dissoconch of Mercenaria mercen- aria, calling them dissoconch with primary shell ridges and dissoconch with secondary shell ridges. These observations agree with ours, since in Brachi- dontes granulata 4 stages in shell formation are also clearly shown. After a yellow prod. II, there is a trans- parent zone bearing only circular striae, and beyond a white shell with the typical radial striae of the adult (Figures 12, 13). Summarizing, we confirm for this species that besides prod. I and prod. II, there is a dissoconch structurally divided into 2 different types of shell: the so-called in- terdissoconch of some authors and the final dissoconch. SUMMARY The description of the larvae and early benthic stages of Brachidontes granulata, employing cultured larvae under laboratory conditions, and intertidal benthic samples from Valparaiso Bay is presented. A partial metamorphosis without attachment to sub- stratum of the cultured larvae is discussed. In the pedi- veliger larvae the velum was lost and a narrow band of dissoconch was formed, but no functional byssal gland was observed. The dissoconch of Brachidontes granulata is structur- ally divided into 2 different zones. One of these should correspond to the so-called interdissoconch of other authors. ACKNOWLEDGMENTS We thank Dr. José Stuardo for his critical review of the manuscript. We also thank Mr. Mario Barahona for his assistance in the laboratory work; Mrs. Nora Aguirre for the photographic prints; Mr. Juan Soto and Mr. Gabriel Lillo for technical aid. Literature Cited Bayne, Brian LEICESTER 1965. Growth and the delay of metamorphosis of the larvae of Mytilus edulis (L.). Ophelia 2 (1): 1-47; 20 text figs. (June 1965) CarrIkER, MELBOURNE ROMAINE 1961. Interrelation of functional morphology, behavior and autoecolo- gy in early stages of the bivalve Mercenaria mercenaria. Journ. Elisha Mitch. Sci. Soc. 77 (2): 168-241; 39 text figs. (Nov. 1961) CHANLEy, Pau 1970. Larval development of the hooked mussel Brachidontes recur- vus Rafinesque (Bivalvia : Mytilidae) including a literature review of larval characteristics of the Mytilidae. Proc. Nat. Shellfish. Assoc. 60: 86-94; 3 text figs. (June 1970) Cuan ey, Paut « Jay D. ANDREWS 1971. Aids for identification of bivalve larvae of Virginia. Malaco- logia 11 (1): 45-119; 51 text figs. (8 October 1971) Cox, Les. REGINALD 1969. General features of Bivalvia. In: Raymond GC. Moore « Curt Teichert (eds.), Treatise on invertebrate paleontology, Part N, Mollusca 6, Bivalvia 1: N2-Nr29; 86 text figs. Geol. Soc. Amer. Inc. & Univ. Kansas De Scuweinitz, EvizaABETH & RicHarp A. Lutz 1976. Larval development of the northern horse mussel Modiolus modiolus (L), including a comparison with the larvae of Mytilus edulis L. as an aid in planktonic identification. Biol. Bull. 150 (3): 348-360; 5 text figs. (June 1976) Hots, P J. e R. H. MiLvar 1963. Abbreviated pelagic life of Chilean and New Zealand oysters, Nature, London 197: 512-513 Jorcensen, C. B. 1946. _Lamellibranchia. In: Reproduction and larval develop- ment of Danish marine bottom invertebrates, with special reference to the planktonic larvae in the Sound (@resund), by G. Thorson. Medd. Komm. Danm. Fisk. Havunderseg., Copenhagen, Ser. Plankton 4 (1): 277-311; 28 text figs. Keen, A. Myra, with the assistance of Jamzs Hamirton McLean 1971. Sea Shells of Tropical West America; marine mollusks from Baja California to Peru. and ed. Stanford Univ. Press, Stanford, Calif i-xiv + 1064 pp.; ca. 4000 figs.; 22 color plts. (1 September 1971) Kuropa, Toxuse!, TADASHIGE HaBE & KaTurA OyYAMA 1971. The sea shells of Sagami Bay. Biol. Labor. Imp. Household, ed.; Maruzen Co. Ltd., Tokyo. Biling. ed.: Japanese 741 pp. + index 28 pp.; English 489 pp.+index 12 pp.; 2 text figs.; 121 color plts. Loosanorr, Victor Lyon 1961. Partial metamorphosis in Anomtia simplex. Science 133: 2070 - 2071 LoosanorFF, Victor Lyon « Harry CHar.es Davis 1963. Rearing of bivalve mollusks. In: FE S. Russell (ed.), Ad- vances in marine biology 1: 1-136; 43 text figs. Acad. Press, Inc., London & New: York Loosanorr, Victor Lyon, Harry Cari Davis & Paut E. CHANLEY 1966. Dimensions and shapes of larvae of some marine bivalve mol- lusks. Malacologia 4 (2): 351-4353; 61 text figs. (31 Aug. 1966) Marincovicy, Lour N., Jr. 1973. Intertidal mollusks of Iquique, Chile. Mus. Nat. Hist. Sci. Bull. 16: 1-49; 102 text figs. OcKELMANN, Kurt W. 1965. | Developmental types in marine bivalves and their distribution along the Atlantic coast of Europe. Proc. 1st Europ. Malacol. Congr. 1962: 25-35; 5 text figs. PADILLA, MicuEL 1973. Observaciones biolégicas relacionadas con el cultivo de Mytilus edulis chilensis en Aysén. Publ. Inst. Fom. pesq. Santiago 54: 1-21; 3 pits. Los Angeles County (February 1973) Explanation of Figures 8 to 13 Early benthic stages of Brachidontes granulata Figure 8: External view of the right valve; length 290 ym Figure g: Internal view of the hinge structure, right valve; shell length 290 pm Figure 10: External view of the right valve; length 440 ym Figure 11: Hinge structure of the right valve; shell length 440 nm Figure 12: External view of the right valve; length 500um Figure 13: External view of the right valve; length 590 nm [Campos « Ramorino] Figures 2 to 13 Tue Veticer, Vol. 22, No. 3 Vol. 22; No. 3 THE VELIGER Page 281 PapILLA, MIGUEL & JULIO OrREGO 1967. La fijacién larval de ostras sobre colectores experimentales en Quetalmahue, 1966-1967. Boln. Cient. Inst. Fom. Pesq. Santiago 5: 1-15; 5 text figs. Ranson, GILBERT 1960. Les prodissoconques (coquilles larvaires) des Ostréidés vivants. Bull. Inst. oceanogr. Monaco 1183: 1 - 41; 136 text figs. (June 1960) Regs, C. B. 1950. The identification and classification of lamellibranch larvae. Hull Bull. Mar. Ecol. 3 (19): 73-104; 4 text figs.; 5 plts. (Nov. 1950) Sois, IvAN 1967. Observaciones biolégicas en ostras (Ostrea chilensts Philippi) en Pullinque. Biol. Pesq. Chile 2: 51 - 82; 6 text figs. (June 1967) Soris, IvAN, Patricia SANCHEZ & SONIA NAVARRETTE 1976. Identificacién y descripcién de larvas de moluscos bivalvos en el plancton del estero Castro. Bol. Soc. Biol. Concepcién 50: 183-195; 12 text figs. WERNER, BERNHARD Soot-Rygn, Tron 1955. A report on the family Mytilidae (Pelecypoda). Allan Han- cock Pacif. Exped. 20 (1): 1-154; 78 text figs.; 10 plts. (Nov. 1955) 1959. Pelecypoda. Reports of the Lund University Chile Expedi- tion 1948-49. Lunds Univ. Arsskr. N. F (2) 55 (6): 1-86; 6 text figs.; 4 plts. (February 1959) Stuarpo, José 1960. Notas sobre ecologia y distribucién de Choromytilus chorus (Molina) (Mytilidae) con una clave adicional para los mitilidos chilenos. Act. y Trab. Prim. Congr. Sudamer. Zoolog. (La Plata, Oct. 1959) 1: 163-174; 3 text figs.; 1 map Watng, PR. 1963. Breeding of the Chilean oyster (Ostrea chilensis Philippi) in the laboratory. Nature, London 197: 676 (February 1963) 1974. Culture of bivalve molluscs. Fishing News (Books) Ltd., Surrey, England, 173 pp.; 38 text figs.; 24 plts. 1939. Uber die Entwicklung und Artunterscheidung von Muschek larven des Nordseeplanktons, unter besonderer Beriicksichtigung der Schalenentwicklung. 22 text figs. Yosuipa, Hirosu1 Zool. Jahrb., Abt. Anat. Ontog. 66: 1-54; (October 1939) 1937. On the pelagic larvae and young of Brachidontes senhaust (Reeve). Venus 7 (3): 121-128; 7 text figs.; 1 plt. (July 1937) Page 282 THE VELIGER Vol. 22; No. 3 Deep Water Collections of Opi sthobranchs in Central California BY ROBERT CHAMBERLAIN Route 3, Box 213, San Luis Obispo, California 93401 AND DAVID W. BEHRENS Biological Research Laboratory, Pacific Gas and Electric Company, P. O. Box 117, Avila Beach, California 93424 (2 Text figures) LitTLE IS KNOWN concerning the biology of opistho- branch mollusks in the deep water continental shelf areas of California. A few papers report the depths and loca- tions of particular deepwater species (SMITH & GorRDON, 1948; HANNA, 1951; CHIVERS, 1966; MAcFarRLAND, 1966; BeRTSCH, 1969; CARLISLE, 1969; LEE « BRopHy, 1969; McDonatp «& NyYBAKKEN, 1975). Many more reports are available, however, describing the ecology of those species residing in shallow inshore waters. This report describes the findings of 6 opisthobranch species from 3 deepwater trawls off Central California. The species, their depths (several representing substan- tial extensions) and associated food items are discussed. METHODS Opisthobranchs were collected in conjunction with nor- mal commercial trawl fishing activities on the vessel Silver Queen. The senior author accompanied the fisher- men to collect any non-commercial specimens brought up in the trawls. Trawl samples were sorted on board and all invertebrate material was preserved in 10% buffered formalin solution and taken to the laboratory for further examination. In the laboratory, the stomachs were opened and flushed with formalin solution and the contents were examined microscopically for food material, which was then identified to the lowest taxonomic level possible. Opisthobranchs were found in 3 of 11 trawls, taken 9.6km off Point Piedras Blancas, California (Figure 1), January 27 to 31, 1978. Trawls ranged in depth from 112 to 360m, and lasted 3 to 5 hours at a speed of 2 knots. 121°37 20/ 20’ 17’ 36°00’ 0 7r V2S ieee 5 nautical miles 45 &: Pt. Piedras 40° Ress ‘ B 35°36" Figure 1 Map of Point Piedras Blancas area showing location of the 3 trawls 121°37" 20/ 20’ 617’ The original map submitted with the manuscript showed depth contour lines labeled in "f". As The Veliger is strictly metric, the measurements were converted to "m'"'. However, after publica- fei tion, we were informed that "f£" stood for ¢ ee O18 Bh oe & "fathoms", not "feet"; the abbreviation should Be nautical miles have been "fm". We present herewith a corrected Deo version of the map with the suggestion that it be pasted over the incorrect map on page 282 of ei Pt. Piedras volume 22. Caution: do NOT use rubber cement. Ae... g00m 720m 540m 360m Ce AY 5 + : Vs ry « Vol. 22; No. 3 OBSERVATIONS anp DISCUSSION In spite of the obvious geographical separation of trawl 4 from 2 and 5 (Figure 1), we have chosen to discuss trawl 2 and 4 together as the first collection because of the similarity of species composition and substrate. Trawl 5 is treated separately as a second collection. Trawl number 2 (Table 1), taken January 27, 1978, occurred at a depth of 236 - 291m. Substrate materials sorted from the trawl were mud, sand and boulders. Trawl number 4, although shallower (107- 218m), produced similar bottom materials. The species composition of live material from the 2 trawls was the same (‘Table 2). THE VELIGER Page 283 A single specimen of Plewrobranchaea californica Mac- Farland, 1966, was collected in each trawl. They meas- ured 100 and 95mm (preserved), respectively. Our col- lections are consistent with Curvers (1977) who reported the bathymetric range of the species to be g to 369m and its substrate to be green mud. Stomach contents analysis of the Pleurobranchaea from trawl 2 produced an interesting assemblage of food items which we feel gives an accurate account of this species’ benthic foraging existence. The stomach material includ- ed 5 specimens of an unidentifiable species of Aglaja. The specimens ranged from 6-8mm in length (preserved). The cephalic shield and mantle were mottled brown in Table 1 Trawl Location and Substrate Data Trawl # Date Depth (m) Latitude Longitude Substrate 236-291 107-218 277-332 Jan. 27, °78 Jan. 28, ’78 Jan. 29, ’78 35°30'17"N 35°40'15"N 35°30'25"N ou ® bo 121°10'47”W 121°20'30”W 121°10'57” W Mud, sand and boulders Mud, sand and boulders Mud, compacted mud boulders Table 2 Species Composition of Trawls Trawl Number Species 2 4 By Cephalaspidea Scaphandridae 1. Acteocina intermedia x Willett, 1928 Aglajidae 2. Aglaja sp. indet. x (from stomach of spec. 4) 3. Chelidonura inermis Xx (Cooper, 1863) Notoaspidae Pleurobranchidae 4. Pleurobrachaea californica Xx x MacFarland, 1966 Nudibranchia Doridacea Cadlinidae 5. Cadlina sp. xX Dendronotidae 6. Dendronotus frondosus Xx (Ascanius, 1774) (from stomach of spec. 4) color and the foot was white. An intact radula of Dendronotus frondosus (Ascanius, 1774) was also found in the stomach of this Plewrobran- chaea. The radula agreed in every respect with that de- scribed in McDona.p (1977). This finding should estab- lish the occurrence of D. frondosus at this depth in Calli- fornia waters. Its deepest occurrence in California is reported to be 19m (SPHON & LANcE, 1968) and 40m at Friday Harbor, Washington (RoBILLIARD, 1970). In the Atlantic, this cosmopolitan species is reported to 400m (SWENNEN, 1961). Also found in this stomach were 2 isopods, possibly juveniles of the suborder Flabellifera, copepod exuvia, many minute fish scales and sand and silt particles. The presence of fish scales would be expected from an animal which feeds by sucking its invertebrate prey off the mud surface. The stomach of the Pleurobranchaea from trawl 4 con- tained several transparent “tunicate-like” organisms. They were too digested to allow for a more precise identi- fication. The only other reports of food preferences of this species are Coan (1964) and Cuivers (1966). However, these were laboratory observations and did not include prey species which would be available to Pleurobranchaea in nature. Page 284 THE VELIGER Vol. 22; No. 3 The second collection, trawl 5 (Table 1), taken January 27, 1978 was in a depth of 277 - 332m. Bottom substrate materials were mud and compacted mud boulders. Three opisthobranch species were collected in the trawl (Table 2). All 3 represent substantial depth extensions. One specimen of Acteocina intermedia Willett, 1928, was collected in this trawl and later identified by Mr. Don Cadien, Marine Biological Consultants, Costa Mesa, California. The shell dimensions were 8 by 2mm, length and breadth, respectively. SmirH « GorDoN (1948) re- port A. culcitella intermedia to be common on sand bot- toms, 18- 55m deep in Monterey Bay, California. Wi- LETT (1928) reports this species in 55m at Catalina Is- land, California. CARLISLE (1969) reports A. intermedia to be frequent in Santa Monica Bay, California, at depths from 37 - 184m. A single specimen of Chelidonura inermis (Cooper, 1863) was also collected in this trawl. The specimen measured 20mm (preserved). Its colors were bright and typical. The specimen was in good condition, but its stomach was empty. PAINE (1963) reported the greatest depth for this spe- cies to be approximately 100 feet [30m]. Described by him as abundant in bays and sloughs, our collection indi- cates that this species occurs offshore in the deep benthos as well. The presence of available food for this highly voracious predator is confirmed by the occurrence, in the same trawl, of 2 other opisthobranch species on which it is known to prey: Acteocina intermedia (previously dis- cussed) and Cadlina sp. (Table 2). The one specimen of Cadlina found in trawl 5 meas- ured about 30mm long and 15mm wide (preserved). This species remains unidentified. We offer the following detail for future authors to use for the purpose of syn- onymy. The dorsum was nearly smooth, bearing very low tubercles. The foot was long and linear and the labial tentacles were short and auriculate. The perfoliate rhino- phores bore 14 lamellae. The entire body color (preserved) was cream, with no distinguishable markings common to Cadlina, such as dark rhinophores or dorsal spotting. The radular formula was 86 X (34 - 36° 1-34 - 36). The rachid- ian teeth bore 5-6 blunt denticles (Figure 2a). The innermost lateral teeth bore 1 - 2 denticles on the inner margin (Figure 2b). The 15" lateral tooth was hook- shaped with 11 - 14 denticles below a long cusp (Figure 2c). The outermost lateral tooth was without a cusp and bore 5 - 7 denticles beginning at its tip (Figure 2d). While this description does not agree with any known California species of Cadlina (McDonatp, 1977), we Figure 2 Radula of Cadlina sp. (from 40% row) a — rachidian; b — 1S lateral tooth; c — 15" lateral tooth d — 33" lateral tooth believe it most nearly agrees with C. flavomaculata Mac- Farland, 1905. The exceptions are the absence of dark rhinophores, and a slight deviation in radular morpho- logy and a number of radular teeth. Cadlina flavomacu- lata is reported to a depth of 220m (Lee & Bropny, 1969). Their specimen, which was trawled off Port Hueneme, Ventura County, California, is no longer available for examination and there are no field notes to assist in confirming that identification and its possible alliance with our specimen. Prior to LEE & BropHy (op. cit.) C. flavomaculata was known as an inshore species found only to 19m (SpHON & Lance, 1968). Due to the apparent difference in morphology, this specimen might well belong to an undescribed species. We are hesitant to formally describe this species without additional ma- terial. Stomach analysis found the gut empty. Vol. 22; No. 3 THE VELIGER Page 285 ACKNOWLEDGMENTS Rice eRe ee natural history of Pleurobranchaea spec. (Gastro- We would like to express our thanks to Mr. Gary Hopkins, skipper of the Silver Queen, for allowing the senior au- thor to accompany him on this trip. Special thanks to Pacific Gas and Electric Company for the use of its Diablo Canyon Biological Research Laboratory and facil- ities for examining the trawl samples, and to Messrs. Bud Laurent and John Warrick for reviewing the manu- script. Thanks also to Mr. Don Cadien for his identifica- tion of Acteocina intermedia and to Messrs. Bob Hender- son and Pat Brophy for their examination of the Aglaja specimen. Literature Cited BrrtscH, Hans 1969. A note on the range of Gastropteron pacificum (Opisthobranchia : Cephalaspidea) . The Veliger 11 (4): 431-433; 1 map (1 Apr.) Caruls.z, Joun G., Jr. 1969. Invertebrates taken in six year trawl study in Santa Monica Bay. The Veliger 11 (3): 237-242 Cuivers, Dustin D. 1966. Observations on Pleurobranchaea californica MacFarland, 1966 (Opisthobranchia, Notaspidea). Proc. Calif. Acad. Sci. 32 (17): 515-521 (22 November 1966) (1 January 1969) poda: Opisthobranchia). Hanna, G DALLAs 1951. A new West American nudibranch mollusk. 65 (1): 1-3; plt. 2 Lze, Ricuarp S. « Patrick BropHY 1969. Additional bathymetric and locality data for some opisthobranchs and an octopus from Santa Barbara County, California. The Veliger 12 (2): 220-221 (1 October 1969) MacFartanp, FRANK Mace 1966. Studies of opisthobranchiate mollusks of the Pacific Coast of North America. Mem. Calif. Acad. Sci. 6: xvi+546 pp.; 72 plts. (8 April 1966) The Veliger 6 (3): 173 (1 Jan. 1964) The Nautilus (27 August 1951) McDonatp, Gary R. 1977- A review of the nudibranchs of the California coast. Unpubl. master’s thesis, Calif. State Univ. Hayward ix+337 pp.; 13 plts. McDonatp, Gary R. & James WILLARD NyBAKKEN 1975. Cerberilla mosslandica, a new eolid nudibranch from Monterey Bay, California (Mollusca : Opisthobranchia). The Veliger 17 (4): 378 - 382; 2 text figs. (1 April 1975) Paring, Ropert TREAT 1963. Food recognition and predation on opisthobranchs by Navanax inermis (Gastropoda: Opisthobranchia) . The Veliger 6 (1): 1-9; pit. 1; 1 text fig. (1 July 1963) Suto, ALLYN GoopwIn & MacKENziIz Gorpon, Jr. 1948. The marine mollusks and brachiopods of Monterey Bay, Cali- fornia, and vicinity. Proc. Calif. Acad. Sci. ( 4) 26 (8): 147 - 245; pits. 3, 4; 4 text figs. (15 December 1948) SpHon, Gate G. & James Rosert LANcE 1968. An annotated list of nudibranchs and their allies from Santa Barbara County, California. Proc. Calif. Acad. Sci. 36 (3): 73 - 84; 1 fig. (25 September 1968) SWENNEN, CHARLES 1961. Data on distribution, reproduction and ecology of the nudi- branchiate Mollusca occurring in the Netherlands. Netherl. Journ. Sea Res. 1 (1-2): 191 - 240 WILLETT, GEORGE 1928. Notes on some Pacific Coast Acteocinas, with description of a new subspecies. The Nautilus 42 (2): 37-38 (25 October 1928) Page 286 THE VELIGER Vol. 22; No. 3 Some Aspects of Food Intake in Octopus joubini Robson BY JENNIFER A. MATHER Department of Psychology, Brandeis University, Waltham, Massachusetts 02154 (3 Text figures) INTRODUCTION ENVIRONMENTAL FACTORS may affect an animal’s food intake in a variety of ways, some of which may be affected by phylogenetic background or mode of life, some of which may be general. For instance, all animals should regulate the amount of food intake at one time by feed- back signals, as DETHIER (1976) has shown for the fly. Yet predators have a set of demands which must affect their intake drastically — the necessity of searching for food, subduing and consuming it, and responding to tem- porary abundance. They may thus, for instance, have a very high “ceiling” for food intake and phylogenetic dif- ferences in their response to increasing density. HOLLING (1966) has postulated a general mode of predator intake dependent on density, and found intake of invertebrates could be described as a negatively accelerated curve rising to a plateau, and that of vertebrates as an S-shaped curve. He admits that these phylogenetic differences should arise because of the capability of vertebrates to learn, and that cephalopods could be similar to mammals. To look at this and other factors affecting food intake, octopuses are thus of interest and the small Octopus joubini, which can easily be kept in captivity, is an ideal species. Information about food intake is available from experi- ments on rats (see HINDE, 1970), flies (DETHIER, 1976), and, in marine species, goldfish (Rozin & Mayer, 1961) and three-spined stickleback. For octopuses, informa- tion on food intake comes from studies of several species in the genus Octopus and is fragmented and sometimes contradictory. ‘Temperature affected food intake in O. briareus; with an increase of 10°C (20° to 30° ), animals ate twice as much (Borer, 1971). Temperature influ- enced food intake of O. vulgaris as well, and again intake increased when temperature was higher (Man- GOLD & VON BoLETZKY, 1973). Deprivation of food raised intake in O. maya (WALKER, Lonco & BrTTERMAN, 1970); intake after 2 days’ deprivation was double that after one day’s. However, deprivation only increased intake of O. briareus minimally (Borer, op. cit.). Octopus briareus was affected by prey density, however. BorER (op. cit.) recorded a linear increase in number of crabs eaten dependent on density, which does not fit HoLiinc’s (1966) model. However, she did not record weight of intake, which would be a clearer measure since octopuses do not always eat all of a killed prey. One factor, matu- ration of female octopuses to the egg laying and brooding stage, influenced animals of several species in the same manner. Female O. cyanea (VAN HEUKELEM, 1973), O. maya (WALKER et al., 1970), O. vulgaris (MANGOLD & VON BoLETzKy, 1973) and O. joubini (OPRESKO & THomMaAS, 1975) all nearly ceased food intake during brooding. As might be expected of predatory animals, octopuses appear to be tolerant of long periods of food deprivation. Rats lost a larger percentage of body weight each day during deprivation (CAMPBELL, TEIGHTSOONIAN & WIL- LIAMS, 1961) than did octopuses. Furthermore, octopuses lost less weight per day over successive days of depriva- tion (Nrxon, 1966; MANcoLD & von BoLETzKy, 1973), whereas rats lost weight steadily. After deprivation and weight loss, Octopus maya were capable of a much in- creased intake of food (WALKER et al., 1970), which would compensate for the period of deprivation. The present study concerns factors affecting feeding and food deprivation in Octopus joubini Robson, 1929. These very small animals are ideal for laboratory investi- gation because they can be kept in small, closed salt water systems and have a short life span (THOMAS & Op- RESKO, 1973). They can also be raised from eggs, as the relatively large young are benthic (von BoLETzKY & von BoLetzkKy, 1969; MATHER, 1972; OPRESKO & THoMAS, 1975). Moreover, since most octopuses have Vol 22; No: 3 THE VELIGER Page 287 the same benthic predatory mode of life, parallels may be drawn between O. joubini and other octopod species. In this study, intake was studied when food was provided ad libitum and under varied conditions of deprivation and prey density, during females’ egg production and brooding. PROCEDURE (a) Maintenance: Octopuses were collected from St. Joseph Bay, Florida (they were picked off the sandy bottom in their shell “homes’’) and flown, still in the shells, to Boston. In the laboratory they were isolated in 18L or 36L capacity aquaria with separate closed sea water systems. Each animal was given a 2.5cm cube plexiglass “home” (4 sides dark, one end clear and one end open). Temperature was controlled separately in each tank and maintained at 23°C (41°). Once each week, salinity was measured and corrected by addition of distilled water and 4 of the water was changed to prevent the accumulation of nitrates. A bright white light provided illumination during the day and a dim red one at night. The changeover was timed to sunrise and sunset in Boston (in order to produce a daylength variation which might have been important for maturation), and corrected weekly. (b) Intake Estimation: Uca fiddler crabs and Nas- saritus mud snails were used as prey. The number of prey provided was always in excess of expected intake esti- mated from previous findings. To ensure an accurate measure of intake, live crabs were first immersed in water, then dried on blotting paper for 10 seconds before being weighed (this procedure was carried out before they were offered as prey and again if they survived, after the octopuses’ feeding period). Live and dead prey were re- moved after approximately 24 hours (Octopus joubini feeds almost exclusively at night, so small differences in amount of day time affected intake little). Bodies of crabs that had been partly eaten (or carapaces and arm bases if the crab had been disjointed) were allowed to drain on blotting paper 10 seconds before being weighed. Live snails were weighed the same way as live crabs, but the empty shells were held for 10 seconds with the siphon notch on blotting paper to ensure adequate drainage. In- take was measured by subtracting weight of prey (and shells) removed from weight of prey offered. (c) Experimental Conditions: (1) Baseline — Four octopuses were offered crabs provided ad libitum for 40 days. Their baseline intake and daily variations were recorded. (2) Deprivation — A group of 4 octopuses was fed crabs after varied periods of deprivation. For 2 replica- tions they were deprived 1, 2 and 4 days in a Latin Square arrangement (data for zero days’ deprivation were col- lected from the ad libitum feeding of the day before and day after trials). As the major intake difference was between zero and one day’s deprivation, the third and fourth replications used deprivation times zero, one, and four days in a Latin Square arrangement. During the third and fourth sequence, weight changes were moni- tored during the 4-day deprivation period. Each octopus’ plexiglass “home” had been weighed damp before he used it. During each day of deprivation and after the octopuses were fed, each was weighed in his home. This procedure included a presumably constant error due to the weight of the water contained in the octopuses’ mantle cavities, but minimized disturbance and produced stable results. The weight of each home was subtracted from the weight of octopus and home to calculate the weight of each octopus. (3) Density — Density of prey species is one way of varying absolute quantity — it can also be varied by varying prey size. Three experimental manipulations were used to separate relative influences of the factors — vary- ing prey size (numbers constant), varying prey number (size constant) and covarying prey size and number so that total prey weight was constant. All 3 tests were carried out using crabs as prey. The last mentioned mani- pulation was done first, and the design corresponds to BoreEr’s (1971) design of alternating low density (4 crabs) with 4 higher densities (8, 10, 12, 16 crabs). Because this design, while a faithful copy of her experimental situ- ation, is impractical for statistical analysis, the other tests were arranged differently. For weight variation (n= 16) 3 levels (weight doubled and redoubled) were used, ar- ranged in a Latin Square arrangement. For numbers var- iation (n=11) there were also 3 levels (number doubled and redoubled) in the same arrangement. Using snails as prey, only tests of size variation (n = 10) and number variation (n==13) were carried out. Numbers were given at the same 3 levels, but size could only be doubled and was thus at 2 levels, because the size variation of the snails is less than that of the crabs. (4) Intake of Females during Egg Production and Brooding — Four females who neared egg laying were provided food ad libitum throughout brooding and until death (#24 was in one 2-week density study prior to egg laying). Six males were also provided food ad lbi- tum as a control. Females’ food intake was charted according to time preceding or following egg laying (they did not all lay eggs at the same time), males’ by the calendar. Page 288 THE VELIGER RESULTS (a) Baseline: Four juvenile octopuses were given excess crab prey and still showed a marked variation in daily intake with no consistent pattern across individ- uals which would suggest environmental influences such as water quality. As Figure 1 shows, intake could be rela- tively constant or cyclic, and could drop to zero for up to 3 days. There was no consistent pattern of intake from day to day but the mean intake of each animal was simi- lar when averaged over a 10-day period, and the mean for all animals rose slowly from o.9g/day in the first 10-day period to 1.2g/day in the 4™ to-day period, as the animals grew. go 12345678910 12345678910 12345678910 Time in Days Weight in grams Figure 1 Some examples of patterns of daily food intake in grams for Octopus joubini provided crab prey ad libitum (n=4) (b) Deprivation: Lack of food for even one day markedly influenced intake. Mean food intake of 4 octo- puses after zero days’ deprivation was 0.212g, after 1 day’s deprivation 0.396g; this difference was significant at the p 3 Bee Effect of Prey Density of Crab and Snail Prey y = on Weight of Food Intake of Octopus joubini = 8. 5 wn 0.50 =| Prey Prey Prey Unit Prey Total Mean Intake 5 Type Number _ Weight Weight Weight Bs, (gm) (gm) (gm) 4 0.56 2.25 0.22 8 0.28 2.25 0.23 Sf 30 = 54 99 =—ot he 89 Ibe Be Crab 10 0.22 2.25 0.18 "FIRST EGGS 12 0.19 2.95 0.17 Tine Work 16 0.14 2.25 0.26 6 0.58 3.50 0.51* en g Crab 12 0.58 7.00 0.81 24 0.58 14.00 1.07 Weight of food intake in grams of female Octopus joubini eating 12 0.25 3.00 0.45** crab prey, averaged over week-long periods measured before and Crab 12 0.50 6.00 0.68 after egg laying (n=4) 12 1.00 12.00 135) Snail 10 0.70 7.00 0.41 10 1.40 14.00 0.56 10 1.10 11.00 0.68 compensate for deprivation shown by O. joubinz is also an onal 20 LO ZED O40 advantage for a predator. These octopuses tolerated 4 40 1.10 44.00 0.96 *ANOVA p < 0.05 **ANOVA p < 0.01 weeks after). Average intake of females rose 3 weeks before egg laying to nearly 1.0g/day, fell the week be- fore to 0.57g/day (this is within the range of male food intake at this time), and then fell to under 0.20g for most of the 6-week brooding period (Figure 3). Average food intake by 6 males during this period ranged from 0.29 g/day to 0.47g/day. DISCUSSION Several aspects of the food intake of Octopus joubini are typical of predatory animals. The wide variability in daily intake despite a constant food supply is also characteristic of predatory goldfish (Rozin & Mayer, 1961), stickle- back (TuGENDHAT, 1960), and blowflies (DETHIER, 1976: 283), but unlike constant intake of omnivorous rats (Cor- BIT & STELLAR, 1961). Octopus vulgaris also varies its intake, from low to high on alternate days (Wodinsky, personal communication). The ability to tolerate and days’ deprivation with a weight loss of only 10%, and the loss per day declined with increasing deprivation. This loss was larger than that of O. briareus for the same de- privation time (6%) (BorER, 1970) and than that of O. vulgaris (4%) for 6 days (MANGOLD & von BoLeTzky, 1973), perhaps because the smaller O. jowbini is meta- bolically more active. Both O. jowbini and O. vulgaris (MANGOLD & VON BoLETzky, op. cit.) compensated for long periods of deprivation by eating so much (up to 20% of body weight for O. joubini) that weight loss was quickly recovered. Food deprivation for even 1 day affects Octopus jou- binz’s food intake. This agrees with TUGENDHAT’s (1960) observations for stickleback and WALKER, LONGO & BITTERMAN’s (1970) observations for O. maya, though intake increased for all 3 species after 1 or 2 days’ dep- rivation. BorER (1970) found only a minimal effect of deprivation on food intake for O. briareus. The data on food intake show two results. First, Octo- pus joubini are consistent in their intake response to ab- solute quantity of crab prey. When crab prey size and weight were co-varied so weight was always 2.252, in- take did not vary significantly. Second, O. joubini were affected by both increased crab size and by increased crab weight. As for O. cyanea (BoucHER-RopDoNI, 1973) and for rats (BALAGURA & HARRELL, 1974), O. joubini Page 290 increased food intake when the weight of units offered was greater. The rise in weight of food intake with in- creased crab prey density for O. joubini was not linear, as Borer (1971) found for number of crabs eaten with O. briareus. Instead, it apparently begins to negatively ac- celerate, as suggested by Hotiinc (1966) for intake of predators with increased prey density. The response to snail prey is less easy to interpret, since snails are less preferred prey for Octopus joubini (MaTHER, 1972) and also time-in-contact (see HoLLinc, 1966) will be less for them than for crabs. This is true because crabs are more active than snails. Wodinsky (per- sonal communication) was able to remove the “density effect” on O. vulgaris by confining crab prey. It is also true because O. joubini spent a long period pulling snails out of their shells before consuming them, but could kill and begin to eat a crab in one minute. Perhaps time-in- contact with snails is small enough that O. jowbini’s in- take was at the base of the S-shape curve suggested by Ho.iinc (1966). Perhaps, as in BEUKEMA (1968), the density effect is complicated by prey preference. The reproductive state of femalesalso affected their food intake. An intake decline before and during brooding is common among octopods, mentioned for Octopus jou- bint by OpresKo & THOMAS (1975), for O. vulgaris by Nrxon (1966), for O. cyanea by VAN HEUKELEM (1973) and for Haplochalaena maculosa by TRANTER & AUGUS- TINE (1973). It is interesting that the decline in food intake began before egg laying - one week before for O. joubini, 2 to 8 weeks before for O. vulgaris (MAN- GOLD & VON BoLetzxy, 1973) and several weeks before for O. cyanea (vAN HEUKELEM, 1973). But no octopod yet studied other than O. joubini has shown increased intake and a growth spurt before egg laying. This pat- tern may be related to O. joubini’s short life span or to its very small size (the weight of eggs becomes a large proportion of body weight) ; it parallels the growth spurt of juvenile female O. vulgaris when control over their optic glands was removed and they matured early (WELLS & WELLS, 1959). The effect may be a hormonal one, high intake being induced by optic gland hormones. Octopus joubini’s food intake and feeding appear fair- ly typical of octopods. Deprivation of food caused elevated intake and weight loss which declined with time, as for other octopods. Density increase and weight increase caused increased intake, a result similar to, but not the same as, that in other Octopus species. Changes in female reproductive state cause first an increase and then a decrease in food intake. Only the latter effect has been previously noted in octopuses, perhaps because O. jou- bini’s short life span compresses and clarifies long-term effects. For this reason, and because these small animals are easy to maintain in the laboratory, O. joubini offers THE VELIGER Vol. 22; No. 3 an excellent opportunity for future research on feeding and food intake. ACKNOWLEDGMENTS I would like to thank Dr. J. Wodinsky and Dr. J. Lackner for their assistance. Literature Cited Baracura, S. « L. E. Harrevy 1974. Effect of size of food on food consumption: Some neurophysio- logical considerations. Journ. Comp. Physiol. Psychol. 86: 658 - 663 BEUKEMA, J. J. 1968. Predation by the three-spined stickleback (Gasterosteus aculea- tus L.), the influence of hunger and experience. Behavior 31: 1-126 Boietzky, SiGuRD von & MARIA VERENA von BoLeTzkKy 1969. First results in rearing Octopus joubini Robson, 1929. Verh. Naturforsch. Gesell. Basel 80: 56 - 61 Borer, K. T. 1971. Control of food intake in Octopus briareus Robson. Journ. Comp. Physiol. Psychol. 75: 171 - 185 Boucuer-Roponl, R. 1973. Vitesse de digestion de l’Octopus cyanea (Cephalopoda: Octo- poda). Mar. Biol. 18: 237 - 242 CampszLL, B. A., R. TEIGHTSOONIAN & R. A. WILLIAMS 1961. Activity, weight loss, and survival time of food deprived rats as a function of age. Corsit, D. J . & E. STELLAR 1964. Palatability, food intake, and obesity in normal and hyperphagic Journ. Comp. Physiol. Psychol. 54: 216-219 rats. Journ. Comp. Physiol. Psychol. 58: 63 - 67 DetuHier, V. G. 1976. The Hungry Fly. Harvard Univ. Press, Cambridge, Mass. Hinpg, R. A. 1970. Animal Behavior. McGraw-Hill, New York Ho.tne, C. S. 1966. The functional response of predators to prey density and its role in mimicry and population regulation. Mem. Entomol. Soc. Canada 48: 5 - 56 MANGOLD, KaTHarINA & SIGURD von BoLeTzKy 1973. New data on reproductive biology and growth of Octopus vul- garis. Mar. Biol. 19: 7 - 12 MATHER, JENNIFER 1972. A preliminary study of the behavior of Octopus joubint Robson. Unpubl. Master’s thesis, Florida State Univ. Nixon, M. 1966. Changes in body weight and intake of food by Octopus vulgaris. Journ. Zool. London 150: 1-9 Opresko, Lez & Ronatp FE THomas 1973. Observations on Octopus joubini: Some aspects of reproductive biology and growth. Mar. Biol. g1: 51 - 61 Rozin, P « J. MAYER 1961. Regulation of food intake in the goldfish. Physiol. 201: 968 - 974 Tuomas, Ronatp EF « Lez Opresxo 1973. Observations of Octopus joubini: Four laboratory reared gen- erations. The Nautilus 87: 61 - 65 Tranter, D. J. e O. AUGUSTINE 1973. Observations of the life history of the blue-ringed octopus Hap- lochalaena maculosa. Mar. Biol. 18: 115 - 128 Tucenpuart, B. 1960. The normal feeding behavior of the three-spined stickleback (Gasterosteus aculeatus L.). Behavior 15: 284 - 318 vAN HEUKELEM, W. 1973- Growth and lifespan of Octopus cyanea (Mollusca: Cephalo- poda). Journ. Zool. London 169: 299 - 315 Wacker, J. J., N. Lonco « M. E. BirTteERMAN 1970. The octopus in the laboratory: Handling, maintenance, training. Behav. Res. Meth. Instrum. 2: 15 - 18 WELLS, M. J. & J. Weis 1959. Hormonal control of sexual maturity in Octopus. Journ. Exp. Biol. 36: 1 - 33 Amer. Journ. Vol. 22; No. 3 THE VELIGER Page 291 Predation by a Rockfish, Sebastes chrysomelas, on Lamellaria diegoensis Dall, 1885 BY DOUGLAS E. HUNT Lockheed Center for Marine Research, 6350 Yarrow Drive, Suite A, Carlsbad, California 92008 Tue LAMELLARIDAE REPRESENT a family of marine mesogastropods that apparently rely upon cryptic adap- tations (GHISELIN, 1964; THOMPSON, 1973) and defen- sive acid secretions (THOMPSON, 1960; 1969) to avoid detection and falling prey to a variety of predatory fish and invertebrates. These gastropods can mimic their own prey organism, compound ascidians, as well as other in- vertebrates not commonly taken by predators, e. g., bar- nacles (GHISELIN, op. cit.; THOMPSON, 1973). These adaptations, as well as the production of strong defensive acidic fluids (approximate pH —1) have been thought to preclude predation of these gastropods by fishes, which THOMPSON (op. cit.) states “... are known to detest food which tastes acidic ...” On August 15, 1977 a female specimen of Sebastes chrysomelas (Jordan « Gilbert), measuring 245 mm, was collected during a food habits study in the vicinity of Diablo Canyon, San Luis Obispo County, California (120°51/23” W; 35°12'44”N). In the stomach contents of this individual was a single specimen of Lamellaria diegoensis Dall, 1885, the only prey item present. Past experience with 80 specimens of S. chrysomelas collected during the present study, as well as observations by Lar- SON (1972), had shown an apparent preference by this rockfish for Octopus spp. and small decapod crustaceans. The occurrence of Lamellaria diegoensis thus represents a unique food item never before documented. This raises several questions as to the location of the lamellarid when it was taken, and the feeding behavior of the fish: 1. Was the lamellarid residing on a tunicate when attacked by the fish? 2. If the lamellarid was mimick- ing anascidian, how was it detected by the fish? 3. Why would the fish feed on a prey so foreign to its usual diet, especially one that is capable of secreting strongly acidic fluids? Field observations indicate that Lamellaria diegoensis was observed most commonly on substrates other than compound ascidians (David Behrens, Diablo Canyon Research Laboratory, personal communication). In these instances the lamellarid failed to mimic the substrate and was Clearly visible. These observations could explain how the lamellarid was observed by the fish; however, the question of why the fish fed on the lamellarid, which can secrete strong acidic fluids, still remains. Continuing lab- oratory and field observations may provide the answer. ACKNOWLEDGMENTS I would like to thank Pacific Gas and Electric Company for permission to release this information. This study was funded through a contract with Lockheed Center for Marine Research. I am also grateful for the critical review of this manu- script by David Behrens and his positive identification of the specimen. Literature Cited GHISELIN, MiIcHAEL TENANT 1964. Morphological and behavioral concealing adaptations of Lamel- laria stearnstt, a marine prosobranch gastropod. The Veliger 6 (3): 123-124; plt. 16 (1 January 1964) Larson, RatpH J. 1972. The food habits of four kelpbed rockfishes Scorpaenidae, Seb- astes, off Santa Barbara, California. M. A. Thesis, Univ. Calif. Santa Barbara, 56 pp. THompson, THOMAS EVERETT 1960. Defensive acid secretion in marine gastropods. Biol. Assoc. U. K. 39: 115-122 1969. Acid secretion in Pacific Ocean gastropods. Zool. 17: 755 - 764 1973- Protective resemblances in British Lamellaria. 28: 75-78 Journ. Mar. Austral. Journ. Journ. Conch. Page 292 THE VELIGER Vol. 22; No. 3 Observations on Spawning in Calliostoma ligatum (Gould, 1849) BY DOUGLAS E. HUNT Lockheed Center for Marine Research, 6350 Yarrow Drive, Suite A, Carlsbad, California 92008 ALTHOUGH THE GENUS Calliostoma occurs commonly along the Pacific Coast of North America, the life histories of these trochids remain virtually unknown. Re- cently, several authors have contributed to the natural history of these snails: KEEN (1975) provided observa- tion on morphology and habits of several Calliostoma from California; PERRON (1975) reported on the feeding behavior of 3 Oregon species of Calliostoma; Lowry, McE roy « Pearse (1974) discussed distribution and habitat preference in a shallow subtidal environment at Monterey, California; SELLERS (1977) and Hunt (1977) discussed behavior and distribution of Calliostoma inhab- iting kelp beds in central California. Spawning behavior of the California species has not been previously docu- mented. On July 18, 1977, a single specimen of Calliostoma ligatum (Gould, 1849) was collected during a shallow water (3.6m) settling plate study at Diablo Canyon in San Luis Obispo County, California (120°51’23”W; 35°12'44” N). The animal was maintained in a container of seawater. The snail immediately moved to the water- air interface and remained stationary for 20 minutes, after which time green eggs were extruded from the right side of the mantle cavity. The eggs were liberated in a mucus sheath in groups of 10 to 12. Within 25 minutes the bottom of the aquarium contained large masses of green eggs. The spawning lasted for 2 hours and 3 min- utes, at which time the total number of eggs spawned was estimated at approximately 3 ooo. Each egg was spherical, 29 - 30 um in diameter. The granular green eggs had a single central yolk sac. Similar spawning behavior has been observed for an- other trochid, Tegula brunnea (Philippi, 1848), collected in Oregon during August (BELCHIK, 1965). Spawning occurred approximately 12 hours after collection in both male and female specimens of T: brunnea. The males were observed to “... discharge puffs of white sperm,” fol- lowed by spawning of moss-green eggs by the females. No male Calliostoma ligatum occurred on the settling plate on which the female specimen was found; however, these snails are common in the area and the possibility of spawning induced by a nearby male does exist. Spawning may also have been induced by the stress of being trans- ferred from the field to the laboratory, or by some other, unknown factor. ACKNOWLEDGMENTS I would like to thank Pacific Gas and Electric Company for permission to release this information. This study was funded through a contract with Lockheed Center for Marine Research. Literature Cited Beorx, Francis P 1965. Note on a range extension and observations on spawning im Tegula, a gastropod. The Veliger 7 (4): 233-234 (1 April 1965) Hunt, Dovatas E. 1977. Population dynamics of Tegula and Calliostoma in Carmel Bay, with special reference to kelp harvesting. M. A. Thesis, San Fran- cisco State Univ.; 81 pp. Keen, A. Myra 1975. On some West American species of Calliostoma. The Veliger 17 (4): 413-414 (1 April 1975) Lowry, Lioyp, A. McE.roy « J. S. Pearse 1974. The distribution of six species of gastropod molluscs in a Cali- fornia kelp forest. Biol. Bull. 147: 386 - 396 PErron, FRANK E. 1975- Carnivorous Calliostoma (Prosobranchia: Trochidae) from the northeastern Pacific. The Veliger 18 (1): 52-54; 1 plt. (1 July 1975) SeLverS, Rospert G. 1977- The diets of four species of Calliostoma (Gastropoda : Trochi- dae) and some aspects of their distribution within a kelpbed. M. S. Thesis, Stanford Univ., 31 pp. Le al Vol. 22; No. 3 THE VELIGER Page 293 NOTES & NEWS Trematodes in Chilean Fissurellid Molluscs MARTA BRETOS anp CLAUDINA JIRON Centro de Investigaciones Marinas Universidad del Norte Sede Iquique Casilla 65, Iquique, Chile MOLLUSCS ARE CLASSICALLY described as intermediate hosts for trematodes. Larval trematodes have been found in a number of gastropod species. In spite of this general statement, one of the authors found adult trematodes in the gonads of Fissurella crassa Lamarck, 1822 in Janu- ary 1976. After this discovery, we investigated the other large-sized Fissurella species living on northern Chilean coasts and we found these trematodes in the gonads of all of them. We have collected specimens of 8 species of Fissurella from several localities in northern Chile, situated between 20°16’S, 70°09/W and 20°41’S, 70°11’/W, from January 1976 until September 1979. ‘The animals examined varied in shell length from 25.3 mm to 111.2mm. The results of this study are summarized in Table 1. Samples of trema- Table 1 Number of analized animals of each species of Fissurrella and percentage of infected specimens. Species N % infected animals Fissurella bridgesii Reeve, 1849 35 71.43 costata Lesson, 1830 29 27.58 crassa Lamarck, 1822 136 13.97 cumingi Reeve, 1849 353, 64.87 latimarginata Sowerby, 1835 46 54.34 limbata Sowerby, 1835 36 47.22 maxima Sowerby, 1835 231 96.97 pulchra Sowerby, 1835 138 77.53 TOTAL 1004 todes from each species of Fisswrella were examined. With the kind assistance of Dr. J. B. Jones of the Min- istry of Agriculture and Fisheries, New Zealand, they were identified as digenetic trematodes belonging to the family Fellodistomidae and to the genus Proctoeces Odh- ner, 1911. The effect of parasitization on the reproduction of the hosts is now being studied in our laboratories. The importance of the present report is to make known the fact that gastropod molluscs can be hosts to adult digenetic trematodes. It is, thus, evident, that not only trematode larvae may be present in molluscs. Fissurella species are economically important in Chile because they are eaten by people. As far as we know, the trematodes mentioned above do not affect man; thus, their presence is not dangerous for man. Nevertheless, it should be noted that only the foot of the fissurellids is eaten; the viscera are always discarded. ERRATA Dr. Howard M. Feder has requested that we call the attention of our readers to the following errors which es- caped the proof readers: page 182, in the first paragraph of “Materials & Meth- ods” change January 1 to January 20; May 7 to May 18 (both 1973) ; and January 20, 1974 to January 26. In Figures 3 and 4 on pp. 185 and 186, respectively, the nuimbers in parentheses below months refer to dates; numbers in parentheses below boxes represent numbers of clams. New Rate Schedules At its regular meeting on October 17, 1979, the Execu- tive Board of the Society, after a careful evaluation of the financial situation, came to the conclusion that an adjustment in the subscription and dues schedules could no longer be avoided. This necessity was underscored by the fact that the volume-year 1978/1979, in spite of several generous donations received and stringent eco- nomizing, brought in effect a deficit of over $9000.00. Consequently, it was decided to increase the membership dues for members residing in the U.S.A. to $18.50; subscription rates for addresses in the U.S.A. are in- creased to $37.50; in both cases a charge of $1.50 for postage must be added. Page 294 THE VELIGER Vol. 22; No. 3 On the other hand, the rates for members and sub- scribers in all foreign countries can be kept the same as that charged for volume 22, 7. e., Swiss Francs 30.00 for dues, Swiss Francs 60.00 for subscriptions, plus, in all cases, Swiss Francs 7.00 for postage. We have learned that it is impossible in some countries to obtain checks in Swiss Francs; in such cases, the equivalent amount at the exchange rate, effective when the remittance is being made, in U.S. $ will be acceptable. However, we must insist that all remittances be made in a form that will not cause any expense to the Society as banks may charge as much as $5.00 for “collecting”. It should be obvious that we are absolutely unable to absorb such fees. We deeply regret the necessity of increasing the dues and subscription rates; our optimism that inflation would be curbed has proven unjustified. We can not fight this trend alone. 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Publication Date of THE VELIGER THE PUBLICATION DATE of The Veliger is the date printed on the index page; this applies even if the date falls on a legal holiday or on a Saturday or Sunday, days when the U.S. Postal Service does not expedite second class mail matter. That the printed date is the actual date of pub- lication under the rules of the International Commission on Zoological Nomenclature is based on the following facts: 1) The journal is delivered to the Post Office on the first day of each quarter, ready for dispatch; 2) at least three copies are mailed either as first class items or by air mail; 3) about 20 copies are delivered in person to the mail boxes or to the offices of members in the Berkeley area; 4) two copies are delivered to the re- ceiving department of the General Library of the Univer- sity of California in Berkeley. Thus our publication is available in the meaning of the Code of the ICZN. The printed publication date, therefore, may be relied upon for purposes of establishing priority of new taxa. REGARDING POSTAL SERVICE We are much disturbed by the steadily increasing num- ber of premature claims for supposedly “missing” issues of our journal. Since we have announced on numerous occasions that our journal is mailed on the dates printed in the issues, z.e., number 1 on July 1, number 2 on October 1, number 3 on January 1 and number 4 on April 1 of each volume year, it is unreasonable to expect delivery of the issues earlier than at least one week after these dates; however, a much longer time must be allowed for delivery to addresses at various distances from Berkeley. Thus, for example, a two weeks lapse is not unusual for as short distances as 500km; and up to 3 and 4 months must be counted on for addresses in the Far East and in Africa. We are faced with the alternative of not replying to what we must consider premature claims or, if the trend con- tinues, we must increase our subscription rates to cover these additional expenses. Our past efforts at keeping the subscription rate as low as possible are, we believe, suffi- cient evidence that we simply cannot afford any other course of action. The postal service causes us enough Vol. 22; No. 3 THE VELIGER Page 205 financial losses. Therefore we urgently request that before a claim is made, the time schedule be carefully checked. We are grateful for the understanding of this difficult situation shown by many librarians and will be grateful to those who, heretofore being perhaps eager to make sure that the library receives what is coming to it, will exercise a little patience. Your harassed Editor. Sale of C. M. S. Publications: Effective January 1, 1978, all back volumes still in print, both paper covered and cloth bound, will be available only from Mr. Arthur C. 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Supplements Supplement to Volume 3: $6.- {Part 1: Opisthobranch Mollusks of California by Prof. Ernst Marcus; Part 2: The Anaspidea of California by Prof. R. Beeman, and The Thecosomata and Gymnosomata of the Cali- fornia Current by Prof. John A. McGowan] [The two parts are available separately at $3.- each] Supplement to Volume 6: out of print. Supplement to Volume 7: available again; see announce- ment elsewhere in this issue. Supplement to Volume 11: $6.-. [The Biology of Acmaea by Prof. D. P. Assorrt et al., ed.] Supplement to Volume 14: $6.-. [The Northwest American Tellinidae by Dr. E. V. Coan] Supplement to Volume 16: $8.-. [The Panamic-Galapagan Epitoniidae by Mrs. Helen DuShane] Orders for any of the publications listed above should be sent directly to Mr. Art West. If orders are sent to us, we will forward them. This will necessarily result in delays. Other supplements: A Glossary of A Thousand-and-One Terms Used in Conchology by WiniFrep H. Arnoip originally published as a supplement to volume 7 of the Veliger has been reprinted and is now available from The Shell Cabinet, Post Office Box 29, Falls Church, Virginia 22046, U.S. A. The cost is US$ 3.50 postpaid if remittance is sent with the order. [A systematic Revision of the Recent Cypraeid Family Ovulidae by Crawrorp NEILL Cate] Supplement to Volume 15: Our stock is exhausted, but copies are still available from The Shell Cabinet, P. O. Box 29, Falls Church, Virginia 22046. [Growth Rates, Depth Preference and Ecological Succes- sion of Some Sessile Marine Invertebrates in Monterey Harbor by Dr. E. C. Haderlie] Supplement to Volume 17: Our stock of this supplement is exhausted. Copies may be obtained by applying to Dr. E. C. Haderlie, U. S. Naval Post-Graduate School, Mon- terey, CA (lifornia) 93940. Supplement to volume 18: $10.50 postage paid. [The Biology of Chitons by Robin Burnett et al.]. (Our supply of this supplement is exhausted; however, copies may be available by making application to the Secretary, Hopkins Marine Station, Pacific Grove, Cali- fornia 93950.) WE ARE PLEASED to announce that an agreement has been entered into by the California Malacozoological Society, Inc. with Mr. Steven J. Long for the production and sale of microfiche reproductions of all out-of-print editions of the publications of the Society. The microfiches are available as negative films (printed matter ap- pearing white on black background), 105mm X 148mm and can be supplied immediately. The following is a list of items now ready: THE VELIGER Vol. 22; No. 3 Page 2096 Volume 1: $1.50 Volume 6: $4.50 Volume 2: $3.00 Volume 7: $6.00 Volume 3: $3.00 Volume 8: $6.00 Volume 4: $4.50 Volume 10: $9.00 Volume 5: $4.50 Volume 11: $9.00 Volume 12: $9.00 Supplement to Volume 6: $1.50; to Volume 18: $3.00 California residents please add the appropriate amount for sales tax to the prices indicated. 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An issue of the journal will be designated as a Memorial Issue in honor of a person from whose estate the sum of $5000.- or more has been paid to the Veliger Endowment Fund. If the bequest is $25 o00.- or more, an entire volume will be dedicated to the memory of the decedent. CALIFORNIA MALACOZOOLOGICAL SOCIETY, Inc. is a non-profit educational corporation (Articles of In- corporation No. 463389 were filed January 6, 1964 in the office of the Secretary of State). The Society publishes a scientific quarterly, the VELIGER. Donations to the Society are used to pay a part of the production costs and thus to keep the subscription rate at a minimum. Donors may designate the Fund to which their contribution is to be credited: Operating Fund (available for current production) ; Savings Fund (available only for specified purposes, such as publication of especially long and signi- ficant papers); Endowment Fund (the income from which is available. The principal is irrevocably dedicated to scientific and educational purposes). Unassigned dona- tions will be used according to greatest need. Vol. 22; No. 3 Contributions to the C. M.S., Inc. are deductible by donors as provided in section 170 of the Internal Revenue Code (for Federal income tax purposes). Bequests, lega- cies, gifts, devices are deductible for Federal estate and gift tax purposes under section 2055, 2106, and 2522 of the Code. The Treasurer of the C. M. S., Inc. will issue suitable receipts which may be used by Donors to substan- tiate their respective tax deductions. Membership open to individuals only - no institutional or society memberships. Please send for membership ap- plication forms to the Manager or the Editor. Membership renewals are due on or before April 15 each year. If renewal payments are made after April 15 but before March 15 of the following year, there will be a re-instatement fee of $1.-. Members whose dues pay- ments (including the re-instatement fee) have not been received by the latter date, will be dropped from the rolls of the Society. They may rejoin by paying a new initiation fee. The volume(s) published during the time a member was in arrears may be purchased, if still available, at the regular full volume price plus applicable handling charges. BOOKS, PERIODICALS, PAMPHLETS The origin and evolution of the gastropod family Pomatiopsidae, with emphasis on the Mekong River Triculinae by Georce M. Davis. 120 pp.; 34 figs; 15 tables. Mono- graph no. 20, The Academy of Natural Sciences of Phila- delphia, 19 and Parkway, Philadelphia, PA 19103. US$12.00 1979 This monograph is a very good work on a very diffi- cult subject. One can not imagine a more difficult gastro- pod group to deal with than snails which are all tiny, less than 5mm long generally, with shells that show no easy identifying features. The anatomy of such small snails is just beyond the point of easy dissection with conven- tional tools and often necessitates serial sectioning. The author is also to be commended for his very modern, up- to-date treatment of this systematic group, taking into account multivariate analysis, plate tectonics, etc. Basically, this work deals with the pomatiopsid sub- family Triculinae, in the southeast Asian river systems, whose members were previously assigned to that frequent dumping ground for unknown small prosobranchs, the THE VELIGER Page 297 Hydrobiidae. In this pomatiopsid subfamily, the author finds that there are 11 genera and 92 species, making it possibly the largest contemporaneous endemic fresh- water fauna of any known lake or river system. This group of organisms was previously considered of the Hydrobiidae and not of the Pomatiopsidae because of the convergence of such structures as the shell, radula, penis, and operculum! (In this list of convergent characters one can see the usual favorite morphological entities em- ployed for most of the known taxonomic studies and as- signments.) Davis found that the amount of convergence toward the Hydrobiidae had been greatly underestimated and is revealed in this group only by a study of the female reproductive system as a whole. The origin of this fauna is now thought to have been Gondwanaland and that it was introduced into Asia by the collision of the Indian land mass with Asia. The sub- sequent Himalayan orogeny, resulting in streamcaptur- ing to form the great river drainages of the southeastern part of Asia, probably caused the ancestral forms of the Triculinae to be introduced into the (apparently) virgin river systems. The net long term result was an apparently rapid and abundant burst of speciation, resulting in to- day’s rich triculine fauna which went unrecognized as such until the present study. There are a few errors — mostly minor — in this treatise. One type of error is exemplified by page roo, in the reference to Krull, 1935, where “An atomische” is used instead of the correct “Anatomische.” In another case, in Appendix IV, the heading “Tribe Lacunopsini, Description of Taxa” is completely omitted, resulting in the inadvertent placement of ““Lacunopsis Deshayes, 1876” within the tribe Triculini) On page 37, the states of Croatia, Slavonia, Dalmatia, Serbia and Bosnia are de- scribed as “districts of Yugoslavia and Hungary,” where in actual fact none of these districts is located in Hun- gary; all are in Yugoslavia. My major complaint is about what is perhaps the most important figure in the entire work, that of figure 3 on page 1o. There is a number of cases where important structures are not labeled in figures C and E. Figure 3E is especially unclear in a number of respects. The relation- ship or connection between the bursa and the sperma- thecal duct (sd) is completely unclear since the duct which is labeled “‘sd” disappears under the oviduct and seminal receptacles. Is there a sperm duct in this figure? Also, there should have been presented here a renoperi- cardial opening since it is essential to the discussion of sperm routes, but no such structure is drawn or indicated. The significance of two tiny, unlabeled protuberances arising from the pericardium is also lost on the reader. Page 208 With respect to figure 3E again, the author states on page 17 that the “major unifying character of the Tricu- linae is the spermathecal duct extending from the bursa copulatrix to the posterior end of the mantle cavity (Fig. 3C-E).” If one looks at figure 3E, this criterion is clearly not satisfied by the Lacunopsini. The entire figure 3 should have been drawn much larger and more accurately with respect to enabling one to trace the various ducts. In another case, figure 18 on page 31 would have greatly benefitted by the inclusion of the bursa copulat- rix in each figure. An extended legend which would ex- plain the various drawings and their relationships, placed immediately next to the figure, would also have been most welcome. There are some general points which the author could have cleared up for the reader. For example, where are the ova fertilized (if known)? On more general terms, a brief statement about the known or presumed func- tions of the various intricate parts of the female repro- ductive complex would have been of tremendous help for a general understanding of what is actually going on in these organisms. Moreover, the reader can not help but wonder what is the author’s opinion about the sig- nificance of such large variety in the female anatomies of the Triculinae, especially in light of the fact that the author states that the male reproductive systems are so similar. If indeed the female genital systems are so varied and involved, is it really possible that the eggs of the various triculine species and genera are “‘so similar” that they can not be told apart at all (page 70) ? One wonders how closely the eggs were examined, especially since there are no illustrations of them. (Even if the eggs are all covered by similar sized grains of sand, this does not preclude the possibility that the egg case shape or sur- face sculpture, etc., could be species specific. ) There are a few highly speculative or weakly argued statements. For example, on page 41, the author states that “Yet other complexities such as spiral-noded sculp- ture or reticulate sculpture might evolve for species re- cognition in sympatry ...” Is there some factual basis for this kind of statement which could be cited or is it merely abstract or unfounded speculation, as it seems to be? An example of a weak argument occurs on page 44, where Davis first states the fact that “because of con- vergent evolution one cannot assume that if two fossil shells look alike they housed closely related animals,” but then later in the same paragraph states that “if several species of an endemic fossil assemblage appear to have survived to the present in the same region, there is a good probability that there is a phyletic connection be- tween the living taxa and the phenetically similar taxa” (italics by me, A. T.). Even in the context of the rest of THE VELIGER Vol. 22; No. 3 the page, this seems to be a circular argument or very close to one. However, I do not mean to be overly critical. Despite my few criticisms, the author is to be congratulated for having done a very significant work on a broad basis. This publication will be of interest not only to the few special- ists in this field, but should attract the attention of all malacologists generally interested in molluscan evolution because of its approach, style and contents. I recommend this work highly in spite of its apparent high price ($12.00) which it commands. Alex S. Tompa Museum of Zoology University of Michigan Ann Arbor, MI 48109 Bivalve mollusks of the western Beaufort Sea by FR. Bernarp. Contributions in Science, Natural His- tory Museum of Los Angeles County, no. 313. 80 pp.; 112 figs. $8.00, plus $1.25 postage and handling. 31 July 1979- The Beaufort Sea comprises Alaska’s Arctic shore. This paper summarizes work by the Oregon State University School of Oceanography, mainly dredgings at depths of 20 to 2500m on the continental slope of that area. It in- cludes also some notice of along-shore collections made by others. A total of 58 bivalve species are recognized, about half being of Pacific origin, half Atlantic. A key to the 22 fam- ilies is given, as also keys to genera and subgenera within some families. All of the species are illustrated by half tone photographs of good quality. Interiors of the right valves for the type species of each genus are shown in line drawings. One new genus, Boreacola, is proposed, in Montacuti- dae. Its type species, B. vadosa, also is proposed as new. A second new species is Axinulus careyt, in Thyasiridae. The list of publications in Literature Cited runs to nearly 12 pages — a good working bibliography on the Arctic Pleistocene and Recent molluscan fauna. An in- genious Appendix gives exact locations, by geographic coordinates and depth, and the repository and catalogue number for each specimen figured. This is an admirably organized and compact report that gives the essential information on morphology and distribution for the taxa discussed. A. Myra Keen oi Vol. 22; No. 1 THE VELIGER Page 19 Anatomy, Ecology and Distribution of the Volutidae and Volutomitridae of the Southern Indian Ocean ( Gastropoda : Prosobranchia ) PATRICK M. ARNAUD Station Marine d’Endoume, 13007 Marseille, France AND JEAN-JACQUES van MOL Université Libre de Bruxelles, 50 Avenue F. D. Roosevelt, 1050 - Bruxelles, Belgique (1 Plate; 10 Text figures) THREE RECENT BENTHIC SURVEYS have been done with the M/S Marion-Dufresne in the subantarctic waters of the Indian Ocean. During cruise MD.03 (1974) several stations were made (including 50 macrobenthic samples) off Kerguelen, Heard and Crozet Islands between shallow water and 4200m. Cruise MD.04 (1975) concentrated on the shelf around Kerguelen Islands, with 231 macro- benthic samples, and cruise MD.08 was mostly concerned with the Marion-Prince Edward shelf (49 macrobenthic samples) and the Crozet shelf (99 macrobenthic samples). We collected 3 species of volutid and volutomitrid gast- ropods at 65 stations, in 73 samples, by trawls, dredges and grabs (Table 1): Provocator pulcher Watson, 1882; Volutomitra curta (Strebel, 1908) and V. fragillima Wat- son, 1882 (Figures 1 to 3). Study of the plentiful speci- mens provides the first thorough insight into these 2 allied neogastropod families in this area. Many faunistic and biogeographic data are recorded and the resulting distri- bution of the 3 species obtained is mapped (Figures 7, 8). Our anatomical studies, made with the help of D. van Weert, have led to discussion of several of their charac- teristics and assessment of their positions in the 2 families involved. The egg capsule of the volutid Provocator pul- cher is also described and figured for the first time. Family Volutidae Provocator pulcher Watson, 1822 Provocator pulcher Watson, 1882: 330, 331; 1886: 260, plt. 13, fig. 5; CarcELLES, 1947: 6; PowELL, 1960: 156; WEAVER & DuPont, 1970: 123, fig. 53 F-G; CLover, 1975a: 10, 1 photo; 1975b: 1, 1 photo Provocator provocator SowERBY, 1887: 305, plt. 18, fig. 176; SMITH, 1942: 62, 63, plt. 13, fig. 95 Zidona (Provocator) pulchra. WENZ, 1943: 1350, fig. 3822 Specimens (empty shells recorded in parentheses) : — Kerguelen Islands MD.03: 7-CP4, 3; 11-CP7, (2); 13-CPg9, 3; 17-CB5, 12 (3); 21-CP14, 5 (2); and 2 egg capsules (incl. 1 empty) ; 24-CB6, 5 (1) MD.04: 7-CP13, 1 (2); 26-CP61, 2; 34-DC88, (1); 35-DC8g, 1; 38-CPg2, 1; 39-DCg3, (1) ; 45-DR106, (2); 7o-CP169, 1 egg capsule; 95-DC233, 1 egg capsule; 118-CP284, 1 (1) — Crozet Islands MD.08: 42-CP197, (1); 75-CP303, (3); 78-CP319, (2) Distribution (Figures 7, 8): The type locality is Kerguelen Islands, West Christmas Harbour, 48°43’S; 69°15’E, 105 fathoms [190m], vol- canic mud. Previously known only by empty shells (Wart- SON, 1882) and by shells from unknown stations off Ker- guelen (CLovER, 1975a, 1975b), the species is here re- Page 20 THE VELIGER. Figure 7 Present records of Volutidae and Volutomitridae off Kerguelen Is- lands and Heard Island: Provocator pulcher (1: living; 2: dead; 3: egg capsule), Volutomitra fragillima (4: living; 5: dead) and Volutomitra curta (6: living; 7: dead) Vol. 22; No. 1 Vol. 22; No. 1 THE VELIGER Page 25 yy aS) Ali u\ ae su am SONG . iG SS: y/ Figure 13 Provocator pulcher: valve of Leiblein; A, frontal section; B, trans- versal section A. Oes. — ariterior oesophagus Gr. — non-cili- ated groove M. Oes. — mid-oesophagus When crossing the nerve ring, the mid-oesophagus has a diameter of about 1.5 mm. Its walls are thin. The mus- cular layer is very weak at its beginning, the fibers have the same disposition as in the anterior part. The external layer of circular muscle fibers is thicker. The inner epi- thelium forms a few low ridges. At a short distance from the beginning, the gland of Leiblein (Figure 11) begins with swellings that appear on the dorsal wall of the oesoph- agus. The gland is progressively differentiated as a tubular, elongate organ coiled upon itself in a compact mass sur- rounding the oesophagus. The lumen is alveolar with mus- cular compartments. The gland is composed of 3 distinct parts. The first is lined with tall columnar gland cells with few mucocytes; the muscular layer is very thin. In the next part, the wall has a thick muscular layer and bears longitudinal and transverse ridges. The glandular epithelium is composed of a higher percentage of muco- cytes. The dead end of the gland forms a sac 2 to 2.5mm wide. Its wall is very thin and lined in mosi of its parts by a flattened epithelium. There are a few scattered low ridges of gland cells comprising some mucocytes. The had fixation of the tissues in this part of the digestive tract prevented better interpretation. Unidentified food remnants have been found in this part of the alimentary canal. Behind the gland of Leib- lein, the posterior part of the mid-oesophagus has a di- ameter of 1.9mm; its internal wall bears many longitudi- nal ridges. The posterior oesophagus is separated from the pre- ceding part by a constriction. Its section increases rapidly to a diameter of 2 to 2.5mm. The muscular layer is very thin, the epithelium consists of ciliated cells and muco- cytes. There are digitated internal ridges. The U-shaped stomach is a very simple sac with an internally plicated wall. The ridges of the dorsal part of the posterior oesophagus reach the single digestive gland aperture. The ventral ridges continue in the intestine without interruption. The intestine runs from the stomach along the right pallial wall. Its diameter is 2 to 3mm. The thin muscular layer is lined with an epithelium which is thickened into longitudinal ridges of columnar, ciliated gland cells with numerous dark granules. The mucocytes, scattered at the beginning, are much more abundant towards the end. The anal gland is a branching tubule derived from an invagination of the renal wall. Its cells are ciliated and granular. Discussion: With these observations, it is possible to determine the systematic position of the genus Provocator. This genus has been classified in the subfamily Zidoninae by CLENcH & TURNER (1964) on the basis of the morpho- Page 26 logy of the radula. This placement is corroborated by the following observations: - uniseriate radula with tricuspid teeth; - accessory salivary glands loosely bound around moder- ately compact salivary glands; - lobes at the bases of the siphon of equal development; - absence of operculum. It should be added that the genus Provocator could be considered more primitive than Alcithoe by the fact that its gland of Leiblein is less detached from the oesophageal wall than in the latter genus (compare the anatomical de- scription of Alcithoe arabica by PoNnDER, 1971). On the specific level, the very close similarity should be noted between the radular teeth of Provocator pulcher (Figure 12A) and P corderoi (cf. plt. 107 in CLENCH & TURNER, 1964) ; the teeth of the latter species are some- what more slender, with more arcuate bases. Family Volutomitridae Volutomitra curta (Strebel, 1908) Paradmete curta STREBEL, 1908: 23, plt. 3, figs. 34a-e; Pow- ELL, 1951: 166; CARCELLES, 1953: 196; PowELL, 1958: 198; 1960: 157 Paradmete longicauda STREBEL, 1908: 24, plt. 3, figs. 36a-b; PowELL, 1951: 165; CARCELLES, 1953: 196, plt. 3, fig. 70 Volutomitra (Paradmete) curta. CERNOHORSKY, 1970: 103, pit. 13, figs. 11-13 Specimens (all from Kerguelen Islands): MD.03: 3-CP2, 6 (1); 3-DS1, 1; 6-CP3, 3; 17-CB5, 70 (9); 21-CP14, 1; 24-CB6, 2 MD.04: 15-DC37, 1; 17-DC39, 1; 24-DR58, 1 (2); 38-CPo92, 1; 47-DC108, 1; 82-CP196, (1); 113-DC 269, (1); 115-DC275, 1 Distribution: The type locality is Shag Rock Bank, 53° 34'S, 43°23’W, 160m, gravel and sand, bottom tempera- ture +2.05°C. It was known from Shag Rock Bank, THE VELIGER Vol. 22; No. 1 South Georgia Island, Enderby Land, Mackenzie Sea. First recorded here (cf. Figure 7) from Kerguelen Islands, a major northward extension of range of this voluto- mitrid. Not obtained at Crozet Islands or Marion/Prince Edward Islands during our surveys. Bathymetric Range: Was known living from 75 to 300 m. Thus it is of interest to point out that this species was obtained living off Kerguelen Islands from 120m to as deep as 650m (present material). Ecology: Rather similar to that of Volutomitra fragil- lima (the 2 species are frequently collected together) but with a lower temperature-tolerance which would account for its absence from both the Crozet and Marion/Prince Edward shelves. Anatomy (Figures 14, 15): Animal rather small; max- imum height of the shell: 28.6mm. Tentacles cylindrical, bearing eyes on their posterior side. Operculum present. The Alimentary Canal: The pleurembolic proboscis is relatively long. The wall of the proboscis sheet is transversely plicate. The strong retractor muscles connect its sides to the lateral walls of the body. The ventral wall of the buccal cavity is pro- tected by a chitinous shield. The accessory salivary gland is an unpaired organ as in other Volutomitridae. This gland is quite small and entirely concealed in the probos- cis. Its thin secretory duct lies just behind the chitinous shield of the buccal floor and opens at the mouth aperture. Histologically the gland consists of a tube lined with cubic epithelium, the cells possessing large nuclei, and an outer layer of circular muscle fibers. Few gland cells are situ- ated in its terminal part. The radula is triseriate. The central tooth is quite large with a single thin lanceolate cusp and 2 long basal processes (Figure 12C). The lateral teeth are small and short. The salivary glands are small; the secretory tubules are composed of one type of cell. The salivary ducts enter the wall of the oesophagus in front of the valve of Leiblein. They open into the buccal cavity close to the junction of Explanation of Figures 1 to 6 Figure 1: Provocator pulcher, height 85mm, from Kerguelen Is- lands, MD. 03-24-CB6 Figure 2: Volutomitra curta, height 15.6mm, from Kerguelen Is- lands, MD. 03-17-CB5 Figure 3: Volutomitra fragillima, height 15.4 mm, from Kerguelen Islands, MD. 04-F51-DC119 Figure 4: Egg capsule of Provocator pulcher on a valve of Mal- letia gigantea measuring 47 X 28 mm; Kerguelen Islands, MD 04-Hg5-DC233 Figure 5: Two egg capsules of Provocator pulcher on an egg cap- sule of skate, Kerguelen Islands, MD. 03-21-CP14 Figure 6: The same capsules, enlarged Vol. 22; No. 1 THE VELIGER Page 27 Figure 14 Volutomitra curta: anterior part of the digestive tract. Abbrevia- tions as for Figure 11. the oesophagus. The ducts are densely ciliated, the cilia having a forward orientation. The anterior oesophagus is relatively narrow. There are 2 lateral ciliated ridges. Ventrally the epithelium is com- posed of flattened, nonciliated cells. Gland cells filled with red granules are in all this part of the digestive tract. The mid-oesophagus begins with the valve of Leib- lein. The tract is rather long and coiled upon itself, a strong muscular bundle is attached between 2 loops. The valve of Leiblein, situated well in front of the nerve ring, is quite typical. The ventral non-ciliated groove of the anterior oesophagus persists as a deep slit into the fold of the valve. This groove, ventral in the anterior part of the valve, moves progressively to the right and becomes dor- E — eye M -—- muscular bundles sal posteriorly. This evidence of torsion thus occupies the same level as in the Volutidae. Two bundles of longitudi- nal muscle fibers are developed into the wall of the valve, below the ridges lining the non-ciliated groove. The histo- logical organisation of the valve corresponds to the one observed in Provocator, except the 2 conspicuous muscu- lar bundles present in its walls. The mid-oesophagus pos- sesses a ciliated groove which corresponds to the pretor- sional dorsal alimentary tract of the anterior oesophagus. At its beginning, the wall of the mid-oesophagus is thin; later on, it becomes thicker by increase of the internal layer of circular muscle fibers. The numerous mucocytes and the red-stained cells that constitute the dorsal epi- thelium of the first part of the mid-oesophagus are pro- Page 28 Rad THE VELIGER CO imaniccnnatt Vol. 22; No. 1 Nhe Figure 15 Volutomitra curta: transverse section through proboscis. B.C. — buccal cavity Oe. — oesophagus gressively replaced by gland cells of irregular shape and filled with brown granules. These latter elements are re- stricted to the dorsal part of the lumen and are separated from the ventral ciliated groove by 2 low ciliated ridges. Further on the dorsal part is completely isolated and forms the gland of Leiblein. At its posterior end, the gland emerges out of the wall of the mid-oesophagus as a small Car. — subradular cartilage Gl. — common duct of accessory salivary glands Pr. — proboscis D. A. Sal. Rad. — radula Pr. S. — proboscis sheet translucent vesicle. Its histology seems to be uniform. The transition from the mid-oesophagus to the posterior oesophagus is visible externally as a rapid decrease in the external diameter of the duct. Nevertheless, the size of the lumen remains constant, as there is a considerable dimi- nution of the muscular layers of the wall. The lumen is lined with ciliated epithelium. THE VELIGER is open to original papers pertaining to any problem concerned with mollusks. This is meant to make facilities available for publication of original articles from a wide field of endeavor. Papers dealing with anatomical, cytological, distributional, ecological, histological, morphological, phys- iological, taxonomic, etc., aspects of marine, freshwater or terrestrial mollusks from any region, will be considered. Even topics only indi- rectly concerned with mollusks may be acceptable. In the unlikely event that space considerations make limitations necessary, papers dealing with mollusks from the Pacific region will be given priority. However, in this case the term “Pacific region” is to be most liberally interpreted. It is the editorial policy to preserve the individualistic writing style of the author; therefore any editorial changes in a manuscript will be sub- mitted to the author for his approval, before going to press. Short articles containing descriptions of new species or lesser taxa will be given preferential treatment in the speed of publication provided that arrangements have been made by the author for depositing the holotype with a recognized public Museum. Museum numbers of the type specimens must be included in the manuscript. Type localities must be defined as accurately as possible, with geographical longitudes and latitudes added. Short original papers, not exceeding 500 words, will be published in the column “NOTES & NEWS”; in this column will also appear notices of meetings of the American Malacological Union, as well as news items which are deemed of interest to our subscribers in general. Articles on “METHODS & TECHNIQUES” will be considered for publication in another column, provided that the information is complete and tech- niques and methods are capable of duplication by anyone carefully fol- lowing the description given. Such articles should be mainly original and deal with collecting, preparing, maintaining, studying, photo- graphing, etc., of mollusks or other invertebrates. A third column, en- titled “INFORMATION DESK,” will contain articles dealing with any problem pertaining to collecting, identifying, etc., in short, problems encountered by our readers. In contrast to other contributions, articles in this column do not necessarily contain new and original materials. Questions to the editor, which can be answered in this column, are in- vited. The column “BOOKS, PERIODICALS, PAMPHLETS” will attempt to bring reviews of new publications to the attention of our readers. Also, new timely articles may be listed by title only, if this is deemed expedient. Manuscripts should be typed in final form on a high grade white paper, 81/2” by 11”, double spaced and accompanied by a carbon copy. A pamphlet with detailed suggestions for preparing manuscripts intended for publication in THE VELIGER is available to authors upon request. A self-addressed envelope, sufficiently large to accom- modate the pamphlet (which measures 5/2” by 81/2’), with double first class postage, should be sent with the request to the Editor. EDITORIAL BOARD Dr. Donatp P. Azssorrt, Professor of Biology Hopkins Marine Station of Stanford University Dr. Warren O. Appicott, Research Geologist, U. S. Geological Survey, Menlo Park, California, and Consulting Professor of Paleontology, Stanford University Dr. Hans BertscH, Curator of Marine Invertebrates San Diego Museum of Natural History Dr. Jerry DonouuE, Professor of Chemistry University of Pennsylvania, Philadelphia, and Research Associate in the Allan Hancock Foundation University of Southern California, Los Angeles Dr. J. Wyatt Duruam, Professor of Paleontology Emeritus University of California, Berkeley, California Dr. Caper Hann, Professor of Zoology and Director, Bodega Marine Laboratory University of California, Berkeley, California Dr. Carote S. Hickman, Assistant Professor of Paleontology University of California, Berkeley, California Dr. A. Myra KEEN, Professor of Paleontology and Curator of Malacology, Emeritus Stanford University, Stanford, California Dr. Victor LoosanoFF, Senior Biologist, Emeritus U.S. National Marine Fisheries Service EDITOR-IN-CHIEF Dr. Rupo.r STOHLER, Research Zoologist, Emeritus University of California, Berkeley, California Dr. Joun McGowan, Professor of Oceanography Scripps Institution of Oceanography, La Jolla University of California at San Diego Dr. Franx A. Pire.xa, Professor of Zoology University of California, Berkeley, California Dr. Rospert Rosertson, Pilsbry Chair of Malacology Depariment of Malacology Academy of Natural Sciences of Philadelphia Dr. Peter U. Roppa, Chairman and Curator, Department of Geology California Academy of Sciences, San Francisco Dr. CrypeE F. E. Roper, Curator Department of Invertebrate Zoology (Mollusca) National Museum of Natural History Washington, D. C. Dr. JupirH Terry Smiru, Visiting Scholar Department of Geology, Stanford University Stanford, California Dr. Ratpu I. Smiru, Professor of Zoology University of California, Berkeley, California Dr. CuHar_es R. STASEK, Bodega Bay Institute Bodega Bay, California Dr. T. E. THompson, Reader in Zoology University of Bristol, England ASSOCIATE EDITOR Mrs. JEAN M. Cate Rancho Santa Fe, California ISSN 0042-3912 THE VELIGER A Quarterly published by CALIFORNIA MALACOZOOLOGICAL SOCIETY, INC. Berkeley, California ‘Ni Oat MARY TOC = = B E s, ‘ VOLUME 22 APRIL 1, 1980 NUMBER 4 CoNTENTS Egg Masses of Mollusca from the Mediterranean Waters of Israel and Notes on Reproduction of the Freshwater Species Theodoxus jordani and Melanoides tuberculata. (14 Plates; 1 Text figure) Av. Baraso & Z. ZENZIPER . . .. » 299 Aligena laterodentata, New Species, from the Caribbean Coast of Honduras nee. via : Leptonacea). (5 Text figures) Harotp W. Harry, JosepH C. Brirron & RopNey NuNLEY .. . . 318 The Lamellariidae of the North Eastern Pacific. (2 Plates; 15 Text four Davm W. BEBRENS . . . : » 323 Predation by the Prosobranch Mollusk Lamellaria ee on es, eae a Colonial Ascidian. (1 Text figure) GrETCHEN LAMBERT. . . Mei ieateuess dar tisects 340 Stone Boring Marine Bivalves from pees Bay, ation. (1 Text figure) E. C. HaperuE REDE Lae Votetben 26 Genetic Variation in Sympatric Sibling ay of Littorina. N. P Wirxins& D.O’RecAN .. . » » 355 Range Extension and Notes on the Feeding of the Nudibranch Okenia ie Puiu NIMESKERN, Jr. . . . . - . . 360 Two New Molluscan Species (Gastropoda : Muricidae) from the Copia Eastern Pacific. (1 Plate) ICEROV/EI" POORMANM Horr) | 2 fol op eng ee deg ied ints) folle)) si ee 6) 1, 2 GOL [Continued on Inside Front Cover] Distributed free to Members of the California Malacozoological Society, Inc. Subscriptions (by Volume only) payable in advance to Calif. Malacozool. Soc., Inc. Volume 23: $37.50 plus $1.50 for postage (U.S.A. only) For ALL foreign countries: Swiss Francs 60.- plus SF 7.- for postage Single copies this issue $22.00. Postage additional. Send subscription orders to California Malacozoological Society, Inc. 1584 Milvia Street, Berkeley, CA 94709, U.S. A. Address all other correspondence to Dr. R. Stoner, Editor, Department of Zoology University of California, Berkeley, CA 94720 Second Class Postage Paid at Berkeley, California ConTENTS — Continued On the Taxonomic Position, the Species and the Paleoecological Significance of the Genera Eubora, Toxosoma and Littoridina (?) in the Pliocene Pebas Forma- tion of the Upper Amazon Region (Gastropoda : Prosobranchia). (17 Text figures) Dietrich Kapotsky .. . 56 oo a Ghai Gastropod Mimicry by Another Pleustid heaouicoale in Central California. (1 Plate) Joun W: Carter's Davip W. BEHRENS ©). 5. 20) Us se) oy ee eee Early Development of Mytilopsis leucophaeata (Bivalvia : Dreissenacea). (1 Plate) Scort EB. SMDAEL) jos a) ce ee Maes oa RT Fish Predation on Pulmonate Limpets. (2 Text figures) Susan BiackFrorpD Cook .. . Ronee . . 380 Intermediate Forms and Range Extension of Pedicularia oe and Pedicularia ovuliformis. (3 Text figures) Rosert W.ScHMIEDER. . . eat 3 jie, ie eee age Helix aperta Introduced in Richmond, California vee Hinoaey (1 Text figure) Barry, RoTH & Dustin 1D, CuIvers! 9) 57-4 05) Spiroglyphus and Stoa, Taxonomic Problems in the Vermetidae. (1 Text figure) Ay MypRa Keen gies a es cel ce, Xen ee sn OG) NOTES & NEWS" ico) jE a eee eee Pee mene to) 5) GIOP Soviet Contributions to Malscoless in ane KENNETH J. Boss & Morris K. JACOBSON Sea Star Predation on Rock-Boring Bivalves. E. C. HaDERLIE BOOKS; PERIODICALS & (PAMPEUCEMS) 79 ems cuieci eo itete i ailec eraser Note: The various taxa above species are indicated by the use of different type styles as shown by the following examples, and by increasing indentation. ORDER, Suborder, DIVISION, Subdivision, SECTION, SUPERFAMILY, Famity, Subfamily, Genus, (Subgenus) New Taxa Vol. 22; No. 4 THE VELIGER Page 299 Egg Masses of Mollusca from Mediterranean Waters of Israel and Notes on Reproduction of the Freshwater Species Theodoxus jordani and Melanoides tuberculata BY AL. BARASH anp Z. ZENZIPER (14 Plates; 1 Text figure) INTRODUCTION A SERIES OF PAPERS on the molluscan fauna of the Medi- terranean coasts of Israel has been published during the last decades. But only a few observations deal with the re- production of Mollusca in this area (BARASH & DANIN, 1973; RAHAT, 1973). The purpose of the present work is to give a preliminary report on egg masses of marine mol- luscs found in the intertidal and infralittoral zones of the Mediterranean Israel and Sinai Peninsula. The Mediter- ranean coast, along which the material discussed here was collected, extends from Rosh-Haniqra in the North (Leb- anon border) to Bardawil (Sinai Peninsula), near the northern entrance of the Suez Canal (see map). Most egg masses were collected during field trips and expeditions, some were brought after months and even years of having been preserved in alcohol; accurate count- ing and measuring thus were often beyond our reach. In several cases, adult parent animals were put into aquaria, where they deposited their spawn, thus confirming the identity of the egg masses found in the sea. These in- cluded: Cerithium scabridum, C. rupestre, Columbella rustica, Bulla striata, Aplysia fasciata, Bursatella leachi savigniana, Elysia timida. The spawn of Rhinoclavis kochi was found only in the aquarium. Egg masses were first sent for identification in 1966 to the late Professor Gunnar Thorson at the Zoological Mu- seum of the University of Copenhagen. Unfortunately, his death interrupted his great scientific work and the materi- al was returned to us containing identified egg masses of 6 species. Dr. Klaus Bandel, Friedrich-Wilhelms University, Bonn, identified egg masses of 5 species, and Dr. Nellie B. Eales, University of Reading, identified the egg masses of 2 species of Aplysiidae. The greatest part of the material is kept in the collec- tion of the Tel-Aviv University, Department of Zoology. The egg masses of Cerithium rupestre and Elysia timida are in the collection of the Hebrew University of Jerusa- lem, Department of Zoology. The localities in which living adult specimens of the species treated in this paper were found are indicated; the vertical distribution of the species is also given. These data should be useful for future investigations of spawn of these species in the area. LIST or SPECIES CONSIDERED GASTROPODA PROSOBRANCHIA MESOGASTROPODA CrrITHIDAE Centthium scabridum Philippi, 1849 Cerithium rupestre Risso, 1826 Rhinoclavis kochi (Philippi, 1848) JANTHINDDAE Janthina ? nitens Menke, 1828 NATICIDAE 4 types, species not determined CassmDAE Cassidaria echinophora (Linnaeus, 1758) Semicassis undulata (Gmelin, 1791) THE VELIGER Vol. 22; No. 4 Page 300 33 Atlit Ma’gan Mikhael Caesarea MEDITERRANEAN SEA 32 Bardawil 31 33° 34° 35° Figure 33 Collecting Localities of Molluscan Egg Masses at the Coasts of Israel and the Sinai Peninsula ToNNDAE Pisania striata (Gmelin, 1791) Tonna galea (Linnaeus, 1758) NASSARDIDAE NEOGASTROPODA Sphaeronassa mutabilis On ee MuricmAzE Trunculariopsis trunculus (Linnaeus SEO EBS : : Fasciolaria lignaria (Linnaeus, 1758) 1758) Murex brandars Linnaeus, 1758 THADDAE ‘Thais haemastoma (Linnaeus, 1767) (Thats carinifera (Lamarck, 1822) CoLUMBELLIDAE Columbella rustica (Linnaeus, 1758) BUCCINIDAE Euthria cornea (Linnaeus, 1758) ConmAE Conus ventricosus Gmelin, 1791 OPISTHOBRANCHIA CEPHALASPIDEA (Bullomorpha) BULLDAE Bulla striata Bruguiére, 1789 Tue VELIGER, Vol. 22, No. 4 [BARASH & ZENZIPER] Figures 1 to 3 =o Vol. 22; No. 4 THE VELIGER Page 301 ANASPIDEA (Aplysiomorpha ) APLYSIDAE Aplysia fasciata Poiret, 1789 Bursatella leachi savigniana Audouin, 1826 SACOGLOSSA ELYsmDAE Elysia timida (Risso, 1818) NUDIBRANCHIA GLOSSODORIDIDAE ? Glossodoris sp. AxOLIDIDAE ? Aeolidiella sp. CEPHALOPODA DECAPODA SEPIDAE Sepia officinalis Linnaeus, 1758 Sepia elegans d’Orbigny, 1826 Cerithium scabridum Philippi, 1849 (Figure 7) Records: Barasy & DANIN, 1973: 309; fig. 7. Israel AyaL, 1978: 114-115. Israel Material: Bat-Yam — 14. X. 1972; intertidal zone, attached to algae Tel-Aviv University, aquarium: 3. X. 1972; 24. X. 1972; 25. X. 1972 Dor, Fisheries Research Station, aquarium: 15. XII. 1973 The egg masses of Cerithium scabridum were found fastened to solid objects (rocks, stones) in the habitat of the adults: rocky shores in the intertidal zone. About 100 specimens of Cerithium scabridum were kept alive in the marine aquarium of Tel-Aviv University. The snails were mostly attached to the vertical walls of the aquarium. The first egg mass was observed on the glass wall of the aquarium on 3. X. 1972. The ribbon-like greyish egg mass consisted of opaque, irregularly coiled strings, 4mm in diameter. The external limiting membrane of the string surrounds a thick gelatinous matrix in which egg cap- sules with transparent walls are embedded. Each capsule contains one egg (diameter 120m). A string contains 5,000 to 20000 eggs. The string laid on 24. X. 1972 was transferred on the same day into a petri dish for closer examination. Many of the eggs were already divided in two; the following days, some of the capsules appeared darker and ciliary movement could be seen in them. On 30. X. 1972, veli- gers appeared, some within the string, enclosed in the thin transparent membrane of the capsule, but most were swimming freely in the dish. The veligers had a distinct shell. Remnants of the disintegrated strings floated in the water. Cerithium scabridum is one of the earlier Indo-Pacific immigrants from the Red Sea into the Mediterranean. The first appearance on the Mediterranean coasts of Isra- el was recorded by PaLLary, 1912: 110; plt. 15. At present they are abundant there. Distribution of adult animals along the Mediterranean coasts of Israel: Akhziv, Shave-Ziyyon, Nahariya, Akko, Qiryat Hayyim, Qishon, Atlit, Caesarea, Dor, Habonim, Mikhmoret, Netanya, Tel-Barukh, Tel-Aviv, Yafo, Bat- Yam, Ashdod, Bardawil. Vertical Distribution: Intertidal zone. Cerithium rupestre Risso, 1826 (Figure 2) Record: BANDEL, 1975: 91. Banyuls-sur-Mer Material: Specimens of Cerithium rupestre taken from Shiqmona were kept in an aquarium of the Hebrew University, Jerusalem. The following data are based on the investigations of Dr. Y. AYAL (1978: 186-187). Spawn was observed in the aquarium during 1976; similar egg strings were deposited in September 1977. The number of eggs in a string varies between 100 and 700; the diameter of an egg is 400 zm. Development is direct; the juveniles hatch after develop- ment of about 15-25 days after spawning. According to the information on spawning of Cerithi- um rupestre at Banyuls-sur-Mer (BANDEL, 1975: 91), Page 302 THE VELIGER Vol. 22; No. 4 the gelatinous strings of the spawn contain 70-100 eggs (?), from which the juveniles hatch after development of about 3 weeks, having already completed metamorphosis within the egg capsules. Distribution of adult animals along the Mediterranean coast of Israel: Akhziv, Nahariyya, Shave-Ziyyon, Akko, Haifa, Bat-Gallim, Shiqmona, Atlit, Dor, Caesarea, Net- anya, Herzliyya. Vertical Distribution: Intertidal zone to 2m depth, rocky shores. Rhinoclavis kochi (Philippi, 1848) [= Cerithium kochi Philippi, 1848] (Figure 3) Record: BarasH & DANIN, 1977: 110; fig. 8. Israel. Material: Tel-Aviv University, aquarium, 28. IIT. 1975 Specimens of Rhinoclavis kochi taken from Haifa Bay were kept in the aquarium of Tel-Aviv University. On 28. HI. 1975 Mr. M. Tom found coiled strings on the sandy bottom of the aquarium. The strings appeared brownish yellow because of the sand grains adhering to them. Mi- nute, round vesicle-like capsules were observed by exam- ination under the microscope. The egg capsules were em-. bedded in a gelatmous matrix and were considerably smaller than the sand grains. Yolky eggs could be seen through the transparent wall of the capsules. Some of the eggs were in early cleavage stages (2 to 4 cells). Distribution of adult animals along the Mediterranean coast of Israel: Nahariyya, Akko, Oiryat-Hayyim, Haifa Bay, Karmel Beach, Shiqmona, Atlit, Dor, Alexander River, Netanya, Tel-Barukh, Rubin River, Ashdod, Ash- qelon, Nizanim, Yunis, Gaza. Vertical Distribution: a) Intertidal zone — rare; b) most- ly infralittoral, 10-101 m depth, sand, sandy mud. Rhinoclavis kochi is one of the most successful Indo- Pacific immigrants from the Red Sea into the Mediter- ranean, and is at present abundant in the Mediterranean waters of Israel. Janthina ? nitens Menke, 1828 (Figure 4) Record: FRAENKEL, 1927: 604-607; pit. III, figs. 1-4; text figs. 5-6 Material: Tel-Aviv, 24. XII. 1944 A snail with the raft and spawn attached to it was found by a student during a field trip to the sea shore. The material preserved in 70% alcohol shrank considerably. Nevertheless, we were able to distinguish the shell of the adult, the soft body, the raft connected with the foot and the brown egg capsules attached to the raft. ‘The egg capsules are attached to the underside of the raft by a mucous string. They are pear-shaped, fixed on the string by their narrow end. Capsules in which veligers are swimming appear brown because of the larval brown shell seen through the transparent capsular wall (fide FRAENKEL, 1927). ‘The only egg mass of the pelagic Janthina was found in a condition which did not allow identification as to its specific affinity because of the shrunken shell preserved in alcohol for years. We surmise that of the 3 species of Janthina known in the Mediterranean malacofauna of Israel, this spawn belongs to J. nitens which is not uncom- mon on our coasts. Another oviparous species, J. globosa (fide ROBERTSON, 1974: 218-219) is represented in the collections of Israel by one shell only. The common species of Janthina in Israel, J. bicolor, is probably ovoviviparous; egg capsules of this species were never found attached to its floating raft. NATICIDAE The flat, coiled ribbon-like egg masses or “collars,” encrusted with particles of the sea bottom, are easily rec- ognized as belonging to Naticidae. The eggs are deposited in regularly arranged egg spaces, the spaces lacking exit holes (the Jarvae are released when the collar becomes brittle and crumbles). Specimens of this family were not observed by the writers during spawning and thus were unable to refer the egg masses collected to definite species. They are divided here into 4 types according to the following char- acters: form of collar, number of whorls, particles of sea bottom, size of egg spaces and eggs. Type A (Figure 5) Material: Palmahim, 23. IV. 1977; cast ashore THE VELIGER, Vol. 22, No. 4 [BARASH & ZENZIPER] Figures 4 to 6b Figure 4 Tigure 5 Figure 6a Figure 6b Vol. 22; No. 4 The arched leathery, dark brown egg ribbon forms an almost complete “collar,” encrusted with fine particles of sand and mud. Measurements of the collar: 9.2 cm peri- phery, 2cm wide; 1.2mm thick. The egg spaces are ar- ranged in one plane with 25 spaces per cm’; each space contains one white oval egg (125 X go pm). Type B (Figures 6a, 6b) Material: Akhziv: 22. IV. 1960; 23. IV. 1966 Qiryat Hayyim: 14. III. 1969 Haifa: 14. III. 1964; 11. V. 1969 Dor: 7. III. 1969 Tel-Barukh: 2. III. 1962 The brittle egg collars were collected in a fragmented state (no complete collars). The colour is grey-pinkish; the egg mass Is 1.2 to 1.4mm thick; the egg spaces are arranged in a single plane relatively widely apart, 9 spaces per cm’. Each space contains one round, white-brownish egg (110X 100mm). Type C (Figures 7a, 7b, 7c) Material: Haifa Bay: 29. TV. 1968; 14m deep; 14. V. 1974; 24m deep Palmahim: 10. V. 1977; 20m deep; 21. IV. 1977, on beach Nizanim: 12. IV. 1977; 20m deep Many parts of collars of different sizes and one com- plete collar were collected, mostly by dredge. The com- plete collar is 10.5cm in periphery, 1.8cm wide, 1.2mm thick. The grey-brown ribbons of this type are encrusted THE VELIGER Page 303 with coarse sand grains. The outer and inner edges of the ribbon lack egg spaces; the outer edge is slightly undu- lated, 4-5.5mm broad; the inner edge is only 2.5-3mm broad. The thin-walled egg spaces are densely arranged in a single plane, bulging from the surface; there are 25 egg spaces per +cm’. Each space contains one white round egg (110X100 um). Type D (Figures 8a, 8b, 8c) Material: Nizanim: 12. IV. 1977; 10. V. 1977 Palmahim: 10. V. 1977. Infralittoral zone, 20 m deep The grey-brown collars were complete, 1.1 mm thick. Unlike the collars of other types, these are spirally twisted in 3 whorls. Similar to the ribbons of type C, the edges are lacking egg spaces and the outer edge is undulated. The outer whorl of the collars measures: 7.2m periphery, 2 cm wide (Nizanim, 10. V. 1977) 6.5 cm periphery, 2 cm wide (Nizanim, 12. V. 1977) I1.0em periphery, 2.5cm wide (Palmahim, ro. V. 1977) The egg spaces are arranged in a single plane, 16 spaces per 4cm’, each space contains a white, small oval egg (90X70 um). The Naticidae are represented in the Israel Mediter- ranean by g species. Very common are: Neverita jose- phina Risso, 1826; Naticarius millepunctatus (Lamarck, 1822) ; Tectonatica flammulata (Requien, 1848). Fairly common is also Naticarius dillwyni (Payraudeau, 1826). The egg masses in our collection may probably be assigned among these species. Type D of the egg masses of Naticidae seems to be similar to the egg masses of Natica (Mamma) mamilla Linnaeus described by GoHar & Ersawy (1967: 114-115; figs. 1, 2) from Al-Ghardaga on the north-western coast Measurements of the egg masses of four types of Naticidae Ribbon Egg spaces Eggs Type thickness granules density? cross section colour form size A 1.2mm 10 pm 25 0.8 X 0.7 mm white oval 125 X 90 um B 1.2-1.4mm 25 um 9 0.8 X 0.9mm white-brown rounded 110 X 100 um Cc 1.2mm 25 um 25 0.9 X 1.0 mm white rounded 110 X 100 um D 1.1mm 20 um 16 0.8 X 0.7 mm white oval 90 X 70 pum (for Plate Explanations see foldout at page 317) Page 304 THE VELIGER Vol. 22; No. 4 of the Red Sea. However, to our knowledge, N. mamilla was not mentioned so far among immigrants from the Red Sea into the Mediterranean (BARASH & DANIN, 1973, 1977). Cassidaria echinophora (Linnaeus, 1758) (Figures ga, gb, 9c) Records: Lamy, 1928: 114 Fioroni, 1966b: 684-688 ; Banyuls-sur-Mer; 60-80 m depth Material: [det. Klaus Bandel]} Caesarea — 21. III. 1962; one egg mass Mikhmoret — 27. III. 1962; 3 egg masses Ashdod — 8. VI. 1971; 1 egg mass The egg masses were found in the intertidal zone. They consist of cup-shaped capsules connected by a dense jelly-like substance, forming irregular clusters. The num- ber of capsules per cluster ranges from 4 to 75. The roundish capsules, 4mm high, contain numerous eggs visible through the translucent walls. “The capsules have no preformed escape aperture. Thus, the hatching snails are forced to burst a hole, fairly large , in the wall. Out of 155-193 eggs in a capsule only 7-23 young develop. ... The juveniles hatch in craw- ling stage.” (Fioroni, 1966b: 684, 839) Distribution of adult animals along the Mediterranean of Israel: Qiryat-Hayyim, Haifa, Shiqmona, Karmel Beach, Atlit, Dor, Caesarea, Hadera, Tel-Barukh, Tel- Aviv, Rubin River, Palmahim, Ashdod, Ashqelon, Gaza, Yunis, Tel-Arish, Bardawil. Vertical Distribution: Infralittoral, 22-220m depth, sandy mud, mud. Semicassis undulata (Gmelin, 1791) [Synonym: Cassts sulcosa (Bruguiére, 1792) ] (Figure 70) Record: Lamy, 1928: 114, Cassts sulcosa (Brug.) Material: [det. Klaus Bandel] Dor - 17. VIII. 1969, 1 egg mass, depth unknown Haifa Bay — 11. VI. 1975, 1 egg mass, depth 32 m Palmahim — 29. VI. 1977, 3 egg masses, depth 20m The egg mass of Semicassis undulata is a tower-like structure composed of pillar-like capsules arranged one above the other in regular layers. Each capsule is about 8mm high, the walls are semi-transparent and numerous yellowish eggs are seen within the capsules. The spawn dredged at Palmahim (29. VI. 1977) con- stitutes a block containing 3 egg masses. The largest com- prises 13 layers. The height of this egg mass is 6.5 cm, the width 5 cm at its lower end (which is attached to an alga), and 3.5cm at the free upper end. Two smaller egg masses adhere to the large egg mass at right angles; one of them consists of 3 layers of capsules, the other of 4. Distribution of adult animals along the Mediterranean coasts of Israel: Nahariyya, Qiryat-Hayyim, Haifa Bay, Karmel Beach, Shiqmona, Atlit, Habonim, Caesarea, Maagan Mikhael, Hadera, Netanya, Herzliyya, Tel- Barukh, Bat-Yam, Palmahim, Ashqelon, off Bardawil. Vertical Distribution: Infralittoral, 1.5-7m depth (live) ; 22-82m depth (shells). Tonna ? galea (Linnaeus, 1758) [= Dolium? galea (Linnaeus, 1758) } (Figure 71) Records: Lo Bianco, 1909: 635 Lamy, 1928: 115 Material: [det. Klaus Bandel] Turkey (Mediterranean coast) — 4. VIII. 1958, 1 egg mass, spaces with eggs Qishon — 14. III. 1964, 1 egg mass, honeycomb-like, egg spaces empty Dor — 15. VIII. 1971, 1 egg mass, spaces with eggs The egg masses form sheets of considerable size (up to 24cm long). It was impossible for us to state with certainty the size of the sheets since they were torn into strips (the measurements given below are approximate). The eggs are within spaces in the gelatinous matrix of the sheet, which is yellowish (Turkey) or pinkish (Dor). The oval, angulate (quadrigonal to hexagonal) spaces are regularly arranged in rows in a single plane. The spaces are clearly separated one from the other, their walls semi-trans- Tue VELIGER, Vol. 22, No. 4 [BARASH & ZENZIPER] Figures 7a to 7c Figure 7a A m ~ 7 ; os t = a - = - a ae 4 ; a ng i S = f = = ay i. Y _ = - ee Vol. 22; No. 4 THE VELIGER Page 305 parent’, numerous eggs are visible in the spaces, even in preserved material. According to THorson (1940: 192), there are 20-40 eggs in a space. Each space has a pre- formed exit-hole on the outer surface of the sheet. BaNnbEL, K. 1975: 94-95, in aquarium, Banyuls-sur- Mer Fioront, P. 1966b: 688, table 2, Banyuls-sur-Mer, to ca. 10m depth and in aquarium Egg mass Length (max.) Thickness Dimensions of egg spaces Egg spaces per cm? Turkey 24cm 2.0 mm 2.5 X 2.0mm 12 Israel: Qishon 4cm 1.4mm 1.0 X 0.8 mm 28 Dor 10 cm 1.6mm 1.5 X 1.0mm 16 It is worth quoting the remark of Dr. Bandel (personal communication, May 1977): “The sample of Turkey is quite like that of Jonna galea 1 have found in the Carib- bean Sea. ... The size of the capsules (spaces) is more dependent on the size of the producing female than on the species of the genus, therefore the three samples may be of the same species.” Egg masses of Tonnidae (Jonna galea) show great similarity to those of Naticidae, but without (sand) en- crustation. They form extensive sheets, the egg spaces are considerably larger than those of Naticidae, easily discerned on the surface. Dr. Bandel, who examined our material of Tonnidae, could not establish with certainty the identity of the spe- cies to which the egg masses in our collection (Turkey, Israel) belong. Since “they are quite like that of Jonna perdix (OSTERGAARD, 1950), Dolium costatum (KNup- SEN, 1940) and Dolium olearum (Gonar & EIsAwy, 1967) — all show more or less the same type of spawn” (personal communication, 1977). The common and almost only species of Tonnidae in the eastern Mediterranean (and in the Mediterranean in general) is Jonna galea (Dolium crosseanum is extremely rare), the egg masses treated in this paper obviously be- long to this species. Trunculariopsts trunculus (Linnaeus, 1758) [= Murex trunculus Linnaeus, 1758] (Figures 12a, r2b, 12c, 12d) Records: Fisuer, P. H. & A. RaFFy, 1933: 1-4; figs. 1, 2. Ban- yuls-sur-Mer, and Endoume ' Knupsen (1950: 98) on Dolium costatum: ‘The wall of the egg spaces consists of two layers — an external gelatinous, trans- parent layer 0.2 - 0.3mm thick, which goes without any inter- ruption from one egg space into the other, and an internal hya- line covering the inside of the egg space.” Material: Dor — 1. VI. 1965; 27. V. 1966; 12. V. 1969 Nahal Poleg — 26. VI. 1976 Herzliyya — 22. I. 1965; 5. VI. 1976; 19. VII. 1976 Tel-Aviv — 6. VII. 1965; 30. VI. 1976 Tel-Barukh — 27. VI.1970; 6.V.1977; 17. VI. 1977; 2. VIII. 1977; 21. X. 1977 Cyprus — 22. VIII. 1970 Egg masses cast ashore (apparently laid in shallow water) are commonly found all year round; according to our observations they are especially abundant in May- July. The egg masses of Trunculariopsis trunculus were reg- ularly collected during 1976 by Z. Zensiper at Tel-Ba- rukh, on the shore; measurements of these egg masses are as shown in small table at top of page 306. The egg capsules are closely attached to each other by their rather broad, branched base and form irregular clusters of several layers. When freshly deposited, their colour is horny and slightly transparent ; they become pale yellowish and less transparent later. The capsule is tongue- shaped, 5-6 mm high and 4-4.5mm wide. One side of the capsule is convex with longitudinal ridges on its perga- meneous wall. The other, the concave side, has a longi- tudinal depression where the escape aperture is found. Before hatching it is covered by a thin membrane. Hatching does not seem to be simultaneous and the same cluster contains full as well as empty capsules. In a cluster collected on 12 May 1976, 38 out of about 400 capsules were empty with distinct open escape apertures. In another cluster collected 2 weeks later, 89 out of about 300 capsules were empty; it is worth noting that a few of them were with ruptured walls and covered escape aperture. The egg mass collected in Cyprus contained juveniles before hatching. In the smooth brown embryonic shell the 2 last whorls and the siphonal canal were clearly seen. (see Figure 12d). Page 306 THE VELIGER Vol. 22; No. 4 Date Clusters Length Width Height 12. V. 1976 23 16.0-7.8 cm 11.2-6.0 cm 6.0-3.2 cm 5. VI. 1976 15 9.1-7.5 cm 6.1-5.2 cm 5.5-3.0 cm 12. VI. 1976 2 8.2-3.6 cm 7.2-2.5 cm 3.4-2.1 cm 19. VI. 1976 8 4.5-2.5 cm 3.1-2.4 cm 4.8-3.7 cm 26. VI. 1976 6 9.1-2.0 cm 5.0-3.2 cm 3.0-0.6 cm 30. VI. 1976 4 4.8-4.1 cm 3.5-2.7 cm 1.8-1.3 cm 12. VIII. 1976 1 3.5 cm 2.1 cm 1.4 cm NOTE on OVIPOSITION oF Trunculariopsis trunculus “Many females participate in forming the egg mass, so that the bulk of the egg mass comprises many hundreds of capsules. From each capsule, after about a month of de- velopment, hatch at the average 15 young snails, which absorbed during their development 20-40 eggs. The juve- niles leave their egg capsules in crawling stage, their metamorphosis was accomplished within the capsule.” (Banvet, 1975: 94). Distribution of adult animals along the Mediterranean coasts of Israel: Trunculariopss trunculus is common along the Israeli coasts, both in the intertidal and infra- tidal zones, from shallow waters to a depth of 84m. They are found mostly on soft substrates: sand, sandy mud, mud, and less on rocks. Intertidal zone (to 5m depth): Akhziv, Nahariyya, Akko, Qiryat-Hayyim, Bat-Gallim, Shiqmona, Dor, Cae- sarea, Mikhmoret, Netanya, Herzliyya, Tel-Barukh, Tel- Aviv, Ashdod. Infralittoral zone (dredged from 15-84 m depths) : Caesarea, Dor, Qiryat-Hayyim, Haifa Bay, Atlit, Bat- Yam, Alexander River, Rubin River, Gaza, Bardawil. Murex brandans Linnaeus, 1758 (Figures 13a, 13b, 13c) Record: Lo Bianco, 1909: 638. Naples Material: {det. Gunnar Thorson] The egg masses of Murex brandaris form irregular clusters of various size (2-15 cm in diameter in our collec- tion). The clusters comprise 20-300 cup-shaped, tough walled capsules. Each capsule is connected by a wide (1-2mm) membranous extension of the wall to 2 or more neighbouring capsules. The measurements of the larger capsules are approximately 3mm high, 4mm wide; the smaller are 2.5mm high, 2.8-3.0mm wide. The exterior side of each capsule is convex, the interior concave. In the centre of the concave side a round exit hole is clearly seen. Through the semi-transparent capsule wall yellowish eggs are visible in fresh spawn. The egg capsules dredged in Palmahim contained shelled veligers (4-9) before hatch- ing, in the crawling stage. The clusters of Murex brandaris are smaller than those of Trunculariopsis trunculus and are found rarely, though adult specimens of M. brandaris are very common, as common as T. trunculus. The scarcity of egg masses of M. brandaris was confirmed by the late Professor Gunnar Thorson (personal communication). Distribution of adult animals along the Mediterranean coasts of Israel: Akko, Haifa Bay, Dor, Zikhron Yaaqov, Caesarea, Alexander River, Tel-Aviv, Bardawil. In the dredge haul of Dor along with the spawn 11 living specimens of Murex brandaris were found. Vertical Distribution: Infralittoral zone, on sandy mud, sand, rarely on rocks or stones; at depths from 9-91 m. Thais haemastoma (Linnaeus, 1767) (Figure 14) Records: BaraSH & DANIN, 1973: fig. 11. Israel Capsules Localities Date Depth Egg masses | number content Haifa Bay 10. VII. 1974 73 m 2 60 many empty Dor 6. V. 1968 64m 1 73 with eggs Tel-Barukh VII. 1965 on rocky shore 1 70 mostly empty Tel-Aviv 6. VII. 1945 on rocky shore 1 about 200 mostly empty Palmahim 29. VI. 1977 80 m 1 250-300 with veligers THE VELIGER, Vol. 22, No. 4 [BarAsH &« ZENzIPER] Figures 8a to gb Figure 8a Figure 8b Figure 8c Figure 9a { si i Ki i 6 os vi if, & + =, € av ‘ é ey Vol. 22; No. 4 THE VELIGER Page 307 BANDEL, 1976b: 18. Santa Marta, Colombia, aquar- ium Material: Tel-Aviv: 6. VIII. 1957, 56 capsules; 8. VIII. 1957, 131 capsules; 2. VI. 1971, 85 capsules; intertidal zone The egg masses of Thais haemastoma consist of cap- sules laid in groups of 56 - 131. The parchment-like semi- transparent walls of the capsules are white, pink or purple. The freshly laid capsules are white, becoming yellow or pink afterwards and purple before hatching of the veliger. The capsules are fixed to a solid substrate (shells, algae), their flattened bases are confluent and have a common base for the group. They are elongated, pillar-like, nar- rowed at the base, broader at the top, 5-12mm_ high, with one side slightly concave. They stand erect side by side, the concave side of one capsule turned toward the _ convex side of its neighbour. The upper side of the capsule is flattened and almost semi-circular, surrounded by a pale narrow rim. A pre- formed round exit hole, covered with a transparent mem- brane, is situated slightly off centre on the upper side. The young leave through the preformed exit hole on the top when the membrane falls off. Distribution of adult animals along the Mediterranean coasts of Israel: Akhziv, Nahariyya, Akko, Qishon, Qiryat- Hayyim, Haifa, Bat-Gallim, Shiqmona, Habonim, Dor, Caesarea, Maagan Mikhael, Mikhmoret, Netanya, Ar- shaf, Herzliyya, Tel-Barukh, Tel-Aviv, Yafo, Bat-Yam, Palmahim, Ashdod, Yunis, Rafah. Vertical Distribution: Intertidal zone up to 4-9m depth. Thais carinifera (Lamarck, 1822) (Figure 14) Records: THORSON, 1940: 202; fig. 19. Iranian Gulf BarasuH & DaNIN, 1973: 313; fig. 10. Israel Material: [det. Gunnar Thorson] Tel-Aviv — 18. VIII. 1961; 21. XI. 1961; 16. VIII. 1965 ; intertidal zone The egg mass of 'Thais carinifera consists of a group of yellowish, pink or purple capsules with parchment-like, semi-transparent walls. The different colours of the cap- sules are due to the stages of the embryos inside them, just as in T: haemastoma. The capsules are fixed to a solid substrate (rock, algae, shell). They stand erect side by side, the broadened bases confluent. In large groups some of the capsules laid earlier may serve as a substratum, thus forming the first step towards an egg cluster (THORSON, 1940). The groups of capsules are of various sizes: the smallest consists of 15 capsules, the largest of about 180 (in our material). The cylindrical falciform capsules are slightly curved, nar- rowed at the top, where the exit hole is situated. The capsules are 8-11.5 mm high; their diameter near the top is 1.5-2 mm. “In this species 95-280 eggs of 0.2-0.22 mm in diameter are found in one capsule, all developing into embryos (no nurse eggs). ... Their size and well developed velum prove that the larva, when hatched will pass through a pelagic stage.” (THORSON, 1940: 204) Distribution of adult animals along the Mediterranean coasts of Israel: Akko, Qiryat-Hayyim, Karmel Beach, Shiqmona, Dor, Atlit, Herzliyya, Tel-Barukh, Tel-Aviv, Bat-Yam, Ashdod, Ashqelon, Yunis, Gaza, Bardawil. Vertical Distribution: Intertidal and infralittoral zones g-26m depth. 'Thais carinifera is an Indo-Pacific immigrant into the Mediterranean Sea and is now a resident in Israeli waters. Columbella rustica (Linnaeus, 1758) (Figure 15) Records: Baccl, 1947: 75-79; fig. 1. Naples KNUDSEN, 1950: 106-107; fig. 15. Cape Verde Isl. BANDEL, 1975: 100. Banyuls-sur-Mer Material: [det. Gunnar Thorson] Bat-Yam - 1. VIII. 1964, 7 capsules; VIII. 1969, 2 capsules; 15. VII. 1976, 43 capsules The egg masses were collected in the intertidal zone attached to algae, e. g., Caulerpa, Pterocladia. The egg masses of Columbella rustica consist of oval, boat-shaped (turned upside down) capsules laid singly and attached to the substrate by their flattened base one by one, often in groups of 2 to 11 capsules. The capsule is 3mm long and 2mm high. The walls are transparent with numerous small ridges and fine striae forming a net-like pattern. An oval exit hole, 1 mm long, is situated Page 308 on the topof the capsule, its thickened edges forming arim. The exit hole is conspicuous in empty capsules. The eggs are greenish with an average diameter of 160-180 um. Some capsules contained 2-19 embryos (which ingested during their development many nurse eggs) ; in 2 capsules a single large embryo with distinct shell was distinguished. “The juveniles escape after development lasting over 4 weeks in crawling stage. They accomplish the metamor- phosis within the capsule long before hatching.” (BANDEL, 1975) Distribution of adult animals along the Mediterranean coasts of Israel: The adult animals live on rocks, usually among algae; adult specimens were collected in the fol- lowing places: Intertidal zone: Akhziv, Akko, Bat-Gallim, Habonim, Dor, Caesarea, Mikhmoret, Netanya, Arshaf, Tel-Barukh, Bat- Yam. — Infralittoral zone: Haifa Bay, Akko, Dor, Alexander River — at a depth of 18-37 m, on rocks and sandy mud substrate. Euthria cornea (Linnaeus, 1758) (Figure 16) Records: Lo Bianco, 1909: 636. Naples Lamy, 1928: 53 BANDEL, 1975: 104. Banyuls-sur-Mer Material: [det. Gunnar Thorson] Akhziv — 21. VI. 1956; intertidal zone, 95 capsules with eggs, 11 empty Haifa — 13. II. 1965, ? , 9 capsules with veligers Haifa — 11. VII. 1974, 74m deep, 20 capsules, most with eggs The eggs of Euthria cornea are laid in capsules ar- ranged in several layers and united by their membranous extensions to clusters. The vase-like capsules are slightly flattened; their dimensions are: 4-5 mm long, 3.5-4.5 mm wide, narrowed at the base to 2.5 mm. The walls are semi- transparent, pergameneous. The number of eggs in each capsule varies from 8 to 22. All eggs will develop and hatch as veliconcha. — ‘The predatory E. cornea secretes in summer (June-July) an egg mass containing about 60 capsules similar to that of Murex. The spawn is composed of capsules containing 10-20 reddish eggs. After a development of over 3 weeks the juveniles that have just accomplished their metamorphosis escape.” (BANDEL, 1975: 104). THE VELIGER Vol. 22; No. 4 Distribution of adult animals along the Mediterranean coasts of Israel: Akhziv, Nahariyya, Akko, Bat=Gallim, Haifa Bay, Shiqmona, Dor, Atlit, Caesarea, Maagan Mikhael, Alexander River, Tel-Aviv, Rubin River, Pal- mahim, Ashqelon, Gaza, Yunis, Bardawil. Vertical Distribution: 1-70m deep; sandy mud; mud Pisania striata (Gmelin, 1791)- [= Pisania maculosa (Lamarck, 1822) | (Figure 17) Records: Fioront, 1966b: 701-704, tab. II. BanyulbsurMer, to 2m depth BANDEL, 1975: 101. Banyuls-sur-Mer Material: [det. Klaus Bandel] b Akhziv — 14. V. 1971; intertidal zone, 4 capsules. _, Akko — 16. VI. 1975; 1-2m deep on Petricola litho- phaga, 5 capsules The cup-like capsule, 3-4.mm in Sess, is flattened with a short peduncle at the base and a round exit hole on the upper side. In 3 capsules numerous small eggs could be seen; the others were filled with embryos. “The capsules contain 488-729 eggs of which 5 - 16 (extreme cases 24) develop” (Fioroni, 1966). The juveniles hatch in crawling stage. BER Distribution of adult animals along the Mediterranean coast of Israel: Akhziy, Akko, Bat-Gallim, Shiqmona, Dor, Atlit, Caesarea, Habonim, Mikhmoret, Netanya, Arshaf, Tel-Barukh, Bat-Yam, Palmahim; Asqelon. Individuals of Pzsania striata dwell on rocks and stones in the intertidal zone to 1-2 m depths. Sphaeronassa mutabilis (Linnaeus, 1758) [= Nassa mutabilis (Linnaeus, 1758) ] (Figures 18a, 18b) Records: Lo Bianco, 1909: 639. Naples Lamy, 1928: 92 FIoRONI, 1965a: 544 Material: [det. Gunnar Thorson] Off Haifa — 29. IV. 1968; dredged, 109 caus on Cymodocea, i in 18-27m Tue VELIGER, Vol. 22, No. 4 [BaRASH & ZENZIPER] Figures gc to 11 Figure 9c s i ™ age et ¢ >a, B92 ' D “% BO a Figure 11 "a a THE VELIGER, Vol. 22, No. 4 [BaRASH & ZENZIPER] Figures 12a to 12d Figure 12¢ Figure 12d icc Vol. 22; No. 4 Atlit — 15. V. 1962; intertidal zone, 10 capsules on algae Tel-Aviv — 28. II. 1963, 7 capsules on Caulerpa, washed ashore Yafo* — 14. IV. 1963. Intertidal zone, 17 capsules on Padina The flattened (antero-posteriorly compressed pouch- like capsules are fixed slightly obliquely to the substrate by their bases. They are arranged side by side in a line with equal spaces between them. The horn-coloured walls are semi-transparent with minute pleats on the sur- face. Each capsule of 3.5-4 mm _ height is narrowed at the top with thin undulated projections there. The egg masses usually comprise 7-17 capsules; the egg mass from Haifa consisted of 109 capsules on the same plant (presumably egg masses laid by several individuals) . “The eggs are more or less rich of yolk, swimming in protein fluid within the capsule. The absorption of pro- tein enables the hatching of veliconcha.” (Fioronq, - 1966: 655) Distribution of adult animals along the Mediterranean coasts of Israel: Nahariyya, Akko, Qishon, Haifa, Atlit, Dor, Caesarea, Habonim, Alexander River, Netanya, Ar- shaf, Tel-Barukh, Tel-Aviv, Yafo, Bat-Yam, Palmahim, Ashdod, Nizanim, Gaza, Yunis, El Arish, Rafah, Barda- wil. i Vertical Distribution: 8-88 m depths, sand, sandy mud, mud. * The egg mass collected at Yafo seems to be of another nassariid species (see Figure 19) THE VELIGER Page 309 Fasctolaria lignaria (Linnaeus, 1758) (Figures 20a, 20b) Records: Bacci, 1947: 79-80; fig. 2. Naples (aquarium) Lamy, 1928: 43 Lo Bianco, 1909: 636. Naples Material: [det. Gunnar Thorson] The egg masses of Fasciolaria lignaria consist of fun- nel-shaped, horn-yellowish capsules arranged in bunch- like groups of 8-33 (in one case a single capsule was found). The capsules are fixed to the hard substrate (rock, stone, shell) by slender peduncles 1.2-1.8 mm long. Their bases are confluent, forming a continuous thin retic- ular membrane covering the substrate. The capsules are 8- 11mm high; the widened upper end is circular, 5 mm in diameter, slightly concave, surrounded by a narrow rim. In the centre of the upper end is an exit hole, 1.5-1.6 mm in diameter, covered by a semi-transparent membrane. Eggs or embryos can be seen clearly through the mem- brane covering the exit hole, and also through the smooth translucent walls. In most of the empty capsules, the exit holes were open. However, in the empty capsules collected in Haifa Bay, the exit holes remained closed, but the walls were torn. Eggs in full capsules were numerous, but could not be counted in preserved material. The capsules collected in Dor (14. VI. 1961) contained embryos with shells (Figure 20b). “The embryo will feed on more eggs and hatch long after the end of metamorphosis.” (Bandel, in litt. 1977) Capsules Locality Date Zone Depth Number in cluster With eggs Empty eee ee I a Haifa Bay 31. VIII. 1974 infralittoral 51m 11 — 1] Atlit 14. VIII. 1965 intertidal tolm 19 12 7 Dor 16. VIII. 1950 intertidal tolm 1 — 1 Dor 14. VI. 1961 intertidal tolm 9 6 3 Dor 24. VII. 1969 intertidal tolm 8 5 3 Tel-Aviv 15.1IX 1961 intertidal tolm 41 33 8 Page 310 THE VELIGER Vol. 22; No. 4 Distribution of adult animals along the Mediterranean coasts of Israel: Nahariyya, Haifa Bay, Bat-Gallim, Kar- mel Beach, Shiqmona, Atlit, Caesarea, Mikhmoret, Net- anya, Tel-Barukh, Tel-Aviv, Palmahim, Ashqelon. Vertical Distribution: shallow waters up to 1-3 m depths and in infralittoral zone, 49-71 m depths Conus ventricosus Gmelin, 1791 (Figure 27) Record: BANDEL, 1975: 106-108. Banyuls-sur-Mer Material: [det. Klaus Bandel] Akhziv, 15. VI. 1971 — 1 group of 5 capsules Caesarea — 2. VII. 1967. 2 groups of 3 and 4 cap- sules Mikhmoret — g. VI. 1964. 3 groups of 3, 5 and 6 capsules All the egg masses were found in the intertidal zone attached to algae. The egg mass consists of white flattened flask-shaped capsules narrowed at their base, forming a group of 3-6. The caps iles are arranged parallel in a row, slightly inclined in on. direction. They are attached to the substrate (algae) by a very short peduncle extended into a broad basal membrane; the membranes are con- fluent into a common base of attachment. The capsules are 3.6-4.2mm high. Each capsule is convex on one side and concave on the other. The convex side of the capsule faces the concave side of its neighbour in the row. The walls are thin and tough, the concave side smooth, the convex side with longitudinal ridges (4 - 6). At the uppermost part of the capsule an elongated slit-like exit hole is covered by a transparent membrane and sur- rounded by a slightly corrugated collar-like border. “{The size of the capsules varies according to the size of the female. Each capsule contains an average of 11 eggs, which all develop. After a period of 3 weeks’ develop- ment the cover of the escape aperture is dissolved and crawling juveniles escape. They have accomplished their development within the egg capsule.” (BANDEL, 1975) Distribution of adult animals along the Mediterranean coasts of Israel: Intertidal zone: Akhziv, Akko, Karmel Beach, Shiqmona, Dor, Caesarea, Tel-Barukh, Bat-Yam, Palmahim. Infralittoral zone: to a depth of 24m, at Haifa Bay OPISTHOBRANCHIA Bulla striata Bruguiére, 1789 (Figures 22a, 22b) Material: Maagan-Mikhael, 19. IX. 1978 — sea water pond Eight adult specimens of Bulla striata were taken from a sea water pond near the shore at Maagan-Mikhael, S of Haifa, together with a flat yellowish-white ribbon-like egg mass. The animals and the egg mass were placed in an aquarium at Tel-Aviv University. After 2 days, on 21. IX. 1978, 4 additional egg masses were found attached by the length of one edge of the rib- bon to the walls of the aquarium. The ribbons were from 4 to 10cm long and 0.6cm wide. The egg capsules em- bedded in the gelatinous matrix were arranged in trans- verse double rows (hence the striated appearance of the ribbon). The number of egg capsules in a single row was 100-120; thus, 1cm length of the ribbon contains about 500 capsules. The capsules are round with smooth walls; their diameter is about 70 ym. In each capsule is one-egg. Pieces of ribbon were put into dishes for observation. On 25. IX. 1978, 4 days after spawning in the aquarium, rotation of embryos within the capsules was observed. Their body was yellowish with a conspicuous orange spot; the latter disappeared later. After 8 days, on 29. IX. 1978, the larvae were released from the capsules into the gelatinous coat of the string, whence they made their way into the water. Numerous veligers moved vividly outside the egg ribbon, their al- most globular transparent shell of 14 whorls clearly vis- ible. Many veligers remained alive and active for about 2 weeks. Gradually the ribbons disintegrated; many empty veliger shells were seen remaining in the dish. Numerous ciliates swarmed around the remnants of the egg ribbon. Similar observations were made on egg masses taken from the same pond on 3. X. 1978. Distribution of adult animals along the Mediterranean coasts of Israel: Live: 1) intertidal zone — Akhziv, Mikhmoret 2) infralittoral zone — Haifa Bay, 8-31m depth Shells: 1) on the beach throughout the coast 2) infralittoral — Akko, Haifa, Atlit, 15-31m depth The abundance of beach shells of Bulla striata in con- trast to the rare occurrence of living animals was very noticeable. THE VELIGER, Vol. 22, No. 4 [BaRASH & ZENZIPER] Figures 13a to 14 Figure 13a Figure 13¢ THE VELIGER, Vol. 22, No. 4 [BaRASH & ZENZIPER] Figures 15 to 17 Figure 17 Vol. 22; No. 4 Aplysia fasctata Poiret, 1789 (Figures 23a, 23b) Record: BEBBINGTON & THOMPSON, 1968: 4-5. Bassin d’Ar- cachon Material: [det. Nellie Eales] Akhziv — 24. IV. 1962; 25. VI. 1962; 8. VII. 1964; 28. VI. 1965; 9. VII. 1974 Netanya — 19. VI. 1969 Tel-Barukh -— 23. V. 1970; 3. V. 1975; 4. V. 1977; 7. V. 1977 Tel-Aviv — 20. V. 1971; 31. VII. 1973 Yafo — 6. V. 1962 Palmahim - 20. V. 1976 The egg masses of Aplysia fasciata were collected during April-July along the coast among algae (Ulva, Enteromorpha).'The spawn forms long cylindrical, loose- ly tangled, yarn-like strings; the largest egg mass con- sisted of about 40m of strings. Their colour is yellow- brown and the diameter is about 2mm. Rounded trans- parent capsules are embedded in the gelatinous matrix; these capsules are arranged 4-6 abreast in transverse rows throughout the entire length of the string. Numerous eggs could be seen through the capsule wall. In 10 capsules examined the numbers of eggs per capsule were as fol- lows: 36, 38, 40, 40, 40, 40, 42, 43, 46, 46, 48 (average ca. 42). Each ovum measured go-114 um. These results agree with those of BEBBINGTON & THOMPSON (1968), who found 118 capsules per centimeter length of the string and 43 ova per capsule; the total number of eggs in the mass 25877 400. Ciliary movement of the veliger was observed in the spawn collected on May 3 and 4, 1977. The embryonic shell of the larvae was of 2 whorls with an almost circular aperture (Figure 23b). In some of the larvae the opercu- lum could be seen on the the rear part of the foot. After hatching, free swimming veligers were found in the ob- servation dish. ““The embryonic period lasts 20-22 days.” (Eales, 1971, in litt.) Distribution of adult animals along the Mediterranean coasts of Israel: Akhziv, Akko, Haifa, Dor, Caesarea, Netanya, Arshaf, Tel-Barukh, Tel-Aviv, Yafo, Bat-Yam. Vertical Distribution: Intertidal zone; infralittoral zone (rarely) 9-241m deep. THE VELIGER Page 311 Bursatella leachi savigniana Audouin, 1826 (Figures 24, 25) Record: BarasH & DANIN, 1973: 329; fig. 13. Israel Material: [det. Nellie Eales] Dor — 6. VI. 1971, Fishery Research Station, salt water pond Dor — 21. III. 1971; 15. VI. 1971, Fishery Research Station, aquarium Maagan Mikhael — 31. I. 1978, salt water pond Tel-Aviv — 26. III. 1971, Tel-Aviv University aquar- ium All egg-laying adult specimens were taken from salt water ponds at Dor and Maagan Mikhael. The egg masses of Bursatella leachi savigniana are laid in gelatinous strings (2-3mm in diameter), that form a tangled green mass. The limiting membrane of the string and the gelatinous matrix are transparent, and the round membranous capsules embedded in the matrix are clearly visible. The capsules are quite regularly arranged along the string; in 1 centimeter of string there are 160-280 capsules. The eggs within the capsules are greenish. The eggs in a string deposited on 26. III. 1971 in a Tel-Aviv University aquarium were seen in their early cleavage stages; after 4 days ciliary movement of the veligers was observed. Two to 3 days later, veligers prior to hatching were seen with light brown shells. The string became brittle and free swimming veligers appeared in the observation dish. Similar observations on Bursatella leachi guineensis were reported by BEBBINGTON (1969: 334-338) : ‘There were from 2 to 6 eggs per capsule, 250 capsules per cm. ... Development within the egg string occupies 6 days. ... The embryos later become cloaked with cilia and they begin to rotate in their capsule (at 4 days) ... the larval shell appears (4-5 days).” Distribution of adult animals along the Mediterranean coasts of Israel: In shallow water, 2-3m depth: Dor, Mikhmoret, Ashdod; in sea water ponds, near the shore: Dor, Maagan Mikhael; infralittoral zone, 12-37 m depth: off Harfa, Bardawil. Page 312 THE VELIGER Vol. 22; No. 4 Elysia timida (Risso, 1818) Material: ‘ (Figure 26) 7. XI. 1976, 3 egg masses in aquaria, Tel-Aviv Uni- versity Record: Rawat, 1976: 186-193; fig. 2. Jerusalem Three adult specimens of Aeolidiella sp. were collected Matenl: at Bat-Yam on 2. XI. 1976 in the intertidal zone (infra- Mikhmoret — depth 0.2-1.0m on rocks covered with vegetation “A total of about 90 Elysia specimens were collected and from these more than 20 egg masses were obtained. ... Fourteen egg masses were followed through to the hatching of free juvenile larvae. ... In some egg capsules two eggs were found, and occasionally empty egg cap- sules were observed. After 2-3 days the eggs developed into a veliger form, which moved around in its egg cap- sule for another 1-2 days. ... Hatching took place after 16-21 days and was completed within 24 hours. Some juveniles left their shells behind, inside the egg mass, while others carried them around for several hours be- fore the shells were discarded.” (RAHAT, 1976: 188, 190) Distribution of adult animals along the Mediterranean coasts of Israel: Akhziv, Akko, Dor, Mikhmoret. Vertical Distribution: intertidal zone, depth of 0.2-1.0m Glossodoris sp. (Figure 27) Material: Mikhmoret — 12. X. 1971, attached to branches of an alga The small egg mass, 4mm wide, forms a flat coiled springlike ribbon folded upon itself in 3 volutions. The gelatinous matrix is transparent with minute white cap- sules of spherical shape densely embedded in it. The cap- sules are arranged in rows along the band, 3-4 capsules abreast in a row seen from above. Remarks: Glossodoris gracilis is one of 3 species of this genus reported from Israel. The egg mass found seems to be similar to that of G. gracilis (HAEFELFINGER, 1963: 706; fig. 1). Aeolidiella sp. (Figures 28a, 28b) Record: Tarby, 1969: 25 - 29; pit. 14, figs. 5, 6 littoral fringe) on rocks. One specimen was about 40 mm long, the 2 others were ca. 18mm. They were put into an aquarium for observation. On 7. XI. 1976, 3 ribbon- like pinkish egg masses were found attached to the walls of the aquarium: one egg mass, the longest one, with 24 coils (width 18mm), the second ring-like, the third not coiled, nearly straight (20mm long). The transparent mucous matrix of the egg mass is lim- ited by a thin membrane. Within the matrix are densely packed many pink-yellowish egg capsules. After 2 days, on g. XI. 1976, ciliary movements were observed within the capsules, and in many of them veligers with larval shells were clearly seen. CEPHALOPODA Sepia officinalis Linnaeus, 1758 (Figures 29a, 29b, 29c) Record: MANGoLp-Wirz, 1963: 101-102. Port Vendres Material: Yafo — 15. II. 1962, bunches of 12, 21, 43 eggs and 8 single eggs Yafo — 3. IV. 1962, 1 bunch of g eggs and 5 single eggs Nizanim — III. 1975, 1 bunch of 52 eggs Palmahim — 21. IV. 1975, 1 single egg Bardawil — 2. XI. 1975, 1 single egg Most egg masses were collected in the intertidal zone during February-April, except one egg taken by dredging in November 1977 at 48m depth off Bardawil. The eggs are large, about 8 mm long, 5 mm wide, black- brown, pear-shaped. The colour derives from the ink secreted on the eggs while they are shed. The eggs are attached to the algal branch (e. g. Cystoseira) by a flat- tened ribbon-like stalk. The stalk is twisted around the “stem” of the alga or around stalks of other eggs, thus forming a group resembling a bunch of grapes. Eight juveniles hatched from the eggs collected at Nizanim in March 1975. They were 10-11 mm long and 4-5 mm wide. The shape of the body was similar to that THE VELIGER, Vol. 22, No. 4 [BARASH & ZENZIPER] Figures 18a to 20b Figure 18 a Figure 18b Figure 19 Figure 20 a Figure 20 b W [BaRASH & ZENZIPER] Figures 21 to 22b Figure 27 Tue VELIGER, Vol. 22, No. 4 9] ot oi o =) 5) 0 ee a.) a ver x Vol. 22; No. 4 THE VELIGER Page 313 aaa aac rrcerertcr reeee een of the adult, the head tentacles and eyes distinct. Cuttle bones of the juveniles found in the preserved material had 8-11 growth lines (Figure 32c). ‘fThe egg masses are deposited in littoral waters on sandy or sandy mud bottoms at a depth of 1-30m. The animals prefer to attach their eggs to solid objects of elongated form. One female lays 200-550 eggs. ... The young Sepia adapts itself immediately to the benthic life of the adults without passing a planktonic phase.” (MaAn- GOLD-Wirz, 1963) Distribution of adult animals along the Mediterranean coasts of Israel: Qishon, Haifa, Caesarea, Atlit, Hadera, Mikhmoret, Alexander River, Netanya, Arshaf Tel- Barukh, Tel-Aviv, Yafo, Bat-Yam, Rubin River, Ashdod, Ashgqelon, Yunis, Gaza, El Arish, Bardawil. Vertical Distribution: 9-183 m depths, sand, sandy mud, rocks. Sepia elegans d’Orbigny, 1826 (Figures 30a, 30b) Records: BouLicAND, 1961: 589-594. Banyuls-sur-Mer Manco.p-Wirz, 1963: 116-117. Port Vendres Knupsen, Rusy G. & J. KNupDsEN, 1972: 83-97. Haifa (3 eggs, 11 juveniles) Material: Palmahim — X. 1977, at 65m, 2 eggs on a tube of a polychaete The eggs of Sepia elegans are deposited and attached singly; they do not form bunches as do those of S. 0 fficina- lis; they are smaller than the latter; they are nearly spherical, 5mm high, 4mm wide; they appear yellow because of the ample quantity of yolk within them as seen through the external transparent white-greyish mem- brane. On the free pole of the egg is found a narrow, unciform projection; similar projections are observed in eggs of many other cephalopod species. A ribbon-like peduncle, white-greyish as the outer membrane arises from the base of the egg and forms a ring around the substrate. The 2 eggs found in Palmahim were attached to a leathery polychaete tube, standing separately one from the other at a distance of a few millimeters. To our knowledge, the finding of an egg mass of S. elegans ona tube of Polychaeta tubicola, as in our case, has not been reported previously. BouLIGAND (1961) indicates Octo- corallia (Alcyonacea, Gorgonacea) as substrate for spawn of S. elegans. Distribution of adult animals along the Mediterranean coasts of Israel: Haifa, Dor, Caesarea, Tel-Aviv, Gaza, Bardawil. Vertical Distribution: in depths of 62-135 m Table 1 Months of collection or observation of the egg masses Species I II Ill IV V VI VII VIII 10.4 X XI XII Cerithium scabridum ; a6 Cerithium rupestre arp + Rhinoclavis kochi 15 ‘ Janthina nitens Naticidae type A F Naticidae type B 3F a1 ay Naticidae type C + 1 Naticidae type D as als Cassidaria echinophora oF ar Semicassts undulata ar oF Tonna galea + + Trunculariopsis trunculus + ats ar a oF a5 Murex brandaris af tp a Thais haemastoma ats ar Thais carinifera ate aF Columbella rustica alg oF Euthria cornea as ats ate Pisania striata ar ete Sphaeronassa mutabilis a" +r ar Fasciolaria ignaria ar 1 ats a Conus ventricosus WF ar Bulla striata af 2 Aplysia fasciata ar + oF AF Bursatella leachi savigniana ou ats a Z Sepia elegans 4 vf Sepia officinalis AF 1 1 Page 314 GUIDE To trot EGG MASSES or GASTROPODA CONSIDERED in tH1s PAPER Characteristic features differentiating the egg masses of marine gastropods studied in this paper are reviewed below: A. Egg masses laid on solid substrata: rocks, stones, al- gae, sand, mud a. Eggs within thin-walled, minute capsules (up to 2.5mm) orwithin spaces embedded in gelatinous matrix. Egg masses forming strings, ribbons, or sheets I. Egg masses encrusted with particles of sea bottom (sand, mud) 1. Egg masses forming convoluted strings, eggs in Capsulespe ees Rhinoclavis kochi (Figure 3) 2. Egg masses forming coiled ribbons - “collars,” EBRS IN SPACES. eeeenen Naticidae (Figures 5-8) II. Egg masses not encrusted with particles of sea bot- tom 1. Eggs within minute capsules (up to 1.5mm). * Egg masses cylindrical, long, coiled and tangled. — Egg strings yellowish-brown ..... Aplysia fasciata (Figure 23a) — Egg strings greenish ..... Bursatella leachi savig- niana (Figure 24) *#Egg masses ribbon-like — Ribbons not coiled, flat, yellowish white Bulla striata (Figure 22) — Ribbons irregularly coiled, greyish .......cccsecesse Cerithium scabridum (Figure 1) — Ribbons spirally coiled, white ..... Elysia timida (Figure 26) 2. Eggs within spaces. Egg masses forming sheets of considerable size ..... Tonna galea (Figure 11) b. Eggs within capsules with tough pergameneous walls, size moderate to large (2.5-13 mm) I. Egg capsules united to clusters composed of sev- eral layers or rows 1. Clusters tower-like; pillar-shaped capsules ar- ranged in regular layers (ti€rS) cscs Semicassis undulata (Figure ro) 2. Clusters irregular, capsules interconnected by pro- jections of their walls * No exit hole, capsule cup-shaped, walls without conspicuous ridges ..... Cassidaria echinophora (Figure 9) ** Exit hole on a side wall of the capsule; conspicu- ous ridges on the wall surface THE VELIGER Vol. 22; No. 4 — Capsules tongue-shaped .......... Murex brandaris and Trunculariopsis trunculus (Figures 12, 13) — Capsules almost triangular, vase-like, narrowed at their base ........... Euthria cornea (Figure 16) 3. Clusters bunch-like, exit hole at the top * Capsules pillar-like, broader and flattened at the topyes ees Thais haemastoma (Figure 14) ** Capsules cylindrical, falciform, narrowed at the topre. eee Thais carinifera (Figure 14) *** Capsules funnel-shaped, attached to the sub- strate by a slenderpeduncley..-3,nana eae Fasciolaria lignaria (Figure 20) II. Egg capsules in groups, standing in a row, side by side, attached by a basal membrane to the sub- strate; exit hole at the upper end 1. capsules pouch-like; exit hole surrounded by thin projections ... Sphaeronassa mutabilis (Figure 18) 2. Capsules compressed, flask-shaped; exit hole slit- likey ee eens Conus ventricosus (Figure 21) III. Egg capsules laid singly; exit hole at upper end 1. Capsules oval, attached by their flattened bases to the substrate. Walls transparent with numer- ous fine ridges ..... Columbella rustica (Figure 15) 2. Capsules cup-shaped, attached to the substrate by a short stalk ...... Pisania striata (Figure 17) B. Egg masses adhering by mucous strings to the under- side of the floating raft produced by the parent, a pelagic snail .......... Janthina nitens (Figure 4) SUMMARY The present paper intends to provide information on egg masses of molluscs from the south-eastern Mediterranean: Israel and North Sinai. The spawn was collected mainly during the warm season - March through August (see Table 1). The great majority of egg masses discussed in this work are of gastropods (over 20 species) ; from other molluscan classes only 2 species of cephalopods are discussed. The egg masses of gastropods belong to groups in which the eggs are protected by a cover of gelatinous material or by tough capsule walls (see list of species). The types of hatching stage in the development of the marine gastropods are: Aplysia fasciata Bursatella leachi Veliger (or crawling) Thais haemastoma Columbella rustica Crawling (or veliconcha) Veligers (pelagic) Cerithium scabridum Rhinoclavis kocht Tonna galea Thais carintfera Bulla striata THE VELIGER, Vol. 22, No. 4 [BarRASH & ZENZIPER] Figures 23a to 25 Figure 23a Figure 23'b Figure 24 7 bi] a 3 Tue VELIGER, Vol. 22, No. 4 [BARASH & ZENZIPER] Figures 26 to 28b Figure 28a Figure 28 b a nae Vol. 22; No. 4 Pisania striata Sphaeronassa mutabilis Fasciolaria lignania Conus ventricosus Elysia timida Cerithium rupestre Cassidaria echinophora Trunculariopsis trunculus Murex brandaris Euthria cornea Among these gastropods are 4 species of Indo-Pacific origin: Cerithium scabridum, Rhinoclavis kochi, Thais carinifera and Bursatella leachi savigniana. These species immigrated from the Red Sea through the Suez Canal and at present common along the Mediterranean coasts of Israel. All 4 species mentioned have free swimming THE VELIGER Page 315 veliger larve. The ability of the larvae to swim and to be carried by currents is obviously of advantage in the process of migration. From western Mediterranean egg masses or veligers of 195 species of mollusks were recorded. The number of species of which egg masses are known from the south-eastern Mediterranean (Israel and North Sinai) is thus but a fraction of the species investigated in the western Mediterranean. Spawn of many more species will likely be found in the course of properly organized in- investigation in the future. (A comparison of the respec- tive numbers is presented in the following table). Prosobranchia West Med. 72 Israel 213 Opisthobranchia Cephalopoda 81 42 6 2 Data based on the following publications: Italy (Naples) : Lo Branco, 1909; FRAENKEL, 1927; PALOMBI, 1939; Baccl, 1947 France (Banyuls-sur-Mer, Endoume, Villefranche) : FisHER, 1933; Franc, 1948; Bouticanp, 1961: HaEFELFINGER, 1969; BAN- DEL, 1975; MANGoLD-Wirz, 1976 Algiers: FRANC, 1948 Israel: BaRASH & DanINn, 1973; RABAT, 1976 Notes on the Reproduction of two Freshwater Gastropods Theodoxus jordani and Melanoides tuberculata Theodoxus jordani (Sowerby, 1832) [= Neritina jordani Sowerby, 1832] (Figures 37a, 3b) Egg capsules of 'Theodoxus jordani were collected near the shore of Lake Tiberias on 1. XI. 1966 and 1g. III. 1973 at a depth of 20-50cm. The capsules were attached to the shells of 'Theodoxus, living on hard substrata (stones) in the lake and to shells of Melanopsis. The capsules are opaque white, semi-globular, some- what flattened; their diameter is 1-1.2 mm. The capsular walls are rather thick and rough, composed of organic matrix covered with small sand grains. The capsule con- sists of 2 halves; the lower half is fastened to the sub- strate, the suture between the two halves is almost at the equator. Most capsules were “open” and empty, the up- per lid broken off, probably when the young escaped. cc “... small sand grains or diatom cases derived from the faeces stored in the rectum in a “Crystal sac” and poured over the capsule as it is laid. ... The capsules contain about 70 eggs each 110-125 um in diameter and uncoloured. Only one egg develops, the rest being used as food and the young snail hatches in crawling stage with a shell about 0.8mm in diameter.” (FRETTER & GraHaM, 1978: 105, on Theodoxus fluviatilis) Distribution of adult animals of Theodoxus jordan: in Israel: Freshwater bodies - Lakes (Hula, Tiberias) ; Rivers (Jordan); Canals (Bet-Shean) ; brooks, springs, ete. Melanoides tuberculata (Miller, 1774) (Figures 32a, 32b) Melanoides tuberculata is one of the most common gastro- pod species in the freshwater bodies of Israel, living on muddy and fine sandy bottoms. The first record on the reproduction of Melanoides tuberculata in Israel (from Page 316 THE VELIGER Vol. 22; No. 4 Lake Tiberias, Sea of Galilee) was published by TcHER- NOV (1975: 173). The snails treated in the present paper were observed in a freshwater aquarium of Tel-Aviv Uni- versity; they were originally taken from ponds in Neot- Hakikar (Dead Sea area). Hundreds of specimens have been living in aquaria for 2-3 years and reproducing there throughout the year. Embryos in the brood pouch were easily detected after crushing the shell of the snail. Shelled embryos were found particularly inside the body whorl; their size ranged from 1.5 to 2.2mm. The ovoviviparity of Melanoides tuberculata is well known (BINDER, 1959; Morrison, 1954) and of particu- lar interest is the phenomenon of parthenogenesis in this species (JAcoBs, 1957). The embryos developing in the mantle cavity leave the “mother” resembling the adult snails. Specimens were taken at random from the population of the aquarium; they were of various sizes, the largest being 18mm high. Almost every specimen of minimum height of 10mm contained embryos. The height of the adult animals and the number of shelled embryos ex- amined in 20 shells were as follows: ance in measurements. Thanks are due to Mr. A. Shoob, Miss L. Mamman and M. Winberg for their efforts to take the photographs, and Mr. S. Shaefer for preparing the map. Literature Cited Ayat, Y. 1978. Geographical distribution, ecological niche and the strategy of reproduction of the colonizer Cerithium scabridum Phil. (Gastropoda: Cerithiidae) as compared with those of some others sympatric non- colonizing congeneric species. Ph. D. thesis, Hebrew Univ. Jerusa- lem Baca, G. 1947. Le capsule ovigere di Columbella rustica (L.) e di Fasetolarta lignarta (L.). Boll. Zool. 14: 75 - 81 BanpeEL, Kraus 1974. Spawning and development of some Columbellidae from the Caribbean Sea of Colombia (South America). The Veliger 16 (3): 271-282; 15 text figs. (1 January 1974) 1975. Das Embryonalgehause mariner Prosobranchier der Region von Banyuls sur Mer. Vie Milieu 25 (1): 83-118 1976a. Observations on spawn, embryonic development, and ecology of some Caribbean lower Mesogastropoda (Mollusca). The Veliger 18 (3): 249-271; 25 text figs. (1 January 1976) 1976b. Morphologie der Gelege und 6kologische Beobachtungen an Muriciden (Gastropoda) aus der siidlichen Karibischen See. Verh. Naturf. Ges. Basel 85 (1/2): 1-32; 20 text figs. (31 March 1976) Adult height in mm 6 8 9 10 10 10 11 11.5 11.5 12 Number of embryos 0 0 0 0 2 3 2 3 4 4 Adults height in mm 12 Ws) 13 13 14 14 15 15 17 18 Number of embryos 5 4 4 4 6 0 5 8 6 13 “As a consequence of their parthenogenicity one indi- vidual suffices to found a new colony.” (STARMUHLNER, 1974: 161) Distribution of adult animals of Melanoides tuberculata in Israel: Hula, Lake Tiberias, Jordan River; freshwater bodies in Dead Sea area (slightly saline) ; rivers in the Mediterranean coastal plains. ACKNOWLEDGMENTS We express our gratitude to Dr. Klaus Bandel and Dr. Nellie Eales for examining a great part of the material treated in this paper. Our thanks and appreciation go to Dr. J. Knudsen for reading the manuscript, instructive remarks and encouraging advice. We are indebted to Professor M. Rahat and Dr. J. Ayal for the useful information on species they have ob- served during spawning. Many thanks to Dr. Ch. Levin- son and Professor L. Fishelson for providing material col- lected on expeditions, and to Mr. Y. Benayahu for assist- 1976c. Morphologie der Gelege und dkologische Beobachtungen an Buccinaceen (Gastropoda) aus der siidlichen Karibischen See. Bonn. zool. Beitr. 27: 98 - 133 1976d. Spawning, development and ecology of some higher Neogastro- poda from the Caribbean Sea of Colombia (South America). The Veliger 19 (2): 176-193; 17 text figs. (1 October 1976) BarasH, Au. & Z. DANIN 1977. Additions to the knowledge of Indo-Pacific Mollusca in the Mediterranean. “Conchiglie” 13 (5/6): 93-110 Barnarp, Keprpe, Harcourt 1963. Contributions to the knowledge of South African marine Mol- lusca. Ann. South Afr. Mus. Cape Town 47 (3): 107-108 BEBBINGTON, ALAN 1969. Bursatella leachi guineensts subsp. nov. (Gastropoda, Opistho- branchia) from Ghana. Proc. Malacol. Soc. London 38: 323 - 341 BEBBINGTON, ALAN & THOMAS EvERETT THOMPSON 1968. Note sur les opisthobranches du bassin d’Arcachon. Act. Soc. Linn. Bordeaux 105 (5): 4-5 Binper, E. 1959. Anatomie et systématique des Mélaniens d’Afrique occidentale (Moll. Gastropoda). Rev. Suisse Zool. 66 (32): 753 Bouticanp, Y. 1961. Le dispositif d’accrochage des oeufs de Sepia elegans sur Alcy- onium palmatum. Vie Milieu 12 (4): 589-594 D’Asaro, Cuarues N. 1970. Egg capsules of prosobranch molluscs from South Florida and the Bahamas and notes on spawning in the laboratory. Bull. Mar. Sci. 20: 414 - 440 Froront, Pio 1965a. Zur embryonalen Entwicklung und zum Schliipfzustand von zwei mediterraneen Nassa Arten. Rev. Suisse Zool. 72 (17): 543 to 568 1965b. Zur embryonalen Entwicklung von Philbertia (Gastropoda, Prosobranchia, Conidae). Verh. Naturf. Ges. Basel 76 (2): 207 to 219 THE VELIGER, Vol. 22, No. 4 [BarasH & ZENZIPER] Figures 29a to 29c Figure 29 4 Figure 29 c THE VELIGER, Vol. 22, No. 4 [BaRASH & ZENZIPER] Figures 30a to 32b Figure 31b Figure 32b Vol. 22; No. 4 f Plate Figures 1966a. Un nouveau cas de rotation des oeufs 7 ropode prosobranche marin. Vie Milieu 172 1966b. Zur Morphologie und Embryogenese des transitorischen Organe bei Prosobranchiern (Mo Rev. Suisse Zool. 73 (44): 621 - 876 FisHer, Paut Henri @ A. Rarry { 1938: La ponte du Murex trunculus L. Monaco No. 618: 1-4 FRAENKEL, Gottrriep S. 1927. Biologische Beobachtungen an Janthina. | Okol. Tiere 7 (4): 597 - 608 | FRETTER, VERA & ALASTAIR GRAHAM 1962. _—‘ British prosobranch molluscs. Ray Si 1964. Reproduction. pp. 127-164 in: K. Yonge, eds. Physiology of Mollusca, vol. 1. A 1978, The prosobranch molluscs of Britain and) Mollusc. Stud. Suppl. 5: 102 - 106 Germain, Louis | 1921. Mollusques terrestres et fluviatiles de Sy 453 - 518 | Gowar, H. A. FE « A. M. Ersawy 1967. The egg masses and development of fo branchs from the Red Sea. Publ. Mar. Biol. HAEFELFINGER, HANS-RUDOLF 1969. Contribution 4 la systématique des Glossq néens (Gastropoda, Opisthobranchia). Rev; to 710 Hyman, Lrssre HENRIETTA 1967. The Invertebrates. 6, Mollusca 1: vii+79 Hill Book Co., N. Y., St. Louis, London | Jacoss, J. | 1957- Cytological studies of Melaniidae (Mollus ence to parthenogenesis and polyploidy. I. Oog¢ genetic species of Melanoides (Prosobranchia-Gi Roy. Soc. Edinb. 63 (II) (16): 341 - 352 | 1958. Cytological studies of Melaniidae (Mollus ence to parthenogenesis and polyploidy. II. Stud} males of the polyploid race of Melanoides it) lineatus. Trans. Roy Soc. Edinb. 63 (II) ( Knupsen, Jorcen 1950. Egg capsules and development of som from tropical West Africa. Dan. Sci. Press Figure 18a: Sphaeronassa mutabilis (Linnaeus, 1758), egg masa Figure 18b: Sphaeronassa mutabilis (Linnaeus, 1758), 2 single capsules Figure 1g: Nassariid species, egg mass Figure 20a: Fasciolaria lignaria (Linnaeus, 1758), @ group of capsules Figure 20b: Fasciolaria lignaria (Linnaeus, 1758), embryos before hatching (taken from a capsule) Figure 21: Conus ventricosus Gmelin, 1791, egg mass Figure 22a: Bulla striata Bruguiére, 1789, egg ribbon Figure 22b: Bulla striata Bruguiére, 1789, capsules with veligera Figure 23a: Aplysia fasciata Poiret, 1789, spawn Figure 23b: Aplysia fasciata Poiret, 1789, veligers Figure 24: Bursatella leachi savigniana Audouin, 1826, adult and spawn Figure 25: Bursatella leachi savigniana Audouin, 1826, pasts of egg string Figure 26: Elysia timida (Risso, 1818), egg mass (after M. Rahat) Figure 27: ? Glossodoris sp., egg mass Figure 28a: ? Aeolidiella sp., adult Figure 28b: ?Aeolidiella sp., egg mass Figure 29a: Sepia officinalis Linnaeus, 1758, eggs Figure 29b: Sepia officinalis Linnaeus, 1758, juveniles (about t week old) Figure 29c: Sepia officinalis Linnaeus, 1758, cuttle bone of juvenile Figure joa: Sepia elegans d’Orbigny, 1826 Figure job: Sepia elegans d’Orbigny, 1826, an egg attached to 2 tube of a polychaete Figure 31a: Theodoxus jordani (Sowerby), egg capsules attached to shells of adults Figure grb: Capsules of Theodoxus jordani (Sowerby) on Melan- opsis praemorsa Figure 32a: Melanoides tuberculata (Miller), adult female Figure 32b: Melanoides tuberculata (Miiller), juveniles taken from female’s brood pouch 5 ; { \ ‘i , ‘ 6 \ ty t ‘ - F a : s on iy ‘ es Vol. 22; No. 4 g6Ss. Un nouveau cas de rotation des oeufs nutritifs chez un gast- : ropode prosobranche mann. Vie Milieu 17: 109-119 1965b. Zur Morphologie und Embryogenese des Darmtraktes und der seeesiterischen Organe bei Prosobranchiern (Mollusca, Gastropoda). Rev. Suisse Zool. 73 (4s): 621-876 Paut Henai & A. Rarry tee “Va ponte du Murex frunculus L, “Monsco No. 618: 1-4 Frasnxsi, Gortrro S. : ; 1937- Biologische Beobachtungen an Jenthina. Zeitschr. Morph. Okol. Tiere 7 (4): 5877608 = petTes, V! Avastair GRAHAM ree Beith pres Ray Soc. London: 386-476 . British prosobranch molluscs. irs Reproduction. pp. 127-164 m: K. M. Wilbur & GC. M. eee eds. Physiology of Mollusca, vol. 1. Academic Press, New York 1978. The prosobranch molluscs of Britain and Denmark. Journ. Mollusc. Stud. Suppl 5: 102-106 Grauary, Louis Bull. Inst. Océanogr. reat. Mollusques terrestres et fluviatiles de Syrie. Paris, vol. 1 455-518 Gomis, HA. Ee A M. Eisawy 1967. The egg masses and development of four taenioglossan proso- branchs fom the Red Sea. Publ Mar. Biol. Sta 14: 109-147 Hagreuriscrs, Haxs-RuporF R 1969. Contribution 4 la systématique des Glossodoridiens meditterran- néens (Gastropoda, Opisthobranchia). Rev. Suisse Zool. 76: 703 t 710 Brus, Lessst Henprerta ; 1957. The Invertebrates. 6, Mollusca 1: viit792 pp.; iust McGraw- Hill Bock Co, N. Y, St Louis, London Melaniidae (Mollusca) with special refer- dy. I. Oogenesis of the partheno- sobranchia-Gastropoda). Trans. 352 y- II. Study of meiosis in the rare les tuberculatus and Melanoides Ezestus. Trans. Roy Soc. Edinb. 63 (II) (20): 433-443 Exvpszn, Jezczn 1950. Egg capsules and development of some marine prosobranchs from tropical West Africa. Dan. Sci Press Copenhagen: 88-116 THE VELIGER Page 317 Lamy, Bpouarp 1998. La ponte chez les gasteropodes prosobranches. ourn. di Conchyl. Paris 79: 25-52; 80-196 J : Lxsour, Manx V. 1937. The eggs and larvae of the British prosobranchs. Journ. mar. Biol, Assoc. U. K. 99: 197-158 Lo Bianco, S. 1909. Notizie biologiche riguardanti specialmente il periodo di ma- tunita sessuale degli animali del golfo di Napoli. Mitteil. a. d. zool. Station Neapel 19 (4): 619-657 MANcaLp- Wirz, KATHARINA 1963. _ Biologie des céphalopodes benthiques et nectoniques de la Mer Catalane. Vie et Milieu, suppl. 1g: 1-285 Morrison, Joseru Paut Evprep 1954. The relationships of old and new world melanians. Proc, U. S. Nat. Mus. 10g (3325): 357-394 Patomai, A. 1939. Uova e larve di Cerithium rupestre Risso (Gastropoda, Proso- branchia). Napoli Stazione Zoolo. 17: 209-212 Rawat, M. 1976. Direct development and symbiotic chloroplasts in Elysia timida (Mollusca, Opisthobranchia). Israel Journ. Zool. 25: 186-193 Ronzrtson, Ropert 1974. Marine prosobranch gastropods: larval studies and systematics. Thalassia Jugosl. 10 (1/2): 213-238 Rusy, G. # Jornozn KNupsen 1972. Cephalopoda from the Eastern Mediterranean. Israel Journ. Zool. 21: 83 - 97 STaARMUBLNER, F. 1974- The freshwater gastropods of Ceylon. Sri Lanca (Ceylon) 25 (1, 2): 161 Taspy, Jean 1969. Etude systématique et biologique sur trois espéces d’Aeolidielles des cdtes européennes (Gastéropodes, Nudibranches). Bull. Inst. Océanogr. Monaco 68 (1389): 1-40; plts. 1-15 Tcuernoy, E. 1975- The molluscs of the Sea of Galilee. Bull. Fish. Res. Sta. Malacologia 15 (1): 171-173 (18 December 1975) Tuorson, GUNNAR 1940 Studies on the egg masses and larval development of gastro- pods from the Iranian Gulf 159-238 Dan. Sci. Invest. Iran prt. IT: Explanation of Plate Figures : Cerithium scabridum Philippi, 1849, adult and egg mass : Cerithium rupestre Risso, 1826, egg mass : Rhinoclavis kochi (Philippi, 1848), egg mass r 2 3 4: Janthina nitens Menke, 1828, adult and egg mass 5: Naticidae, type A, egg ribbon (collar) 6a: Naticidae, type B, collar fragment 6b: Naticidae, type B, eggs za: Naticidae, type C, collar 7b: Naticidae, type C, egg spaces 7c: Naticidae, type C, eggs 8a: Naticidae, type D, collars 8b: Naticidae, type D, egg spaces 8c: Naticidae, type D, eggs ga: Cassidaria echinophora (Linnaeus, 1758), egg mass gb: Cassidaria echinophora (Linnaeus, 1758), a group of capsules gc: Cassidaria echinophora (Linnaeus, 1758), 2 single cap- sules 10: Semicassts undulata (Gmelin, 1791), egg mass 11: Tonna ? galea (Linnaeus, 1758), egg mass 12a: Trunculariopsis trunculus (Linnaeus, 1758), clusters of egg capsules rab: Trunculariopsis trunculus (Linnaeus, 1758), egg cap- sules sac: Trunculariopsis trunculus (Linnaeus, 1758), 2 single capsules sad: Trunculariopsis trunculus (Linnaeus, 1758), embryos before hatching (taken from the capsule) 1ga: Murex brandaris Linnaeus, 1758, egg mass 13b: Murex brandaris Linnaeus, 1758, groups of capsules 1jc: Murex brandaris Linnaeus, 1758, 2 single capsules 14: Thais haemastoma (Linnaeus, 1767), left, egg mass Thats carinifera (Lamarck, 1822), right, egg mass 15: Columbella rustica (Linnaeus, 1758), egg capsules 16: Euthria cornea (Linnaeus, 1758), egg mass 17: Pisania striata (Gmelin, 1791), egg capsules Figure 18a: Sphaeronassa mutabilis (Linnaeus, 1738), egg mass Figure 18b: Sphaeronassa mutabilis (Linnaeus, 1758), 2 smge capsules Figure 1g: Nassariid species, egg mass Figure 20a: Fasciolaria lignaria (Linnaeus, 1758), 2 group of capsules Figure 20b: Fasciolaria lignaria (Linnaeus, 1758), embryos before hatching (taken from a capsule) Figure 21: Conus ventricosus Gmelin, 1791, egg mass Figure 22a: Bulla striata Bruguiére, 1789, egg ribbon Figure 22b: Bulla striata Bruguiére, 1789, capsules with veligers Figure 23a: Aplysia fasciata Poiret, 1789, spawn Figure 23b: Aplysia fasciata Poiret, 1789, veligers Figure 24: Bursatella leachi savigniana Audouin, 1826, adult and spawn Figure 25: Bursatella leachi savigniana Audouin, 1826, parts of egg string Figure 26: Elysia timida (Risso, 1818), egg mass (after M. Rahat) Figure 27: ? Glossodoris sp., egg mass Figure 28a: ? Acolidiella sp., adult Figure 28b: ?Aeolidiella sp., egg maas Figure 29a: Sepia officinalis Linnaeus, 1758, eggs Figure 29b: Sepia officinalis Linnaeus, 1758, juveniles (about r week old) Figure 29c: Sepia officinalis Linnaeus, 1758, cuttle bone of juvenile Figure joa: Sepia elegans d’Orbigny, 1826 Figure job: Sepia elegans d’Orbigny, 1826, an egg attached to 2 tube of a polychaete Figure 31a: Theodoxus jordani (Sowerby), egg capsules attached to shells of adults Figure 3rb: Capsules of Theodoxus jordani (Sowerby) on Melan- opsis praemorsa Figure 32a: Melanoides tuberculata (Miiller), adult female Figure 32b: Melanoides tuberculata (Miler), juveniles taken from female’s brood pouch Page 318 THE VELIGER Vol. 22; No. 4 Aligena laterodentata, New Species, from the Caribbean Coast of Honduras (Bivalvia : Leptonacea ) BY HAROLD W. HARRY ', JOSEPH C. BRITTON?, anp RODNEY NUNLEY? (5 Text figures) THE ACCOUNT OF THE LEPTONACEA (CHAVAN, 1969) in the Treatise on Invertebrate Paleontology (Moore, Ed., 1969) was in press when the review of the genus Aligena was published by Harry (1 January 1969). Chavan’s con- cept of Aligena, both with regard to the shell dentition and family relationships, differs from the concepts of ge- neric and family limits as construed by Oxsson (1961), Harry (1969), KEEN (1971) and others. KEEN (1971) re- allocated a species from the Pacific coast of Middle Amer- ica, Aligena obliqua Harry 1969, to the genus Orobitella Dall, 1900. She noted that a second species, Aligena cerritensis Arnold, 1903, — at least the Recent members of this species as interpreted by Harry, — is probably a junior synonym of Orobitella trigonalis (Carpenter, 1857). An additional species of Aligena, recently found on the Caribbean coast of Honduras, contributes significantly to elucidating the limits and relationships of supraspecific taxa in the Leptonacea. Aligena laterodentata Harry, Britton and Nunley, spec. nov. (Figures 1-5) Shell white, thin, translucent, inflated, large for the genus: the holotype, a left valve, is 8.5mm long, 7.2mm high and 3.1mm semi-diameter. It is sub-reniform in profile. The prominently inflated umbos touch each other, are pro- sogyrous, and located midway the length of the shell. A persistent protoconch is 0.21 mm long. Anterior and poste- rior margins of the adult shell are evenly rounded, of equal 1 4612 Evergreen Street, Bellaire, Texas 77401 2 Biology Department, Texas Christian University, Fort Worth, Texas 76129 curvature and length; the ventral margin is deeply and broadly sinuate in the middle. A prominent, wide sulcus, vaguely defined, extends nearly to the umbo and ends at the marginal sinus. There is no lunule, escutcheon or cor- selet. The ligament is not visible externally. The outer surface is swollen, somewhat more so before than behind the sulcus. The surface is evenly sculptured with promi- nent growth striae, closely spaced and regular. The interior is very smooth, non-nacreous. The shell margin is thin and without sculpture. The sub-umbonal cavity is very deep. The two adductor muscle scars are about equal in size, sub-rectangular, and joined with a pallial line without a sinus (the line not evident in the holotype). The hinge plate is divided into anterior and posterior parts, completely separated below and slightly behind the Figure 1 Aligena laterodentata Harry, Britton « Nunley, spec. nov. External view of left valve of the holotype, 8.5mm long Vol. 22; No. 4 Figure 2 Aligena laterodentata Harry, Britton & Nunley, spec. nov. Dorsal view of the same shell as in Figure 1 umbo. The posterior part contains the resilium in an elon- gate resilifer groove, which slopes downward and back- ward from the anterior end of this part of the plate. The posterior plate is widest just behind the resilifer, where it ' projects ventrally as an obtuse point. Behind this point there is a single, elongate, posterior lateral tooth in the left valve with a deep groove margining it above, for the reception of the upper of two posterior lateral teeth in the right valve. The latter teeth are separated from each other by a very deep groove forming a socket for the left lateral tooth. The upper lateral of the right valve is much smaller than the lower one. No anterior lateral teeth are present in either valve. There are two cardinal teeth in the left valve, the larger at the hind end of the front hinge plate, the smaller sep- Figure 3 Aligena laterodentata Harry, Britton « Nunley, spec. nov. Internal view of the same shell as in Figure 1 THE VELIGER Page 319 arated from it by a nick in the plate. Both cardinals are finger-shaped, projecting across the midline. The right valve has only a single cardinal tooth, similar in size and shape to the larger one of the left valve. On the dorsal margin of the left valve a deep, narrow groove extends forward from the gap between the two parts of the hinge plate to the umbo. No comparable gap is present in the right valve, but a large boss, elongate antero-posteriorally, sub-cylindrical, with rounded ends, seems to be attached to the hind end of the front part of the hinge plate and to extend backward, covering the gap between the two parts of the plate. This does not seem to be a tooth in the strict sense. Seven right and 7 left unmatched valves have been ex- amined. They are only slightly worn, but lack periostra- cum. The holotype is a left valve, number 781442 of the molluscan collection of the U.S. National Museum of Natural History. Three paratypes are number 781443 of that museum. Other paratypes are in the molluscan col- lection at Texas Christian University. The type locality is Calabash Bight (Manatee Bight), along the southern shore of Roatan Island, off the Carib- bean coast of Honduras. Specimens were collected in fine- grained sediments associated with Thalassia beds at a water depth of approximately 3 m. The species is relatively common as shell material in the type locality, occurring in densities calculated to be 20 to 30 specimens per square meter. No living specimens have been recovered. NUNLEY (1979) presents a comprehensive review of the type locality and its molluscan fauna. In size, shape and sculpture Aligena laterodentata is similar to A. cokeri Dall, 1909, from the west coast of Middle America, and to A. salamensis (Jaeckel and Thiele, 1931) from the Indopacific faunal realm. The similarity is so great that removing it to another genus or subgenus because it has an extra cardinal tooth and posterior lateral teeth would at present be unwarranted. No other species of Aligena is known to have lateral hinge teeth, or more than a single cardinal tooth in each valve, CHAvAN’s (1969) interpretation of the genus notwithstanding (see below). Otsson (1961) and KEEn (1962, 1971) admit that Orobitella Dall, 1900, is very similar to Aligena, and they include both genera in the Montacutidae. The characters which separate the two genera are relative differences, that is, differences in degree of contmuous variables. The most prominent character whereby the two genera have been separated is the position of the umbos. They are midway the length of the shell or only slightly behind that point in Aligena, which is thus equilateral or nearly so. The umbos are distinctly behind the midpoint of the shell Page 320 THE VELIGER Vol. 22; No. 4 Figure 4 Aligena laterodentata Harry, Britton « Nunley, spec. nov. Enlargement of the hinge of the same shell as in Figure 1 length in Orobitella, so that the anterior end of the shell is somewhat longer than the posterior end. If the two species included by Harry (1969) in Aligena are transferred to Orobitella,as KEEN (1971) has done, the limits of form of the species relegated to each genus achieve more precision. Very distinctive of most species remaining in Aligena is the median sulcus on the surface of the shell, resulting in a sinuate ventral margin and transforming an essentially quadrate profile into one that is slightly or distinctly reniform. In the type species, the Miocene A. aequata (Conrad, 1834), as well as in the Recent species from the coast of New England, A. elevata (Stimpson, 1851), the sulcus is absent. It was not found in the holotype of A. nucea Dall, 1913, of the Gulf of Cali- fornia, but it occurs in other specimens of that species as interpreted by Harry (1969). The sulcus is present in Figure 5 Aligena laterodentata Harry, Britton « Nunley, spec. nov. Right valve, enlargement of the hinge Vol. 22; No. 4 THE VELIGER Page 321 A. texasiana Harry, 1969, A. cokeri Dall, 1909, A. salamensis (Jaeckel and Thiele, 1931), and it is very prom- inent in A. laterodentata spec. nov. ROSEWATER (1983) has proposed that behavioral differ- ences may be correlated with the prominence of the sulcus among species of Aligena. He noted that on the Pacific coast of Panama, A. cokeri lives firmly attached to the exterior of the tube of the polychaete, Mesochaetopterus alipes Monroe, 1933. On each side of the attachment there is a hole through the tube, apparently serving as incurrent and excurrent passages, respectively, for water circulating through the mantle cavity of the bivalve. He further sug- gested that this unusual arrangement may be correlated with the prominent sulcus of the shell, which coincides with the position of the byssus. In support of this inter- pretation he noted that Aligena elevata, the life habits of which were extensively described by Gace (1968, as Montacuta elevata), has only a flimsy and temporary bys- sal attachment to the exterior of the tube of the polychaete with which it lives, Clymenella torquata (Leidy, 1855), and A. elevata lacks the furrow and sinus on the exterior of the shell. The peculiar rounded appendage on the hinge of the right valve of A. laterodentata, in the space between the two parts of the hinge plate, may represent a calcified part of the ligament. Only a fragment of this structure was present in the other specimens examined. This may be the same structure which Keen (1962) found in Orobitella (Isorobitella) singularis Keen, 1962, from San Quentin Bay, west coast of Baja California. The hinge appendage of A. laterodentata fits her description of it in O. sengularis very well, but the figure she presented is of little help (op. cit., p. 324; fig. 4c, reproduced by CHavan, 1960, p. N532, fig. E34, 2b). Incidentally, O. singularis Keen, 1962, was not included in her monumental work on the Sea Shells of Tropical West America (KEEN, 1971) because it was found north of the limit of the area covered. She did mention it briefly in conjunction with another species (of. cit., p. 144). How- ever, O. singularis seems to be identical with “Orobitella sp.” from Sechura, Peru, figure 4 of Plate 35 of OLsson (1961), which is not dealt with in the text of that work, nor apparently noticed elsewhere. Cuavan’s (1969:N523 of volume 2) cryptic account of the hinge dentition of the genus Aligena is as follows: “Hinge with small 3a, stout 1, curved 2a in prolongation of an anterior lateral, with small 2b behind it; wide oblique ligarnent and single strong posterior lateral on each valve.” By decoding this cryptogram according to the apparent key, found on pages N53-N56 of the first volume of this part of the Treatise (Moore, 1969, Editor), the following translation results: Hinge with (a) small (cardinal tooth and a) stout (one before it in the right valve, a) curved (cardinal tooth) in prolongation of an anterior lateral (tooth), with (a) small (cardinal tooth) behind it (in the left valve); etc. This is inconsistent with the description of the Kelliidae (CHavan, 1969: N522) in which family Chavan placed Aligena, for of the family he says there are “no distinctly elongate anterior laterals . . .” The figure of “Aligena aequata (Conrad), numbers 7a and 7b of figure E27 on page N524, have the hinge structure poorly shown, but they seem to indicate a single, long, posterior lateral tooth in each valve, plus two cardinal teeth in the left valve, the larger of which is a continuation of a long anterior lateral lamella; whether cardinal teeth or an anterior lateral tooth are present in the right valve (his fig. E27, No. 7a) is a matter of interpretation of a very ambiguous drawing. His figures are apparently originals, not previously published, but the source of the specimens is not stated. They depict a much more quadrate shell, with straight, truncated an- terior and posterior margins and more inflated umbos, than the topotypic Aligena aequata (Conrad, 1834) which Harry (1969: 165-166, figs. 1-3) studied at the US. National Museum of Natural History. Chavan’s species is evidently not the same one. ACKNOWLEDGMENTS Funds for travel to Roatan Island, Honduras, were pro- vided by Texas Christian University Research Founda- tion; the Department of Biology, and several Tropical Biology classes of Texas Christian University. We are par- ticularly indebted to Kevin and Tammy O’Kane, Tom Odom, Christine Miller and Larry Champagne, who con- tributed significantly to the acquisition of the samples from which this new species was obtained. Literature Cited Cravan, ANDRE 1969. Superfamily Leptonacea, pp. N518-N537 in: Raymond C. Moore, ed., Treatise on invertebrate paleontology, Part N, Mollusca & Bivalvia, vol. 2. Geol. Soc. Amer. and Univ. Kansas Press ii+N4or to N592 Gace, JoHN 1968. The mode of life of Montacuta elevata, a bivalve ‘commensal with Clymenella torquata (Polychaeta). Canad. Journ. Zool. 46: 877 - 892 Harry, Harotp WILLIAM 1969. A review of the living leptonacean bivalves of the genus Ali- gena. The Veliger 11 (3): 164-181; 40 textfigs. (1 Jan. 1969) - Page 322 THE VELIGER Vol. 22; No. 4 Keen, ANGELINE Myra 1962. A new West American subgenus and new species of Montacu- tidae (Mollusca: Pelecypoda), with a list of Mollusca from Bahia de San Quentin. Pacif. Natural. § (9): 321 - 328 1971. Sea shells of tropical West America: marine mollusks from Baja California to Peru, 274 ed. Stanford Univ. Press, Stanford, Calif. i-xiv+ 1064 pp.; ca. 4000 text figs.; 22 col. plts. (21 September 1971) Moorz, Raymonp C. (ed. 1969. ‘Treatise on Invertebrate Paleontology. Part N. Mollusca 6, Bivalvia. Geol. Soc. Amer., 2 vols.; 952 pp.; illust. NuNLEy, RopNEY 1979. A level-bottom molluscan fauna from Calabash Bight, Isla de Roatan, Honduras. M. Sci. Thesis, Texas Christ. Univ. 86 pp.; 6 plts. Oxsson, Axe, ADOLF 1961. Mollusks of the tropical eastern Pacific, particularly from the southern half of the Panama-Pacific faunal province (Panama to Peru). Panamic-Pacific Pelecypoda. Paleo. Res. Inst., Ithaca, New York, 574 pp.; 86 plts. (10 March 1961) RosEWwaTER, JOSEPH 1976. Some results of the National Museum of Natural History Smith- sonian Tropical Research Institute survey of Panama. Bull. Amer. Malacol. Union for 1975: 48-50 Vol. 22; No. 4 THE VELIGER Page 323 The Lamellariidae of the North Eastern Pacific BY DAVID W. BEHRENS Pacific Gas and Electric Company, Biological Research Laboratory, P.O. Box 117, Avila Beach, California 93424 (2 Plates; 15 Text figures) INTRODUCTION THE FAMILY LAMELLARDDAE (Orbigny, 1841) has been in taxonomic disarray since the description of its first species. Characterized around Lamellaria perspicua (= Helix perspicua Linnaeus, 1758) and Lamellaria latens (= Bulla latens Miiller, 1776), modern taxonomists are even in dis- _ agreement concerning the status of the type species. The specific designation Lamellaria proposed by Montagu (1815) has even drawn protest (BERGH, 1886b) because of its resemblance to that of an algal genus, Laminaria. It is not the intention of this paper, however, to review the higher taxonomic nomenclatural problems of the fam- ily, but rather to address the specific designations of the species occurring in the North Eastern Pacific. The spe- cific designation Lamellaria Montagu, 1815, is used in good faith over Marsenia (Leach) Oken 1823, due to its recent and seemingly wide acceptance (including: THIELE, 1931; Wenz, 1938; BurRcH, 1946; LARocQuE, 1953; FRETTER & GRAHAM, 1962; Marcus & Marcus, 1967; McLean, 1969; KEEN, 1971; THOMPSON, 1973, 1976; YoncE & THOMPSON, 1976). The North Eastern Pacific lamellariids are abundant, although comprising only a few species. Often overlooked, they include some of the most fascinating deceivers in the phylum Mollusca. While trying to identify several species, it became ap- parent to me that the existing nomenclature of the family is confusing and sometimes contradictory. Due to a paucity of information concerning body form, coloration, radulae, and due to the unavoidable reliance on shell descriptions, many of which are not diagnostic, this family has been overlooked by most molluscan guides or handled in a per- functory manner in the few references that acknowledge it. Historically, the family has been studied with opistho- branchs due to their shell-less appearance and resemblance to certain dorid nudibranchs (BERGH, 1908; Marcus & Marcus, 1958, 1959, 1960, 1969). Of ecological interest and receiving some attention, is their close resemblance to compound ascidian tunicates upon which they feed and live (HERDMAN, 1893; ANKEL, 1935; GHISELIN, 1964; THOMPSON, 1973), and their ability to secrete acid from the mantle to fend off predators (THompson, 1960, 19609, 1976; YONGE & THOMPSON, 1976). A review of the diagnostic features of the group has proven some features to be more useful and reliable than others and also presented a question as to the validity and placement of several species. This paper presents a syn- thesis of the North Eastern Pacific lamellariids with com- ments concerning their identification, an artificial key and notes on their natural history. Presented also is a taxonomic reallocation. MATERIALS Materials from the following museums were examined: California Academy of Sciences, San Francisco, California (CASIZ); Los Angeles County Museum of Natural His- tory, Los Angeles, California (LACM); Santa Barbara Museum of Natural History, Santa Barbara, California (SBMNH); as well as the private collections of Donald B. Cadien (DBC), R. Henderson (RH), Gary McDonald (GM) and Carol Skoglund (CS). Due to the preserved state of most of the above material, little information could be gleaned concerning the form and color of the living animals. Fortunately, live specimens of all species except Lamellaria perspicua and L. inflata were available for examination. Animals were studied in their original habitat, using SCUBA as well as in laboratory aquaria. Color transparencies and preserved voucher specimens have been deposited for future reference at CAS, LACM and SBNHM. Localities referred to in the text are shown in Figure 1. Page 324 THE VELIGER Vol. 22; No. 4 Sitka Sound Coos Bay e ie Cape Arago Humboldt Bay 42. San Francisco Bay +r eh Carmel Monterey Bay Cayucos = esBlaness <2. Morro Bay Diablo Canyon-€:-_~ Santa Barbara Palos Verdes _ fe - Los Angeles = = ~ San Diego Cholla Bay 7 [a Puerto Pefiasco Pinel Dees Bahia San Luis Gonzaga ®% We Cape Tepoca se ae Guaymas Figure 1 Map of the North Eastern Pacific Vol. 22; No. 4 NOMENCLATURAL PROBLEMS Of the original 7 species included in the family Lamel- lariidae on the Pacific Coast, 4, Lamellaria stearnsu, L. rhombica, L. orbiculata and L. inflata, were described by their various authors solely on the basis of shell morphol- ogy, with no description of mantle or radulae. For only one species, Lamellaria perspicua, was the radula de- scribed (Marcus & Marcus, 1967). Datu (1871) described two species and a subspecies from the same locality. The major differences between the three taxa were the presence of a dorsal pore in the mantle of one and size differences between the remaining two. Supplemented by notes from R. E. C. Stearns, the collector, the description of Lamellaria stearnsit was based upon two beach-washed shells; L. stearnsi var. orbiculata on 5 specimens; and L. rhombica on two specimens. Purely qualitative, Dall’s description gives little diagnostic infor- mation to separate the 3 species, and at one point he states, “. .I should have hesitated about describing them as more than varieties, if it were not that differences appear in the soft parts also” (DALL, 1871: 123). These differences were not observed by Dall, but communicated to him by Stearns who had the benefit of fresh material. In a later paper, Dati in Orcutt (1885) suggests that Lamellaria stearns and its variety L. s. orbiculata might only be sexually distinct, and he retains L. rhombica as a valid spe- cies. Due to the overlapping ranges of the two subspecies, I follow Dat in Orcutt (op. cit.), BurcH (1946), SmirH & Gorpon (1948) and TaLmancE (1966) in synonymizing them. Dati in Orcutt (op. cit.) states that L. rhombica “proves” to be a Marsenina Gray (1850) and should take the name Marsenina rhombica. He gives no data or rea- son, however, for this placement. Examination of the type specimens of all 3 species by Dr. James H. McLean, LACM, revealed the shells to be indistinguishable (personal communication, James H. McLean). During my investigation, I collected hundreds of animals whose shells could fit both descriptions. At variance with Dall’s description, each of these specimens had a pore in the mantle revealing the shell. This point was reported previously by GHISELIN (1964). I believe Dall’s original intention for separating Lamellaria stearnsu and L. rhombica was based upon the presence of a pore in the mantle; however, in the absence of an animal lacking a pore, his L. stearnsii will require a redefinition. Concerning the heritage of the names of the two similar California species and their proper generic placement, descriptions and proposed placement follow. THE VELIGER Page 325 To confuse the nomenclature further, Lamellaria perspicua, L. stearnsu and L. diegoensis were assigned to the genus Marsenia (Leach) Oken 1823, a junior synonym of Lamellaria (Montagu), by BERcH (1886b). Bercu (1886b), MacGinitie & MacGrnrtiz (1949), Marcus & Marcus (1967) and Keen(1971) carry Lamellaria orbiculata raised to species level, although Dati in Orcutt (1885) dubbed it nomina dubia. I be- lieve the report of L. stearnsii orbiculata from the tropical waters of the Gulf of California (MacGiniTizE & Mac- GINITIE, 1949; KEEN, 1971) to be in error as L. stearnsii is a temperate species. The species they refer to as a “white lamellariid” is most probably L. diegoensis which is commonly collected on white tunicates in the northern Gulf. An unexplained version of the spelling of Lamellaria stearnsit appears in YATES (1890) and later in Howarp (1972) where it appears as Lamellaria stearnsiana. A more recent report even multiplies problems within the family. Appotr (1974) suggests that Marseniopsis sharonae is a dwarf or juvenile of Lamellaria rhombica. This suggestion is presented without supporting evidence and, I believe, is unfounded. LAMELLARODAE The shell is internal, concealed or fully enveloped in the mantle, which is not retractile. Naticoid in form, the shell is few whorled, with a periostracum. The aperture is large, the suture impressed. An operculum is lacking. Mantle shape, texture and color are highly variable. Members of the family have shown remarkable concealing ability, the mantle resembling closely the compound ascidian species upon which they prey. Species are represented by both monoecious and dioecious forms. The Lamellariidae are also characterized by the larval shell, a type referred to as echinospira (FRETTER & GraHAM, 1962). The echinospira shell is double, one shell within the other. This adaptation is reported to aid the buoyancy of the swimming veliger (McCLoskey, 1973). GASTROPODA MESOGASTROPODA LAMELLARDDAE Lamellariinae Lamellaria Montagu, 1815 Marsenina Gray, 1850 Marseniopsis Bergh, 1886 Page 326 GENERIC SEPARATION Three factors have proven valuable in separating the Eastern Pacific genera of the family: 1) radula, 2) reproductive type and 3) presence of a dorsal fissure or pore in the mantle: see Figure 2 (modified from Bercu, 1886 a&b). Reproductive Dorsal Radula Type Pore Genus @)? i Mat 2579) oh AY Lamellaria Dioecious Absent I, POT OTE Ost Pp) Marseniopsis Monoecious Present Marsenina Figure 2 Separation of Genera SPECIES DESCRIPTION For the purpose of clarity, the following descriptions are limited strictly to the diagnostic characteristics unique to the individual species, such as mantle shape, texture and color, radula, etc. Shell descriptions, although unreliable for species identification, have been included. Lamellaria Montagu, 1815 The genus Lamellarta in the Eastern Pacific can be char- acterized by separate sexes, the absence of marginal teeth on the radula and a shell entirely enveloped in a non- retractive mantle. Type Species: Lamellaria tentaculata Montagu, 1815 (= Bulla latens) (fide WinckworTH, 1932; Burcu, 1946) THE VELIGER Vol. 22; No. 4 References and Synonymy: Lamellaria perspicua (Linnaeus, 1758) (Figures 3, 4) Helix perspicua Linnaeus, 1758. Lamellaria perspicua (Linnaeus, 1758). THIELE, 1931: 266. KEEN, 1971: 483. ABBOTT, 1974: 145. Marsenia perspicua (Linnaeus, 1758). BERGH, 1886b: 13. Lamellaria perspicua perspicua (Linnaeus, 1758). MARCUS & Marcus, 1967: 145. Lamellaria perspicua mopsicolor pUBoIs-REYMOND MARCUS, 1958. MARCUS, 1959: 14; 1963: 51. GHISELIN, 1964: 123. Marcus & MARCUS, 1967: 148. KEEN, 1971: 483. ABBOTT, 1974: 145. Material Examined: I) 1 specimen, intertidal, Puerto Penasco, Mexico; leg. D. B. Cadien, 31 March 1972. 2) 1 specimen, 9- 135m, Taboga Island, Panama Bay, Panama (80°35 N , 79°30’ W ); Jeg. E. Bergeron and J. H. McLean, 9 June 1965 (LACM 65-25). 3) 3 specimens, intertidal, Cholla Bay, Sonora, Mexico; leg. C. Skoglund, 15 October 1978. Nomenclatural Comments: Reported worldwide, Lam- ellaria perspicua was first recorded from the Pacific Ocean (southern Chile) by Marcus (1959). Since that report, Marcus & Marcus (1967) assigned subspecific status to two variations of L. perspicua collected in Puerto Penasco, Sonora, Mexico. Due to the geographical overlap of the two forms, and the number of specimens examined, sep- arate status does not seem justifiable. With further study, similarities between the Lamellana perspicua described by Marcus & Marcus (1967) and L. diegoensis may prove them identical. Sufficient mate- rial was not available here to carry out such comparisons. Description: I was unable to find or examine any living specimens of this species. The following description is Explanation of Figures 16 to 19 Figure 16: Lamellaria diegoensis. Isla Vista, Santa Barbara County, California 32mm Figure 17: Lamellaria diegoensis. Isla Vista, Santa Barbara County California 45 mm Figure 18: Marsenina stearnsi. Diablo Canyon, San Luis Obispo County, California 15mm Figure 19: Marsenina stearnsii. Diablo Canyon, San Luis Obispo County, California 15mm THE VELIGER, Vol. 22, No. 4 [BEHRENS] Figures 16 to 19 Figure 16 Figure 17 Figure 19 a — Vol. 22; No. 4 based upon 5 preserved specimens (listed above) and Marcus & Marcus (1967) for their animals collected in Puerto Penasco, Sonora, Mexico. External Morphology and Coloration: Lacking a dor- sal pore or fissure (Figure 3), Marcus & Marcus (1967) describe it as more or less translucent, the male with the whitish hue, the female with the pinkish hue. Glandular inclusions of the skin opaque white in the male, vitreous in the female, the anterior border of the foot is opaque white. After preservation, the male exhibited black pig- ment spots standing out from the gray ground color, the female being more uniform. The sole of the foot is gray, the head lighter and the penis white. The specimen re- ferred to as Lamellaria perspicua mopsicolor was a dark bluish gray. The cutaneous bosses in the mantle had dark centers, surrounded by lighter rings. Figure 3 Drawing of living Lamellaria perspicua (after Marcus & Marcus, 1967: 9) The specimen collected by D. Cadien was reported to be yellowish-beige with brown markings, while alive. The C. Skoglund specimens were pinkish-brown with liver colored specks and small yellowish papillae. After preser- vation, the specimens faded to a translucent hue with brown and white specks. A color photograph of a Euro- pean specimen appears in YONGE & THOMPSON (1976, pit. 9). Shell: White, growth lines distinct. Apex prominent. Marcus & Marcus (1967) distinguish this species from THE VELIGER Page 327 Lamellaria diegoensis by the absence of malleations on the shell surface. All of the D. Cadien, C. Skoglund and LACM specimens thought to be L. perspicua showed very slight malleations in the shell surface. Without the entire animal, it would appear that this species could not be recognized as L. perspicua from the shell alone. Radula: (from Marcus & Marcus, 1967). Male has 60 rows, the female 79 rows of teeth. The left leg of the rachidian is slightly longer. This difference is more pro- nounced in the female. The male rachidian has 4-5 strong denticles on the right, 6-7 finer ones on the left; the female 5-8 and 8-10, respectively. The lateral tooth of the male has 4-8 strong inner and 7-12 finer outer den- ticles; the female 5-8 and 4-11, respectively. The specimen referred to as Lamellania perspicua mopsicolor displayed 4-5 denticles on the right side of the rachidian only, while the laterals bore 4 - 6 coarse, sharp, inner and 8 smaller outer denticles (Figure 4). Figure 4 Radular teeth of Lamellaria perspicua a — rachidian tooth b — lateral tooth The specimens I examined had 55 rows (DBC & LACM) and 41-45 rows (CS) of teeth. The rachidians bore 4-6 right, and 3-7 left denticles, the left set being finer. The laterals bore 3-5 strong inner, and 7-12 fine outer den- ticles. The left leg of the rachidian did not seem noticeably longer in any of the specimens. This description bears a striking similarity to that of Lamellaria diegoensis, particularly because of similarity in shell surface textures and radular dentition (not pre- viously described in L. diegoensis). Size: Marcus & Marcus (1967) report their specimens to measure 15 by 1omm and 18 by 19mm, length by breadth. The largest specimens I examined were 18 by 15 mm (DBC) and 20 by 202mm (LACM), preserved. Page 328 THE VELIGER Vol. 22; No. 4 Distribution: Worldwide. Pacific Ocean: Magellanic region and southern Chile; Puerto Pefasco, Sonora, Mex- ico; Panama. Lamellaria inflata (C. B. Adams, 1852) (Figures 5, 6) References and Synonymy: Stomatella inflata C. B. Apams, 1852: 216, 440. Lamellaria inflata (C. B. Adams). STRONG & HERTLEIN, 1939: 186, 296. TURNER, 1956:56. SHASKY, 1961: 23. DUSHANE & POORMAN, 1967: 427. Marcus & Marcus, 1967: 148. DUSHANE & SPHON, 1968: 242. BERTSCH & SMITH, 1970: 172. KEEN, 1971: 483. ABBOTT, 1974: 145. Material Examined: 1) 6 specimens, intertidal, Jervis Island, Galapagos (0°25'S,, 90°42’ W ); leg. Ameripagos Expedition, 24 March 1971 (LACM 71-68) 2) 2specimens, intertidal, Punta Alfaro, Isabela Island, Galapagos (0°25'20"S , 90°57’10” W ); leg. Ameri- pagos Expedition, 25 March 1971 (LACM 71-70) 3) 2 specimens, 15-30m, Academy Bay, Santa Cruz Island, Galapagos (0°25’S, 90°15'38” W ); leg. Ameripagos Expedition, 18 March 1971 (LACM 71-55) 4) 1 specimen, Guaymas, Mexico; no leg., no date (LACM HH-53) 5) 1 specimen, 18m, Tenacatita Bay, Jalisco, Mexico; leg. G. Willett, no date (LACM A. 375) 6) 2 specimens, Puertecitos, Baja California (30°25’N, 114°38’ W ); leg. R. Mucke, 26 December 1964 (LACM 67-15) 7) 2specimens, 9-20m, Banderas Bay, Jalisco, Mexico; leg. C. Snell and B. McMillen, 14 February 1967 (LACM 67-15) 8) 1 specimen, 4.5-20m, Cuastecomate, Jalisco, Mexico (19°13'44"”N , 104°44’53” W ); leg. J. McLean and P. Oringer, 13-21 October 1968 (LACM 68-41) 9) 1 specimen, 72-126m, Guatulco Bay, Oaxaca, Mex- ico; leg. G. Willett, 7 March 1938 (LACM 38-9) 10) 1 specimen, Cape Tepoca, Lebos, Sonora, Mexico; leg. R. Poorman, March 1975 (identified from LACM color transparency) 11) 1 specimen, intertidal, San Luis Gonzaga, Baja Cal- ifornia; leg. W. E. Harvey, 6 February 1966 (identi- fied from color transparency ) 12) 1specimen, Cholla Bay, Pelican Point, Sonora, Mex- ico; 16 February 1969 (CAS) 13) 1 specimen, intertidal, Cholla Bay, Sonora, Mexico; leg. D. W. Taylor, 2 November 1968 (CAS) Description: I was unable to examine any living animals of this species; however, preserved material was examined from LACM along with a series of color transparencies of living specimens from various contributors. External Morphology and Coloration: Lacking a dorsal pore or fissure. Textured entirely by a series of small pores. Yellow in color, the mantle is marked by a brown saddle lying just posterior to the center of the dorsum, as well as a brown patch on either side of the anterior siphon (Figure 5). Neither the saddle nor the patches reach the margin of the dorsum. The coloration, although faded, is retained in the preserved animal. Figure 5 Drawing of Lamellaria inflata from color transparency of a live animal Shell: White, thick, opaque. Fine irregular growth striae, without spiral striae. Suture well impressed. Similar in shape and character to Marsenina stearnsi. The reliance in the literature upon the description of the columellar side of the aperture being “abruptly arcuated” is, I think, unjustified. I found the angle of arcuation to be indistin- guishable from that of the other species. Vol. 22; No. 4 Radula: About 50 rows of teeth. The rachidian teeth are without any denticulation. The lateral has 3 strong inner denticles and 3-4 lesser outer denticles (Figure 6). Figure 6 Radular teeth of Lamellaria inflata a — rachidian tooth b — lateral tooth Size: From the specimens examined, this species seems to be of smaller size than the other species. The largest specimen measured 6 by 5mm, length and breadth, re- spectively. Distribution: Cape Tepoca, Puerto Lobos, Sonora, Mex- ico to Panama, as well as Bahia San Luis Gonzaga to La Paz, Baja California and the Galapagos Islands. Lamellaria diegoensis Dall, 1885 (Figures 7, 8, 9, 16 and 17) References and Synonymy: Lamellavia diegoensis Da.t, 1885: 538. DALL in Orcutt, 1885: 538. OLDROYD, 1927: 738. WILLETT, 1939: 124. Burcu, 1946: 2. Morris, 1966: 76. MARcus & MARCcus, 1967: 147. MCLEAN, 1969: 38. KEEN, 1971: 483. HUMAN, 1972: 28, 1973: 21. McCLOSKEy, 1973: 157. BISHOP & BisHOP, 1973: 149. ABBOTT, 1974:145. Marsenia diegoensis (Dall). BERGH 1886(b): 13. Lamellaria orbiculata Dall, 1871. MacGinitie & MAcGin- ITTE, 1949: 372. Marcus & Marcus, 1967: 147. KEEN, 1971: 483. Material Examined: I) 1 specimen, 15m, Chace Reef, Lover’s Point, Mon- terey Co., California; Jeg. Lloyd Gomez, 15 August 1969 (CAS) THE VELIGER 2) Page 329 I specimen, 12-15m, Del Monte shale beds, Mon- terey Bay, California; leg. D. Powell, 7 March 1965 (CAS) I specimen, 180m off Soberanes Point, California; leg. D. D. Chivers, 20 July 1971 (CAS) I specimen, Asilomar, Monterey Co., California; leg. G. McDonald, April 1971 (M71-4-33-1) I specimen, intertidal, Garcia’s Ranch, Cayucos, San Luis Obispo Co., California; leg. G. McDonald, 22 January 1970 (M70-1-22-3) I specimen, Asilomar, Monterey Co., California; leg. G. McDonald, 14 October 1970 (M70-10-14-13) I specimen, 18m, Del Monte shale beds, Monterey Bay, California (36°37’N , 121°52.5’W ); leg. J. H. McLean, 1960-62 (LACM 60-22) I specimen, 200 ft., Los Angeles County Sanitation District - T3; leg. D. Cadien, 9g December 1976 I specimen, 135m, Los Angeles County Sanitation District-T5; leg. D. Cadien, 14 December 1977 I specimen, 22.5m, Palos Verdes, California; leg. D. Cadien, 11 October 1977 I specimen, Orange County; Jeg. D. Cadien, Sum- mer 1974 I specimen, 135m, Palos Verdes, California; leg. D. Cadien, 18 May 1977 I specimen, 21m, Huntington Beach, RUNST #2, California; leg. D. Cadien, 6 February 1978 I specimen, 24m, Engel’s Bank, California; leg. D. Cadien, 8 February 1976 7 specimens, 6m, Isla Vista, Santa Barbara Co.; California; Jeg. R. Henderson, 24 March 1977 I specimen, intertidal, Diablo Cove, San Luis Obispo Co., California; leg. L. L. Laurent, 23 September 1977 I specimen, 6m, Seal Haul Out, Diablo Canyon, San Luis Obispo Co., California; leg. D. W. Behr- ens, 23 September 1976 I specimen, Morro Bay, California; leg. D. W. Behrens, 7 February 1978 I specimen, Morro Bay, California; leg. D. W. Behrens, 8 March 1978 I specimen, 5m, Intake Cove, Diablo Canyon, San Luis Obispo Co., California; Jeg. D. W. Behrens, 27 April 1977 I specimen, intertidal, Seal Haul Out, Diablo Can- yon, San Luis Obispo Co., California; Jeg. L. L. Laurent, 25 May 1978 1 specimen, Isla Vista Beach, Santa Barbara Co., California; Jeg. R. Henderson, January 1978 5 specimens, Punta Diggs, San Felipe, Baja Califor- nia; leg. L. O. Miles, 15 April 1952 (CAS) Page 330 THE VELIGER Vol. 22; No. 4 24) 1 specimen, Santa Barbara, California; Jeg. D. Olsen, 1961 (SBMNH 23600) 25) 1 specimen, Goleta Point, Santa Barbara, Califor- nia; leg. D. Olsen, 6 December 1961 (SBMNH 23618) 26) 1 specimen, 4 mi so. of Gaviota, Santa Barbara County, California; Jeg. K. Newson, 11 November 1967 (SBMNH 24993) 27) 1 specimen, Boulder Beach, Summerland, Santa Barbara County, California; Jeg. S. Anderson, 26 June 1967 (SBMNH 24650) 28) 6specimens, intertidal, Cholla Bay, Sonora, Mexico; leg. C. Skoglund, 15 October 1978 Description: EXTERNAL MORPHOLOGY AND COLORATION: Dorsal fissure or slit lacking. Mantle texture and color extremely variable, yet at the same time distinctive. Color white through purple and brown; mottled, blotched and radially marked. Texture from smooth to velvety (Figure 7b) or papillated (Figure 7a) with branched projections. When present, mantle markings and papillations follow one of several specific formats: a dorsal saddle, as in Lamellaria inflata; a mid-dorsal horseshoe pattern with its opening situated anteriorly; or some variation of a radial pattern (Figure 8). Papillations usually white, brown and/or gray resembling tufts of hydroid or bryozoan growth (Figures 16 and 17). Underside of mantle speckled with yellow and white flecks. The foot is truncate ante- riorly; rounded, posteriorly. Shell: The most diagnostic feature of the shell is its mal- leated (hammered) surface texture. This character sep- arates it from all other species. Columella without thick- ened callus. Figure 7 Drawing of Lamellaria diegoensis a) a highly papillose individual, Santa Barabara, 24 March 1977 b) Specimen with low tubercles, Diablo Cove, San Luis Obispo County, 25 May 1978. Both drawn from color transparencies of the animals alive Vol. 22; No. 4 THE VELIGER Figure 8 Common color pattern formats of Lamellaria diegoensis Radula: Sixty rows of teeth. The rachidian is stout and nearly flat, not forming a hook, as in L. perspicua and Page 331 L. inflata (Figure gb). It has 3-4 denticles on each side. The dentition on the lateral teeth begins farther from the tip of the tooth (Figure gc). They have 4-5 outer denticles and 8-11 inner denticles. Lateral teeth overlap in an ar- rangement shown in Figure ga. Size: ‘The largest of the Eastern Pacific lamellariids, it reaches 45mm in both length and breadth. Distribution: Moss Landing, California to Baja Cali- fornia and Sonora, Mexico. Intertidal to 139m. Marsenina J. E. Gray, 1850 The genus Marsenina in the Eastern Pacific is separated from the other lamellariid genera in that it is hermaphro- ditic (BeRcH, 1853, 1886 a & b), and its mantle exhibits a dorsal fissure or pore and is retractile. Like Marseniopsis, the radula possesses marginal teeth (Gray, 1850; BercH, 1886 a&b). Type Species: Lamellaria prodita Lovén, 1846 (by M) (= Oxynoe? glabra Couthouy, 1838) Nomenclatural Comments: In the eastern Pacific, this genus is represented by two species: Marsenina stearnstt (Dall, 1871) and Marsenina rhombica (Dall, 1871). Orig- inally, both were placed in Lamellaria by Datu (1871). Figure 9 Radular teeth of Lamellaria diegoensis a) juxtaposition of lateral teeth (rachidians not shown) b) rachidian teeth c) lateral teeth Page 332 Dati in Orcutt (1885) suggested placement of L. rhom- bica in Marsenina Gray, 1850, but failed to give any ex- planation for the reallocation. Had the radula been exam- ined, L. stearnsii would have been placed there also. The two species are indistinguishable based on Dall’s original descriptions, which were based solely upon shell characteristics, and an examination of the types which are dry shells. Two morphologically different species do exist, however. Because of this, I am somewhat at a loss concern- ing which of the two names applies to the species found along this coast. Dall originally described a dorsal pore in the mantle of Lamellaria rhombica, but not in L. stearnsit. As previously mentioned, no poreless animal, aligned with this genus, has since been reported. GHISELIN (1964) was the first to note this discrepancy. In his report, he also provided photographs of a specimen with a dorsal pore, which he called Lamellaria stearnsu. Considering the dis- tinct difference in the external morphology of the live animals, none of which had been reported until publica- tion of Ghiselin’s photographs, I propose that the prec- edent has been set for Marsenina stearnsu, leaving the name M. rhombica for the remaining species. Marsenina stearnsii (Dall, 1871) comb. nov. (Figures 10, 11, 78 and 19) References and Synonymy: Lamellarta depressa Dall, mss, 1866. Lamellaria stearnsit DALL, 1871: 122. ORCUTT & DALL, 1885: 539. TRYON, 1886: 63. KEEP, 1892: 47. WILLETT, 1919: 26. OLDROYD, 1924: 164; 1927: 737. STRONG & HANNA, 1930: 19. ROGERS, 1936: 146. WILLETT, 1939: 124. BURCH, 1946: 2. SMITH & GORDON, 1948: 199. GHISELIN, 1964: 123. TALMADGE, 1966: 840. Marcus & Marcus, 1967: 147. BrooxsHirE, 1968: 6. McCLoskEy, 1973: 155. BisHop & BisHop, 1973: 149. ABBOTT, 1974: 146 Marsenia stearnsii (Dall, 1871). BERGH, 1886(b): 13. Lamellaria stearnsi Dall, 1871. THompson, 1973: 75 Lamellaria stearnsii orbiculata DAL, 1871: 122. ORCUTT & DALL, 1885: 539. OLDROYD, 1927: 738. BURCH, 1946: 2. ABBOTT, 1974: 146. Marsenia orbiculata (Dall, 1871). BERcH, 1886(b): 13. Lamellaria stearnsiana Dall, 1871. YATES, 1890: 44. Howarp, 1972: 23. THE VELIGER Vol. 22; No. 4 Material Examined: 1) 7 specimens, Cape Arago Park, Oregon; no leg., 19 July 1947 (CAS) 2) 3 specimens, Yankee Point, Monterey Co., Califor- nia; leg. A. G. Smith, 1955 (CAS) 3) 1 specimen, San Juan Island, Puget Sound, Wash- ington; leg. L. Andrews, September 1964 (CAS) 4) 1 specimen, Portuguese Beach Park, Sonoma Co., California; leg. L. Andrews, 6 June 1963 (CAS) 5) 3 specimens, 9-12m, Del Monte shale beds, Mon- terey Bay, California; Jeg. David Powell, 2 October 1964 (CAS) 6) 1 specimen, Pacific Grove, California; no leg., no date (CAS) 7) 4 specimens, Point Pinos, Pacific Grove, California; no leg., July 1910 (CAS) 8) 1 specimen, intertidal, Garcia’s Ranch, Cayucos, San Luis Obispo Co., California; leg. G. McDonald, 22 January 1970 (M7o-1-22-1) 9) 1 specimen, intertidal, Field’s Ranch, San Luis Obispo Co., California; Jeg. G. McDonald, 29 May 1968 (M68-5-29-1) 10) 3 Specimens, intertidal, Spooner’s Cove, San Luis Obispo Co., California; Jeg. G. McDonald, 2 No- vember 1971 (M71-11-2-5) II) 7 specimens, intertidal, Ethelda Bay, Hecate Strait, British Columbia (52°58.2’N, 129°31.2’W); leg. D. B. Quayle and F. R. Bernard, 22 October 1968 (LACM 68-74) 12) 5 specimens, 6m, Intake Cove, Diablo Canyon, San Luis Obispo Co., California; Jeg. D. W. Behrens, 20 September 1976 13) 1 specimen, Diablo Cove, San Luis Obispo Co., Cal- ifornia; leg. J. L. Kelly, 29 September 1977 14) I specimen, 6m, Intake Cove, Diablo Canyon, San Luis Obispo Co., California; Jeg. D. W. Behrens, 15 May 1978 15) 4 specimens, 6m, Seal Haul Out, Diablo Canyon, San Luis Obispo Co., California; /eg. D. W. Behrens, 20 September 1976 16) 3 specimens, 8m, Seal Haul Out, Diablo Canyon, San Luis Obispo Co., California; leg. D. W. Behrens, 27 April 1978 Explanation of Figures 20 to 293 Figure 20: Marsenina rhombica. Point Piedras Blancas, San Luis Obispo County, California 30 mm Figure 21: Two Marsenina rhombica. Point Piedras Blancas, San Luis Obispo County, California 23 and 35 mm, respectively Figure 22: Marseniopsis sharonae. Morro Bay, California, sub- strate, Botrylloides sp. 15 mm Figure 23: Marseniopsis sharonae. Morro Bay, California, sub- strate, Botrylloides sp. 15 mm THE VE icrR, Vol. 22, No. 4 [BEHRENS] Figures 20 to 23 Figure 22 Figure 21 Figure 23 Vol. 22; No. 4 Description: EXTERNAL MORPHOLOGY AND COLORATION: Mantle with a dorsal slit, pore or fissure which may be retracted to expose the shell (Figure 10). Mantle smooth. Color white to pinkish-white with a pattern of evenly- spaced small pores (Figures 18 & 19). These pores may be seen from beneath, through the mantle. Where the shell is exposed, internal organs show through the shell and are salmon colored. GHISELIN (1964) published black and white photographs of the species, on and off its tunicate substrate. THE VELIGER Page 333 teriorly, hooked anteriorly with none or one denticle flank- ing the medium spur. In plane view, they appear nearly square (Figure 11b). The lateral teeth are similar to the rachidian in dentition. There are two long, thin, hook- shaped marginal teeth, as in Marseniopsis. Each marginal tooth has one denticle situated to the outside of the pair of teeth (Figure 11d). Not included in the original descrip- tion, the radula agrees closely with that described for Marsenina in Gray (1850), and later in Bercu (1886b). Figure 10 Drawing of Marsenina stearnsii from color transparency of live animal. Diablo Canyon, San Luis Obispo County, California Shell: Smooth, thick, opaque white. Indistinguishable from those of Lamellaria inflata and Marsenina rhombica. Growth lines even and smooth. Dati (1871) described the characteristic of microscopic fine revolving striulae which later (1885) he described as distinctive. At my request, Dr. Harold Rehder, USNM, examined the holotype and re- ports to me that he was unable to find such a character in the shell surface. I too, have yet to find this character in any North Eastern Pacific lamellariid shell. Radula: Differs from Lamellaria by possessing marginal teeth (Figure 11a). The rachidian teeth are truncate pos- Figure 11 Radular teeth of Marsenina stearnsii and Marsenina rhombica a) juxtaposition of radular teeth b) rachidian tooth c) lateral teeth d) pair of marginal teeth Size: Moderate, reaching 18 by 16mm in length and breadth. Distribution: British Columbia, Canada to San Diego, California. Its occurrence at Maria Madre Island, Mexico Page 334 (Stronc & HANNA, 1930) is questionable. Intertidal to 8m subtidally. Marsenina rhombica (Dall, 1871) (Figures 11, 12, 20 and 27) References and Synonymy: Lamellaria rhombica Dall, 1871: 122. DALL in OrcuTT, 1885: 539. OLDROYD, 1924: 164; 1927: 137. ROGERS, 1936: 147. WILLETT, 1939: 124. BURCH, 1946: 2. SMITH & GoRDON, 1948: 199. GHISELIN, 1964: 123. MARCUS & Marcus, 1967: 147. McCLosky, 1973: 155. ABBOTT, 1974: 145. Marsenina rhombica (Dall, 1871). ORCUTT & DALL, 1885: 539. BERGH, 1886(b): 14. ROGERS, 1951: 492. LAROcQUE, 1953: 157- Material Examined: 1) 1 specimen, Carmel Point, Monterey Co., Califor- nia; leg. G. McDonald, 2 December 1971 (M71-12- 2-4) 2) 2 specimens, Carmel Point, Monterey Co., Cali- fornia; leg. G. McDonald, 3 November 1971 (M71-11-3-3 & 4) 3) 1 specimen, Spooner’s Cove, San Luis Obispo Co., California; leg. G. McDonald, 2 November 1971 (M71-11-2-5) 4) 2 specimens, 112-4%2m, Halibut Point, Sitka Sound, Baranof Island, Alaska (56°06’N, 135°24’ W); leg. J. H. McLean, 25 and 26 July 1973 (LACM 73-13) 5) 4 Specimens, intertidal, Ethelda Bay, Hecate Strait, British Columbia (52°58.2’N, 129°31.2’W); leg. D. B. Quayle & F. R. Bernard, 22 October 1968 (LACM 68-74) 6) . 1 specimen, Shelter Lane, California; no leg., 30-31 May 1946 (CAS) 7) 1 specimen, intertidal, Point Piedras Blancas, San Luis Obispo Co., California; Jeg. D. W. Behrens, 20 November 1976 8) 2 specimens, intertidal, Point Piedras Blancas, San Luis Obispo Co., California; leg. D. W. Behrens, 11 November 1977 9) 5 specimens, intertidal, Point Piedras Blancas, San Luis Obispo Co., California; leg. R. Henderson, 13 November 1977 10) 1 specimen, intertidal, Point Piedras Blancas, San Luis Obispo Co., California; Jeg. D. W. Behrens & S. Anderson, 30 December 1978 II) 2 specimens, 6 to 13.5m, Zaikof, Montague Island, Alaska; leg. R. Fay, 9 September 1978 (RH) THE VELIGER Vol. 22; No. 4 EXTERNAL MORPHOLOGY AND COLORATION: As in Marsenina stearnsii, a dorsal pore in the mantle exposes the shell (Figure 12). Mantle texture varies from smooth to warty. Color is variable, translucent to gray or yellow and orange with various degrees of spotting, speck- ling or blotching (Figures 20 & 27). Points of color appear to penetrate deep into the mantle, as bosses. Several speci- mens had a black swath posteriorly adjacent to the dorsal pore. Description: Figure 12 Drawing of Marsenina rhombica from color transparency of live animal. Point Piedras Blancas, San Luis Obispo County, California Shell: Generally indistinguishable from Marsenina stearnsit. A very thick, strong growth mark was observed in several specimens. This feature was not seen in any M. stearnsit. Radula: See description above (Figure 11). Not distinguishable from Marsenina stearnsit. Size: Larger than Marsenina stearnsit, reaching 40 x 35 mm in length and breadth. Distribution: Montague Island, Alaska to Diablo Cove, San Luis Obispo County, California. Intertidal to 60 meters subtidally. Vol. 22; No. 4 Marseniopsis Bergh, 1886 The genus Marseniopsis in the Eastern Pacific can be char- acterized by separate sexes, naticoid shell, hexahedral body shape and a distinctive radula. Type Species: Marseniopsis pacifica Bergh, 1886 (fide WENz, 1938). Marseniopsis sharonae (Willett, 1939) (Figures 13, 14, 15, 22 and 23) References and Synonymy: Lamellaria sharoni WILLETT, 1939: 123. Lamellaria sharonae Willett, 1939. BuRcH, 1946: 1. CATE & Care, 1962: 91. DUSHANE & SPHON, 1968: 235, 242. Mc- LEAN, 1969: 39. KEEN, 1971: 485. BISHOP & BISHOP, 1973: 149. ABBOTT, 1974: 145. Lamellaria sp. MacGINiITIE & MACGINITIE, 1949: 115. Marseniopsis sharonae (Willett, 1939). SPHON, 1975: 95. Material Examined: 1) 2specimens, Morro Bay State Park, Morro Bay, San Luis Obispo Co., California; Jeg. R. A. Roller, 2 February 1968 (M68-2-2-1) 2) 1 specimen, intertidal, Puerto Penasco, Sonora, Mex- ico; leg. D. Cadien, 1 April 1972 3) 1 specimen, 20m, Oyster Point, San Mateo Co., San Francisco Bay, California; Jeg. D. D. Chivers, 17 August 1971 (CAS) 4) 29 specimens, 5m, Morro Bay, California; leg. D. W. Behrens, 10 February 1977 5) 15 specimens, 1.5m, Morro Bay, California; leg. D. W. Behrens, 20 April 1977 6) 19specimens, 3m, Morro Bay, California; Jeg. D. W. Behrens, 27 July 1977 7) 9 specimens, 3m, Morro Bay, California; Jeg. D. W. Behrens, 18 August 1977 8) 13 specimens, 3m, Morro Bay, California; Jeg. D. W. Behrens, 7 September 1977 9) 21 specimens, 1.5m, Morro Bay, California; leg. D. W. Behrens, 24 October 1977 Nomenclatural Comments: Its original placement in Lamellaria by Willett was apparently done hesitantly, as reported by Burcu (1946). SPHON (1975) later reassigned it to the genus Marseniopsis without data or basis given for the reallocation. However, on the basis of radular mor- phology (BERcH, 1886b), I concur with Sphon’s generic placement. THE VELIGER Page 335 Description: EXTERNAL MORPHOLOGY AND COLORATION: The shape of the mantle is its most diagnostic character (Figures 13, 22 and 23). Lacking a dorsal fissure or slit, it is divided into six areas by low ridges that commence from a raised hexagonal area in the center of the dorsum. In the preserved state, the shape remains as a tight six-sided bead. Color in life is highly variable, white through red, purple and brown. The ridges are ornamented with a series of specks and irregular spots. A single spot is usually found in the center of the dorsal hexagon and on each side. As a rule, light colored specimens have dark spotting while darker specimens are ornamented with light spot- ting. A color photograph appears in KEEN (1971, plt. XVI). See Jn eS ~ 6 rT a 4 oF EE et Figure 13 Drawing of Marseniopsis sharonae from color transparency of live animal Morro Bay, California Shell: Naticoid in form, much more so than any other lamellariid (Figure 14). Very thin, transparent. Columella brown. Pinkish periostracum. Page 336 Figure 14 Drawing of the shell of Marseniopsis sharonae, 8mm a) dorsal view b) ventral view Radula: Distinctive in dentition from other lamellariids, 2.1.1.1.2 (Figure 15). The rachidian teeth are not-forked- posteriorly as in Lamellaria, but truncate. The rachidian dentition is best described as having five strong terminal denticles. Two denticles are lateral to and slightly behind the center denticle. The laterals are similar to the rachid- ian in strength, having 3 equal strong terminal denticles and two weak outer denticles. There are two marginal teeth in the radula, as in Marsenina. They are long, thin and forked at their anterior end. The outer tooth appears to cross over the inner tooth. This aligns the denticles of each in a lateral series with each other and those of the adjacent lateral tooth (Figure 15a). Such a radula agrees closely with that described in Marseniopsis in BERGH (1886a & b). Size: Of moderate size, it reaches 13 mm in length and breadth. THE VELIGER Vol. 22; No. 4 z ~ are Figure 15 Radular teeth of Marseniopsts sharonae a) juxtaposition of radular teeth b) rachidian tooth c) lateral tooth d) pair of marginal teeth Distribution: Coos Bay, Oregon (personal communica- tion, James Lance, La Jolla, California) to Puerto Penasco, Sonora, Mexico, and Bahia San Luis Gonzaga, Baja Cali- fornia. Intertidal to shallow (5m) subtidal. MIS-ALLOCATED SPECIES Other mollusk species have, on occasion, been incorrectly allocated to Lamellariidae, most notably the Pleuro- branchidae. In the North Eastern Pacific, OLpRoyp (1927), WiLLETT (1939) and AssotT (1974) refer to a Lamellaria digueti Rochebrune (1895). Originally Pleuro- branchus digueti, it was allotted to Lamellaria by DALL (1871). BurcH (1946) was the first to note the misplace- Vol. 22; No. 4 THE VELIGER Page 337 ment. Marcus & Marcus (1967) report this allocation to be untenable. Key to the Species of the Lamellartidae of the North Eastern Pacific (based on mantle characteristics) 1. Mantle with dorsal slit, pore or fissure, which may expose the shell when the mantle is retracted........ 2 - Mantle whole, without a dorsal pore or fissure........ 3 2. Mantle smooth, white to pinkish white, with a pat- tern of evenly spaced transparent acid glands MESCINPLING ASIAN POLES este ce teste oe tcesene attests oe ee ee Marsenina stearnsii (Dall, 1871) - Mantle smooth to warty. Color variable, translu- cent or grey to yellow and orange, with or without various degrees of spotting or blotches.................... Weare cel intl Marsenina rhombica (Dall, 1871) 3. Mantle divided into six areas by low ridges that commence from a raised hexagon in the center of the dorsum. Color variable, white through orange, red, purple and brown. A single spot in center of dorsal hexagon and on each SIE..........cessssssssssssesseneeen sci n a Marseniopsis sharonae (Willett, 1939) - Mantle not divided into six areas by low ridges POFININY AT aISEG, NEXAQOM oe csceseses tect cevessensncioeeeconses 4 4. Underside of mantle with white or yellow specks. Mantle smooth to velvety or papillated. Color Wartablen nc cc.--s-r Lamellaria diegoensis Dall, 1885 - Underside of mantle without white or VIG WASPECKS eee cere eee ccacsescsarccsnsesseesensnbscanstosessey 5 5. Mantle smooth, opaque yellow, with a brown sad- dle across dorsum, as well as a brown patch on either side of the anterior siphon. Remainder of mantle covered with a pattern of evenly spaced poresse = Lamellaria inflata (C. B. Adams, 1852) - Mantle translucent, white, yellow or pinkish brown. Glandular inclusions on the skin glassy or white to brown........Lamellaria perspicua (Linnaeus, 1758) NATURAL HISTORY AND PROTECTIVE RESEMBLANCE Substrate preference and the question of protective re- semblance have received a great deal of discussion in the literature. HERDMAN (1893) and ANKEL (1935) reported an association between Lamellaria perspicua and the Atlantic ascidian Didemnum candidum (=Leptoclinum maculatum) which they considered as protective to the lamellariid. Tuompson (1973, 1976) collected Lamellaria from southern Britain, none of which resembled an ascid- ian host, but instead resembled other substrata. He found Lamellaria latens which resembled the hard marine sub- stratum upon which it was collected and L. perspicua which resembled an assemblage of barnacles. Of the latter, two specimens were collected at Helford, Cornwall, with barnacle-like markings on their dorsa. So life-like was the resemblance, a hand lens was required to show otherwise. This fascinating phenomenon is documented with color photographs in YoncE & THompson (1976). On the Pacific Coast at Corona del Mar, California, MacGinitig & MacGrnitie (1949) report an unidentified yellow lamellariid feeding upon the yellow sponge, Lissodendoryx noxiosa and a brick-red lamellariid (Marseniopsis sharonae, their figure 200) feeding upon a sponge of the same color. At Puerto Pefiasco, Sonora, Mex- ico, they report Lamellaria orbiculata (probably L. diegoensis) as occurring on the underside of rocks en- crusted with either a white sponge or white tunicate, which it closely resembles. GHISELIN (1964) discusses the mor- phological and behavioral relationships of the resemblance of Marsenina stearnsii (= Lamellaria stearnsit) to the com- pound ascidian, Trididemnum opacum (Ritter, 1907) upon which it lives. McCLoskey (1973) concurs with Ghiselin that Marsenina rhombica (= Lamellaria rhombica) also resembles a tunicate colony, but occurs on other substrata as well. McCloskey collected his specimens on the solitary tunicate Ascidia paratropa (Huntsman, 1912). Although I collected many animals on barren rock sur- faces, my observations support the contention that all lamellariid species are, for the most part, tunicate-substrate specific. Of the genus Lamellaria, only L. diegoensis was observed in the field. Excluding the 8 animals collected on barren rock, 15 specimens were collected on Aplidium sp., Cystodytes lobatus (Ritter, 1900), Eugyra sp., Poly- clinum planum (Ritter & Forsyth, 1917) and Trididemnum opacum. Two specimens found on T. opacum were highly cryptic, white animals marked similarly to Marsenina stearnsit. A single specimen found in beach wash algae on Isla Vista Beach, Santa Barbara County, California, was marked with two pink coralline algae-colored areas on a brown background; this observation is most likely inci- dental and inconclusive. Seven individuals of Lamellaria diegoensis collected from a single salmon-colored colony of Aplidium near Isla Vista Beach, California, although cryptic, exhibited -7 Page 338 THE VELIGER Val. 22; No. 4 separate color and texture phases. Marked typically, indi- viduals included yellow, orange, brown, lavender and gray color regimes. I found Marsenina stearnsii to possess the very strict substrate specificity and resemblance reported by Gui- SELIN (1964). All specimens were collected on Trididem- num opacum, and were almost indistinguishable from the tunic of the host. Marsenina rhombica was collected from several tunicate species. In each case, the gastropod’s color and texture closely resembled the host substrate. Those tunicate hosts identified included Aplidium sp., Ascidia sp. and C’ysto- dytes sp. Marseniopsis sharonae was found only on Botryolloides sp., an orange, quiet-water species. Despite the wide color range of M. sharonae, it was often difficult to locate indi- viduals sunken in the surface or in folds of this encrusting tunicate species. The exact substrate preferences for Lamellaria per- spicua and L. inflata are unknown; however, color notes accompanying specimens, when available, report the an- imals closely resembled their tunicate host. PREDATOR DETERRENT AND EPIFAUNAL RELATIONSHIPS Some members of the Lamellariidae possess a predator deterrent in the form of acid secretions from glands in the mantle (THOMPSON 1960, 1969, 1976; YONGE & THOMP- SON 1976). THOMPSON (1960) studied such secretions in Lamellaria perspicua from the Atlantic. This feature has not been investigated in any other Pacific species. YONGE & THompsoN (op. cit.) presume that the acid secretion functions to dissuade a sharp-sighted predator which has not been fooled by the snail’s protective resemblance. In the laboratory, I have observed the opisthobranchs Triopha catalinae (Cooper, 1863) and Hermissenda crassicornis (Eschscholtz, 1831) to retreat hastily from the contact with the mantle of Marsenina rhombica. This contact-retreat behavior was repeated several times by the same animal. Another Pacific species_of lamellariid shows no such deterrent, and in contrast, allows a variety of fouling or- ganisms to attach to its mantle. Until this study, I was aware of only two examples of molluscan mantle epi- fauna: these were the growth of barnacles on the mantle of the keyhole limpet, Megathura crenulata (John Carter, Lockheed Center for Marine Research (LCMR), personal communication), and hydroids on the mantle of the gum- boot chiton, Cryptochiton stelleri (Douglas Hunt, LCMR, personal communication). I am not aware, however, of any published reports on such associations on living meso- gastropod mantles. I noted epifaunal associations on Marseniopsis sharonae in Morro Bay. In 9 instances, I observed live barnacles, Balanus sp., attached to the living non-retractile mantle tissue of the host. The barnacles measured up to 1.5mm diameter. A minute hydroid, Plumularia sp., was found on 6 specimens. Like the barnacles, the hydroids were firmly attached to the mantle tissue. The third species growing on mantle tissue was the scud, Corophium sp. These were found with their secreted mucous tubes attached to the mantles of five specimens. On the most heavily encrusted specimen, the Corophium mat covered nearly one-half of the exposed mantle. A subsequent association, most certainly due to the scud encrustation, was the occurrence of the boring clam Hiatella arctica. Two to 4 individuals were found within the mucous mat encrusting all 5 lamellariids. Hiatella and Balanus were also found living on the tunic of the tunicate Botryolloides sp. on which the Marseniopsis were collected. ACKNOWLEDGMENTS I wish to express my appreciation to Pacific Gas and Elec- tric Company, Department of Engineering Research for the use of their laboratory facilities during the study. Many thanks to Dustin Chivers (CAS), Gale Sphon and James McLean (LACM), Erick Hochberg (SBMNH), Don Cadien, Bob Henderson, Gary McDonald and Carol Skog- lund for their loan of specimens. Thanks also to Jim Lance, Brita Larsson, Harald Rehder, Virginia Waters, Jim Carl- ton and Steve Long for their assistance and support; and to Bud Laurent, John Warrick and the referees for their critical review of the manuscript. Literature Cited Assort, Rosgrat Tvoxer 1974. American Seashells. and ed., Van Nostrand Reinhold Go., New York; 663 pp., 4000+ text figs.; 24 plts. (in color) Apams, Cartes BAKER 1852. Catalogue of shells collected at Panama with notes on synonymy, station and habitat. Ann. Lyc. Nat. Hist. New York 5: 229 - 296 (June); 297 - 549 (July) [reprinted: 1852, R. Craighead, New York. viii +334 pp.] ANKEL, WuLFF EMMO 1935. Das Gelege von Lammellaria perspicua. Okol. Tiere go: 635 - 647 Beroxu, Lupwio SopHus RupoLr Zeitsch. Morphol. 1853. _Bidrag til en monografi af Marseniaderne. Dansk. Viden. Selsk. Skr. (5) 3: 1-119; plts. 1-5 1886a. Die Marseniaden, eine vorlaufige Mittheilung. Zool. Jahrb. Syst. 1 (1): 165-176 Vol. 22; No. 4 THE VELIGER Page 339 a ttn 3886b. Report on the Marseniadae collected by H. M.S. Challenger during ae years 1873-1876. Rep. Sci. Res. Challenger, Zool. 15: 1-25; pit. 1 1908. The Opisthobranchiata of South Africa. Soc. 17: 1-110 BsrtscH, Hans « AvBEric A. SMITH 1970. Observations on opisthobranchs of the Gulf of California. The Veliger 13 (2): 171-174 (1 Qctober 1970) BrsHor, M. J. « S. J. BisHopr 1973. A census of marine prosobranch gastropods at San Diego, Cali- fornia. The Veliger 16 (2): 143-152 (1 October 1973) BrooksHIRE, JACK 1968. Mollusca of the San Luis Obispo County area. Part 1 — Gastro- poda. The Tabulata 1 (4): 5-6 (1 April 1968) Burcu, Jonn Quincy 1946. (no title) in Minutes Conchol. Club So. Calif. 57: 1-2 Cate, JEAN McCrezry & Crawrorp NEILL Cate 1962. The types of Lamellaria sharonae Willett, 1939 (Gastropoda). The Veliger 5 (2): 91 (1 October 1962) Dari, Wituiam Hearey 1871. Descriptions of sixty new forms of mollusks from the west coast of North America and the North Pacific Ocean, with notes on others already described. Amer. Journ. Conch. 7 (2): 93-160; plts. 13-16 (2 November 1871) DuSuane, Heven « Gaz G. SPHON 1968. A checklist of intertidal mollusks for Bahia Willard and the southwestern portion of Bahia San Luis Gonzaga, State of Baja Cali- fornia, Mexico. The Veliger 10 (3): 233 - 246; plt. 35 (1 Jan. ’68) DuSwHanez, Heren & Roy PoorMAN 1967. A checklist of mollusks for Guaymas, Sonora, Mexico. The Veliger 9 (4): 413-441 (1 April 1967) Frerrer, VERA & TAIR GRAHAM 1962. ‘British prosobranch molluscs, their functional anatomy and eco- logy. London, Ray Soc. xvi+755 pp.; 316 figs. Guisziin, MicuazEL TENANT 1964. Morphological and behavioral concealing adaptations of Lamel- laria stearnsit, a marine prosobranch gastropod. The Veliger 6 (3): 123-124; pit. 16 (1 January 1964) Gray, Maria Emma 1850. Figures of molluscous animals selected from various authors, etched for the use of students. vol. 4, iv+219 pp., Brown, Green & Longmans, London Herpman, W. A. 1893. Mimicry of Lamellaria perspicua. to 130 Howarp, Fayz B. 1972. Then and now. Human, Vernon L. 1972. | The marine Mollusca of Newport Bay, California. The Ta- bulata 5 (4): 10-12 & 23-24 (1 October 1972) 1973. | Marine Mollusca in kelp holdfasts at Paradise Cove, California. The Tabulata 6 (3): 12-15 & 21-23 (1 July 1973) Kuzn, A. Myra, with the assistance of Jamzs Hammtton MoLzan 1971. Sea Shells of Tropical West America; marine mollusks from Baja California to Peru. and ed. Stanford Univ. Press, Stanford, Calif i-xiv + 1064 pp.; ca. 4000 figs.; 22 color pits. (1 September 1977) La Rocauz, AURRLE 1953. Catalogue of the Recent Mollusca of Canada. Mus. Canada 129: xi+ 406 pp. MacGiniriz, Gzorce Ener & NeETTiz MACGINITIE 1949. Natural history of marine animals. New York. pp. i- xii + 1-473; illust. Mascus, Ernst ; 1959. Lamellariacea und Opisthobranchia. Reprt. Lunds. Univ. Chile Exped. 1948-49. No. 36 Lunds Univ. Arssk., N. FE (2) 55 (9): 1 to 133; figs. 1 - 196 1963. | Opisthobranchs from the Lesser Antilles. ragao 19 (79): 1-76 Maacus, Eve.ing pu Bois-ReymMonp & Exnst Marcus 1960. | Opisthobranchs from American Atlantic warm waters. Bull. Mar. Sci. Gulf and Caribb. 10 (2): 129-203 1967. American opisthobranch mollusks. Stud. trop. Oceanogr. Miami 6: 8+256 pp.; 1 plt.; 250 text figs. (December 1967) 1969. Opisthobranchian and lamellarian gastropods collected by the “Vema.” Am. Mus. Novit. No. 2368: 1 - 33 (24 April 1969) Trans. So. Afr. Phil. The Conchologist 2: 129 The Tabulata 5 (3): 17-26 (1 July ’72) Bull. Nat. McGraw-Hill Book Co., Stud. Fauna Cu- McCiosxey, L. R. ‘ 1973. Development and ecological aspects of the echinospira shell of Lamellaria rhombica Dall (Prosobranchia; Mesogastropoda). O- phelia 10 (2): 155-168 (October 1973) McLean, James HamILTon 1969. Marine shells of southern California. Los Angeles County Mus. Nat. Hist. Sci. Ser. 24, Zool. 11: 104 pp.; 54 figs. Monracu, GzorcE 1815. An account of some new and rare marine British shells and animals, Trans. Linn. Soc. London 11 (2): 179-204; pits. 12-14 Morris, Percy A. 1966. A field guide to shells of the Pacific Coast and Hawaii. Houghton Mifflin, Boston, 2d ed., 297 pp. Oxproyp, Ipa SHEPARD 1924. Marine shells. of Puget Sound and vicinity. Univ. Wash. Puget Sound Biol. Sta. 4: 272 pp.; 49 plts. (March 1924) 1927. The marine shells of the west coast of North America. Stan- ford Univ. Press, Stanford, Calif. 2 (3): 605-941; plts. 73 - 108 [Chitons: 848 - 925] 1885. Notes on the mollusks of the vicinity of San Diego, Cal., and Todos Santos Bay, Lower California. Proc. U.S. Nat. Mus. 8: 534 - 544 (10 October) ; plt. 24 (21 October) ; 545 - 552 (26 October) Rocers, Jutia ELLEN 1936. The shell book. Charles T. Branford Co., Boston 1951. ‘The shell book (rev. ed.) Charles T: Branford Co., Boston, xxi+503 Pp.; 77 plts. Suasxy, Donatp R. 1961. Notes on rare and little known Panamic mollusks. The Veliger 4(1): 22-24; part plt. 4 (1 July 1961) SsatH, ALtyn Goopwin « MacKznziz Gorpon, Jr. 1948. The marine mollusks and brachiopods of Monterey Bay, Cali- fornia, and vicinity. Proc. Calif. Acad. Sci. (4) 26 (8): 147-245; pits. 3, 4; 4 text figs. (15 December 1948) SpHon, Garg G. 1975. Marsentopsis sharonae (Willett, 1939), comb. nov. The Nautilus 89 (3): 95 Stronc, ArcHIBALD McCiure « G Darras HANNA 1930. Marine Mollusca of the Tres Marias Islands, Mexico. Proc. Calif. Acad. Sci. (4) 19 (3): 13-22 (4 June 1930) Stronc, ArcHIBALD McCiure e# Lzo Gzorcz HERTLEIN 1939. Marine mollusks from Panama collected by the Allan Hancock Expedition to the Galapagos Islands, 1931-1932. Allan Hancock Pacif. Exped. 2 (12): 177-245; pits. 18-23 THr.z, JoHANNES 1929. Handbuch der systematischen Weichtierkunde. Erster Teil, Lori- cata; Gastropoda. I: Prosobranchia (Vorderkiemer). pp. 1 - 376, Jena, Gustav Fischer 1934. | Handbuch der systematischen Weichtierkunde. 2(3): 779 to 1154; figs. 784 - 893 THompson, THOMAS EVERETT 1960. Defensive acid-secretion in marine gastropods. Journ. Mar, Biol. Assoc. U. K. 39: 115-122 1969. Acid secretion in Pacific Ocean gastropods. Austral. Journ. Zool. 17: 755 - 764 ; 1973. Protective resemblances in British Lamellaria. Journ. Conch. 28 (2): 75-78 1976. Biology of opisthobranch molluscs. Ray Soc. London. vol. 1: 207 pp.; pits. 1-8, 10-21; 106 text figs. Wenz, WILHELM 1938-1944. Gastropoda. Allgemeiner Teil und Prosobranchia. In: O. H. Schindewolf (ed.), Handbuch der Palaozoologie 6 (1): i- xii; 1 - 1639; illustr. Gebr. Borntraeger, Berlin WrtettT, Gzorcz 1939. Description of a new mollusk from California. lus 52 (4): 123-124; plt. 9, figs. 1, 1a, 1b WIncKworTH, RONALD 1932. The British marine Mollusca. 21) - 252 Yates, Lorenz G. 1890. The Mollusca of Santa Barbara County, California. Bull. Santa Barbara Soc. Nat. Hist. 1 (2): 37-45 Yoncse, Cnartes Maurice & THomas Everett THOMPSON 1976. Living marine molluscs. Wm Collins Sons, St. James’ Place, London, 288 pp. The Nauti- (1 April 1939) Journ. Conch. London 19 (7): Page 340 THE VELIGER Vol. 22; No. 4 Predation by the Prosobranch Mollusk Lamellaria diegoensis on Cystodytes lobatus, a Colonial Ascidian BY GRETCHEN LAMBERT Department of Biological Science, California State University, Fullerton, California 92634 and Hopkins Marine Station of Stanford University, Pacific Grove, California 93950 (1 Text figure) INTRODUCTION THE LAMELLARID GASTROPODS are highly specialized for predation on ascidians (HERDMAN, 1893; FRETTER & GRAHAM, 1962; GHISELIN, 1964). All the lamellariids are cryptically colored; some resemble their prey while others resemble the general substrate of their surroundings (THompson, 1973). Their overall biology is poorly under- stood, however, because collections have been of only one or a few specimens at a time, and lengthy studies have not been carried out. The present paper describes the predator- prey relationship between Lamellaria diegoensis Dall, 1885, and the compound ascidian Cystodytes lobatus (Ritter, 1900). The results of several feeding experiments and calorimetric measurements of the ascidian colonies and mollusk feces are included, with a partial energy budget for L. diegoensis computed from these data. All observations and experiments were carried out on a single L. diegoensis because although C. lobatus is very common in the low intertidal and subtidal zones in central Cali- fornia, only one L. diegoensis was collected despite a con- certed effort to find more over a six-month period. In spite of the drawbacks of the small sample size, this work was carried out because it may well be that the only way we shall ever gain a fairly comprehensive knowledge of the biology of the lamellariids is from the compilation of many separate studies. METHODS One Lamellania diegoensis was collected on March 7, 1978, on a piece of Cystodytes lobatus colony from the low intertidal zone during a -18 cm tide at Pt. Pinos, Pacific Grove, California. The piece of C. lobatus colony and ad- ditional colonies were maintained in an aquarium with running sea water at the Hopkins Marine Station; the mollusk was not actually observed on the colonies until March 23, even though the colonies had been examined daily in conjunction with another study (LAMBERT, 1979). The mollusk was maintained solely on a diet of C. lobatus from March 7 until August 31, 1978. For most of this period it was kept cool on the water table in a Pyrex baking dish loosely covered with black plastic. This partial darkness was provided because like Lamellaria stearnsii Dall, 1871, studied by GHIsELIN (1964), L. diegoensis re- acted strongly and negatively to bright light. The water was changed once a day and a new piece of ascidian colony offered every two to three days. During July and August, a total of 15 daily fecal pellet counts, wet weights and dry weights were made with the aim of ultimately using these values to determine the amount of prey consumed and therefore the calories in- gested. To this end, pieces of fresh ascidian colonies and the Lamellaria fecal pellets were dried to constant weight at 55°C (Table 1), and the caloric content determined using a Parr model 1411 semimicro bomb calorimeter (Table 2). Ash determinations were made with a muffle furnace for the C.. lobatus colonies; there were not enough mollusk feces for this procedure, so the percent ash given in Table 2 was calculated from the bomb residue. In both cases the ash content has been corrected for CaCO, endo- thermy according to PAINE (1966, 1971) because of the large quantity of ascidian spicules. No acid corrections were made because of the large amount of naturally oc- curring acid in these animals. Vol. 22; No. 4 THE VELIGER Page 341 Table 1 Wet and dry weights (mg) of Cystodytes lobatus colonies and Lamellaria diegoensis fecal pellets. Cystodytes lobatus Lamellaria diegoensis fecal pellets/24 hrs Wet Dry Ratio Date # Pellets Wet Dry Ratio 2877 294.5 0.102 7/13/78 207 88.0 22.8 0.259 3277 318.5 0.097 7/14 162 84.9 23.3 0.274 3393 327.0 0.096 7/17 147 80.6 18.3 0.227 2786 253.0 0.091 7/20 128 58.9 11.8 0.200 3573 331.5 0.093 7/26 186 88.7 26.1 0.294 3462 297.0 0.086 8/1 134 81.4 18.9 0.232 3210 284.5 0.089 8/6 172 95.1 22.6 0.238 3299 275.0 0.083 8/10 104 49.4 12.6 0.255 3261 290.4 0.089 8/11 150 81.3 19.5 0.240 3418 284.6 0.083 8/14 112 62.5 15.9 0.254 8/15 177 87.1 21.5 0.247 x 3256 295.6 0.091 8/16 165 84.0 17.6 0.210 +0.006 8/19 132 62.6 1B}. 0.209 8/20 136 64.8 13.1 0.202 8/21 162 78.9 WAI 0.153 152 76.5 17.9 0.233 x + 28 +13.4 + 4.7 +0.035 Table 2 Average energy values (calories/g ash-free dry wt) of Cystodytes lobatus colonies and Lamellaria diegoensis fecal pellets. x cal g—! ash- % ash % ash free dry wt{+SD) # det. (bomb det.) (muffle furnace det.) # det. Cystodytes lobatus 3679 + 244 7 62.9 63.3 3 whole colonies Lamellaria diegoensis 5578 + 107 5 78.8 — fecal pellets RESULTS ann DISCUSSION EXTERNAL MorpHotocy oF Lamellaria diegoensis The following description of the living mollusk adds to BEHRENS’ (1980) redescription (this issue) of this species. The shell was completely internal, with no opening on the mid-dorsal mantle surface to expose the shell. The pale pink mantle glistened with many shiny refractile spots. Flecks of black pigment were scattered everywhere, but they were especially numerous on the variable-sized con- tractile tubercles, which were largest mid-dorsally. Small tubercles extended right to the mantle edges and were also pale pink. Orange pigment could be seen inside some of the largest tubercles; its color closely matched the orange yolk of tadpole larvae developing in the ascidian colonies. White material was concentrated at the base of some of the tubercles; it resembled the white calcareous spicules that occur in large numbers in Cystodytes lobatus and form sacs that surround each zooid abdomen (Lam- BERT, 1979). Many yellow pigment spots occurred on the inside of the mantle and on the top and bottom of the foot that were not visible in a dorsal view. The foot, visible posteriorly only when the animal was moving, also had a large number of red pigment spots. Like all the lamellariids (FRETTER & GRAHAM, 1962; THOMPSON, 1969), the man- tle contained numerous acid glands that produced sulfuric acid, as indicated by barium sulfate precipitation when barium chloride was added to a drop of mantle fluid. In this respect it again resembled its prey, for the test of C. lobatus is packed with numerous sulfuric acid filled Page 342 THE VELIGER Vol. 22; No. bladder cells (Abbott « Newberry, in press) 35 to 50um in diameter (LAMBERT, 1979). The anterior end of the mantle formed a siphon that was usually greatly extended when the mollusk was mov- ing. When it was resting or feeding on Cystodytes, its siphon and dorsal tubercles were considerably shortened and its body much flattened. The lateral mantle edges be- came spread out, giving the animal an almost circular Figure 1 Lamelleria diegoensis at rest on a colony of Cystodytes lobatus scale bar 2cm outline (Figure 1). The mantle edges became thin, trans- lucent and nearly colorless, blending in with the bladder cell filled tunic of C. lobatus so well that the outline of the mollusk was difficult to distinguish. FEEDING ON Cystodytes lobatus Cystodytes lobatus occurs primarily in two color forms, translucent white and pale pink. The Lamellaria diegoensts blended somewhat better on the pink colonies and was originally collected on a pink colony but showed no feeding preferences between the two color types. The method of feeding has been described in detail by FRETTER & GRAHAM (1962): The animal rasps through the ascidian test and sucks up the zooids, leaving a hole with scalloped edges. In Cystodytes the zooid abdomens are surrounded by spic- ular sacs made up of closely overlapping disc-shaped cal- cium carbonate spicules. Inserting a fine probe into the tunic ruptures some of the acid-filled bladder cell mem- branes beyond the spicular sac, resulting in immediate dissolution of nearby spicules and release of COz bubbles. The L. diegoensis ate right through the tunic, ingesting large numbers of spicules along with the zooids, and many of the spicules appeared in the feces almost undamaged in appearance, indicating that the mollusk was somehow able to neutralize the bladder cell contents quickly. The feces were either white or pink depending on the color of the tunicate colony being eaten. The feces had a striped appearance, with layers of spicules and tunic material alternating along the length of the pellet. This might re- flect the alternating order in which these parts were in- Table 3 Results of Lamellaria diegoensis (L.d.) feeding on Cystodytes lobatus (C.1.) Beginning wet wts. (g) End wet wts. (g) Difference (g) Fecal Pellets Total Control Control Control # of wet Date L.d. C1. CL. Ld. CL. CLL. L.d. CL Cl. pellets wt. (g) 8/4 beg 6.528 10.286 8.999 5.154 9.219 9.079 —1.374 —1.067 +0.08 266 0.147 8/6 end C.l. Net loss = —0.987 = 133/day = 0.074/day (2 days) = —0.4935 g/day 8/8 beg 5.17 13.65 16.044 5.145 11.696 15.532 = 025 —1:954 > —0.512 366 0.188 8/12 end C.l. Net loss —1.442 = 92/day —0.047/day (4 days) = —0.3605 g/day | 8/18 beg 5.672 20.864 34.696 5.986 18.08 33.01 +0314 —2.784 ° —1.686 521 0.250 8/21 end C.l. Net loss = —1.098 = 174/day =0.083/day (3 days) = —0.366 g/day x=128 x=0.065 + 38/day + 0.017/day Vol. 22; No. 4 gested. In Table 1 are the wet and dry weights for 10 pieces of C’. Jobatus colonies and for 15 daily fecal pellet accumulations for L. diegoensis. The calorimetric analysis of these dried materials is given in Table 2. Several feeding experiments were carried out (Table 3) to determine the average amount of food eaten per day and correlate this with the number and weight of fecal pellets produced per day. Unfortunately the handling, blotting dry and weighing of Lamellaria had a definite negative effect. The results indicate, however, that the animal was acclimating (compare beginning and end weights for each of the feeding runs), and better results would probably have been obtained if the experiments had been continued longer. The feeding experiments ap- pear so variable as to be unusable, but internal checks show a surprisingly close correlation. For example, the ratio of the number of fecal pellets produced per day dur- ing the feeding experiments (128) to the average number from Table 1 (152) is nearly the same as the ratio of the weight of dry feces produced per day during the feeding trials (15mg) to the average dry wt. from Table 1 (17.9 mg). These two sets of numbers were obtained indepen- dently by direct measurements and counts. Between May 3 and August 4, 1978, the 3 months prior to the feeding trials, the animal increased in weight from 2.2 to 6.5g, indicating a steady gain. Assuming that Lamellaria had maintained a constant weight during the feeding experi- ments, the average net weight loss of Cystodytes was 0.392 .058g wet wt/day (all losses added together and divided by 9 days total feeding time). This value is equiv- alent to 0.036 g dry wt/day, or 0.013 g ash free dry wt/day after subtracting for the 63.3% ash content (Table 3). The result is 48.6 calories ingested per day. The lamellariid produced an average of 0.065g wet weight of feces per day during the feeding experiments. Multiplying this by the dry/wet wt ratio from Table 1 results in a value of 0.015 g dry feces/day, or 0.003 g ash free dry wt/day after subtracting the 78.8% ash content. This is equivalent to 17.9 calories/day lost in the feces. Thus, the net caloric intake for Lamellaria diegoensts was 30.7 calories/day (=48.6-17.9 cals/day). Given that the Lamellaria weighed 6.53 g at the start of the feeding ex- periments, on a per gram basis its daily intake would be 4.7 calories/g body wt/day. This value is probably low when compared to the average number of fecal pellets produced per day when the animal is undisturbed (Table 1). Using this latter value, one can compute that the aver- age net caloric intake would be 36 calories/day when the animal is undisturbed, or 5.5 calories/g body wt/day. THE VELIGER Page 343 To determine feeding preferences, a few of the ascidian species growing next to Cystodytes colonies in nature were offered to Lamellaria diegoensis. The mollusk was starved for 3 days, then placed with Aplidium sp., Polyclinum planum (Ritter & Forsyth, 1917) or didemnids (unid.) for 24 hours. In no case were any of these ascidians eaten. When the Lamellaria was then offered Cystodytes it im- mediately began to feed, even in rather bright light before its dish was covered with black plastic. SUMMARY Lamellaria diegoensis, a prosobranch gastropod, was col- lected intertidally on the compound ascidian Cystodytes lobatus at Pacific Grove, California. A description is given of the mollusk’s external morphology and coloration in order to compare it with its prey. Lamellaria diegoensis closely resembles its prey in color and texture, and certain behavioral traits enhance its cryptic nature. It was main- tained in the laboratory on a diet of only C. lobatus for 6 months; during this period its weight nearly tripled, from 2.2 to 6.5 grams. Several feeding experiments were per- formed, and these results together with bomb calori- meter values for the ascidian colonies and mollusk feces were used in computing a partial energy budget for L. diegoensis. During the feeding experiments, the animal ingested 48.6 calories/day, of which 17.9 calories/day were lost in the feces, resulting in a net caloric intake of 30.7 calories/day, or 4.7 calories/g body wt/day. The caloric values given here are only preliminary; they are included in the hope that if this work is repeated they can be used on a comparative basis. All the lamellariids share a similar mode of existence; therefore their energy budgets should be comparable. Because their diet is so re- stricted, if a reliable conversion can be made between number of feces and calories ingested, a variety of feeding experiments are possible with little perturbation to the mollusk. ACKNOWLEDGMENTS I wish to thank the staff at the Hopkins Marine Station for providing space for me, in particular Dr. Donald Abbott in whose laboratory this work was carried out. Dr. Abbott’s help and encouragement are greatly appre- ciated. David Behrens identified the Lamellaria, and he and Dr. Roger Seapy made valuable comments on the Page 344 THE VELIGER Vol. 22; No. 4 manuscript. I also thank Dr. C. E. Jones and Larry Colin, who assisted me with the bomb calorimeter, and Faylla Chapman, for her willing assistance in many ways. J. Cooper, Y. Fadallah, C. Harrold, A. Hines and R. Sellers aided in subtidal collection of Cystodytes lobatus. I grate- fully acknowledge the constant support, technical assist- ance and companionship of my husband Charles through- out this work. Literature Cited Assott, DonaLD PuTNaM & ANDREW Topp NEWBERRY In press. Urochordata: the tunicates. In: R. Morris, D. PB. Abbott « E. C. Haderlie, eds., Intertidal invertebrates of California. Stan- ford Univ. Press Bexnrens, Davp W. 1980. The Lamellariidae of the North Eastern Pacific. 22 (4): 323 - 339; 2 plts.; 15 text figs. The Veliger (1 April 1980) Frerrer, Vera e ALAsTAR GRAHAM 1962. British Prosobranch Mollusks. i- xvit+755 pp., illust. Ray Soc., London GuiseLin, MicHAEL TENANT 1964. Morphological and behavioral concealing adaptations of Lamel- laria stearnsii, a marine prosobranch gastropod. The Veliger 6 (3):123-124; pit. 16 (1 January 1964) Herpman, W. A. 1893. ‘Mimicry’ of Lamellaria perspicua. LAMBERT, GRETCHEN 1979. | Early post-metamorphic growth, budding and spicule forma- tion in the compound ascidian Cystodytes lobatus. Biol. Bull. 157: Orcutt, Cartes Russe (with comments by WittiAm Heatey Dat) 1885. Notes on the mollusks of the vicinity of San Diego, Cal., and Todos Santos Bay, Lower California. Proc. U. S. Natl. Mus. 8 (536): 534-552; plt. 24 (so September 1885) Paring, Rosert TREAT 1966. | Endothermy in bomb calorimetry. 126-129 1971. The measurement and application of the calorie to ecological problems. Ann. Rev. Ecol. Syst. 11: 145 - 164 THompPson, THomAS EVERETT 1969. Acid secretion in Pacific Ocean gastropods. Zool. 17: 755 - 764 1979. Protective resemblances in British Lamellaria. Journ. Conch. 28: 75-78 Conchologist 2: 129 - 130 Limnol. Oceanogr. 11: Austral. Journ. Vol. 22; No. 4 THE VELIGER Page 345 Stone Boring Marine Bivalves from Monterey Bay, California BY E. C. HADERLIE Department of Oceanography, Naval Postgraduate School, Monterey, California 93940 (1 Text figure) INTRODUCTION MonTEREY Bay HAS BEEN A CENTER for shell collecting by conchologists for more than a hundred years. Included in many of the early collections were representative bivalve borers of the families Mytilidae and Pholadidae, and Mon- terey Bay is the type locality for several species. SMITH & _ Gorpon (1948) summarized these early studies and pre- sented a list of all boring and nestling bivalves reported from Monterey Bay up to World War II. For many years after the Smith & Gordon paper appeared, little work was published on stone boring animals in Monterey Bay. In recent years, however, there has been renewed interest in this important group of animals and several theses and published papers have been partially or totally devoted to them (BoorH, 1972; BuRNETT, 1972; CLARK, 1978; Donat, 1975; HapERLIE, 1976, 1977, 1979; HADERLIE & Donat, 1978; Haver et al., 1974; MINTER, 1971). Since 1970, a continuous long term study on marine bivalve stone borers by students and staff of the Naval Postgraduate School in Monterey has been underway. This study has consisted of two parts. First, the horizontal and bathymetric distribution and substrate preference of living borers within Monterey Bay have been investi- gated. This has involved extensive shore collecting, shal- low water diving, and shipboard dredging operations. Second, experimental studies aimed at determining repro- ductive seasons, settling times, growth rates, and longevity of individual borers have been carried out. Studies have made use of experimental stone panels placed in the sea at various times and depths and for varying periods and then recovered and examined for evidence of bivalve borer settlement and growth. This paper presents the results to date of the first part of this study, namely the identity, distribution, abundance, and substrate preference of stone boring bivalves of the families Mytilidae and Pholadidae and associated nestlers in Monterey Bay. ACKNOWLEDGMENTS I want to acknowledge the help of many people, but espe- cially Captain Reynolds and the crew of R/V Acania for their unfailing cooperation and help during many arduous hours of dredging operations over many years in all weather conditions on Monterey Bay. Also, I want to thank my colleagues and students who assisted in the hundreds of shallow water Scuba dives for making observations and collections, photography, or recovery of lost gear such as dredges. These include G. C. Booth, G. W. Clark, W. Donat, J. C Mellor, C. S. Minter, and J. Norton. In the early part of the study Dr. Ruth Turner of Harvard as- sisted in the identification of borers and has given advice on many phases of the work. Drs. R. S. Andrews and W. C. Thompson of the Naval Postgraduate School and Dr. H. G. Greene of the U.S. Geological Survey helped in sorting out the complex geology of Monterey Bay, and Florence Lee-Wong of the U.S. Geological Survey made the petrographic analyses of thin sections of various rock samples. The office of Naval Research, Oceanic Biology Program, has provided long term financial support for this project, and the Naval Postgraduate School Research Foundation Program has assisted in the intertidal part of the project. TOPOGRAPHY ann GEOLOGY oF MONTEREY BAY Monterey Bay is a large open embayment along the central California coast some 115 km south of San Francisco (see Figure 1). The mouth of the Bay is about 37km in width when measured from Point Santa Cruz in the north to Point Pinos in the south. The shoreline at the southern- most part of the Bay, on the Monterey Peninsula, is a rocky headland composed of Santa Lucia granodiorite of Cretaceous age. From Monterey eastward and north to Page 346 THE VELIGER Vol. 22; No. 4 : Rae i One! Terrace Pt. Pt. Santa Cruz SEB EP ~ i 2 , _ Soquel Pt. .X.7 SS Natural Bridges fee Vd 27 ve \ EES fi Elkhorn Slough MONTEREY (ia so Pt. Pinos J: Fort Ord y, A (~~ Del Monte Beach e * Cabrillo Pt. {+ . Dredging Transects Cannery Row Monterey Harbor | Cc Pe bli Bo Monterey Vol. 22; No. 4 Soquel Point there are broad sandy beaches backed by dune fields in the south and cliffs in the north. The shore- line in the vicinity of Santa Cruz is composed of steep bluffs with flat-topped terraces. These sea cliffs are Qua- ternary marine terrace deposits which overlie Pliocene Purisima Formation at Point Santa Cruz. This latter for- mation is exposed as yellow sandstone and siltstone in the intertidal zone. To the west of Point Santa Cruz the sea cliffs are composed of Pliocene Santa Cruz mudstone over- lying layers of chert of Miocene Monterey Formation. This latter formation extends as broad terraces or finger reefs interspersed with sandy pockets into the intertidal and subtidal zones. The subtidal topography and geology of Monterey Bay is diverse and complex. The sea bottom relief is dominated by the Monterey Submarine Canyon which originates less than 2km west of Elkhorn Slough and extends westward for over 90 km as a deep, V-shaped cut in the continental shelf. The remainder of the Bay bottom is a relatively flat, gently sloping continental shelf, covered with unconsoli- dated sediments and interrupted by a few rocky reef out- crops. GALLIHER (1932) was the first to publish informa- tion on the location and lithologies of these rocky outcrops. SHEPARD & EMERY (1941), MARTIN (1964), and GREENE (1977) have extended these observations in many details and have also reviewed the extensive literature on the geology of Monterey Bay. During these previous studies, and from dredge hauls made during the present investi- gation, rock samples from many of these outcrops and from the walls of the Monterey Submarine Canyon have been obtained. Samples of porphyritic biotite granodiorite have been dredged from off Point Pinos and from the south wall of the Monterey Canyon. This granitic rock that forms the basis of the Monterey Peninsula is apparently also the dominant basement rock under Monterey Bay. The sedimentary rocks dredged from the bottom of the Bay are mudstones, siltstones, sandstones, cherts, and con- glomerates derived from the Monterey Formation of mid- dle Miocene age and from Pliocene Purisima Formation. AREA or STUDY This study has been limited to Monterey Bay which is here defined as the area to the east of a line running from Point (< on facing page) Figure 1 Map of Monterey Bay Area THE VELIGER Page 347 Pinos on the Monterey Peninsula to Terrace Point west of Santa Cruz in the north. The area of most intense study, and that from which we have the most complete data, is the shallow subtidal zone off Del Monte Beach to the east of Monterey harbor at the most southerly end of Monterey Bay (Figure 1). This area is roughly defined by the exten- sive kelp bed made up primarily of Macrocystis pyrifera (Linnaeus, 1771) whose holdfasts are attached to the dis- continuous outcrops of shale of the Monterey Formation. The water depth in the area of shale outcrops is from 10 to 20m. In most of the area the bottom consists of rela- tively flat smooth shale, sometimes covered with sand to varying thickness, at other times completely exposed ex- cept for low lying sand pockets. In roughly the middle of the study site off Del Monte Beach, the topography is much more rugged with hummocks and ledges distributed parallel to the trend of the Tularcitos Fracture Zone which enters the Bay from the southeast. Some of these ledges are continuous ridges rising 2m or more above the adjacent bottom and running several hundred meters seaward. The shale in this area has been examined and sampled re- peatedly over the past 10 years by Scuba diving, and the benthic fauna and flora, including the distribution of sev- eral species of boring bivalves, have been investigated (BooTH, 1972; BURNETT, 1972; HapDERLIE, 1976; HapeEr- LIE et al., 1974; MINTER, 1971). The present investigation has continued these studies and extended them seaward by using shipboard dredging techniques to sample the exposed shale in deeper water off Del Monte Beach and throughout Monterey Bay. Some dredge hauls were made in areas previously studied by shallow water diving in order to recover larger samples of rock substrate than can be collected conveniently by divers. During this investigation, we have successfully collected bottom samples on 40 dredge hauls in Monterey Bay (Figure 1). In addition to the subtidal areas of Monterey Bay de- scribed above, we have surveyed the intertidal zones for sedimentary rocks which might harbor bivalve borers. At the southern end of Monterey Bay along the shore of the Monterey Peninsula only granitic rock is exposed in the intertidal zone and no bivalve borers can penetrate this hard basement material. From Monterey harbor around the Bay to Soquel Point in the north, the shore consists of sandy beaches. But from Soquel Point westward to Ter- race Point, sedimentary rocks of both the Purisima For- mation and the Monterey Formation are exposed in the intertidal zone as reefs or sea cliffs. We selected two of these reefs to the west of Point Santa Cruz for intensive study of bivalve borers. Page 348 METHODS ann EQUIPMENT Our objectives in this study were to determine the identity, density of numbers of individuals, distribution horizontally and with depth, and substrate preference of mytilid and pholad borers in sedimentary rocks in and around Mon- terey Bay. As these borers are hidden, sometimes deeply, in the substrate, with only the distal ends of the siphons projecting out of the burrow, in order to find, identify, and count the borers it was usually necessary to mechan- ically break the rocks apart and remove the animals one at a time from their burrows. By removing and keeping representative living mytilid and pholad borers in sea water aquaria we were able to study the size, morphology, and color of the extended siphons after the animals had been positively identified using shell characteristics. With this information we were able, while diving in shallow water, to identify and quantify many of the bivalve borers found in the Monterey shale off Del Monte Beach. A photographic catalogue of typical siphons with the animals in situ in subtidal rock has been prepared (see Boor, 1972, HaDERLIE e¢ al., 1974, and MINTER, 1971, for details of techniques and representative photographs). One major aim of this investigation was to sample sedi- mentary rock for borers in water below diving depths in Monterey Bay and to collect large rock samples from shal- low water that were impossible to recover by diving. The Naval Postgraduate School’s R/V Acania was used for most dredging operations using a heavy duty chain bag dredge with a mouth opening measuring 45cm by 85cm. Attempts were made to locate and sample most of the sedimentary rock outcrops in Monterey Bay. In some cases specific areas were re-visited where earlier workers had recovered rock while dredging. This was particularly true of the dredging stations on the south wall of the Monterey Canyon. In other areas, such as in the shallow water off Del Monte Beach and off Santa Cruz, parallel tows were made through the kelp beds at progressively deeper loca- tions. Beyond the kelp beds, where we had no idea if any solid substrate was exposed, random tows were made some- what parallel to one another and running generally from southwest to northeast. In this way, a substantial part of the bottom was sampled particularly in the southern bight of the Bay south of 36°38’30" N (a line running east from Point Pinos to Fort Ord). After deciding on the specific locality to be sampled, the dredge was towed slowly over the substrate. If the behav- ior of the towing cable indicated that the dredge had been hitting solid substrate and breaking off pieces of rock so that the chain bag was full or nearly so, the dredge was recovered after having passed over 100m or less of bot- THE VELIGER Vol. 22; No. 4 tom. In other cases the dredge was towed for up to 500m, sometimes picking up rock at the very end of the tow, sometimes coming up empty. If the dredge collected a sample early in the tow it was recovered and then lowered again and the tow continued along the same transect. Thus, the samples of rock recovered came from various places along the transect, but we were never sure where the dredge actually collected the sample. On the south wall of the Monterey Canyon the usual dredging technique was to position the ship over the deep part of the Canyon, lower the dredge to the bottom, then allow the ship to drift slowly south or east so that the dredge would be dragged upward over the wall to the edge of the shelf. The location of various dredging transects is shown in Figure 1, and Table 1 lists the various dredging operations and the recovered samples. The position of the ship at the beginning and end of each transect was determined by visual bearings when close along shore or by Loran C when offshore. The depth of water and amount of dredging cable out was then used to estimate the path of the dredge along the bottom. This was then plotted on a standard No. 18685 (formerly C&GS No. 5403) chart of Monterey Bay. This chart with the plotted transects is on file in the Department of Oceanography, Naval Postgraduate School. In Figure 1, and in the location column of Table 1, only the general locality and water depth of the transects are given. In order to learn more about the type of stone pene- trated by borers, representative samples of bored sedimen- tary rock recovered during this study were analyzed by thin section to determine their petrographic nature. In addi- tion, CLarK (1978) subjected representative bored rock samples to carbonate analysis using a Leco W 12 Carbon Determinator (Model 761-100). RESULTS The following section will present results of (1) observa- tions and collections made by shallow water diving on the (adjacent column —) Explanation of Table 1 In column Substrate Sample: Concr. = Concretions In columns Relative Abundance, Vacant Bore Holes; and Relative Abundance, Living Borers: A = abundant F = few O = none Page 349 THE VELIGER No. 4 d Vol. 22; SS 3G € Pp G I a AI | 000'T Woy) | Yyorag auo0y [PC JJO “WZ | 6L6I IB T q G 6 9 & 6 V Rt 008°S woyD| yoveg ajuoy [eq JJO ‘We GT| 661 uel ZT G I 9 ie Vv I | 000% BULISLINg | ZNID BURS JULIO JJO “WET | B61 JVW OT & 8 T 14 6 € € Vv I | 0OL auo spn] WUIOd sovVIIBT JO “Ug | Q/6T WA EZ 6 I at 6 L 14 i G A Rt 008 auoyspnyy| sesplig jeanjeN jyo “wi / 861 WA &% 3G 14 A HI | 009 Woy} zn41D eyuKg JUIO jjo “Wt / 8L6I1 49H €Z 3G I 9 A 1 VV | OOr's auo pny | ZN4- BUNS JULIO JJO ‘WY | 8/61 42M OT l 8 T bp 6 69G—COG 6 6 6 6 VV | 006'b eUISIIN jurog Jenbog jjo ‘uy | 8/61 Gea OT O V_ | 006 BULISIAN ejoudey jjo ‘w OT | gZ6T 42a OT & 69 @ Gh 8. 06 @ VV | 0002 PULISIINg ejoudey jo ‘wy, | 8/61 42a OT O oO | 009 euusting| uodueD Aorajuoy ‘Ww Oog | 8Z61T 42d ¢ — pnw weq| uoduey Aasajuoy ‘uw ggt| 8261 Ga T _ pnw yeq| uokueyp Aosajuoy ‘wi ogT | 8261 Gd 1 — pny; eq uodue Asiraquoyy ‘Wi 06] 8/61 928A T 9 Or (nS & 86 G&G Vv V_ =| 000°0S2 WOTJINUOD | YRag a}U0P [eq JJO ‘Ui 0Z| LL6I das gz 9 G I al I LI @l 2 8 if Vv A | 006° yWOUOD ‘yWeYyD| YyoReg aUoW [eq Jo ‘Ww OZ} LL61 des 1z — pny {jaarryg] uoduey Aaraquoy ‘wi Qzt | 2261 82V IT I 6 L I I I VV! 00S*Z ‘WoUOD ‘WaYyD | yoeag auo; [eq JJo ‘wW pz] L/61 8NV g € I G I 3G 8 61 Vv A | 008'FT SuOo!}o19U0TF) moy Arouuey Jjo ‘WwW OP} 1161 8ny 7 }b a V_ | O0F'8 woyD WIO SIOAOT JJO ‘ur Gg} 7/61 uN[ 1Z G L if L G 3G Vv A | OOS‘OT | WUD ‘yeYD | yowag ajuoW Jaq JJo ‘Ww GZ} L161 dv 9 (6 I Ol 8ti O¢ Vv A | OO¢'ST | woUD suoyD |yeeg ajuoy [[9q Jo ‘we GT | 2261 Ady 9% O A | 0OL woyD moy Arouuey jjo ‘Ww og | £261 Je OF G A V | 006'F “IIUOD 194 WULO, O[[lUqeD JJO “W099 | LL6I IA 6 oO V | 00r'9 YOlg Woy WUIOd O[[qey JJO “We PG} LL6T 49 6 Vv V_ | 0066 ‘I9UO0D +WeYD WUIOg O[[1aqeD Jyo “we Zq| LZ6T uel GZ O a 009 suonesu07y | uokuey Aoiojuoyy ‘Wl Opt} ZZ6T uel Z = pny Aery| uokuen Aosraquoy ‘ugg, | zZz6T uel 2 O O | 006 Woy) sould JUlog Jjo “WW 06 | L261 uel Z & if I & p G T 66 Vv ad 00¢°9 oUOD +3494 moy Asouuey jjo ‘wg | 9/61 29d 9 9 & 3G & I G 18 Vv A | 0086 WOUOL) SWaYD | Yoreg aju0W [aq JJO “UW ZT | OL6T AON OT v 3G if G 6 v I I OTs Vv V 00°38 suonjasuoD | Yowog aUOW] JEC{ JJ ‘Wt GT | 9/61 ABW 8T & & & G F v Gp 06 6G T & I V A OOPS WoyD | yorag aUoW [Pq JJO “WITT | 9461 IPI € 8 I T VVC}: OO8'T SyDO]q WoYD | yoRag aUOW [eq JJO “W ZZ | GL6T AON 8T O Vv | 0056 syoo]q Woy, PAO NOY JJO “We OG | GL6T AON 8T G A Vv OOF T “OUOTD + }OYD WIOg O[[liqeyD JjO “Wi ZG | GL6T AON 8T I if ¢ I G V A | 000°9 WoyD | yoeag ajuoW [eq JjO “WET | GL6T PO 06 4 I II if IT 6 Or 6 Vv A 006'T weyD | yoeoeg aju0y [aq Jjo ‘WIT | Gz6t des 6T 61 66 COUL IT 06 i OF th @ if 6 09 = «6G Vv A 008°F9 90][q W9YD | yoRog aUOW] [eq JJO ‘W FZ | GLET IEW 61 v ¢ vI OO I Or @ Vv A | 000°SZI OIG WoYD | yovog aUoW [eC JjO “W ZZ | PLET IEW 1d SS}. tt a BY x > Sy Oye ay eee Oh ee ee a @) 1s > = ams 7 (,Uud) ajdures uoneo0y 2» yydaq aed . of Sf eS |S & 8 8 8 F 8 8 So = Leas a pe - omens o = ES pay Se a nS mectay Sa ES 1S) Spies Salley trade 5 5 3 8 2 = = > > 3 g a > > Q Sic < | °l S Go eS ee ee ee ee are te sy Ses Se EG aay ee eee Bete A i eee a i lame eee Se cl oaltfiee & 2 is) So > s. — 3 iS 2 Re Sf S 3 Ae (= Se ye Og eS we ae eS Se ae. e | RIa & 8 Se Se Ge = 2 S| > g = = Se 2. 5p) 3 5 : B| 8 --— SIO]ISON S010 g peda][OD SispISAON puv sisaio0g puv ‘pasaaoooy safdurvs ‘spneZy Surspoiq T 91981 Page 350 Vol. 22; No. 4 THE VELIGER Monterey shale off Del Monte Beach, (2) dredging opera- tions, (3) intertidal work near Santa Cruz, and (4) car- bonate and petrographic analyses of representative rock samples. 1. DrIvinc OBSERVATIONS ON MONTEREY SHALE oFF Det Monte BEeacH The paper by Haperute et al. (1974) included a review of all diving operations and observations made in the kelp bed off Del Monte Beach up to that time. Since then, additional diving work has been done to sample parts of the exposed shale bottom that had not been examined previously. Methods and techniques employed were the same as in the earlier studies. The following species of bivalve borers have been found while diving on the shale outcrops of the Monterey For- mation off Del Monte Beach: Adula californiensis (Phi- lippi, 1847), A. falcata (Gould, 1851), Lithophaga plumula Hanley, 1843, Barnea subtruncata Sowerby, 1846, Chaceia ovoidea (Gould, 1851), Netastoma rostrata ( Valenciennes, 1846), Parapholas californica (Conrad, 1837), Penitella conradi Valenciennes, 1846, P. gabbii (Tryon, 1863), P. penita (Conrad, 1837) and Zirfaea pilsbryy Lowe, 1931. In addition the following molluscan nestlers have been found in vacant pholad holes: Crepidula perforans (Valenciennes, 1846), Hiatella arctica (Linnaeus, 1767), Irus lamellifer (Conrad, 1837), Kellia laperousi (Deshayes, 1839), and Petricola carditoides (Conrad, 1837). As will be pointed out below in the section on results from dredg- ing, one additional species of pholad, Penitella fitchi Tur- Ner, 1955, was found in shale in slightly deeper water off Del Monte Beach. Some of the borers listed above can readily be identi- fied in situ by a diver, provided the animals are fairly large and the siphons are extended and exposed. These include Barnea subtruncata, Chaceia ovoidea, Parapholas californica, and Zirfaea pilsbryi. The siphons of all others are so small that, although they can be seen clearly pro- jecting from a bored rock sample kept in an aquarium, they are exceedingly difficult to detect in the field under average diving conditions. The siphon tips of members of the genus Penitella can usually be distinguished from all other genera, but species determination in the field usually is impossible. Likewise, although the siphons of the my- tilids Adula californiensis, A. falcata, and Lithophaga plumula are distinguishable from those of pholads, they are not sufficiently distinctive from species to species to allow for identification in the field. Identification and attempts to quantify densities of pop- ulations of borers by observing the borers in place, while diving, is further hampered by shifting sand at the Del Monte Beach diving site. Along this beach there is con- siderable onshore-offshore sand movement with seasons. Following the first storm waves striking the beach in No- vember and December each year much of the sand is combed off the beach and distributed in subtidal waters out to about 10m depth. Sand covers much of the flat shale outcrops, sometimes up to 30cm or more in thick- ness, for several months of the year. Yet, as will be pointed out later, some of the bivalve borers in the shale survive this seasonal burial. Others that cannot tolerate periodic burial are limited in distribution to the projecting ledges and ridges that remain sand-free throughout the year. As part of this over-all study, BoorH (1972) attempted to determine the distribution and density of boring bi- valves that could be identified in situ along two transects running seaward for 500 m off Del Monte Beach. He found a discontinuous distribution of species along each transect, and considerable variation between the transects. Booth noted that Parapholas and Zirfaea were best able to toler- ate periodic sediment cover and that Chacera was most commonly found boring horizontally into shale ledges. Ad- ditional diving operations since 1972 have confirmed these observations. Booth was unable to detect Adula, Barnea, Lithophaga, or Penitella species in the deeper water along the transects he studied and concluded these borers were restricted to shallow water. In other areas off Del Monte Beach, we have not only made observations while diving, but have recovered shale samples by excavating the sub- strate and have found representatives of all these genera except Barnea out to far beyond the ends of Booth’s tran- sects. Booth also concluded that variations in hardness and carbonate content of the exposed Monterey shale were the major factors influencing the inhomogeneous distribution of bivalve borers along his transects. However, recent dredging operations on the Del Monte Beach shale out- crops have allowed us to recover large blocks of Monterey Formation, particularly hard chert, and in this dense homogeneous rock we have found representatives of most of the genera of bivalve borers living side by side. 2. DREDGING OPERATIONS Table 1 summarizes the results of the dredging operations carried out over a 5 year period from March, 1974, to March, 1979. The 40 dredge hauls listed in Table 1 are those where a sample of the bottom was recovered success- fully. Many more hauls, over 60 in fact, were made where the dredge came up empty. The general location of each haul is given in Table 1 and Figure 1. Vol. 22; No. 4 In the majority of the successful dredge hauls, sedimen- tary rock was recovered, but on most of the hauls made on the south wall of the Monterey Submarine Canyon only mud, clay, or gravel came up in the dredge. We made many other attempts to dredge rock from the canyon walls but collected no sample at all. This was a disappointment, for one of our objectives in this study was to sample the rocky wall of the canyon to determine if living stone borers were present and if they played a role in causing erosion and deepening of the canyon. In southern California, WarME, SCANLAND & MarsHALi (1971) found that Parapholas californica, Netastoma rostrata, Adula cali- forniensis, and Lithophaga plumula bored intensely into the rocks of the rim and upper walls of the Scripps Sub- marine Canyon, and that in some areas these organisms were more important as eroders of rock than physical and chemical processes. Even though the walls of the Monterey Submarine Canyon are steep, it is apparent from our re- sults that a sticky layer of clay covers most areas. GREENE _ (1977) succeeded in recovering rock samples from both the north and south walls of the Monterey Canyon. In some cases these were granite samples, in others siltstone or sandstone. Many of the non-granitic rocks Greene re- covered showed bore holes made by pholads and possibly mytilids, but no living borers were found. We must tenta- tively conclude, therefore, that living marine bivalve borers are not causing extensive erosion of the walls of the Monterey Canyon at the present time. In the shallow subtidal water off Santa Cruz relatively few dredge hauls were successful in recovering samples. Off Capitola and Soquel Point a few successful hauls re- covered samples from the Purisima Formation, and off the terraces to the west of Point Santa Cruz some samples of chert and mudstone were recovered. Most of these samples showed evidence of stone borer activity. In the shallow water at the southern end of Monterey Bay, we were much more successful in recovering rock samples and living bivalve borers as is indicated in Table 1. In some cases the chain bag dredge would come up with one large block of chert that had been broken off a ledge, or a large flat concretion broken out of a cherty matrix. In other cases it picked up loose pieces of chert or rounded calcareous concretions that had been lying free on the bottom. Most samples collected off Del Monte Beach con- sisted of a mixture of these two rock types, and most samples showed extensive borer activity leading to con- siderable erosion of the rock. We have collected two species of borer bivalves from subtidal shale off Del Monte Beach while diving, yet have THE VELIGER Page 351 never recovered either of them in dredged samples. Zirfaea pilsbry: is a larger pholad commonly found boring into stiff clay or hard mud at Elkhorn Slough. Off Del Monte Beach, it is relatively abundant, occupying vertical burrows in the softer shale and mudstone. The distinctive siphons made identification easy. This species is often found in densities of 5 animals per m’. Zirfaea excavate burrows up to 60cm deep in the shale that forms the flat bottom between elevated reefs. Dredging is therefore un- likely to recover rock samples containing Zirfaea. Diving observations have indicated that large animals of this species can project their siphons up through as much as 30cm of sand cover. A second species not collected by dredging, Barnea subtruncata, is somewhat smaller than Zirfaea. It too has distinctive siphons and lives in soft flat shale off Del Monte Beach but in numbers far fewer than Zirfaea. Most of the boring bivalves found during this investi- gation had been collected earlier in Monterey Bay, and the subtidal shale off Del Monte Beach is the type locality for several species. One pholad, however, had never been collected before north of southern California (HADERLE, 1979). Penitella fitcht was described from specimens col- lected from intertidal rock at Bahia San Bartolomé, Baja California (TURNER, 1955). Additional specimens have been found at Redondo Beach, La Jolla, and San Diego. Kennedy (1974) reported P. fitchi as a fossil in Pleistocene deposits from southern California and Baja California. During the present study, P. fitchi was found on four occasions. Single living animals were recovered on each of the dredge hauls numbers 20, 22 and 23. The animals were from 5.0 to 6.5 cm in shell length, were in the post- boring stage, and occupied burrows in hard chert. On Dredge Haul No. 25, a single set of valves (4.0cm long) of a dead specimen was found in a burrow in chert. One of the objectives of the dredging part of this study was to determine the bathymetric distribution of boring bivalves in Monterey Bay. As will be pointed out below, some species occur in the intertidal zone at Santa Cruz as high as 2m above MLLW (mean lower low water or zero tide level). Subtidally our dredging operations have shown (Table 1) that living bivalve borers are common down to depths of 50m in the southern part of the Bay. Below a depth of 50m very few living animals have been found in recovered rock samples, although many rocks brought up from these deeper waters were riddled with burrows that were identical to those made by living pholads in shallower water. Many of the bored rock sam- ples from deeper water had been broken off by the dredge, so these samples had not been transported to deeper water. Page 352 Perhaps these bore holes were made by pholads in the past when the level of Monterey Bay was lower than at present, or when temperatures were different. The vacant holes do not appear to be geologically old, however, for they were not filled with compacted sediment and were often not even occupied by nestlers. Experiments now in progress (to be reported on later) have shown that very few boring bivalve larvae settle on or bore into experimen- tal rock panels exposed in water depths exceeding 70m in Monterey Bay. 3. INTERTIDAL REEFS NEAR SANTA CRUZ One of the earliest reports on living bivalve borers in the intertidal zone at Santa Cruz was in the original edi- tion of RicKETTs & CaLvin (1939). Ricketts had observed Platyodon cancellatus (Conrad, 1837) in enormous numbers in banks of stiff blue clay and noted their erosive influence along the shore. He also found siphons of Parapholas californica projecting from rocky reefs in the intertidal zone at Santa Cruz, but the exact location of the reefs was unspecified. In this investigation we have concen- trated on the rocky reefs in the area to the west of Point Santa Cruz and have not observed large numbers of living Platyodon nor Parapholas. The few living Platyodon cancellatus observed appeared to be nestlers in vacated pholad burrows. In the blocks of Purisima Formation in the cliffs some 5m above sea level at Santa Cruz, how- ever, there are many Pliocene fossils of P. cancellatus to be seen as was reported by AppicotT (1966). Most of the results of the investigation being reported on here from the intertidal zone at Santa Cruz were in- cluded in a thesis by CLark (1978). This past year, studies have continued, particularly on the terraces west of Nat- ural Bridges State Park. On one large intertidal reef composed primarily of Monterey shale west of Point Santa Cruz, Clark found the following species of bivalve borers: Adula californiensis, A. falcata, Lithophaga plumula, Netastoma rostrata, Penitella gabbii, P. penita, and Parapholas californica. Of these, Penitella penita was by far the dominant species of borer and was found in rock ranging from the hardest chert to soft mudstone, from 0.7 to 2.0m above MLLW, and in population densities of more than 10 mature indi- viduals per 75cm” surface area. Most individuals were found boring horizontally into ledges on the reef, partic- ularly in pot holes where concretions had been displaced. Approximately half the P. penita were in the boring stage, half were mature with a fully-formed callum. The largest individuals had a shell length (exclusive of siphonoplax) of 5.7m. THE VELIGER Vol. 22; No. 4 The mytilid Adula californiensis was the second most common borer found on the reef at Santa Cruz. The larg- est of these had a shell length of 3 cm. All the other borers were present in much smaller numbers. Nestling bivalves occupying vacant pholad holes included Semele rupicola Dall, 1871, Hiatella arctica, Protothaca staminea Conrad, 1837, Petricola carditoides, Kellia laperousii and Platyo- don cancellatus. C.ark (1978) also studied the borers in a transect across a flat, gently sloping terrace of chert and mudstone lo- cated west of Natural Bridges State Park. He found that here, too, Penitella penita was the dominant borer with Adula californienss, Penitella gabbii, and Netastoma rostrata present in smaller numbers. The highest level where any of these occurred was at 1.0m about MLLW where a few Penitella penita were found. The broad terraces found between Natural Bridges State Park and Terrace Point are broken periodically by wide channels which cut through the terraces all the way up to the base of the sea cliff some 40m shoreward from low water level. These channels have a floor of sand which varies in thickness throughout the year. Where these chan- nels have cut through the terraces they have left vertical walls on each side, some 2m or more high in some places. During this past year these exposed vertical sections of the terraces have been examined for stone borers. Shifting sand along the lower part of these walls erodes the rock very fast and many of the vertical walls are severely undercut. These regions harbor relatively few stone borers, mainly Penitella penita and Netastoma rostrata, and all individuals recovered were small and immature. It is pos- sible that borers cannot survive long enough to reach maturity in this substrate being rapidly abraded by mov- ing sand. In the vertical walls above the area of major sand movement, however, many Penitella borers were found and about half of these were large mature animals. Penitella penita again was the dominant species observed, but P. gabbit was also common. Netastoma rostrata and Adula californiensis were also present, but in small num- bers. Chaceia ovoidea was also found boring horizontally into these intertidal rock walls. Frreu (1953) reported Chaceiaas being commonat Santa Cruz, but Clark (1978) did not observe this species during his work on the inter- tidal reefs and terraces. During this past year many speci- mens of C. ovoidea have been collected from the walls of the surge channels near Terrace Point at levels of 0.5 to 1.0m above MLLW. All of the Chaceta so far observed have been small, up to 1.7cm shell length, and in the immature boring stage, and no large Chaceia burrows have been seen, as are common in subtidal waters off Mon- terey. This would indicate that even the upper walls of Vol. 22; No. 4 the surge channels through these intertidal terraces erode away and expose the Chaceia before these long-lived borers become mature. 4: CARBONATE AND PETROGRAPHIC ANALYSES OF ROCK SAMPLES The literature on rock boring organisms extends back for more than 200 years. Yet, many of the problems consid- ered in these published studies and observations remain unresolved. At times in the past it has been fashionable to divide stone borers into two large categories, those that appear to dissolve the rock by chemical means, and those that abrade the rock mechanically. Among the bivalve borers, the mytilids, such as Lithophaga and Adula, have been considered chemical borers despite the fact there is no direct evidence to support the contention. Pholads as a whole have been considered to be mechanical borers, a conclusion based primarily on the functional morphology of these animals and their shells. The investigation being reported on here from Monterey Bay does not answer any of the lingering questions regarding the specific method or methods used by bivalves in burrowing into solid rock substrates. These studies have shown, however, that meth- ods used for boring may be far more complex than we have suspected, and any one borer may be able to use a variety of methods for excavating burrows into various rocks having different physical and chemical properties. The fact that Lithophaga plumula (usually considered to be a chemical borer living primarily in calcium carbonate substrates) and various species of the genus Penitella (usually considered to be mechanical borers living pri- marily in soft rock) can live side by side and reach max- imum size while boring into exceedingly hard, dense, siliceous chert in the shallow subtidal and intertidal zone of Monterey Bay indicates that we have much to learn about the fundamental mechanisms of rock boring in marine animals. The dominant rock types where living bivalve borers have been found in the present investigation in Monterey Bay fall into three main categories: (1) silty biogenic cherts and siliceous shales of the Monterey Formation, (2) calcareous concretions of various shapes and sizes associated with the chert beds or derived from them and lying free on the bottom, and (3) Purisima Formation dredged from shallow water at the north end of the Bay. We have attempted to learn something about the physical and chemical nature of the first two types of these rocks into which many species of bivalves so regularly erode sizable burrows. THE VELIGER Page 353 The amount of CaCOs in any of the samples from the Monterey Formation (except for concretions), be they relatively soft mudstone or exceedingly hard chert, was extremely low, varying from 0.03 percent in some samples to a maximum of 0.74 percent in others. Analysis of thin sections of chert from the Santa Cruz reef and from sub- tidal outcrops off Del Monte Beach showed it to be pri- marily (90-95%) a ground mass of siliceous biogenic hash (radiolaria, sponge spicules, diatoms, etc.) with 5-8 percent clasts of silt-sized quartz, feldspar, biotite, mag- netite, hematite, and microcrystalline chert. No cementa- tion was present. Chemical analysis of the concretions, both from the subtidal waters in Monterey Bay and from the reefs in the intertidal zone at Santa Cruz, gave a CaCOs content of 80-85 percent. Thin sections demonstrated that the concretions were recrystallized fossiliferous limestone with a ground mass of muddy carbonate now recrystallized to sparry calcite. The fossils were recrystallized foraminifera and siliceous tests replaced by calcite. SUMMARY 1. This paper presents the results of a 10 year study on the distribution of bivalve mollusks that bore into rocky substrates in Monterey Bay. 2. Living borers were found in all types of sedimentary rocks from 2m above MLLW to depths of over 50m. 3. The Monterey shale exposed under the kelp beds in southern Monterey Bay has been examined by divers and 11 species of bivalve borers have been identified including the mytilids Adula falcata, A. californi- ensis, Lithophaga plumula, and the pholads Barnea subtruncata, Chaceia ovoidea, Netastoma rostrata, Parapholas californica, Penitella conradi, P. gabbit, P. penita, and Zirfaea pilsbryt. 4. Over 100 dredge hauls were made at various places in Monterey Bay from the shallow water off Santa Cruz to deep water in the Monterey Canyon. Forty of these hauls were successful in recovering a bottom sample. In water down to 50m deep in the southern part of the Bay, rock samples recovered carried the same borers as were found in the diving operations, with the exceptions of Adula californiensis, Barnea subtruncata, and Zirfaea pilsbry. In addition, the dredging recovered a species not previously reported from Monterey Bay, Penitella fitchi. Dredging in shal- low water off Santa Cruz recovered samples of Puris- ima Formation with a variety of borers. Page 354 5. In deeper water of Monterey Bay, recovered rock samples showed evidence of bivalve borer activity, but no living borers were found. 6. Few samples of sedimentary rock were recovered from the walls of the Monterey Canyon and none carried living borers. 7. The intertidal reefs at Santa Cruz were populated, from low tide level to 2m above MLLW, with Adula californiensis, A. falcata, Lithophaga plumula, Cha- ceia ovoidea, Netastoma rostrata, Parapholas califor- nica, Penitella gabbii and P. penita. Of these, Peni- tella penita was the dominant species. 8. Chemical and petrographic analyses of various rock samples indicated that most of the bivalve borers in Monterey Bay bore into both siliceous rocks of various hardness and into calcareous rocks. Literature Cited Appicotr, WARREN OLIVER 1966. Late Pleistocene marine paleoecology and zoogeography in cen- tral California. U.S. Geol. Surv. Prof. Paper 523C: 1-21; 4 plts. 6 text figs. BooTH, GREGORY SEELEY 1972. The ecology and distribution of rock-boring pelecypods off Del Monte Beach, Monterey, California. Unpubl. Master’s Thesis, Naval Postgrad. School, Monterey, Calif. (June 1972) Burnett, Nancy ANN 1972. The ecology of the benthic community of bivalve mollusks in the shale at the Monterey sewer outfall. Unpubl. Master’s Thesis, San Francisco State Univ., San Francisco, Calif. CriarK, GERALD WAYNE 1978. Rock boring bivalves and associated fauna and flora of the inter- tidal terrace at Santa Cruz, California. Unpubl. Master’s Thesis, Naval Postgrad. School, Monterey, Calif. (September 1978) Donat, WinFiztp III. 1975. Subtidal concrete piling fauna in Monterey Harbor, California. Unpubl. Masters Thesis, Naval Postgrad. School, Monterey, Calif. (September 1975) THE VELIGER Vol. 22; No. 4 Fitca, Joun E. 1953. | Common marine bivalves of California. Fish Bull. go: 1 - 102; 63 text figs. GauiHeR, E. W. 1932. Sediments of Monterey Bay, California. Minerol. 28: 42 - 71 GreENE, H. Gary 1977. Geology of the Monterey Bay region. Geol. Surv. Open-file Reprt. 77-718: 1-347 Haperug, Euvcenge CLinTon Calif, Fish & Game, Reprt. Calif. State U. S. Dept. Interior, 1976. Destructive marine wood and stone borers in Monterey Bay. Proc. 34 Intern. Biodegrad. Symp., J. M. Sharpley & A. M. Kaplan, eds. Applied Sci. Publishers, London, pp. 947 - 953 1977. Fouling communities in the intertidal zone of wooden and con- crete pilings at Monterey, California. pp. 229-239 in: V. Romanov- sky (ed.), Proc. 4th Internat. Congr. Mar. Corrosion and Fouling 1979. Range extension for Penitella fitchi Turner, 1955 (Bivalvia: Pholadidae). The Veliger 22 (1): 85 (1 July 1979) Haveruiz, EuGeNe CLinton & WINFIELD Donat III 1978. Wharf piling fauna and flora in Monterey Harbor, California. The Veliger 21 (1): 45-69; 1 plt.; 10 text figs. (1 July 1978) Haperuiz, Evcene Cuinton, J.C. Merior, C.S. Minter III « G.C.Bootu 1977- Fouling communities in the intertidal zone on wooden and con- crete pilings at Monterey, California. pp. 241-251 in: V. Romanov- text figs. (1 October 1975) Kennepy, Georoe L. 1974. West American Cenozoic Pholadidae (Mollusca: Bivalvia) San Diego Soc. Nat. Hist. Mem. 8: 1 - 128 Mart1n, B. D. 1964. Monterey submarine canyon, California: Genesis and relatton- ship to continental geology. Ph. D. Thesis, Univ. So. Calif, Los Angeles, Calif. 249 pp. MINTER, CHARLES STAMPS, IIT 1971. Sublittoral ecology of the kelp beds off Del Monte Beach, Mon- terey, California. Unpubl. Master’s Thesis, Naval Postgrad. School, Monterey, Calif. (September 1977) Ricketts, Epwarp F. e Jack Carvin 1939. Between pacific tides. 320 pp.; illust. Sueparp, Francis ParRKER & K. O. EMERY 1941. Submarine topography off the California coast. Amer. Spec. Paper No. 31: 103 - 112 Surra, ALLYN Goopwin « Mackenziz Gorpon, Jr. 1948. | The marine mollusks and brachiopods of Monterey Bay, Cali- fornia, and vicinity. Proc. Calif: Acad. Sci. ( 4) 26 (8): 147-245; plts. 3, 4; 4 text figs. (15 December 1948) Turner, RutH Drxon 1955. The family Pholadidae in the western Atlantic and eastern Pacif- ic. Part II — Martesiinae, Jouannetiinae and Xylophaginae. Johnsonia 3 (34): 65-160; plts. 35-93 Warme, Joun E., THomas B. ScaNLanp & Net KF MarsHati 1971. | Submarine canyon erosion: Contribution of marine rock bur- rowers. Science 173: 1127-1129 Stanford Univ. Press, Stanford, Calif Geol. Soc. Vol. 22; No. 4 THE VELIGER Page 355 Generic Variation in Sympatric Sibling Species of Littorina BY N. PR. WILKINS ann D. O’REGAN National University of Ireland, Department of Zoology, University College, Galway, Ireland INTRODUCTION A NUMBER OF STUDIES on marine invertebrates have cor- related spatial heterogeneity in allele frequencies at cer- tain enzyme gene loci with the dispersal ability of organ- isms (W1UM-ANDERSEN, 1970; ScHOPF & GoocH, 1971; Goocu et al., 1972; SNYDER & GoocH, 1973; BERGER, 1973, 1977; GAINES et al., 1974; WILKINS et al., 1978). In general, these studies have indicated that the differ- ence in allele frequencies between geographically sepa- rated populations is greater in species which lack a plank- tonic stage in the life cycle than in those with planktonic eggs or larvae, or both. Site-specific, or region-specific, alleles are also more common in the former than in the latter. These conclusions have, to a large extent, been drawn from studies on littorinids or from comparisons of other taxonomically unrelated species with littorinids. At the loci encoding esterases (BERGER, 1973) phosphoglucose isomerase and phosphoglucomutase (WILKINS et al., 1978) intersite allele frequency variations are greater and site specific alleles more common in Littorina saxatilis (Olivi, 1792) which is ovoviviparous than in L. littorea (Linnaeus, 1758) which has planktonic eggs and larvae. Littorina obtusata (= L. littoralis) which has benthic eggs is inter- mediate between these two. Results such as these on littorinids may be influenced by the taxonomic status of the species investigated. If, for ex- ample, Littorina saxatilis were composed of two or more distinct species with characteristic allele frequencies, the observed geographic variation might reflect differing dis- tributions or proportions of these species at the various sites. The most recent evidence (HELLER, 1975) indicates that in the European area at least, “L. saxatilis’ may comprise four separate, non-interbreeding species viz. L. rudts (Maton, 1797), L. nigrolineata Gray, 1839, L. neglecta Bean and L. patula Jeffreys. Likewise, two distinct species comprise the “L. obtusata (= littoralis)” complex viz. L. obtusata (Linnaeus) and L. mariae Sacchi « Rast. (Saccut & RASTELLI, 1967). We have examined over 3.000 winkles, separated mor- phologically into these species, from g sites in Ireland and one site in France. We present here the results of the anal- ysis of genic variability at the phosphoglucose isomerase and phosphoglucomutase loci in these samples. The results indicate that genic variability does correlate with dispersal capability for the species Littorina rudis, L. obtusata and L. littorea, but extrapolation from these to other species is not warranted. The significantly greater degree of genic variability in L. rudis is unexpected when the potential for inbreeding in this ovoviviparous species is considered. MATERIALS anp METHODS Samples were collected from rocky shores at 6 sites (Silver- strand, Barna, Spiddal, Carna, Doolin, Aranmore island) in or near Galway Bay on the west coast of Ireland, and from one site near each of the localities Cork (south coast), Carnsore (southeast coast), Dublin (east coast) and Brest (west coast of Brittany, France), as indicated in Table 1. Table 1 Numbers of individuals of the six species of winkles analyzed from the various sites. SIL., Silverstrand; BAR, Barna; SPI, Spideal; CAR, Carna; ARA, Aran Island; DOO, Doolin; COR, Cork; CAN, Carnsore; DUB, Dublin; BRE, Brest. Littorina Littorina Littorina Littorina Littorina Littorina Site littorea nigro. neglecta rudis obtusata martae SIL. 56 77 90 60 72 101 BAR. 90 118 66 127 157 79 SPI. 146 — = — 171 — CAR. 60 = 54 = 77 = ARA. 45 = = — 86 — DOO. 55 52 = 75 — — COR. 58 61 — 25 96 —_ CAN. 73 = 71 79 83 = DUB. 90 = =— 60 160 - BRE. 75 — = 74 78 = TOTALS 748 308 281 500 980. 180 Page 356 THE VELIGER Vol. 22; No. 4 Not all individuals from each site were analyzed for both enzymes; the figures in Table 1 are the maximum number analyzed at one or other gene locus from each site, Species were identified and separated using the criteria described by HELLER (1975) for the Littorina saxatilis complex and those of SaccH1 « RasTELLI (1967) for the L. obtusata complex. We were unable to identify winkles of the L. patula type in any of the samples. Individuals were main- tained alive in sea-water aquaria until analyzed by the usual electrophoretic technique (WILKINS, 1977). Allele frequencies in the different populations or species, or both, were compared by X? analysis of 2xj tables of observed allele numbers, with one or two degrees of free- dom as appropriate. RESULTS The electrophoretic patterns observed were consistent with those previously reported (WiLkins et al., 1978). Phospho- glucose isomerase (Pgi) patterns indicated a dimeric en- zyme encoded at a single locus in all the species, with 2, 3 or 4 alleles expressed in each species. Mobilities of the F, F', S, and S' isozymes were similar in all species except for Littorina littorea in which all Pgi isozymes migrated more anodally than in the other species. Phosphoglucomu- tase patterns indicated that this enzyme is monomeric, en- coded at a single locus with 1, 2 or 3 alleles in each species. Isozyme mobilities were similar in all species except L. littorea in which all isozymes migrated more slowly than in the other species. Table 2 summarizes the results obtained for all 6 species at both loci. The 3 species of the saxatilts complex did, in- deed, have characteristic allele frequency distributions: at both loci all 3 species differed significantly (P 0.05), it is interesting that these brackish water bivalve larvae were capable of growth to metamorphosis at near-oceanic salinities (32%). Characteristics of the hinge line are used to differen- tiate bivalve species both as larvae and adults (REES, 1950; Lutz & JABLONSKI, 1978; SDALL, 1978). Serial develop- ment of the hinge line in Mytilopsis leucophaeata is shown in Figure 7. Using the terminology of CHANLEY & ANnpREwS (1971), M. leucophaeata develops “knobby” Explanation of Figures 1 and 2 Figure 1: SEM photomicrographs of hinge lines of Mytilopsts leu- cophaeata at two times prior to metamorphosis: a) 2 days after fertilization (DAF); shell length = 74 nm; b) 6 DAF; shell length = 180m; and at three times subsequent to metamorphosis: c) 12 DAF: maximum shell dimension = 270ym; d) 18 DAF: maximum shell dimension = 375m; e) 28 DAF: maximum shell dimension = 500 pm; showing shelf under the umbo Figure 2: SEM microphotograph of juvenile Mytilopsis leucophae- ata (0.5cm maximum dimension) (a) with pits randomly dis- persed anterior to pallial line; and (b) a magnified view of pit (3 um in diameter) THE VELIGER, Vol. 22, No. 4 [StpDALL] Figures 1 and 2 4 Figure Ic Figure rd Figure re Figure 2b Vol. 22; No. 4 umbones which became “skewed” anteriorly following metamorphosis. At 26° C, larvae fed a mixed phytoplank- ton diet (Isochrysis galbana, Monochrysis lutheri and Tetraselmis suecica) metamorphosed 6 to 8 days after fer- tilization at a mean shell length (parallel to the hinge line) of 210um (n=25). Neither larvae nor juveniles of M. leucophaeata possess hinge teeth at any stage. The “tri- angular tooth” referred to in ABBott’s (1974) description of M. leucophaeata is a projection of the shelf near the beak, not to be confused with hinge teeth. Because the adult ligament is broad, the shell presumably does not require teeth for stabilization of the opening and closing motion of the valves. For the adult population sampled, mean maximum shell dimension was 2.2 cm (n = 125). Figure 2 shows a typical juvenile (0.5 cm maximum shell dimension). Evenly dispersed and uniformly shaped tu- bules penetrate the inner calcareous crossed-lamellar layer typical of the Dreissenacea (KENNEDY, TayLor & Hatt, 1969a). These tubules range in diameter from 3 to 5 um and should not be confused with the very small tubule- like structures occurring in mytilids (an order of magni- tude smaller). The tubules of Mytilopsis leucophaeata are uniformly distributed in, and restricted to, the older regions of the shell bounded by the pallial line. This dis- tribution suggests a secondary origin for the tubules (KENNEDY, TAYLOR & HALL, 1969b). In their brief review of tubulate bivalve shells, Kennepy e¢ al. (1969b) state that these tubules penetrate the periostracum. Such was not the case in the material examined in the present study. The tubules of M. leucophaeata first appear in metamor- phosed juveniles (0.5mm maximum shell dimension) and are seen in individuals of all larger sizes. To summarize: (1) in the population of Mytilopsts leucophaeata studied, gamete release consistently occurred as salinities de- creased during the annual rainy season; THE VELIGER Page 379 (2) settlement of spat peaked 2 weeks after gamete re- lease in natural populations; in the laboratory at 26° C, well fed larvae metamorphosed in 6 to 8 days; (3) Mytilopsis leucophaeata larvae and post-larvae are euryhaline, capable of development to metamorphosis at 10 to 32%; (4) neither larvae nor older stages of Mytilopsis leuco- phaeata possess hinge teeth; (5) shells of juvenile and older Mytilopsis leucophaeata are tubulate; tubules are restricted to the older shell regions bounded by the pallial line and do not pene- trate the periostracum. Literature Cited Aszott, Ropgrr Tuoxgr 1974. American seashells. a=d ed.; 663 pp; 4060+ figs.; pli. 1 - 24 (in color). Van Nostrand Reinhold Co., New York Carmiczer, MeLBouRNE ROMAINE 1979. Ultrastructural effect of cleaning molluscan shell with sodium hypochiorite (Clorox). The Nautilus 93 (2-3): 47-49 Cuan.ey, Paur &# Jay D. ANDREWS 1971. Aids for identification of bivalve larvae of Virginia. Malaco- logia 11 (1): 45-119; 51 text figs. (8 October 1971) Emerson, Wituiam Kerrn & Morris KaRLMANN JACOBSON 1976. Guide to shells. Alfred A. Knopf, Inc., New York, xviii+ 482 pp.; 97 pits. [of which 16 colored]; 6 text figs. Kennepy, Wittiam James, JouN Davm Tayitor & ANTHONY Hai 1969a. Environmental and biological controls on bivalve shell mineral- ogy. Biol. Rev. 44: 499 - 530 1969b. The shell structure and mineralogy of the Bivalvia. Introduction — Nuculacea — Trigonacea. Bull. Brit. Mus. (N. H.) Zool. Suppl. 3: 1-125; 29 plt. LoosanoFF, Victor Lyon « Harry Cart Davis 1963. Rearing of bivalve mollusks. | In: Advances in marine biology. I: 1-136; 43 text figs. EF S. Russell (ed.). Academic Press, London Lutz, R. A. & D. JaBLonsx1 1978. Cretaceous bivalve larvae. Rzzs, C. B. 1950. The identification and classification of Jamellibranch larvae. Hull Bull. Mar. Ecology 3: 73 - 104 Swoatt, S. E. 1978. The development of the hinge line in tropical mussel larvae of the genus Perna. Proc. National Shellfish. Assoc. 68: 86 Science 199: 439 - 440 Page 380 THE VELIGER Vol. 22; No. 4 Fish Predation on Pulmonate Limpets SUSAN BLACKFORD COOK Department of Zoology, The Ohio State University, Columbus, Ohio 43210 (2 Text figures) FISHES MAY BE IMPORTANT PREDATORS on invertebrates in tropical marine systems (Bakus, 1969) either because they inadvertently remove invertebrates while browsing on algae (RANDALL, 1974; VINE, 1974; Day, 1977) or because they focus specifically on invertebrate prey (HIATT & StraspurG, 1960). In this note I present indirect evi- dence that loss of 21% of a population of the intertidal pulmonate limpet Siphonaria normalis at Enewetak Is- land, Enewetak Atoll, Marshall Islands resulted from se- lective removal by fishes during a spring high tide. Mor- tality was size-specific and altered the size structure of the population. Siphonariid limpets occupy specific home sites on rocks (Cook, 1969) when they are not grazing on microscopic algae (Voss, 1959). Home sites of individuals on soft rocks are marked by “scars,” depressions whose dimen- sions are the same as limpet basal dimensions. Because Stphonaria normalis are always “at home” at dead low tide (1 homing failure out of 133 grazing excursions) and limpets which do wander stay close to scars and can be readily recognized by visible gaps between shell margins and the rock, the presence of empty scars at low tide coupled with a lack of “extra” limpets on the rock is evi- dence of mortality. Such a combination of events occurred in a Siphonaria normalis population located on beachrock on the lagoon side of Enewetak Island. During one late afternoon spring tide (1.5m above o datum, October 6, 1975), four of the 19 original residents of a 25 x 25 cm quadrat disappeared. On the next daytime low tide, I found no “extra” limpets and discovered that empty scars were distinctively marked with grooves extending from the adjacent rock down into scars (Figure 1). Within a nearby 50 x 50cm area, similar marks were associated with the empty scars of 10 of 49 Figure 1 Two adjacent empty limpet scars marked by prominent grooves; each scar is about 8mm in length residents, while all of another 50 empty scars on the rock outside of these two areas were marked. Marks were rare on the rock between scars (<10% of the total number found). Bits of limpet tissue present in two of the marked scars confirmed that limpet mortality had indeed occurred. Al- though the rock in the vicinity of two additional animals was also marked, these animals escaped removal. All empty scars and marks were obliterated by algal growth within 2-4 days. Marks were of an appropriate size and spacing to have been made by the canine teeth of a large wrasse such as Coris aygula (J. Randall, personal communication). This species is found at Enewetak in shoal areas, as well as deeper water, and often eats mollusks (J. Randall, pers. comm.). Two other possible predators, the golden plover Pluvialis dominica (A. Kohn, pers. comm.) and the muri- cid gastropod Thais armigera (Menge, 1973), can be ex- cluded. Plovers are not likely to dive to reach limpets sub- merged during spring high tides. Predatory snails were not Vol. 22; No. 4 seen in the area either before or after limpet loss and in any case would not have left grooves on the rock. Large limpets were clearly more affected than were smaller individuals. The mean length of empty scars (and therefore of former residents) was significantly greater than the mean length of survivors (Figure 2, t-test, P< 0.001). Proportions of scars in the size categories 4-7 mm and 8-12mm were significantly different from proportions expected if limpets had been removed according to initial abundance (chi-square goodness of fit, P P>o.02). 20 C1 Intact n=50 @ Missing n=14 Number of Limpets g O Per Shell Length in mm g 10 11 12 Figure 2 Size comparisons of intact limpets on scars and unoccupied grooved limpet scars following fish predation Information on how often fish predation affects tropical limpet populations over the long term is lacking. Although such predation is one explanation for the scarcity of large THE VELIGER Page 381 Siphonaria normalis at Enewetak (less than 1% larger than 8mm, MENGE, 1973; less than 1% larger than 8mm at another study site, Cook, unpublished), alternative ex- planations are possible. These include size-specific preda- tion by Thats (MENGE, 1973) or shorebirds, or both, as well as periodic catastrophic mortality of all limpets fol- lowed by recruitment and growth of recruits into small to medium sized limpets. ACKNOWLEDGMENTS I thank Drs. C. B. Cook and R. A. Stein for editorial com- ment and Dr. J. Randall for valuable advice on fishes. This work was made possible by a grant from the Energy Research and Development Administration to the Univer- sity of Hawaii and the Mid-Pacific Marine Laboratory and the Irma E. Voigt Fellowship from the American Association of University Women. This note is a Contri- bution from the Mid-Pacific Marine Laboratory. Literature Cited Baxus, Geratp Joszeru 1969. Energetics and feeding in shallow marine waters. Rev. Gen. & Experim. Zool. 4: 275 - 369 Coox, Susan BiackForp 1969. Experiments on homing in the limpet Siphonaria normelis. Anim. Behav. 17: 679 - 682 Day, R. W. 1977- Two contrasting effects of predation on species richness in coral reef habitats. Mar. Biol. 44: 1-5 Hiatt, R. W. & D. W. Strassurc 1960. Ecological relationships of the fish fauna on coral reefs of the Marshall Islands. Ecol. Monogr. 30: 65 - 127 Mencz, Bruce A. ; 1973. Effect of predation and environmental patchinesson the body size of a tropical pulmonate limpet. The Veliger 16 (1): 87-92; 8 pit.; 3 text figs. (1 July 1973) RANDALL, JoHN E. 1974. The effects of fishes on coral reefs. Coral Reef Sympos. 1: 159 - 166 In: Yntern. In: Proc. Second Intern. Ving, P J. 1974. Effects of algal grazing and aggressive behavior of fishes Poma- centrus lividus and Acanthurus sohal on coral reef ecology. Mar. Biol. 24: 131 - 136 Voss, Nancy A. 1959. Studies on the pulmonate gastropod Siphonaria oe (Lin- naeus) from the southeast coast of Florida. Bull. Mar. Sci. Gulf Caribb. 9: 84-99 Page 382 THE VELIGER Vol. 22; No. 4 Intermediate Forms and Range Extension of Pedicularia californica and Pedicularia ovuliformis ROBERT W. SCHMIEDER Sandia Laboratories, Livermore, California 94550 (3 Text figures) Two ForMS OF Pedicularia are known from the California coast: P. californica (Newcomb, 1864) and P ovuliformis (Berry, 1946). In comparison, the former is slightly lar- ger, irregular, with simple outer lip, and lighter in color. The latter is uniform in size and shape, with a heavy dentate outer lip, crenulate outer margin, and a rather intense rose-pink color. Heretofore, no specimens inter- mediate between these two forms or their existence north of the Farallon Islands have been known. The October 1978 scuba expedition to Cordell Bank, Figure 1 Outlines of the 1978 Cordell Bank Pediculeria, viewed dorsally. The grid spacing is 1 mm, and the numbering corresponds to the entries in Table 1 Vol. 22; No. 4 THE VELIGER Page 383 30km W of Point Reyes, carried out by the author and colleagues (SCHMIEDER, 1979) yielded 13 specimens of Pedicularia, most collected live among abundant hydro- coral (Stylantheca porphyra), thus extending the range to Latitude 38° N. The specimens were collected in clear water (visibility ~18m, temperature ~14° C) at depths of 39.6 to 46.8m, from a tiny pinnacle at Latitude 37°- 5930”, Longitude 123°24’30”, now known as Craines Point. Figure 1 shows outlines of the 13 specimens. Among the specimens are several of the 2 well-known forms, plus several of intermediate character. One in particular exhibits both the classic simple lip of Pedicula- Figure 2 Drawings of two specimens, Pedicularia ovuliformis (No. 5), and the intermediate form (No. 6). Drawn from photographs ria californica, and, within the aperture, the classic den- tated lip of P ovuliformis. Remnants of the crenulate outer margin can be seen on the exterior, well below the aperture. The exterior color is very similar to that of P ovuliformis, except that the shell added beyond the crenu- lations is lighter, close to that of P californica. Figure 2 shows 2 of the specimens, one clearly P ovuliformis and the other the intermediate form. Other specimens in the collection show similar but less complete suggestions of both forms. Several have an easily identified smaller, darker “core,” of the size and color of Pedicularia ovuliformis, surmounted by a lighter, more open, simple labrum, giving them the overall ap- proximate appearance of P californica. Microscopic ex- amination shows evidence of overgrowth of a flat, heavy labrum, probably dentate, and possible remnant of inte- rior overhang of the flat lip of P ovuliformis. Table 1 gives the significant characters of these specimens and in Figure 3 is plotted the width vs length, which is seen to be approximately linear. It is therefore not unreasonable to conclude that there is probably only one species of Pedicularia in Northern California, that P ovuliformis is a phase leading to P californica by partial resorption of the flat dentate lab- rum and simultaneous extension of a simple labrum, with size increase and reduced pigmentation. This possibility was also considered by Berry (1946), who rejected it in favor of defining P ovuliformis as a new species. A possible explanation for these facts is the following: In the colder water further north, the growth rate is probably slower than in more southerly areas where Pedi- cularia is more commonly found. Thus, more time would Table 1 Pedicularia from Cordell Bank 1978 No Length (mm) Width (mm) State 1 5.2 3.4 live 2 5.8 3.7 dead 3) 6.0 3.6 4 6.1 4.0 5 6.2 3.5 6 6.9 4.3 live 7 6.9 4.9 dead 8 7.9 5.0 live 9 8.5 5.6 live 10 9.1 5.9 dead 11 10.0 6.3 dead 12 10.4 6.0 live 13 12.4 8.6 live Dentation Crenulation Remarks rudimentary none partial none partial rudimentary P. ovuliformis complete partial P. ovuliformis complete complete P. ovuliformis complete residual Intermediate form ambiguous ambiguous White; aperture unusual none none P. californica ; none none P. ovul. core, w/chg. of growth at junction rudimentary none ambiguous partial Lips somewhat thickened none none Lips slightly thickened; has none none P. ovul. core Lips extremely thickened; very heavy, several phases. Page 384 Width (mm) w Length (mm) Figure 3 Width vs. length of the 13 specimens be spent in making the transition from P ovuliformis to P californica, and the probability of finding the interme- diate form would be correspondingly higher, accounting for its discovery at Cordell Bank. The individual reaches maturity as P ovuliformis, spending some time on its color- encrypting substrate Stylantheca. Then, in response to an environmental change (which may actually be a change THE VELIGER 15 Vol. 22; No. 4 in the substrate occasioned by a more primitive cause such as temperature), it suddenly puts on a growth spurt. In order to contain the animal, the outer lip is rapidly extended, leaving the flat, dentate margin as an interior shelf, which is then resorbed to clear the passage (resorp- tion is known in other species of Pedicularia) . Irregularity in form is due to the rapidity of growth — if conditions returned to the earlier state, it might well construct an- other flat, heavy, dentated outer lip. There is some evi- dence of this last, since the largest (and presumably oldest) specimens in the Cordell Bank collection have greatly thickened lips, and retain irregular forms. These specimens have been deposited in the collection at the California Academy of Sciences, San Francisco, California, accession number 60363. ACKNOWLEDGMENT It is a pleasure to acknowledge the considerable help in this study by Barry Roth of the California Academy of Sciences. Literature Cited Berry, Samug, STmLMAN 1946. Californian forms of Pedicularia. Leafl. Malacol. 1 (1): 1-2 Newcoms, W. 1864. Description of a new species of Pedicularta. Proc. Calif Acad. Nat. Sci. 3 (2): 121-122 Scumizper, Ropert W. 1979. Cordell Bank Expedition 1978. Unpubl. Reprt. available from the author and at Calif; Acad. Sciences, San Francisco. Vol. 22; No. 4 THE VELIGER Page 385 Helix aperta Introduced in Richmond, California (Mollusca : Pulmonata) BARRY ROTH Department of Geology, California Academy of Sciences, San Francisco, California 94118 DUSTIN D. CHIVERS Department of Invertebrate Zoology, California Academy of Sciences, San Francisco, California 94118 (1 Text figure) In Marcu 1979, Mr. Gordon E. Bergman and his daugh- ter Katherine discovered an infestation of the European and North African snail, Helix aperta Born, 1778, in an industrial district of Richmond, Contra Costa County, California. During a period of intermittent rains, the snails were observed to be numerous, crawling actively during the day. Also in March, Mr. Sterling Bunnell brought specimens collected by Mr. and Miss Bergman to the California Academy of Sciences for identification. From early April to May 1979 the authors investigated the site of infestation - a region of artificial fill along the south side of Point Isabel, on the east shore of San Fran- cisco Bay in the southern part of Richmond. We found Helix aperta to be common south of Central Avenue (Figure 1), from near the base of the Central Avenue overpass over Hoffman Boulevard to the vicinity of a sewage pumping station approximately half the dis- tance along the south shore of Point Isabel, and from the south edge of the paved roadway to the margin of the filled land. The infested area is entirely on artificial fill emplaced between 1949 and 1968, according to editions of the U. S. Geological Survey Richmond Quadrangle (7.5 Minute Series, Topographic). The ground is flat and weedy with large clumps of the shrub, Baccharis pilularis var. consanguinea (DC.) C. B. Wolf. The soil is clayey and gravelly, rather extensively tunnelled by ro- dent burrows. Helix aperta shells were present among weeds and in the litter under Baccharts clumps. Living individuals were most common on and under rubbish on the ground. The greatest concentration was in the area south of the foot Point Isabel, Richmond, California. Crosshatching indicates presence of Helix aperta of the Central Avenue overpass, which is a site of casual dumping of garden clippings and household refuse. A few living specimens were observed in rodent burrows. During a visit May 4, living juveniles (to about 1 cm diameter) were found on the leaves of thistle (Cirsium) and sow- thistle (Sonchus) as much as 30cm above the ground. Dead snails in various stages of decomposition indicate that the species may have been established for more than one year. Associated mollusks included Helix aspersa Miil- ler, 1774, Milax gagates (Draparnaud, 1801), Deroceras Page 386 caruanae (Pollonera, 1891), and D. reticulatum (Miiller, 1774), all introduced; and Punctum conspectum (Bland, 1865), native to California but probably introduced to this site. No Helix aperta were observed north of Central Ave- nue in a weedy field with fennel plants (Foentculum vul- gare Mill.), in a slough with pickleweed (Salicornia) north of the overpass, nor along the Bay shore east of the Point Isabel peninsula. A region of Baccharis bushes near the northwest end of Point Isabel yielded only Helix as- persa. The largest specimen collected, an empty shell, meas- ures 26.4 mm in height and 30.4 mm in maximum diame- ter, equal to the size attained in Europe; it has 3.25 whorls. Shells of most living specimens collected are about 20mm in height and 22mm in maximum diameter, with about 3.25 whorls, and probably represent a single age class. Helix aperta was first reported in California by Gam- MON (1943); he described an infestation discovered in San Ysidro, San Diego County, in April 1941, where the snails were abundant on streets after showers. Preliminary sur- veys in 1941 showed the species to be present over at least 10 blocks and many acres of pasture and river bottom land in San Ysidro, Further survey during the following winter and spring revealed it in several additional blocks in San Ysidro and over many acres around dumps in the vicinity. Gammon estimated that approximately 500 acres were then infested. Some damage to home gardens was re- ported. The species was still present and was causing agri- cultural damage in 1951 (CAMPBELL, 1953). The San Ysidro infestation was the only occurrence reported by Hanna (1966) in his survey of introduced mollusks of western North America. We were advised by Dr. Tokuwo Kono (in litt. 26 April 1979) of the California Depart- ment of Food and Agriculture that that agency has no other records of the species’ establishment in California. DuNDEE (1974) noted museum lots of Helix aperta from New Orleans, Louisiana, and Nahant, Massachu- setts. She stated that, as of the date of her writing, the species was no longer present in New Orleans. Harry (1948) reported that living, imported H. aperta were sold in open baskets in Louisiana but that the species was not known to have established itself. Helix aperta is native to the Mediterranean region. It is said to be chiefly a ground snail, rarely climbing to any height on plants (FROMMING, 1954). In its countries of origin it is found principally under foliage and stones, often on cultivated land. Gammon (1943) stated that it is very active during the rainy season but soon burrows in soil to depths as great as 6 inches (15cm). Estivation THE VELIGER Vol. 22; No. 4 takes place in an earthen cell, behind a thick, convex epi- phragm, with re-emergence following autumn rains. The burrowing habit makes it difficult to determine the extent of an infestation and hampers eradication. ‘Two modes of introduction to the Richmond site seem possible: (1) the species was brought in with soil used for artificial fill, or (2) it was present on garden debris dumped in the area. The burrowing habit of the snail makes transport in soil a possibility. On the other hand, the observations that, as far as could be determined, dis- tribution is limited by a paved road and density is greatest near a dumping site suggest a more recent introduction, probably with horticultural rubbish. This implies that H. aperta may be established in gardens in one or more of the neighboring communities. There is a continuing potential for introduction. Be- tween July 1, 1973, and September 30, 1977, Helix aperta was intercepted 17 times by the U. S. Department of Agri- culture Animal and Plant Health Inspection Service (U. S. Department of Agriculture, 1979). The species was intercepted at ports in Louisiana, New York, and Texas, in or on baggage, stores, and cabbage. Infested shipments originated in France, Greece, Italy, and Spain. Between 1964 and 1972, additional interceptions were made in car- goes originating in Argentina, Australia, the Azores, Bel- gium, England, Haiti, Ireland, Mexico, Netherlands, and Peru, as well as most of the countries around the Mediter- ranean (DUNDEE, 1974; R. D. Munkittrick, personal com- munication, May 1979). In 1974, permits were issued to 3 importers allowing entry of H. aperta into the New York City metropolitan area for local sale (R. D. Mun- kittrick, personal communication, May, 1979). Point Isabel was formerly a locality for the native snail Helminthoglypta contracostae (Pilsbry, 1895) (PitsBRY, 1927, 1939). ALLYN G. SmrrH (in Prssry, 1939: 102) reported H. contracostae as “‘a shell of the mud flats along the shore on the east side of San Francisco Bay in the general vicinity of San Pablo. ... At these places definite colonies are to be found under low bushes, in tall grass, and under Salicornia.” The nominal species Epiphrag- mophora arnheimi Dall, 1896 -— now regarded as a synonym of H. contracostae -— was based on specimens from the San Pablo area. No Helminthoglypta were ob- served in the course of this investigation, and, given the extent of habitat alteration along this part of the Bay shore, we consider it unlikely that H. contracostae sur- vives in the Point Isabel area. SmrrH (1970: 41) stated, “the habitat for the race arnheimi on the east side of San Francisco Bay has been completely destroyed by indus- trial expansion.” Vol. 22; No. 4 THE VELIGER Page 387 ACKNOWLEDGMENTS We are grateful to R. D. Munkittrick, U. S. Depart- ment of Agriculture, for supplying literature and inter- ception records and T. Kono, California Department of Food and Agriculture, for data on the distribution of Helix aperta in California. Literature Cited Campsett, R. E. 1953. [Damage caused to cabbage near San Ysidro, California, by Helix aperta). Coop. Econ. Insect Reprt. § (6): 75 (6 Feb. 1953) Dunoez, Dez SAUNDERS 1974. Catalog of introduced molluscs of eastern North America. Sterkiana 55: 1-97 (September 1974) Frémuano, Ewatp Gamwon, Earte T. 1943. Helicid snails in California. Bull. Calif. Dept. Agricult. $2 (3): 173-187; 3 text figs. (July-Sept. 19493) Hanna, G Datias 1966. Introduced mollusks of western North America. Oceas. Pap. Calif. Acad. Sci. 48: 1 - 108; plts.1 - 4; 85 text figs. (16 February 1966) Harry, Harotp WILLIAM 1948, Notes on the foreign land snails of Louisiana. The Nautilus 62 (1): 20-24 (22 July 1948) Prrssry, Henry Auoustus 1927. Land mollusks (Helicidae) from central and northern California. Proc. Acad. Nat. Sci. Philadelphia 78: 477 - 488; plts. 36-40 (7 Feb.) 1939. | Land Mollusca of North America (north of Mexico). Acad. Nat. Sci. Philadelphia Monogr. 3; 1 (1): i-xvii, 1-573, i- ix; text figs. 1-377 (6 December 1939) Sinrz, ALtyn GoopwIn 1970. American Malacological Union CS aerene rare and endan- gered mollusks. 6. Western land snails. Malacologia 10 (1): 39-46 (14 November 1970) U. S. DEPARTMENT OF AGRICULTURE 1979. List of imtercepted plant pests (pests recorded from July 1, 1973, through September 30, 1977). Animal and Plant Health In- spection Service [APHIS] Publ. 82-5. 568 pp. (February 1979) 1954. Biologie der mitteleuropaischen Landgastropoden. Berlin: Duncker 2 Humblot. 404 pp.; 60 text figs. wtih CEMENT Page 388 THE VELIGER Vol. 22; No. 4 Spiroglyphus and Stoa, Taxonomic Problems in the Vermetidae BY A. MYRA KEEN Department of Geology, Stanford University, Stanford, California 93405 (1 Text figure) In 1961, WHEN I was pPRoposiNc a reclassification of the family Vermetidae, I could not come to a clear resolution of the problems with two generic names. Reviewing those problems now, I am reminded of the words of a Dutch zoologist, M. M. ScHEpMAN (1908: 183), who said caustically of the vermetids that they are “the most dis- agreeable among gastropods to deal with. The extreme variability ... and the often vague descriptions render ... impossible ... safe identifications.” This applies, indeed, to Spiroglyphus Daudin, 1800, and Stoa De Serres, 1855. I advised, in 1961, setting Spzroglyphus aside as a genus dubium, although it was a taxon long con- sidered molluscan, for I found there was a possibility the original material might have been annelid instead. I felt the name should be revived only if authentic type material could be brought to light or a plausible neotype specimen selected. The second name, Stoa, I dismissed as not ap- plying to the Mollusca, on the advice of the paleonto- logist Dr. B. F Howell, who was revising the fossil Annel- ida for the ‘“Treatise on Invertebrate Paleontology.” He thought that Stoa was undoubtedly a tubicolous annelid. However, when that volume of the “Treatise” was pub- lished, Howell did not list the name Stoa because he found no fossil records of it. More recently, in a system- atic reference work on polychaete annelid genera, Favu- CHALD (1977: 152) cited Stoa under the serpulid family Spirorbidae, but only as a genus that is “invalid” and “indeterminable.” The problem of Spiroglyphus remains today as enigmatic as ever, and Stoa floats ghost-like, unclaimed in either Mollusca or Annelida. This paper is an attempt to review the problems for both of these taxa and to assess the consequences of alternative courses of action. The generic name Spiroglyphus Daudin, 1800, com- prised two species, S. politus and S. annulatus. Only the latter was figured (see Figure 1). Both were said to have tubular shells, irregularly coiled, entrenched on the sur- face of other shells, S. politus on pectens and pinnas in Figure 1 Figures 28 and 29 from Daupin, 1800, “Recueil de mémoires et de notes sur des espéces inédites ...” Figs. 28-29. Spiroglyphus annulatus 28. Grandeur naturelle, sur une fissurelle 29. Grossi, et détaché. the Indian Ocean area and S. annulatus on patellas and fissurellas, also of the Indian Ocean. The figure shows a specimen attached to what appears to be a strongly ribbed keyhole limpet. The entrenched shell is well portrayed, showing the lamellar growth increments that are char- acteristic of many vermetids. However, the initial whorls are shown as straight, a characteristic of tube worms, whereas vermetid gastropods have helical coiling in the initial whorls. The shell to which Daudin’s specimen was attached is rather sketchily drawn. He had not seen the soft parts of either of the two species, and his discussion shows that he was only dimly aware of how tube worms and mollusks might differ. The name Stoa was proposed for a new genus of tubi- colous annelids by De Serres in 1855. He had three spe- cies, all illustrated, but he had only the shells for the first two, §. ammonitiformis and S. spirulaeformis. For the Vol. 22; No. 4 THE VELIGER Page 389 third one, S. perforans, he figured also an operculum. He characterized the genus as having an irregularly coiled shell, the last whorl often detached from the first and even prolonged into a straight tube; the aperture oval, ending in a solid, calcareous, conic operculum formed of three small circular bands, concave externally, diameter 2 to 3mm. The illustration of this operculum shows it as having a steeply conic profile, with a notch on the outer edge, unlike that of any known species of vermetid. De Serres’ statement, made twice, that the operculum is calcareous led Dr. Howell and later authors to infer that Stoa was indeed an annelid. Tryon (1886) re- printed De Serres’ figures (plt. 51, figs. 48-49; plt. 52, fig. 54; plt. 54, figs. 84-86). Gray in 1840, in a synopsis of the contents of the British Museum, picked up Daudin’s Spiroglyphus for use in Mollusca, family Vermetidae, alongside Vermetus, which Daudin had also described and had correctly considered as gastropod. The first malacologist to make extensive studies of the family was O. A. L. Morch, who published a series of papers in the years 1859 through 1862. Al- though in general Morch was a careful observer, his dis- cussions were sometimes ambiguous. He recognized Spiro- glyphus as a genus, coordinate with a genus Siphonium, which he dated from Gray, 1850, and he made Stoa a synonym of both. The fortunes of S7phonium, a name that was invalid because preoccupied by Siphonium Link, 1807, need not be pursued here. In synonymizing Stoa under Siphonium, Morch commented: “‘M. de Serres has established a genus of ‘Annelides sedentaires tubicolés,’ under the name Stoa, chiefly on account of their habit of burrowing a bed in the surface of other shells, exactly corresponding to the genus Spiroglyphus of Daudin. Both authors have mixed together species of two very different genera, the one with a concave, the other with a thick, convex operculum; but as Daudin has selected for illustration of his genus a species with an operculum of the latter kind, I regard the represented species as the type ...” (MO6rcu, 1861: 152). On Stoa he said (2bid., p- 153): “A conical operculum, which is solid and cal- careous, seems ... unnatural and without analogy among the Mollusca and Annelida ... . The last whorl of the operculum terminates abruptly at the edge ... which I have never seen so strongly expressed, neither have I seen an operculum of the high conical shape figured. ...” Apparently, though, he did accept Stoa as molluscan rather than annelid. Despite having synonymized Sioa, Morch used thename subgenerically under Siphonium for 6 species. Nowhere did he categorize the difference between Szphonium, s. 5., and Stoa nor cite a type species for either (in fact, the selection of S. annulatus as type of Spiroglyphus was one of the few times in all his published work that he used the word “type”). His basis for including species under Stoa is not clear; he reported the operculum as unknown in two, nearly flat in one, and concavely conic in three. The first species he listed was Siphonium (Stoa) politum (Daudin), under which he synonymized, because of sim- ilar wording of the descriptions, Stea perforans De Serres. It is this latter that has the steeply conic operculum. In a discussion of the genus Spiroglyphus, Morcu (1862a: 326) again cited Stoa as a synonym and said, “The shell ... is so similar to that of Stoa that it can only be distinguished by the operculum, which is convex outside, flat inside, with a central cylindrical wart. ... The colour varies ... from bright purple to nearly black, and the surface from nearly flat to very convex.” The next malacologist to deal with these names was Tryon (1886: 163-ff.). As Tryon was a lumper, he de- moted many of Morch’s units to lower rank but followed his arrangement in the main. He, too, synonymized Stoa with Siphonium (‘in part”) and Spiroglyphus (“in part’). Morch’s subgenus Dendropoma he ignored as a “sec- tion.” The name Stoa then virtually disappeared from the literature until 1939, when J. R. leB. Tomlin used it sub- generically for a South African vermetid, Vermetus (Stoa) corallinaceus (reallocated by later authors to Dendro- poma). Tomlin did not diagnose or cite a type species for the subgenus. Spiroglyphus, however, continued in use in Mollusca — although confused with annelids by cer- tain Californian paleontologists — until 1960. This, then, is the historical background for my recom- mendations in 1961. No one since has put in a brief for resuscitating Stoa, but at least two authors (Morrison, 1968; ABBOTT, 1974: 99 - 101) have risen to the defense of Spiroglyphus on the basis of prioritv. I would like here to point out the various alternatives and consequences. First and most obvious step is search for original type material. When I inquired of Dr. Edouard Fischer- Piette of the Paris Museum, who had been successful in re- covering Adanson’s lost collection, whether the Daudin specimens might be at Paris, he replied that not only were they not in the Paris Museum but that he had no idea of how to start a search. Obviously, an outsider would stand little chance of making this discovery. De Serres’ specimens were stated to be in the collections of the Université de Montpellier. Through a French colleague, M. Jacques Laborel, I was put in touch recently with the curator there, Mlle. F Cassagne-Mejean, who promised not only to search for the material but to send photo- graphs if it were found. Nothing, however, was forth- coming, and we must conclude that this material is lost. The problem of Spiroglyphus is more complex than ap- pears on the surface. Not only is there an open possibility Page 390 that the original specimens were annelids but also that the concept that developed of it as a mollusk might have been unjustified. Authors have taken Morch’s state- ments at face value without realizing that he was making judgments based only on his personal experience. Discus- sing S. annulatus, which he made the type species, he quoted Daudin’s statement that it occurred on patellas and fissurellas of the Indian Ocean, and then said (1862a: 330 - 331): “The represented species is probably Frssur- ella barbadensis Gm.” Later authors (e. g., MorRISON, 1968: 45) disregard the “probably” and identify the shell on which the supposed vermetid is shown as F barbadensis Gmelin, 1791, a Caribbean species. One won- ders whether they tried actually laying a Caribbean Fis- surella alongside Daudin’s figure. Either Daudin omitted the entire anterior half of the supporting fissurellid and showed only the part behind the orifice or else he drew the shell tilted but the entrenched specimen normally. In either case, he failed to show the radiating ribs accurately, and the proportions seem inconsistent. Had not Daudin cited “‘fissurelles,’” I would even suggest that the support- ing shell might be one valve of a bivalve that had been perforated by a drill-shell. Although fissurellids are more common in the Caribbean, there are some species of that group in the Indo-Pacific area, and to my eyes a figure labelled ““Diodora jukesiu (Reeve)” from northwest Aus- tralia, in “Selected Shells of the World Illustrated in Co- lour,” by SHIKAMA & HorIKOSHI, 1963: plt. 5, fig. 12, isa better match than any figure or shell of a Caribbean fis- surellid that I have been able to find. I feel that Daudin’s stated locality should be taken seriously unless there is compelling evidence to the contrary. Morch’s characterization of the morphology of Spiro- glyphus annulatus really is based on his observation of Caribbean material. He was aware of only one species of “Spiroglyphus” there, with a convex purple-black oper- culum, flat on its inner surface except for a central mamil- la. Under S. annulatus he synonymized two Caribbean forms, Vermetus irregularis and V. corrodens, both of Orbigny, 1842 — described from Martinique, the type material now in the British Museum. Actually, there are in the Caribbean three species of entrenching vermetids, each with a distinctive operculum, only one of which is convex. The second Caribbean form has a flat operculum, and the third has one that is concave and horn-colored, with an orange spot on the central mamilla. What the Indo-Pacific counterpart has for an oper- culum is an open question. I have seen material from Vietnam that, for shell features, matches Daudin’s figure well. However, I have not yet detected a specimen with an intact operculum. There is a lot from the Riu Kiu Is- lands in the Stanford University collection that has a conic, THE VELIGER Vol. 22; No. 4 reddish operculum. The coiling resembles Daudin’s figure, but the shells are sculptured spirally, not transversely. PRA- SHAD & RAo in 1933 had an excellent description of Ver- metus (Spiroglyphus) andamanicus, a vermetid that bur- rows in the surface of large Trochus in the Indian Ocean area. The operculum in this is somewhat but not markedly convex. Morch’s discussion implied (although he did not actu- ally state) that Daudin described and figured an oper- culum. What Morch took to be a basic pattern for the genus was what he had observed in one Caribbean form. As Morton (1965: 627) has shown, however, there are at least six fairly consistent opercular patterns in the en- trenching vermetids. No worker, at present, is ready to split the group into this many subgroups, but establishing for each species which pattern the operculum takes is important for future reference. Before any further split- ting is done, we need more work on local populations in areas of abundance to assure that groupings made on the basis of hard-part morphology (shell and operculum) are consonant with what can be observed as to the soft parts, growth habits, reproduction, etc. Trying to be impartial on the matter of Spiroglyphus, both Dr. Michael Hadfield — specialist on Pacific Ver- metidae — and I have been keeping watch for specimens that might qualify for designation as neotypes. We have not yet found one that meets all requirements. Nor have we found a satisfactory replacement specimen for Stoa. An alternative to designating fresh material as replace- ment would be to designate the holotype of some already satisfactorily documented species as neotype of the cryp- tic taxon. Skillfully enough done, this might be a way of filling three of the open niches for opercular types. For Stoa it might be done by designating the lectotype of Ser- pula maxima Sowerby, 1825 (in the British Museum) as the neotype of one of De Serres’ species, preferably Stoa perforans, which would then be given status, but only as a junior synonym. However, this would be to disregard basic habits of these mollusks — Sowerby’s species is as- sociated with corals or on reef rocks and does not attach to bivalves as Stoa does. The name would be sal- vaged at the expense of consistency. Spiroglyphus might be reinstated by designating as neotype the holotype of Vermetus andamanicus Prashad & Rao, 1933. This latter is extant and in good condition in the museum of the Zoological Survey of India, as reported recently by the curator, Dr. N. V. Subba Rao (letter dated July, 1979). However, the type specimens for this species are much smaller than what Daudin indicated and not on a fissur- ellid of any sort. I would feel reluctant to set aside a well- proposed name such as this is in order to validate an equivocal earlier one. Thus, I would reject the option of Vol. 22; No. 4 THE VELIGER Page 391 making arbitrary neotype selections. Rather, it seems preferable to jettison both Spiroglyphus and Stoa. Declaring Spiroglyphus and Stoa as genera dubia would leave the field open for Dendropoma, the first name for entrenching vermetids that was based upon authentic type material. During the last two decades that name has been used in a substantial number of papers (at least 16 titles), not only by malacologists but also by earth scientists and others who are beginning to recognize how useful these mollusks are as markers of shoreline conditions. Living as they do attached to rocks and large shells in the upper intertidal zone, they provide good evidence for any changes in shoreline levels. Therefore, in view of this practical incentive for nomen- clatural stabilization and with the support of Dr. R. Tucker Abbott (personal communication), who on fur- ther study has changed his opinion about conserving Spiroglyphus, 1 am asking Dr. Michael Hadfield to join me in a petition to the International Commission on Zoological Nomenclature requesting that Spiroglyphus and Stoa be suppressed as generic names founded on unidentifiable species. Literature Cited Assott, Ropert Tucker 1974. American seashells. aed ed, 663 pp.; 24 col. pits Van Nostrand Remhold, New York Dauoin, FE M. 1800. Recueil de mémoires et de notes sur des espéces inédites ou peu connues de mollusques, de vers, et de zoophytes. Paris, privately printed. pp. 1-50; 2 plts. Dz Serres, MaRcEL 1855. Note sur un nouveau genre d’annélide tubicole perforant. Ann. Sci. Nat., Paris, Zool. (4) 4: 238-243; 1 pit. FAUCHALD, KRISTIAN 1977- The polychaete worms: definitions and keys to the orders, fami- lies, and genera. Los Angeles County Nat. Hist. Mus., Science Ser. no. 28: 187 pp.; 42 figs. Keen, A. Myra 1961. A proposed reclassification of the gastropod family Vermetidae. Bull Brit. Mus. (Nat. Hist.), Zool., 7 (3): 183-212; 2 pits. Morcu, Otro ANDREAS Lowson 1859-1860. Etudes sur la famille des vermets. Journ. de Conchyl. 72 342 - 360 (1859); 8: 27- 48 (1860) 1861-1862. Review of the Vermetidae. I. Proc. Zool. Soc. London for 1861 (no. 2): 145-181, September 1861; II. ibid. (no. 3): 326 - 365; plt. 25, April 1862; III. ibid. for 1862 (no. 1): 54-83, June 1862 Morrison, JosepH Paut ELDRED 1968. Spiroglyphics: a study in species association. Amer. Malac. Union, Ann. Reprt. for 1968: 45-46 Morton, JoHN Epwarp 1965. Form and function in the evolution of the Vermetidae. Bull. Brit. Mus. (Nat. Hist.), Zool., 1: (9): 585-630; 15 figs. ScHEPMAN, MatrHeus Marinus 1908-1913. The Prosobranchia of the Siboga Expedition. Leiden, E. J- Brill Co., Monogr. 49, Siboga Expedition, 484 pp.; 32 plts. Tryon, Gzorce WASHINGTON, Jr. 1886. | Manual of Conchology, Philadelphia, ser. 1, vols. 1-17 (1879 to 1916). Vermetidae, 8: 163 - 191; plts. 48-58 (July 1886) Page 392 NOTES & NEWS Soviet Contributions to Malacology in 1978 BY KENNETH J. BOSS AND MORRIS K. JACOBSON Museum of Comparative Zoology, Harvard University Cambridge, Massachusetts 02138 INTRODUCTION As IN PAST YEARS, herein are presented the titles of Soviet malacological literature as abstracted in the Refer- ativnyy Zhurnal during the year 1978 (see The Veliger 21 (4): 490 for last year’s listing and reference to those published earlier). Hopefully, this delineation of mainly Russian contributions to the field of molluscan studies will continue to be useful. Generally, we have followed the editors of the Referativnyy Zhurnal in the arrange- ment of categories and often we have appended a short summary of the contents of an individual paper. To molluscan systematists the most important Soviet work continues to be the often detailed, revisionary mono- graphs which sometimes introduce new taxa and frequent- ly suggest fundamental alterations in accepted systems of classification. Special attention, thus, should be called to: 1) Shileiko’s study on the naticoid gastropods as well as his major treatise on the helicacean stylommatophor- ans of the USSR and to his series of papers on the type species of the genera of the orthurethrous pulmonate fam- ily Buliminidae (= Enidae), which introduces a classi- ficatory scheme quite different from that of Zilch; 2) Moskalev’s revision of lepetid prosobranchs in which new genera are established and those previously poorly known given more extensive coverage; 3) Ivanova’s monograph on the anomalodesmacean verticordiid bi- valve Policordia; 4) Kudinskaya’s and Minichev’s description of a new species of the acochlidiid opistho- branch Hedylopsis along with a key to all known species; THE VELIGER Vol. 22; No. 4 5) Nesis’ review of the enoploteuthid cephalopod sub- family Ancistrocheirinae; and 6) Nigmatullin’s re- view of the systematics of the ommastrephid squids which might be compared to the recently published American work of Wormuth (1976. The biogeography and numerical taxonomy of the oegopsid squid family Ommastrephidae in the Pacific Ocean. Bull. Scripps Inst. Oceanogr. vol. 23). Several notable items dealt with specific geographical areas or faunas: 1) Golikov and Kusakin published a detailed important work on the shell-bearing littoral snails of the USSR; 2) Golikov and Gul’bin exten- sively treated the prosobranchs of the Kurile Islands, and 3) Gul’bin and Fedchina discussed the closely juxta- posed sublittoral gastropods of the Commander Islands. Other noteworthy contributions included: 1) Goru- chev’s discussion of the evolution of the buccinoid gastro- pod Neptunea in the Bering Sea which might well aug- ment the recent research of Nelson (1978. The Veliger 21 (2): 203-215); 2) Nesis and Shevtsov’s citation of the first data on the abyssal cephalopods in the Sea of Okhotsk; and 3) Golikov’s description of the new ge- nus Costaria, the northernmost fasciolariid. Several of the physiological and biochemical papers indicated an intense interest in electrophoretic techniques, especially in distinguishing species. Different authors pointed out that previously recognized forms or ‘species’ were in reality only variants of single species and thus to be accorded the oblivion of synonymy. One item which should be of especial interest to Ameri- can workers doing research on the marine fauna of the north Pacific, north Atlantic or Arctic Oceans was Ka- fanov’s discussion on the evolution of the malacofauna of the north temperate-boreal zones, for which we include a more extensive abstract than usual. We thank Dr. Joseph Rosewater of the Smithsonian Institution who aided us in obtaining copies of the Refer- ativnyy Zhurnal from the Library of Congress. Mrs. Mary Jo Dent carefully typed the manuscript. Abbreviations and acronyms we have used are: AN — Akademiya nauk (Academy of Science) Biol. Morya — Biologiya Morya (Marine Biology) ES - English summary Mekh. — Mekhanizmy adaptatsii zhivykh organizmov k vliyaniyu faktorov sredy (Adaptive mechanisms of living organ- isms to environmental factors), Leningrad NDVS - Nauch. Dokl. Vyssh. Shkol. Biol. Nauk. (Scientific Re- ports of the Higher Educational School for Biological Sciences) Vol. 22; No. 4 SID - Sbornik Rabot Instituta biologiya morya Dal’nevost. nauch. tsentr. Akad. Nauk. SSSR. - (Papers of the Marine Biological Institute of the Far Eastern Scientific Center, Academy of Science, USSR). Viad. — Perv. Vsyes. konf. po mor. biol., Vladivostok. Tezisy dokl. (The first All-Union conference on marine biology, Vladivostok, Thesis Reports) VNK _ - Vses. nauch. konf. po ispol’z promyslov bespozvonochn. na pishchev. kormov. i tekhn. tseli, Odessa, Tezisy dokl. (All-Union scientific conference for the commercial use of invertebrates for food, fodder and technical purposes, Odessa, Thesis Reports) ZEBP - Zhurnal Evolyutsionnoi biokhimii i fiziologii. (Journal of evolutionary biochemistry and physiology). ZZ — Zoologicheskii Zhurnal (Zoological Journal) GENERAL Beroer, V. Ya. 1977. ‘The functional organization of the system of salt water adaptations in euryhaline marine mollusks. Vlad.: 15 - 16 BERZHBINSKAYA, N. A. 1977. The regulation of glycolysis at the systemic and molecular levels in molluscan tissues. Vlad.: 26 - 27 [Enzymes involved in glycolysis in chitons, bivalves, gastropods and cephalopods were investigated] GorBATENKO, S. A., S. D. NiKo.agv « S. V. Popov 1975. Factors which influence the isotonic oxygen-carbonate composition of the shells in Caspian mollusks. SID, no. 4: 159 - 166 [Analysis shows that molluscan shells grow in isotopic equilibrium with the '8O content of the water of the Caspian Sea, virtually in- dependent of variations in temperature, suggesting the suitability of paleoisotopic studies of Pontic Pliocene-Quaternary mollusks] Icnar’ev, A. V.« G. A. YEVSEEV 1977. The temperature of growth of marine mollusks and their geographical distribution. Vlad.: 57 - 58 [18 species of stenohyaline bivalves from the shelf zone of the Sea of Japan and the Sea of Okhotsk were studied; correlations were noted between the geographic distribution of each species and lim- iting thermal factors] Karanov, A. I. 1978. On the centers of origin and several characteristics of the ecological evolution of the cold and temperate water marine malacofauna of the northern hemisphere. Biol. Morya, No. 1: 3-9 [The development of modern northern coldwater marine malaco- faunas is intimately related to 3 main Paleogene zoogeographic regions: Northern-Japanese-Sakhalin, Western North American (Columbian) and Boreal-Central Tethyan (northeastern Atlantic), all genetically derived from a more remote tropical source fauna, that of the extremely rich, highly endemic Tethys of the Mesozoic. In the Cenozoic, there were 3 isolated tropical faunas; Indo-West THE VELIGER Page 393 Pacific, Central American and West Indo-African as well as a small subtropical unit — the Lusitanian-Mediterranean. The source fauna of the cold-temperate boreal was largely the northern periphery of the tropical areas, especially Tethyan relicts in the North Pacific (Northern Japanese-Sakhalin) derived from the Indo-Malayan Region. Today, the number of North Pacific endemic species is 6 to 8 times greater than in the North Atlantic (Boreal-Central Tethyan). This faunistic evolution, with the con- comitant development of strong climatic provincialism, began with the global decline in temperatures in the Cenozoic and the extinc- tion or southward displacement of less adjustable elements. With less competition, survivors in the northern extremes radiated poly- typically, forming new but closely related taxa, usually with broad ecological tolerances or eurytopy and dispersed widely through the Boreal and Arctic regions. These elements were unable to reinvade the tropics or subtropics with their high species diversity and filled niches. ] Karpevicu, A. F « A. T. SHuRIN 1977. ‘The role of manganese in the metabolic processes of Bal- tic Sea mollusks. Biol. Morya, No. 6: 50-57 (ES) [Increasing concentrations of manganese found in tissues of 3 spe- cies of bivalves (Macoma balthica, Mya arenaria, and Dreissena polymorpha) as ambient levels of Mn were raised until, beyond 20 mg/L, death ensued. Shells in Mn solutions for 2 months became blackened as a result of accretion of granules of Mn, and the authors suggest that this may be one manner in which Mn concretions are formed] Kuoxkuutkin, I. M. & A. I. LAZAREVA 1977. Adaptive polymorphism in populations of terrestrial mol- lusks. Mekh., p. 151 LuKanin, V. V. 1977. The complex effects of temperature and salinity on the adaptive reactions of some White Sea mollusks. Mekh., p. 84 Reznik, Z. V. 1977. Zoogeographical features of the terrestrial molluscan fau- na of the Stavropol Heights. Fauna Stavropol’, No. 2: 37-43 [75 species were found, 42 from the Mediterranean subregion (of which 5 are found in the hilly regions of Europe, 12 from the Mediterranean and Pontic subregions, and 25 Caucasian endemics) ; 13 species from the Europe-Siberian subregion, and 19 species of wide Palearctic distribution] ZOLOTAREV, V. N. 1975. Molluscan shell structure and paleotemperature analysis. SID, no. 4: 114 - 158 [Careful analysis of shell structures offers the possibility of evalu- ating ambient environmental temperatures prevalent at the time of the formation of various parts of the shell] Zo.oTaREV, V.N. & A. V. IGNAT’EV 1977. Seasonal changes in basic shell layer thickness and the temperature at time of growth in marine mollusks. Biol. Morya, No. 5: 40-47 (ES) [Mass-spectrometric analysis of isotopic-oxygen in the shell structure of 5 species showed correlation with growth intervals and tempera- ture at time of growth. Annulations in the shell usually indicate Page 394 reduced shell deposition during the winter at minimal tempera- tures] GASTROPODA, GENERAL Avyakrinskaya, I. O. 1978. Sexual dimorphism and age differences in morpho-bio- chemical characteristics of some gastropods. ZZ 57 (3): 359- 363 (ES) [In some gastropods sexual dimorphism is externally apparent in the different coloration or shape of male and female individuals. However, it is expressed also in lesser known internal ways: in differ- ent concentrations of respiratory pigments, total proteins, myoglo- bin and the specific weight of the hemolymph] Goutxov, A. N. « O. G. Kusakin 1978. The shell-bearing littoral gastropods of the USSR. Nauka, Leningrad. 292 pp.; illust. [This useful handbook discusses 148 species placed in 70 genera, 41 families, 14 orders and 5, subclasses] Guv’sin, V. V. & I. Yu. FEpCHINA 1977. Sublittoral prosobranch gastropods of the Commander Islands. Vlad., p. 41 [A biogeographical characterization, including data on vertical dis- tribution, is given for the 88 species of prosobranchs occurring a- round the Commander Islands] PROSOBRANCHIA ALYAKRINSKAYA, I. O. « S. N. Dotcova 1977. Quantitative contents of the respiratory pigments of Ra- pana thomasiana Crosse in the Black Sea. Dokl. AN SSSR 236 (6): 1500-1502 [The mg per g of body weight is given for the respiratory substan- ces, hemocyanin, tissue hemoglobin and myoglobin] Goxtxov, A. N. 1977- On the family Fasciolariidae in temperate waters. Issled. fauny moryei, 21/29: 102-104 (ES) [Costaria borealis, n. g., n. sp. was taken at 414m to the east of Iturup Island in the Kurils, a first record of a fasciolariid discovered in the temperate waters of the northern hemisphere] Gouixov, A. N. « V. V. Guv’BIn 1977. Prosobranchs of the Kuril Island shelf. II. Order Hami- glossa-Homoestropha. [In] Fauna pribrezhn. zon. Kuril. ostro- vov (Fauna of the littoral zone of the Kuril Islands), Moscow, pp. 172 - 268 [Of 241 species and subspecies of prosobranchs, 128 were noted in the orders Hamiglossa, Toxoglossa, Heterostropha, Ptenoglossa and Homoestropha. Species of Mohnia, Sipho, Aulacofusus, Sulcosinus, Retifusus, Buccinum (including 2 new buccinid genera), Trophon- opsis, and Oenopota are delineated. The fauna of the southern part of the islands belongs to the Pacific-boreal subregion and differs THE VELIGER Vol. 22; No. 4 from the northern Kumi Islands; a special Kuril subprovince is recognized, especially in the central islands] 1978. Prosobranchs of the Kuril Island Shelf. I. Order Doco- glossa-Entomostoma. Zhivotn. i rastitel’n mir shel’fovikh zon Kurilisk. Ostrovov (Fauna and flora of the Kuril Island shelf zone), Moscow, pp. 159 - 223 [Of 241 species and subspecies of prosobranchs collected, 113 be- longed to the orders Docoglossa, Fissobranchia, Anisobranchia, Protopoda, Discopoda, Canalifera, Echinospirida, Aspidofora, and Entomostoma. 22 species and subspecies belong to Puncturella, Scissurella, Margarites, Homalopoma, Lacunitunica, Onoba, Ris- soella, Trichotropis, Torelia, Cerithiopsis and Furukawaia. One new genus of Lacunidae and a new subgenus of Trochidae are proposed. Egg capsules of Capulus nobilis, Lunatia pila, and Cryptonatica janthostoma are described and figured for the first time] GorucuHev, V.N. 1978. The gastropod genus Neptunea Réding in the Bering Sea. Nauka, Moscow, 90 pp. [Data on the biology, relationships and distribution of the species of Neptunea in this area augment a new taxonomic treatment of the genus] In’, L. B. 1975. On the growth changes of Arcularia gibbosula. SID, No. 4: 167-173 [A study of the successive growth changes from immature to mature stages in the nassid gastropod Arcularia gibbosula clearly indicates the systematic relationships of the species] KarabBel.t, O. Z. & B. M. LocviINENKO 1977. ‘The distribution of the genus Turricaspia, section Laeui- casfra, in the Caspian Sea. Vestn. Moscow State University, Biologia, No. 3: 57 - 63 [On the basis of benthic sampling in the central and southern parts of the Caspian (321 stations), the vertical and horizontal ranges and substrate preferences of the 4 species of Laevicaspia were established] 1978. The distribution of Theodoxus, family Neritidae, in cen- tral and southern parts of the Caspian Sea. NDVS, No. 2: 42 - 46 [321 bottom stations were investigated and the ranges of Theodoxus established along with its depth distribution and substrate prefer- ences] Kartavizev, Yu. F. 1976. Color polymorphism in the shells of the gastropod mol- lusk, Littorina brevicula (Philippi). SID No. 5: 93 - 98 [3 color forms of this species occur in Vostok Bay of the Maritime Province] MoskALey, L. I. 1977. Revision of Lepetidae of the World Ocean. Tr. In-ta okeanol. AN SSSR 108: 52-78 (ES) Vol. 22; No. 4 [Two new genera, Limalepeta and Bathylepeta are established and diagnoses and further information given on Lepeta, Iothia, Propili- dium, Cryptobranchia, Notocrater, Maoricrater and Punctolepeta] Suiizixo, A. A. 1977. Material on the morphology of the Naticoidea and prob- lems of the taxonomy of the superfamily. Tr. In-ta okeanol. AN SSSR 108; 79 - 97, 9 figs. (ES) [17 representatives of the superfamily were studied and the families Polinicidae and Naticidae, with Naticinae and Sininae, recognized. The Polinicidae are distinguished by having: 1) the cephalo- podial portion of the male gonoduct as an open groove; 2) a simple, uniform penis and 3) a horny operculum. The Naticidae have a closed male gonoduct, a variable, complicated penis and a corneous or calcareous operculum; the Sininae have the shell reduced. The new genus Scarlatia (type-species, Natica fortunei Reeve) is described] Suustov, A. I. 1978. An analysis of the correlative structures of morphological features of the Bithynia leachi group. Zhiznen tsikly, ekol. i morfol. gelmintov zhivotnikh Kazakhstana (The life cycles, ecology and morphology of helminths of Kazakstan), Alma- Ata, pp. 52 - 61 [The conchological features of the Bithynia leachi complex were studied statistically, showing that B. I. troscheli and B. l. inflata are mere varieties and not real subspecies] OPISTHOBRANCHIA Kuprnsxaya, E. V.& Yu. S. MINnICHEV 1977. Psammological Essays. 1. The organization and systematic platement of Hedylopsis murmanica sp. n. (Acochlidiida) . Trudy Petergof. biol. in-ta LGU (Leningrad State University) , No. 6: 69 - 86 (ES) [The new species is an interstitial opisthobranch. A key to all spe- cies of Hedylopsis as well as a discussion of the phylogenetic relation- ships of the Acochlidiida are provided] OsuKuova, E. V. 1978. The functional morphology of the epithelium of the di- gestive glands in the nudibranch Coryphella rufibranchialis. Trudy Petrogof. biol. in-ta LGU (Leningrad State University), No. 26: 118-131 (ES) [A light-optical and ultrastructural examination reveal 3 types of cells: 1) non-differentiated cells, 2) calcium (lime) cells with an albumen-synthesizing apparatus for active ion transport, 3) digestive cells, the cytoplasm of which is filled with hetero- morphological vacuoles depending on the stage of the digestive cycle] TERRESTRIAL PULMONATA ALYAKRINSKAYA, I. O. 1977. Seasonal variations in the buffering capacity of the hemo- THE VELIGER Page 395 lymph in Helix pomatia. Dokl. AN SSSR 236 (5): 1264- 1266 [In periods of summer aestivation, winter hibernation, and nest dig- ging, when normal aerobic respiration is disturbed, there is an in- crease in the buffering capacity of hemolymph] Martek, P. V. & V. M. MaKEEVA 1977. Polymorphic esterases and the distribution of Bradybaena fruticum Mill. Zh. obshch. biol. 38 (6): 908-913 (ES) [Electrophoretic analysis of esterases in 3 populations of Brady- baena fruticum near Moscow showed that the populations are gen- etically distinct; esterase polymorphism was also correlated with phenotypic characters such as shell-banding] Suiterko, A. A. 1977. ‘The structure and systematic relationships of the species of the genus Siraphorus Lindholm 1925 (Enidae). NDVS, No. 9: 40 - 46 [Previously 2 species were assigned to Siraphorus: entoptyx Lind- holm, now recognized as the sole species in Siraphorus, which is herein accorded subgeneric rank in Pseudonapaeus; and moltschan- out Likharev and Rammel’meyer, for which a new and separate genus is erected, Sirapharoides. The structure of the reproductive organs of Central Asiatic enids differs from those of the Caucasus, and the generic name Jaminia is improperly applied to enids from Central Asia, especially the region of Tyan-Shan] 1978. A study of type-species of certain generic taxa in the Buliminidae (=Enidae). 1. Species of Middle and Central Asia. ZZ 57 (3): 344-358 (ES) [The external appearance and internal reproductive structures of 12 taxa are described: Pupinidius, Serina, Subzebrinus, Styloptychus, Chondrulopsina, Mastoides, Parachondrula, Laeonapaeus, Pseudo- petraeus, Laevozebrinus, Chondrulopsis, and Pseudonapaeus] 1978. A study of the type-species of the generic group taxa of the Buliminidae (=Enidae). II. Species of Europe, Near East and Asia Minor. ZZ 57 (4): 512-522 (ES) [The genital anatomy and shells are described and figured for 10 taxa: Ena, Zebrina, Buliminus, Pene, Jaminia, Antichondrus, Chona- rula, Eubrephulus, Geminula, and Pseudochondrula| 1978. A study of the type-species of the generic group taxa of the Buliminidae (=Enidae). III. Species of the Crimea and Caucasus. Problems of familial systematics. ZZ 57 (6): 834- 850 (ES) [The groups, Caucasicola, Peristoma, Thoanteus, Brephulopsis, Ra- musculus, and Retowskia are characterized by their conchological and reproductive features. A new classification and phylogeny of the family is proposed wherein 8 subfamilies are recognized: Chond- rulinae, Eninae, Retowskiinae, Multidentulinae, Bulimininae, Jami- niinae, Chondrulopsininae and Pseudonapaeinae] Page 396 1978. ‘Terrestrial mollusks of the superfamily Helicoidea. Fauna of the USSR, Mollusks vol. 3, prt. 6, 384 pp.; 6 figs. Nauka, Leningrad [This important revision of the superfamily, with lengthy intro- duction on the biology and morphology of helicacean snails, dis- cusses 166 Recent and 15 fossil Soviet species, giving figures of the shells and genital anatomy; several alterations of previously accep- ted suprageneric classifications are introduced] AQUATIC PULMONATA GoroxkHov, V. V. 1978. ‘The ecological adaptation of Lymnaea truncula to man- made alterations to its normal habitat. Rastitel’n. i zhivotn. naselenie Moskvy i Podmoskov’ya (Floral and faunal popula- tions in Moscow and vicinity), Moscow, pp. 126 - 127 [An account of the occurrence of populations of this species in disturbed environments in the vicinity of Moscow] Kamaroin, N. N. « V. A. SoKoLov 1977. A study of the structure and functional characteristics of the osphradium of Lymnaea. Mekhan sensor retseptsii (The mechanism of sensory receptivity), Leningrad, pp.183 - 188 Potanina, N. V, V. 1. Starostin & M. A. LANGE 1977. Autoradiographic examination of amoebocytes in the blood of Lymnaea stagnalis. Zh. obshch. biol. 38 (6) : 940 - 944 (ES) Soxo.ov, V. A. « N. N. KaAMARDIN 1977. On the possibility of the differentiation of osmotic and chemical components during irritation of the osphradial sys- tem of the pond snail. Mekh., pp. 127 - 128 ZAITSEVA, O. V. 1978. Histochemical examination of the osphradium in Lym- naea stagnalis. Morfol. osnovy funkts. evolutsii (Morphologi- cal survey of functional evolution), Leningrad, pp. 18 - 23 [The osphradium, a chemoreceptor, is an isolated epithelial chan- nel innervated by numerous sensory fibers having their endings on the epithelial surface. Cytochemical study reveals 2 moieties in the epithelial mucus, an acid mucopolysaccharide and a glycoprotein] ZaiTSsEVA, O. V., V. A. Kova.ev « L. S. BocHAROVA 1978. A study of the morpho-functional interrelationships of the sensory epithelium and statoconia in the statocysts of Lymnaea stagnalis. ZEBP 14 (3) : 307 - 309 [The statoconia have an egg-like shape (from 4.5X2.4 to 11.0X 6.6 wm) and lie freely in the statocyst cavity, in contact with the cilia of the sensitive cells. These cilia are about 7.5 ym long and are placed about 1.4 to 1.5 wm apart, with supporting microvillary structures scattered among them. The position of the animal is monitored by the constant functional polarization of the sensitivity of the statocysts via the presence or absence of contact between the statoconia and the cilia] THE VELIGER Vol. 22; No. 4 BIVALVIA ALYAKRINSKAYA, I. O. 1978. Biochemical adaptation to desiccating conditions in bi- valves of Kurshsky (Courland) Bay in the Baltic Sea. ZZ 57 (1): 136-138 (ES) [During desiccation, calcium ions, provided by the dissolution of the crystalline style and the shell itself, are incorporated as buffers into the hemolymph to offset increased acidity in Anodonta com- planata, Unio tumidus, and Dreissena polymorpha] Burovina, I. V., O. A GoncHarEvSKAYA & N. B. PrvovAROVA 1978. X-ray microanalysis of the elementary composition of hemolymph in the mussel, Mytilus edulis. ZEBP 14 (3): 241- 245 [In a study of the ions of Na, K, Cl, Ca, P, S, and Mg, in mussels in waters of low salinity, it was shown that K was accumulated in the hemolymph at higher concentrations than that of the environ- mental medium] Goromosova, S. A. 1977. Elements of carbohydrate metabolism of mussel (Myti- lus) in connection with adaptations to environmental ex- tremes. Ekol.-fiziol. issled. v prirode i eksperiments. Frunze, PP- 155 - 169 Ivanova, V. L. 1977. New data on the composition and distribution of the deepwater genus Policordia Dall, Bartsch, and Rehder. Tr. In- ta okeanol. AN SSSR 108: 173 - 197; 18 figs. (ES) [2 new subgenera with 8 new species are recognized anatomically: Angustebranchia with 5 species in the ultra-abyssal Kurile-Kam- chatka Trench and Latebranchia with 8 species, mostly of bathial- abyssal occurrence in the Pacific Ocean] Karpenko, A. A. 1977. Cardiac rhythm and the physiological condition of the littoral scallop (Patinopecten yessoensis). Vlad.: 65 - 66 [Cardiac rhythm is a good indication of the level of activity of metabolism in bivalves. In this scallop, it rises with rising tempera- tures, increases after swimming and in the proximity of predators like starfish, and rises sharply with lowered salinity] Kartavtsev, Yu. F, S. M. Nixirorov « A. I. PupovKIn 1977. The taxonomic position and genetic variability of Creno- mytilus grayanus. VNK, pp. 41 - 42 [Electrophoretic analysis fails to support the contention of Smirnov that there are 2 taxonomically separable forms of this species] Kuosor’ey, V. G. 1978. On the ability of unionids to regulate the ionic composi- tion of water. Biopovrezhdeniya materialov i zashchita ot nikh (Bio-damages of materials and the defense against them), Moscow, p. 11 [Freshwater bivalves are indicators and regulators of alkaline con- centrations] Vol. 22; No. 4 Krasnov, YE. V. 1977. Non-specific adaptation by marine mollusks to the process of calcification. Materiali 2-go Simpoz. Protsessi adaptsii i biol. aktivn. veshchestva, Vladivostok, 1975, pp. 42 - 46 Kukin, A. F 1976. The reproductive cycle of Swiftopecten swifti. SID No. 5: 122-130 [Normally gonochoristic (only 1 hermaphrodite found in more than 200 individuals), Swiftopecten swifti in Vostok Bay spawns from August through the first half of September at 21-22° C. Histologi- cal studies document changes in the gonad] Locvinenxko, B. M., O. P. Kopotova « O. Yu. PRABDUKHINA 1979. Redkollegiya Biol. Nauki (Editorial Board, Biological Sci- ence), Moscow, 17 pp. (manuscript deposited in VINITI) [Electrophoretic analysis of populations of Mytilus galloprovincialis and Ostrea edulis in the Black Sea] LyuTsKanov, D. 1975(1976). A biometrical study of Dreissena polymorpha Pal- las in Shablessky Lake. Nauch. trudy Plovdiv. Univ., Biol. 13 (4): 227-236 (Bulgarian; Russian and French summaries) [Analysis of 2 populations from 2 depths showed adaptive differ- ences in both conchological and anatomical parameters] MarcHeEnko, A. A. & A. L. PoLENov 1977. On the participation of Gomori-positive neuro-secretory cells in Mytilus edulis L. in adapting to hypocsin. Ekol.-fiziol. issled. v prirode i eksperimete. Frunze, pp. 223 - 224 Marcus, B. A. & G. P PinaEv 1977. The composition of the muscle albumens and its system- atic significance in bivalve classifications. Vlad.: 99 [Adductor muscle extracts from 25 species of bivalves from the Seas of Japan and Okhotsk were examined electrophoretically and a phenogram of relationships constructed, which validated the classi- fication of Nevesskaya and her colleagues] Mepvepeva, L. A. 1976. The reproductive cycle of Spisula sachalinensis. SID No. 5: 131 - 135 Nevesskaya, L. A. 1975. Ontogenetic development of the shell and various modes of speciation in bivalves. SID No. 4: 17 - 34 [Analysis of ontogenesis of the shells of 30 Miocene and Quater- nary species of bivalves revealed the occurrence of the evolution of some species by neotony] Nixirorov, S. M. 1977. On the systematics of the oysters of the southern littoral of the maritime province. Vlad.: 108 [Electrophoretic analysis of various tissue extracts shows that Crass- ostrea gigas is a single species, not 3 distinct ones] Nixiroroy, S. M. « A. I. PuDovKIN 1977. Genetic and morphometric variability in Crassostrea gigas. VNK: 60-61 THE VELIGER Page 397 [Electrophoretic data indicate that only a single species of oyster, Crassostrea gigas, inhabits Peter the Great Bay] Nistratova, S. N., V.N. Danitova « I. S. NrKow’sKAYA 1978. Seasonal changes in the sensitivity to acetylcholine in the cardiac muscle of Anodonta complanata and A. piscinalis. ZE BP 14 (4): 358 - 364 (ES) [The greatest change in sensitivity occurs at spawning when gametes are transferred and fertilization takes place with retention of zygotes in the ctenidia and before development of the glochidia in the mar- supium] Pinaev, G. P 1978. Levels of organization in the retractive apparatus of the adductor muscles of bivalves. 3-ya vsyes. konf. po biokhimi myshch (Third All-Union conference on the biochemistry of muscles), Moscow, pp. 13 - 15 [Electrophoresis of muscle proteins in 26 species of bivalves con- firmed their systematic placements and established a correlation between the contractile system and the quantities of actin, myosin and paramyosin] PuiseTsKaya, E. M. 1977. Insulin in mollusks and its role in the regulation of the carbohydrate metabolism. ZEBP 13 (5): 600-606 (ES) [Insulin or an enzyme very similar to it is produced in intestinal cells of the freshwater mussels Anodonta cygnea and Unio pictorum] Premincer, N. K. 1977. Protein composition of the contractile apparatus of muscle tissue in the mussel, Crenomytilus grayanus. Biol. Morya, No. 5: 82 -84 [Electrophoretic analysis showed high paramyosin and low myosin in anterior byssal retractor muscle and 2 parts of the adductor muscles, but the reverse obtained in muscle tissues of the mantle, heart and foot] Punin, M. Yu. 1978. ‘The cellular composition of the digestive gland of White Sea lamellibranchs. Trudy Petergof. biol. in-ta LGU (Lenin- grad State University), No. 26: 132-144 (ES) [Morphological and histological studies were undertaken on Mya arenaria, M. truncata, Cyprina islandica, Cardium ciliatum, Serri- pes gronlandicus, Musculus discors and Modiolus modiolus. A great deal of similarity was discovered in the histological structure of their digestive glands] RomanENKo, V. D., O. M. Arsan, V. D. Sotomatina, S. P Vesev’sxn, V. P Pryapxo « N. Yu. YEVrUSHENKO 1977. Tonic exchange and the role of calcium in Anodonta cyg- nea. Gidrobiol. zh. 13 (5): 115-119 Sxuv’sxn, I. A., N. B. Prvovarova, T. I. Ivanova, V. G. LEonT’EV, I. V. Burovina & A. E Feporov 1977. The adaptation of neuro-muscular cells in mussels to de- creases in salinity. Vlad.: 129-130 Page 398 STADNICHENKO, A. P 1978. Mollusks of the family Sphaeriidae in the Crimea. Report I. Vestn. zoologii 1: 67 - 73 [4 species were found in 40 stations from Simferopol to Feodosia: Euglesa casertana, E. personata, Musculium lacustre v. major, and E. henslowana v. inappendiculata, the last being new to the Crimea. The first 3 are rheophilic while the last prefers slow water and is found only in Salgira Creek at Pereval’noye] SVESHNIKOV, V. A. 1977. Aspects of the life cycle of the Far Eastern mytilid species Crenomytilus grayana. Dokl. AN SSSR 226 (4): 1028 - 1031 [Interacting periods of the life cycle may be recognized: The pelagic larvae are attracted to the benthic adults, settle and at- tach selectively by their byssal threads; initially the immatures live in the lower tier, sheltered somewhat from predators and grow rapidly; subsequently growth is hindered as competition increases for space and resources] VarAKsyYN, A. A. 1977. On the neurosecretions of the mytilid (Crenomytilus gray- anus) and the pectinid (Patinopecten yessoensis). Biol. Morya. No. 4: 58-65 (ES) [Morphometric and cytological investigations show that neurosecre- tory activity in the cerebropleural, visceral and pedal ganglia of the males and females of this species alters during the course of the year, presumably reflecting a correlation with sexual activity] Vasiu’Eva, V. S. 1978. Heat retention of cells in the upper littoral sand dwelling infaunal bivalves of Peter the Great Bay (Sea of Japan). Biol. Morya. No. 1: 53-57 [8 species are dealt with. Heat retention in these bottom dwelling, infaunal invertebrates depends on their vertical and geographical distribution as well as on the ambient temperature] YAKovLev, Yu. M. 1978. The reproductive cycle of the giant oyster (Crassostrea gigas) in the Sea of Japan. Biol. Morya, No. 3: 85 - 87 (ES) [Histological study of gametogenesis shows, during the course of the year, specific seasonal stages in the gonad: spawning, post- spawning, reduction, growth, maturation and prespawning] Yavnoy, S. V. 1977. Secretion of growth layers in the internal structure of the shell in some species of the Mactridae. Vlad.: 156 - 157 [Spisula sachalinensis, S. voyi, and Mactra sulcalaria were studied; the outer shell layer is cross-lamellar and has 2 sublayers; the inner shell layer is complex-cross-lamellar; growth intervals, for as short a time as a week, can be detected] YeEvseEEy, G. A. 1975. Characteristics of the ontogenesis of the pallial smus in Macoma balthica. SID, No. 4: 75 - 82 [Slight differences in the configurations of the pallial sinus, apparent- ly correlated with salinity, separate western (Arctic-Baltic) and eastern (Pacific) populations of Macoma balthica] THE VELIGER Vol. 22; No. 4 1976. The origin of Vostok (East) Bay in the Sea of Japan and the history of its bivalve fauna. SID, No. 5: 23 - 62 [The formation of Vostok Bay began 8000-9000 years BP during a cold boreal period when 22 species of bivalves, mostly cold water forms, were found there. Postglacial transgression (8000-2000 years BP) found 45-49 species with the % of cold water species declining. Greatest diversity occurred during the last transgression when the sea level was up to 1 m higher than at present and 60 bi- valve species were present, most of them warm water elements] Zorina, I. P. 1978. New bivalve species of the Gulf of Tonkin (South China Sea). Tr. zool. inst. AN SSSR 61: 193 - 203 [From the region of the delta of the Red River, Fai-Tsi-Long Archi- pelago and Hainan Island are described as new with figures, di- mensions, distributional and ecological data: Phacoides scarlatoi, Pillucina vietnamica, Codakia golikovi, Pitar levis, P. sulcata, Dip- lodonta gurjanovae, Raeta lactea, Moerella fragilis, and Gari tonkinensts] CEPHALOPODA Buruxovskn, R. N., G. V. Zuev, Cu. M. NIGMATULLIN & M. A. TsyMBAL 1977. A methodical basis for recognizing the extent of maturi- ty in the female reproductive system of squids, using Stheno- teuthis pteropus as an example. ZZ 56 (12): 1781 - 1791 [Several stages of the continuous process of maturation are recog- nized and suggested to be typical of all squids. These are separable mainly into those associated with oogenesis along with the develop- ment of accessory reproductive organs, and those connected with the accumulation of mature eggs in the oviduct] Dusinina, T. S. 1977. Characteristics of the morphology of the larvae of Onych- oteuthidae in the tropical Atlantic. VNK, pp. 34 - 35 [The larvae of 4 species are described, falling into 2 types: 1) the more primitive Onykia including Onykia and Moroteuthis with mantle length of 8mm; and 2) the more advanced Onychoteuthis, including Onychoteuthis and Ancistroteuthis with mantle lengths between 13-15mm. Allometric growth patterns are noted in the arms, tentacles, and penis] Nests, K. N. & G. A. SHEvTsov 1977. Preliminary information on the abyssal cephalopods in the Sea of Okhotsk. Biol. Morya, No. 5: 76-77 (ES) [In bottom dredgings of up to 3400m in June, 1975, 7 specimens of 5 species were taken: Gonatopsis borealis, G. okutani, Grimpo- teuthis albatrossi, Belonella borealis, Benthoctopus profundorum. All specimens, except those of the first species, were mature or matur- ing, their dimensions close to or exceeding the maximum reported. It was noted that Belonella pacifica pacifica is a synonym of B. belone and that B. p. borealis is an independent species] Vol. 22; No. 4 THE VELIGER Page 399 Nesis, K. N. 1978. The conference on the study of cephalopods. Kaliningrad, October 1976. Biol. Morya, No. 1: 83 - 84 [Summary of papers presented at meeting] 1978. The subfamily Ancistrocheirinae (Enoploteuthidae). ZZ 57 (3): 446-450 (ES) [The subfamily previously included Thelidioteuthis alessandrinit, known from a larvae and juvenile, and Ancistrocheirus lesueuri, known only from adults. Recently collected specimens show that Thelidioteuthis is a junior synonym of Ancistrocheirus though both alessandriniu and lesueuri are good species, the former being cosmo- politan in the tropics; the range of the latter is unknown] NicmaTuLun, Cu. M. 1977. Relationships and ecological structures in squids of the family Ommastrephidae. VNK, pp. 55 - 56 [Taxonomic characters grade from most primitive in the subfamily Tllicinae, to intermediate in Todarodinae and Ornithoteuthinae to most advanced in Ommastrephinae. Ecological specializations par- allel this trend: more primitive representatives are sublittoral, most- ly partly benthic forms which lay eggs on the bottom, intermediate forms are found mostly over the slope, and the most advanced, or evolutionarily youngest, include oceanic forms with pelagic eggs and considerable development of photophores] Petrov, O. A. 1977. On the fertility of the New Zealand squid, Notodarus sloanit sloani. VNK: 63 - 64 [The gonads of 40 females, collected in February-April, were ex- amined. 3-4 generations of eggs, from 0.1- 1.5mm in diameter, were discovered in the ovaries in various phases. Most oocytes were on the surface of the ovary, spherical or oval in shape, and from yellowish to amber in color. Although no clear relationship between the size of the female and the diameter of the ova was noted, there was a direct correlation between the size of the female and the number of ova: as the length of the mantle increased to 1 cm, the quantity of eggs grew to an average of 80000 each; females larger than 38cm had as many as 550 000] Rozeneart, E. V, S. P. SHevrsova « L. M. EpSHTEIN 1977. ‘The properties of cholinesterase in the optic ganglia of the Teuthidae [sic]. VNK: 76-77 [A large amount of cholinesterase is contained in the tissues of the optic ganglia, and its properties vary, as a correlate of the life his- tory of the animal, in different species, such as Ommastrephes bart- rami and Illex illecebrosus] Suevtsova, S. P, A. P. BrestTxkIn, K. N. Nesis « E. V. RoZENGART 1977. On the identity of the affinities of cholinesterase in the optic ganglia of Ommastrephes bartrami from the South At- lantic and from the Great Australian Bight. Okeanologia 17 (6): 1102-1106 ZaLyYGALIN, V. P, G. V. ZuEv « Cu. M. NicMATULLIN 1977. Characteristics of spermatophore production and male fertility in Sthenoteuthis pteropus (St.). VNK: 37 - 38 [A close correlation between mantle length and spermatophore length (of spermatophores in the spermatophoric sac) was estab- lished in a sample of over 70 male specimens; evidence for size dimorphism in spermatophores in the mantle cavity was presented] Zuev, G. V.& Cu. M. NicMATULLIN 1977. Basic elements of the internal structure of Sthenoteuthis pteropus (St.) in the northern part of the tropical Atlantic. VNK: 38-40 [Variable dimensions of the mantle are correlated with geographic positions and climatic conditions] POLY PLACOPHORA Smenxo, B. I. 1976. Chitons of East Bay in the Sea of Japan. SID, No. 5: 87 - 92 [17 species are recognized with the following geographical affinities: 64% low-Boreal, 24% subtropical-low Boreal, and 12% pan-Bore- al] CALIFORNIA Matacozoo.ocicat Society, Inc. is a non-profit educational corporation (Articles of In- corporation No. 463389 were filed January 6, 1964 in the office of the Secretary of State). The Society publishes a scientific quarterly, the VELIGER. Donations to the Society are used to pay a part of the production costs and thus to keep the subscription rate at a minimum. Donors may designate the Fund to which their contribution is to be credited: Operating Fund (available for current production) ; Savings Fund (available only for specified purposes, such as publication of especially long and signi- ficant papers); Endowment Fund (the income from which is available. The principal is irrevocably dedicated to scientific and educational purposes). Unassigned dona- tions will be used according to greatest need. Contributions to the C. M.S., Inc. are deductible by donors as provided in section 170 of the Internal Revenue Code (for Federal income tax purposes). Bequests, lega- cies, gifts, devises are deductible for Federal estate and gift tax purposes under section 2055, 2106, and 2522 of the Code. The Treasurer of the C. M. S., Inc. will issue suitable receipts which may be used by Donors to substan- tiate their respective tax deductions. Page 400 Sea Star Predation on Rock-Boring Bivalves BY E. C. HADERLIE Department of Oceanography Naval Postgraduate School, Monterey, California 93940 THE MonTEREY Formation shale which forms extensive outcrops under kelp beds in southern Monterey Bay is penetrated by several species of boring bivalves of the families Mytilidae and Pholadidae. Divers examining these shale outcrops can see the protruding siphons of many of these borers and in some cases can make positive identification of the species by siphon morphology (HapER- LIE et al., 1974). Most borers retract the siphons into the burrow when touched and a few can retract the siphons within the shell. None have methods of blocking the bur- row entrance once the siphons have been withdrawn. The sea star Pisaster brevispinus (Stimpson, 1857) is a common predator in the shallow subtidal water in Mon- terey Bay. It is seen on soft bottoms as well as on the exposed shale reefs. On soft bottoms the sea star feeds on a variety of buried bivalves by digging down to them or by extending the central tube feet downward into the sediment for as much as 17 cm and lifting the clams up to the mouth (VAN VELDHUIZEN & PuiLups, 1978). On rocky reefs the sea star often is seen over pholad and bor- ing mytilid burrows, and when examined has its stomach everted and pressed into the opening of the burrow. When large pieces of shale are excavated by divers, or broken free by a heavy dredge, and brought ashore for examination, it is found that at least 50% and sometimes 100% of the bivalve boreholes are empty or contain only the valves of dead borers. Some of these valves are from young immature animals. Direct evidence of predation by Pisaster brevispinus on boring bivalves has been obtained recently. In an attempt to determine the growth rates of boring bivalves, a num- ber of animals in the young, active boring stage have been removed from their natural burrows in shale, identified and measured, then re-introduced into “artificial” bur- rows drilled into shale similar to that from which the animals came. Species used in these experiments include Lithophaga plumula Hanley, 1843; Adula falcata THE VELIGER Vol. 22; No. 4 (Gould, 1851) ; Chaceia ovoidea (Gould, 1851) ; Neta- stoma rostrata (Valenciennes, 1846); Parapholas califor- nica (Conrad, 1837); Penitella penita (Conrad, 1837) and Penitella gabbi (Tryon, 1863). The experimental burrows are cylindrical (not the normal pear-shape) and fit the bivalves snugly. The open end of the burrow of each is plugged with a perforated stopper allowing the siphons to extend out into the water but preventing the animal from falling out of the burrow. The panels of shale containing the borers are then put in racks and placed on the bottom in approximately 1om of water; each month the shale panels are recovered and x-rayed to determine growth rates of the borers. The radiation does not appear to harm the experimental animals for they remain alive and continue to bore at the same rate as control animals. During the past year mortality among all of these experimental animals has been high but the cause was unknown until racks were recovered where several Pisaster brevispinus were found on the shale panels in the process of feeding on representatives of all the borers named above. Careful observation revealed how the bivalves were attacked. In some cases the sea star had extended its central tube feet through the hole in the stopper, attached the tube feet to the shell of the borer, then pulled the bivalve up against the stopper until this plug popped out and the borer (and stopper!) could be enveloped by the sea star’s stomach. When this tactic failed, presumably because the stopper could not be dis- lodged, Pisaster everted its stomach and inserted it through the hole in the stopper into the burrow. In these cases the stomach did not appear to enfold the borer, but digestive enzymes apparently were secreted over the borer and the bivalve digested in its own burrow. In nature, where the shape of the burrow precludes removal of the animal in one piece, this latter method of feeding is perhaps the one used and would account for the clean, empty shell valves found in many burrows. The experiments are continuing, but now the shale panels are housed in cages that exclude predatory sea stars. Literature Cited Haverue, Evceng Cuiinron, J.C. Mzeitor,C.S. Minter III « G.C. Boorn 1974. The sublittoral benthic fauna and flora off Del Monte Beach, Monterey, California. The Veliger 17 (2): 185-204; 3 plts.; 9 text figs. (1 October 1974) Van VeELDHUIZEN, H. D. « D. W. PHILuIPs 1978. Prey capture by Pisaster brevispinus (Asteroidea: Echinoderm- ata) on soft substrate. Mar. Biol. 48: 89-97 Vol. 22; No. 4 Generous Donation by the Conchological Club of Southern California On the same day that the January issue was delivered to us by the printer, a very generous contribution to the Veliger Endowment Fund from our friends, the Concho- logical Club of Southern California, arrived in the mail. We are very grateful for the continued support, which serves a twofold purpose: firstly, it helps us in our effort to keep the cost of the journal at its — in these days of galloping inflation — unusually low price; and secondly, it encourages us in what we hope to be a worthwhile undertaking. ERRATUM Mrs. Helen DuShane has informed us that Dr. Joseph Rosewater of the National Museum of Natural History, Washington, D. C., has advised her that the museum num- ber of Epitonium appressicostatum is in error. Lines 14 ff. in the second column on p. 107 of volume 22 of our jour- nal should read: “|... 2 specimens, as Epitonium appressicostatum (USN M 678703), ...” — not (USNM 59334). Mrs. DuShane regrets the error. Publication Date of THE VELIGER THE PUBLICATION DATE of The Veliger is the date printed on the index page; this applies even if the date falls on a legal holiday or on a Saturday or Sunday, days when the U.S. Postal Service does not expedite second class mail matter. That the printed date is the actual date of pub- lication under the rules of the International Commission on Zoological Nomenclature is based on the following facts: 1) The journal is delivered to the Post Office on the first day of each quarter, ready for dispatch; 2) at least three copies are mailed either as first class items or THE VELIGER Page 401 by air mail; 3) about 20 copies are delivered in person to the mail boxes or to the offices of members in the Berkeley area; 4) two copies are delivered to the re- ceiving department of the General Library of the Univer- sity of California in Berkeley. Thus our publication is available in the meaning of the Code of the ICZN. The printed publication date, therefore, may be relied upon for purposes of establishing priority of new taxa. REGARDING POSTAL SERVICE We are much disturbed by the steadily increasing num- ber of premature claims for supposedly “missing” issues of our journal. Since we have announced on numerous occasions that our journal is mailed on the dates printed in the issues, 7.¢e., number 1 on July 1, number 2 on October 1, number 3 on January 1 and number 4 on April 1 of each volume year, it is unreasonable to expect delivery of the issues earlier than at least one week after these dates; however, a much longer time must be allowed for delivery to addresses at various distances from Berkeley. Thus, for example, a two weeks lapse is not unusual for as short distances as 500km; and up to 3 and 4 months must be counted on for addresses in the Far East and in Africa. We are faced with the alternative of not replying to what we must consider premature claims or, if the trend con- tinues, we must increase our subscription rates to cover these additional expenses. Our past efforts at keeping the subscription rate as low as possible are, we believe, suffi- cient evidence that we simply cannot afford any other course of action. The postal service causes us enough financial losses. Therefore, we urgently request that before a claim is made, the time schedule be carefully checked. We are grateful for the understanding of this difficult situation shown by many librarians and will be grateful to those who, heretofore being perhaps eager to make sure that the library receives what is coming to it, will exercise a little patience. Claims for defective or missing pages must reach us within 60 days from the publication date. We will not respond to claims for missing issues made less than 30 days by domestic addressees, or less than 60 days by foreign addressees after the publication date of our journal issues. Your harassed Editor. We are willing to accept requests for expediting our journal via AIR MAIL; however, in that case we must ask for an additional payment of US$8.00 in all cases where the Veliger goes to domestic addresses, and a depos- it of US$18.00 for all foreign addresses (including PUAS). Page 402 THE VELIGER Vol. 22; No. 4 Of course, we will carry forward as a credit toward the postage charges of the following year any amount over the actually required postage charges. We have read in January this year that at a meeting of the U. P. U. (Universal Postal Union) the decison was made that all international postage rates will be doubled in 1981; that, however, in view of the very low rates in effect in the U. S. A. (due to the depreciation of the dol- lar), this country would be allowed to increase its foreign rates in October 1980. We have, however, not been able to get any confirmation from the postal authorities in this area. If it turns out later to be true, we may be forced to ask our members and subscribers in foreign countries for a supplemental payment to cover the increase. This will be especially true in the cases where the journal is being sent by Air Mail. Moving? If your address is changed it will be important to notify us of the new address at least six weeks before the effective date, and not less than six weeks before our regular mailing dates. Because of a number of drastic changes in the regulations affecting second class mailing, there is now a sizeable charge to us on the returned copies as well as for our remailing to the new address. We are forced to ask our members and subscribers for reimbursement of these charges; further, because of increased costs in connection with the new mailing plate, we also must ask for reimbursement of that expense. The following charges must be made: change of address - $1.- change of address and re-mailing of a returned issue — $2.75 minimum, but not more than actual cost to us. We must emphasize that these charges cover only our actual expenses and do not include compensation for the extra work involved in re-packing and re-mailing returned copies. At present we are charged a minimum fee of $15.00 on each order for new addressograph plates. For this rea- son we hold off on our order until 6 weeks before mailing time, the very last moment possible. If, for any reason, a member or subscriber is unable to notify us in time and also is unable to make the proper arrangement with the Post Office for forwarding our journal, we will accept a notice of change of address, accompanied by the proper fee and a typed new address on a gummed label as late as 10 days before mailing time. We regret that we are absolutely unable to accept orders for changes of address on any other basis. In view of the probable further cur- tailment in the services provided by the Postal Service, we expect that before long we may have to increase these time intervals. Endowment Fund In the face of continuous rises in the costs of printing and labor, the income from the Endowment Fund would materially aid in avoiding the need for repeated upward adjustments of the membership dues of the Society. It is the stated aim of the Society to disseminate new infor- mation in the field of malacology and conchology as widely as possible at the lowest cost possible. At a Regular Membership meeting of the Society in No- vember 1968 a policy was adopted which, it is hoped, will assist in building up the Endowment Fund of the Society. An issue of the journal will be designated as a Memorial Issue in honor of a person from whose estate the sum of $5000.- or more has been paid to the Veliger Endowment Fund. If the bequest is $25 000.- or more, an entire volume will be dedicated to the memory of the decedent. WE CALL THE ATTENTION orf our foreign correspondents to the fact that bank drafts or checks on banks other than American banks are subject to a collection charge and that such remittances cannot be accepted as payment in full, unless sufficient overage is provided. Depending on the American banks on which drafts are made, such charges vary from a flat fee of $1.- to a percentage of the value of the draft, going as high as 33%. Therefore we recommend either International Postal Money Orders or bank drafts on the Berkeley Branch of United California Bank in Berkeley, California. This institution has agreed to honor such drafts without charge. UNESCO coupons are N OT acceptable except as indicated elsewhere in this section. Regarding UNESCO Coupons We are unable to accept UNESCO coupons in payment, except at a charge of $4.25 (to reimburse us for the ex- penses involved in redeeming them) and at $0.95 per $1.- face value of the coupons (the amount that we will receive in exchange for the coupons). We regret that these char- Vol. 22; No. 4 ges must be passed on to our correspondents; however, our subscription rates and other charges are so low that we are absolutely unable to absorb additional expenses. Sale of C. M. S. Publications: Effective January 1, 1978, all back volumes still in print, both paper covered and cloth bound, will be available only from Mr. Arthur C. West, P.O. Box 730, Oakhurst, CA(alifornia) 93644. The same applies to the supple- ments still in print with certain exceptions (see below). Prices of available items may be obtained by applying to Mr. West at the address given above. Volumes 1 through 8 and to through 12 are out of print. Supplements not available from Mr. West are as fol- lows: Supplements to vol. 7 (Glossary) and 15 (Ovulidae) are sold by “The Shell Cabinet,’ P O. Box 29, Falls Church, VI(rginia) 22046; supplement to vol. 18 (Chitons) is available from “The Secretary, Hopkins Marine Station, Pacific Grove, CA(lifornia) 93950. Supplements Supplement to Volume 3: [Part 1: Opisthobranch Mollusks of California by Prof. Ernst Marcus; Part 2: The Anaspidea of California by Prof R. Beeman, and The Thecosomata and Gymnosomata of the Cali- fornia Current by Prof. John A. McGowan] Supplement to Volume 6: out of print. Supplement to Volume 7: available again; see announce- ment elsewhere in this issue. Supplement to Volume 11: [The Biology of Acmaea by Prof. D. P. Aszorr et al., ed.} Supplement to Volume 14: [The Northwest American Tellinidae by Dr. E. V. Coan] Supplement to Volume 16: [The Panamic-Galapagan Epitoniidae by Mrs. Helen DuShane] [Growth Rates, Depth Preference and Ecological Succes- sion of Some Sessile Marine Invertebrates in Monterey Harbor by Dr. E. C. Haderlie] Supplement to Volume 17: Our stock of this supplement is exhausted. Copies may be obtained by applying to Dr. E. C. Haderlie, U. S. Naval Post-Graduate School, Mon- terey, CA (lifornia) 93940. THE VELIGER Page 403 WE ARE PLEASED to announce that an agreement has been entered into by the California Malacozoological Society, Inc. with Mr. Steven J. Long for the production and sale of microfiche reproductions of all out-of-print editions of the publications of the Society. The microfiches are available as negative films (printed matter ap- pearing white on black background), 105mm X 148mm and can be supplied immediately. The following is a list of items now ready: Volume 1: $1.50 Volume 6: $4.50 Volume 2: $3.00 Volume 7: $6.00 Volume 3: $3.00 Volume 8: $6.00 Volume 4: $4.50 Volume 10: $9.00 Volume 5: $4.50 Volume 11: $9.00 Volume 12: $9.00 Supplement to Volume 6: $1.50; to Volume 18: $3.00 California residents please add the appropriate amount for sales tax to the prices indicated. Please, send your order, with check payable to Opistho- branch Newsletter, to Mr. Steven J. Long, 792 Laurie Avenue, Santa Clara, CA 95050. Volumes and Supplements not listed as available in microfiche form are still available in original edition from Mr. Arthur C. West, P.O. Box 730, Oakhurst, CA(lifornia) 93644. Orders should be sent directly to Mr. West. Single Copies of ““The Veliger”’: We have on hand some individual copies of earlier issues of our journal and are preparing a list of the various issues available with the prices. Some issues are present in only one or two copies, while others may be present in 10 or more copies. As we are anxious to make room, we will offer these numbers at an exceptionally low price. This list may be obtained by sending a self-addressed, stamped envelope to the Veliger, 1584 Milvia Street, Berkeley, CA(lifornia) 94709. Foreign correspondents should en- close one international postal reply coupon. Requests for the list, for which return postage is not provided, will be ignored. Membership open to individuals only - no institutional or society memberships. Please send for membership ap- plication forms to the Manager or the Editor. Membership renewals are due on or before April 15 each year. If renewal payments are made after April 15 Page 404 but before March 15 of the following year, there will be a re-instatement fee of $1.-. Members whose dues pay- ments (including the re-instatement fee) have not been received by the latter date, will be dropped from the rolls of the Society. They may rejoin by paying a new initiation fee. The volume(s) published during the time a member was in arrears may be purchased, if still available, at the regular full volume price plus applicable handling charges. Backnumbers of the current volume will be mailed to new subscribers, as well as to those who renew late, on the first postal working day of the month following receipt of the remittance. The same policy applies to new members. Tue VELIGER is not available on exchange from the Cali- fornia Malacozoological Society, Inc. Requests for re- prints should be addressed directly to the authors con- cerned. We do not maintain stocks of reprints and also cannot undertake to forward requests for reprints to the author(s) concerned. OPEN POSITION ror MALACOLOGIST FreELD Museum oF NaTurAL History announces a search for an Assistant Curator of Invertebrates in the Depart- ment of Zoology. Appointment of the successful candi- date, who must have a Ph. D. or substantially completed all degree requirements, will take place about January 1, 1981, for a three year term. Preference will be given to workers on the systematics of freshwater or marine mol- lusks. Research will be of the candidate’s choice, with 50% time available. Further information can be obtained from: Dr. Robert K. Johnson, Chairman, Invertebrates Search Committee, Divison of Fishes, Field Museum of Natural History, Roosevelt Road at Lakeshoe Drive, Chicago, Illinois 60605, U.S. A. The search will close in September 1980. Field Museum of Natural History is an Equal Opportu- nity Employer. IMPORTANT NOTICE If the address sheet of this issue is PINK, it is to indicate that your dues remittance had not arrived at the time the mailing was prepared (2. e., by March 1, 1980). We wish to take this opportunity to remind our Members that a reinstatement fee of one dollar becomes due if member- ship renewals have not been received by C.M.S., Inc. THE VELIGER Vol. 22; No. 4 by April 15, 1980. However, in view of the unreliability of the postal service, members should not be alarmed by this notice as their remittances may be received between the first of March and the date of mailing this issue on April 1. From overseas addresses we must allow a minimum of 6 weeks for surface mail. On the other hand, it is possible that the envelope and dues notice enclosed between pages 292 and 293 of the January issue have escaped your atten- tion. If so, now is the time to use them to avoid interrup- tion in the delivery of this periodical. BOOKS, PERIODICALS, PAMPHLETS Note on a Panamic Province Cone ‘Two papers in a new conchological publication, The Shell Collector, discuss the synonymy of Conus brunneus Wood, 1828. Under the title “The Conus bartscht/brun- neus controversy now over,” Alex Kerstitch shows that the two forms are separable and that they occur in slightly different habitats. The spire of C. bartschi is smooth, whereas that of C. brunneus has 2 to 6 incised grooves. There is also a difference in range, and C. andrangae Schwengel, 1955, is shown to be a synonym of C. bartschi Hanna and Strong, 1949. A companion note by John K. Tucker in this journal reinforces Kerstitch’s view, with further illustrations. KerstircH, Axex [title cited above]. The Shell Collector, no. 2, pp. 38-41; 4 color figures. 1979. Tucker, JoHN K. “Conus bartschi and Conus brunneus: two closely related Eastern Pacific cone shells.” zbid., PP. 42-43; 13 figs. A. Myra Keen Malacological Review P.O. Box 420, Whitmore Lake, Michigan 48189, U.S.A. vol. 12 (1-2), iv+186 pp.; illust. As most of its predecessors, this volume contains several original research articles, followed by several “Brief Com- munications.” On pp. 97 - 104 are found announcements Vol. 22; No. 4 of past and future events of interest to malacologists. And on page 105 starts, “Contents of Current Malacological Periodicals.” Among the research articles, the one by R. I. Johnson, entitled “The types of Unionacea (Mollusca: Bivalvia) in the Museum of Zoology, The University of Michigan,” is of special interest to taxonomists dealing with this fresh- water group. In the interest of stabilizing taxonomy, the whereabouts of extant type specimens is of singular im- portance. Many similar lists have been published either for certain groups or for material described by a particular worker. For reasons this reviewer does not understand, the price of this work to individual subscribers is still only $10.- ($10.50 for subscribers in all foreign countries). R. Stohler A Field Guide to the Land Snails of Britain and North-west Europe by M. P Kerney « R. A. D. Cameron, illustated by Gorpon Rirey. 288 pp.; 649 illustrations (408 in color on 24 plates) ; 392 distribution maps. Collins, Publishers, 14 St. James Place, London, SW1A 1PS. £5.50. 7 June 1979 This remarkably well organized, relatively small (pock- et size) book is one this reviewer wishes he had had available in his younger days when he was studying Swiss mollusks. Now, some 50+ years later, it is pleasant to renew acquaintance with some “old friends.” The introductory pages, as is not uncommon in books of this type, contain hints on the use of the book, explain the format adopted, discuss biology, habitat and related topics; a brief section deals with collecting and identi- fying land snails. A glossary of “shell terms” concludes the “preamble.” A systematic check-list of all species treated in this book occupies pages 39 to 47. We are especially pleased that the authors list in this part in what we consider the only fully correct manner; that is, complete with author and year of original description. As far as memory will serve, the color figures appear true to life. The quality of the line drawings is high. The text is clear and easy to understand. An interesting detail of some of the distributional maps is the indication that the particular species is spreading or receding, as the case may be. If we give the impression that we like the book — all we have to say is: we do. R. Stohler THE VELIGER Page 405 Indexes to the Nautilus Geographical (Vols. 1-90) and Scientific Names (Vols. 61-90). Edited by R. Tucker Azpsorr. American Malaco- logists, Inc.., Publ. P O. Box 2255, Melbourne, Florida 32901. 238 pp.; US$24.00. (26 October 1979) The geographical index of the first 90 volumes of the Nautilus was compiled by Mrs. Margaret Crockett Tes- key, who, for many years, was the Secretary of the Ameri- can Malacological Union. This index occupies 80 pages. It is arranged alphabetically by country, subdivided, in the case of the United States, by individual States. A fur- ther subdividing has been made under each of the catego- ries mentioned into ‘Land, Freshwater,’ ‘Marine’ and ‘Fossil.’ Under these final subheadings, the titles are ar- ranged strictly alphabetically. The index to the scientific names in the Nautilus, com- piled by Mrs. Louise Burrell Hastings, occupies the final 158 pages of the volume. It is stated in a sort of Foreword that this index includes 24 000 entries. We did not check the accuracy of this statement! This list is also strictly alphabetical. Each species is listed under its trivial name and also under the genus name. In sampling the book, we did not detect any obvious typographical errors. But even if there are such errors or possible inadvertent omissions, they will not materially detract from the value of the compilation. R. Stohler Pacific Coast Subtidal Marine Invertebrates by DantEt W. GoTsHALL & LAURENCE L. LAuRENT. Sea Challengers, Los Osos, California 93402. 112 pp.; 160 color illustrations on 40 plates; 2 pp. line drawings; 5 maps. US$10.87 paperback; $14.20 hard cover, mailing and handling included. 1979 This book will be welcomed by the diving enthusiasts, who, with the aid of modern SCUBA equipment, can explore the shallower portions of the nearshore waters of the Pacific Coast of North America. This area is still a very rich one, in spite of the seemingly determined efforts of modern man to destroy it. We can think of no more powerful way of convincing people that the conser- vation and protection of this rather fragile environment is worth every effort than to have as many people as possible visit and study the subtidal area. Such visits will reveal an unexpected wealth of fas- cinating and, sometimes, bizarre forms of living beings, Page 406 THE VELIGER sometimes startlingly colorful and yet, at the same time, often difficult to see because of their cryptic habit. The well-printed reproductions of masterfully taken color photographs in this book will satisfy the curiosity not only of the novice diver, but also that of a more experi- enced diver. We must applaud the authors for their choice of “shots” for reproduction, although we think that many users would like to have several dozen other species in- cluded. There are “only” 44 molluscan species represen- ted out of a possible 700 or 800 and the especially color- ful nudibranchs are represented “only” by 14 species. It must have been a painful job to eliminate dozens and dozens of excellent shots from consideration for inclusion. R. Stohler Cephalopods from Deepwater Dumpsite 106 (Western Atlantic): Vertical Distribution and Seasonal Abundance by C. C. Lu & Cryne E E. Roper. Smithsonian Contri- butions to Zoology, No. 288. 36 pp.; 11 text figs.; 5 tables (2 of these in the appendix). 1979 Dumpsite 106 is an area about 160km east of Cape Henlopen, Delaware. In size it is 20’ by 30’. Water depth ranges from 1550m to 2750m. “The dumpsite is used for the disposal of acid waste, industrial chemicals and radio- active wastes, ....” The objectives of the study were to determine species composition, vertical distribution, seasonal occurrence, relative abundance, spawning tendencies, and relationship to water types of 36 species of cephalopods (octopus and squids). The results obtained on 4 cruises are detailed in the 2 tables in the appendix, occupying 11 pages. R. Stohler Vol. 22; No. 4 THE VELIGER is open to original papers pertaining to any problem concerned with mollusks. This is meant to make facilities available for publication of original articles from a wide field of endeavor. Papers dealing with anatomical, cytological, distributior.al, ecological, histological, morphological, phys- iological, taxonomic, etc., aspects of marine, freshwater or terrestrial mollusks from any region, will be considered. Even topics only indi- rectly concerned with mollusks may be acceptable. In the unlikely event that space considerations make limitations necessary, papers dealing with mollusks from the Pacific region will be given priority. However, in this case the term “Pacific region” is to be most liberally interpreted. It is the editorial policy to preserve the individualistic writing style of the author; therefore any editorial changes in a manuscript will be sub- mitted to the author for his approval, before going to press. Short articles containing descriptions of new species or lesser taxa will be given preferential treatment in the speed of publication provided that arrangements have been made by the author for depositing the holotype with a recognized public Museum. Museum numbers of the type specimens must be included in the manuscript. Type localities must be defined as accurately as possible, with geographical longitudes and latitudes added. Short original papers, not exceeding 500 words, will be published in the column “NOTES & NEWS"; in this column will also appear notices of meetings of the American Malacological Union, as well as news items which are deemed of interest to our subscribers in general. Articles on “METHODS & TECHNIQUES” will be considered for publication in another column, provided that the information is complete and tech- niques and methods are capable of duplication by anyone carefully fol- lowing the description given. Such articles should be mainly original and deal with collecting, preparing, maintaining, studying, photo- graphing, etc., of mollusks or other invertebrates. A third column, en- titled “INFORMATION DESK,” will contain articles dealing with any problem pertaining to collecting, identifying, etc., in short, problems encountered by our readers. In contrast to other contributions, articles in this column do not necessarily contain new and original materials. Questions to the editor, which can be answered in this column, are in- vited. The column “BOOKS, PERIODICALS, PAMPHLETS?” will attempt to bring reviews of new publications to the attention of our readers. Also, new timely articles may be listed by title only, if this is deemed expedient. Manuscripts should be typed in final form on a high grade white paper, 812” by 11”, double spaced and accompanied by a carbon copy. A pamphlet with detailed suggestions for preparing manuscripts intended for publication in THE VELIGER is available to authors upon request. A self-addressed envelope, sufficiently large to accom- modate the pamphlet (which measures 5/2” by 8/2”), with double first class postage, should be sent with the request to the Editor. EDITORIAL BOARD Dr. Donatp P. Assort, Professor of Biology Hopkins Marine Station of Stanford University Dr. Warren O. Appicott, Research Geologist, U. S. Geological Survey, Menlo Park, California, and Consulting Professor of Paleontology, Stanford University Dr. Hans Bertscu, Curator of Marine Invertebrates San Diego Museum of Natural History Dr. Jerry DononvE, Professor of Chemistry University of Pennsylvania, Philadelphia, and Research Associate in the Allan Hancock Foundation University of Southern California, Los Angeles Dr. J. Wyatt Duruam, Professor of Paleontology Emeritus University of California, Berkeley, California Dr. Capet Hanp, Professor of Zoology and Director, Bodega Marine Laboratory University of California, Berkeley, California Dr. Carore S. Hickman, Assistant Professor of Paleontology University of California, Berkeley, California Dr. A. Myra KEEN, Professor of Paleontology and Curator of Malacology, Emeritus Stanford University, Stanford, California Dr. Victor LoosanorF, Senior Biologist, Emeritus U.S. National Marine Fisheries Service EDITOR-IN-CHIEF Dr. Rupo.tr SToOHLER, Research Zoologist, Emeritus University of California, Berkeley, California Dr. Joun McGowan, Professor of Oceanography Scripps Institution of Oceanography, La Jolla University of California at San Diego Dr. Frank A. Pitetka, Professor of Zoology University of California, Berkeley, California Dr. RoBert Rosertson, Pilsbry Chair of Malacology Department of Malacology Academy of Natural Sciences of Philadelphia Dr. PETER U. Roppa, Chairman and Curator, Department of Geology California Academy of Sciences, San Francisco Dr. Ciype F. E. Roper, Curator Department of Invertebrate Zoology (Mollusca) National Museum of Natural History Washington, D. C. Dr. JupirH Terry SmitH, Visiting Scholar Department of Geology, Stanford University Stanford, California Dr. Ratpu I. Smiru, Professor of Zoology University of California, Berkeley, California Dr. Cuares R. STASEK, Bodega Bay Institute Bodega Bay, California Dr. T. E. THompson, Reader in Zoology University of Bristol, England ASSOCIATE EDITOR Mrs. Jean M. 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A Zz = wo & 3 & 5 =e rs) a 2 NVINOSHLINS S3/IYVve & se a Sos \HS¥Ay = \\\ SS = Ore > * Na 4 = = : a \ a= 1s : m . = Ssh wm "s = SMITHSONIAN INSTITU < g z= & NSS = S Ws WF ZN YS w D W 2 = = 3 z poatava rr be Aes = fa Awe (= ss oE =: + - y ra i= hy A 4 < os = 3) > — Wp — \oRabsr EVE C3 > WO EF i] 2 a Uj pile Ae (Mw ff 4 P \e st Ee ta is} 2 SS iS BRS m = LIWS S JlyVe EQYA 20 easy G Nekabye/ — Zz ” NOILNLILSNI” NVINOSH o in Up — “Sam 2 ee IAN INSTITUTION NOI = = Zz We a UIT LIBRARIES, SMITHSONIA ca AN Z ‘S = yy — Ssa1uY ior @ ITUT 2 a4 = \ 5 a Sue |BRARIES, SMITHSONIAN _ INST = ined a ; = NVINOSHLIWS SalYVYaIT_L ef hls ph ee Myf = 2 ON NOILOLILSNI ap) Zz ale ga ee = | wo ONIAN _INSTITUTION 2 Ss S "FYE BRARI BSSMuTHs = W q AS ax =| < Lee, S Wd 0G ez = Nass < c = tip a x. = < iS NN a 5 a ¢ = Saruyy: << 5 ta 3 BS = NOLLALILSNI AY NSH ENS a5 5 = Zz N” INSTITUTION NO Tipe 2) = Yi = 117” LIBRARIES~ SMITHSONIA Tas z = = at z = = > -NVINOSHLINS_ S31uvua Ee o fj, > = 2 F aps fe) ow = Gly, ay s x ie _ = Yt 4. > = a x Yy \ 2 = = Ui hy - a Cee alte NSTITUT > SS = z ae m Sei orrae SMITHSONIAN | INS = “Wve m = o uaia Lit a = - WN = HUWS S3I1YV Ze = => Wr, w "INSTITUTION NO z = iy = Ort AY Te ate INS g = = 5 ws = AN % Zz = wt, : : 2 NX 2 =z XS = Sie = Yip, & a + 2 =. aN = S Ny = J fs 9 3 E SS Fe A Qs WAS fe) by: Ao ote = = = S SHLINS S3I1YVY 1S = Wr 2 7 = z ON NOILALILSNI_ NVINO o Zz = IM >" “a INSTITUTI Zz Ww = = aS 2 SMITHSONIAN _ an n s “yunosuuis@s31uvual1-Lipraries® wn, ® zs = a Sab oO Zz Es Yd yp a "ee z 2 g : fy 3 < 5 E : E : & MO fos é : eau 2 2 =| 0 - ) a S_ SMITHSONIAN INST Ss = 2 LIBRARIE = eS S - Saluvudiy_ a = 2) ea = | NVINOSHLIWS iF fe) wo : NOILNLILSN z ce IAN_ INSTITUTION ie teas S ow = 2 HRARIES_ SMITHSONIAN _ S os = 2 =e _ S +o = 5 = 2 B rm B hy ed 7 ee) ie m = saiuvuE FY Gy a - = z i NLILSNI n a “Gj; — ” ™ a NOIL = = 3 Vif rm z SMITHSONIAN B = =a i My ~ NVINOSHLIWS z = = yy fg Ss D ha INLILSNI_ NVI % < = 2 Be a 9 Ly 2 | ef fy B z ? 2 “Wy = Sy 3 z ae ". 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