TELE. ee PLIGCER A Quarterly published by AO |( ISSN 0042-3211 CALIFORNIA MALACOZOOLOGICAL SOCIETY, IN@. Berkeley, California R. Stohler (1901-2000), Founding Editor Volume 48 November 2, 2006 CONTENTS New Late Cretaceous Mytilid and Tellinoidean Bivalves from California RUCEIARID) JL, SQUIRES, AINTO) ILOWWIBIUDA IRs SYNUIL cog oud Gules bio din eisic ooo a nieinis elcid cite Bin 121 A New Genus of Indo-West Pacific Turridae (Gastropoda: Prosobranchia) JNINTTONIO JXONTIAIEO) ANID) MUNGO) IMIORASSE 45's bolsid oo Beso 6 ob ooo clad so bon cle omar 136 Growth and Activity Patterns in a Backyard Population of the Banana Slug, Ariolimax colum- bianus ZANE KEG REARSON OLIVER PEARSON )AND PETER. RALPH ©... ./.)... 2552-25-50 5--s 143 Lower Eocene Gastropods from the El Bosque Formation, Central Chiapas, Mexico MARIA DEL CARMEN PERRILLIAT, JAVIER AVENDANO, FRANCISCO J. VEGA, AND JESUS SOLE.. 151 Redescription of Two Antarctic Species of Cuspidaria: C. concentrica Vhiele, 1912 and C. minima (Egorova, 1993) (Bivalvia: Cuspidariidae) IDIBEO G. ZBLANA ANID GIGI IGONRGNE, dco canadoudaaeodoes Dena enue OHM oe eee 170 Shedding Light onto the Genera (Mollusca: Nudibranchia) Kaloplocamus and Plocamopherus with Description of New Species Belonging to These Unique Bioluminescent Dorids MV ONINEAVALTESFAN DMI ERREN GE Vin GOSEINER™ oo dat nes uane diel geen ties ce ces 178 A New Species of Lipidochitona (Mollusca: Polyplacophora) from El Salvador CEDAR Il, CARGUANOS 9 oscobecd ode coe deca culg son eur ts 6 a UlOt ls Seana aie eee 206 » On the Occurrence of Rhomboidella prideaux (Leach, 1815) (Mollusca: Bivalvia: Mytilidae) in the Eastern Mediterranean BILAL OZTURK, JEAN-MAURICE POUTIERS, MESUT ONEN, AND ALPER DOGAN .........- 215 CONTENTS — Continued The Veliger (ISSN 0042-3211) is published quarterly in January, April, July, and October by the California Malacozoological Society, Inc., % Santa Barbara Museum of Natural History, 2559 Puesta del Sol Road, Santa Barbara, CA 93105. 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THE VELIGER © CMS, Inc., 2006 The Veliger 48(3):121—135 (November 2, 2006) New Late Cretaceous Mytilid and Tellinoidean Bivalves from California RICHARD L. SQUIRES Department of Geological Sciences, California State University, Northridge, California 91330-8266, USA AND LOUELLA R. SAUL Invertebrate Paleontology Section, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007, USA Abstract. One new subfamily, one new genus, and two new species of Late Cretaceous warm-water, shallow-marine bivalves from California are described and named. The mytilid Xenomytilus fons, gen. et sp. nov., from middle to uppermost Maastrichtian strata in central and southern California, represents the type genus of Xenomytilinae, subfamily nov. This new subfamily is comprised of Lycettia Cox, 1937, and Xenomytilus. Lycettia is an Old World Tethyan bivalve that ranges from the late Early Jurassic (Toarcian) to the Late Cretaceous (Campanian). Xenomytilus is known only from California, but future studies might reveal it to be more widespread. Xenomytilus fons is from Maastrichtian strata in central and southern California and is locally moderately common. It inhabited siliciclastic nearshore, warm waters and most likely had an epifaunal mode of life, with attachment by byssus to hard substrate. Specimens were commonly transported by turbidity currents into deeper waters. The other new species is the tellinoidean /canotia californica, sp. nov., from the upper Turonian upper part of the Baker Canyon Member and lower part of the Holz Shale Member of the Ladd Formation in the Santa Ana Mountains, Orange County, southern California. It lived in inner shelf waters and was infaunal in silts and very fine sands. /canotia is a relatively rare, warm temperate and Tethyan, Cretaceous (Valanginian to Maastrichtian) bivalve known predominantly from Europe. Rare dispersals of Jcanotia into the New World were, most likely, by westward-flowing equatorial currents that coincided with high stands of sea level. Its occurrence in southern California is its only record on the Pacific slope of North America. INTRODUCTION Stewart (1930:87) mentioned that the holotype of Mytilus pauperculus Gabb (1864) is from Martinez (perhaps in- This paper concerns the recognition of a new subfamily, dicating Maastrichian age), California, the University of genus, and species of mytilid bivalve, and a new species California, Museum of Paleontology (Berkeley) label of tellinoidean bivalve. Both are warm-water taxa and with this specimen says “Curry’s,”’ which would make occur in shallow-marine, Upper Cretaceous rocks in Cal- the type Albian or Cenomanian in age. Whiteaves (1879: ifornia. The new mytilid bivalve, Xenomytilus fons, gen. 167, 1903:394, unfig.) and White (1889:34, 38, unfig.) et sp. nov., is of Maastrichtian age and from central and tentatively reported this species from Vancouver Island. southern California, whereas the new tellinoidean bi- For a discussion of the members of the lower Redding valve, Icanotia californica, sp. nov., is of late Turonian Formation, Members I-III, see Squires & Saul (2003a). age and from southern California. For a discussion of the ““Roof of Coal” stratigraphic unit, Although specimens of Cretaceous mytilid bivalves in see Squires & Saul (2003b). Lithodomus Cuvier, 1817, is the rock record of the Pacific slope of North America are the junior synonym of Lithophaga Réding, 1798. Stewart usually not well preserved, they are moderately common. (1930:103—104) reported that the holotype of the “*Cre- The mytilids and possible mytilids previously reported nella’ concentrica might be a crushed specimen of an from Cretaceous strata in this area are listed in Table 1. astartid. Much work is needed to get all of the listed bivalve The modioliform “‘Modiolus’? major Gabb (1869:191— names tied to definite genera and to fully establish the 192, pl. 32, fig. 88; Stanton, 1895:48, pl. 3, fig. 1) from chronologic and geographic ranges of the species. None Hauterivian and Albian strata in the Great Valley Group of the previously named or reported mytilids or possible of northern California is not a mytilid, according to Stew- mytilids from Cretaceous strata in the study area belongs art (1930:104, pl. 4, fig. 1), who placed this bivalve in to the new genus of mytilid described here. the carditid genus Myoconcha J. de C. Sowerby, 1824. The following comments pertain to Table 1. Although Campbell et al. (1993) reported that Gabb’s species is part The Veliger, Vol. 48, No. 3 Table 1 Mytilids and possible mytilids (names enclosed in parentheses) previously reported from Cretaceous strata of the north- eastern Pacific region. Taxon Age Locale(s) Mytilus pauperculus Gabb, 1864 “Mytilus lanceolatus’> Sowerby of White- Albian? eaves, 1884 Modiolus persistens Whiteaves, 1900 Albian? “Lithodomus maudensis” Whiteaves, 1884 Albian? Modiolus siskiyouensis Gabb, 1864 Lithophaga oviformis Gabb, 1864 Turonian Inoperna bellarugosa Popenoe, 1937 Late Turonian Lithophaga sp. Ludvigsen & Beard, 1994 and 1997 Modiolus teleus Stewart, 1930 Lithodomus nitidus Whiteaves, 1903 Brachidontes bifurcatus Popenoe, 1937 Mytilus? quadratus Gabb, 1869 Maastrichtian “Modiolus siskiyouensis” of Whiteaves 1903. Cretaceous “Modiolus (Brachidontes)” sp. indet. of Cretaceous Whiteaves, 1903 “Mytilus” sp. indet. of Anderson, 1958 Cretaceous “Crenella” concentrica Gabb, 1864 Cretaceous? Albian or Cenomanian Turonian?, Coniacian?, or Campanian? Late Santonian or early Late Santonian? and early Campanian Early Campanian Late middle Campanian Mt. Diablo, Contra Costa Co., northern California and Vancouver Is., British Columbia Shingle Bay, eastern Skidegate Inlet, Moresby Is., British Columbia East end of Maude Is., Queen Charlotte Islands, British Columbia East end of Maude Is., Queen Charlotte Islands, British Columbia Siskiyou Mountains, Jackson Co., southern Oregon Lower Redding Fm., Members I-III, Cow Creek, Shasta Co., Calif. Ladd Formation, lower Holz Shale Mbr., Orange Co., Calif. Haslam Formation, Nanaimo, Vancouver Is., British Co- lumbia Chico Fm. at Tuscan Springs, Tehama Co. & at Pentz, Butte Co., California Pender Fm. (“Roof of Coal,’’ Nanaimo), Vancouver Is., British Columbia Ladd Formation, upper Holz Shale Member, Orange Co., Calif. “Chico Group” at Martinez, Contra Costa Co., northern Calif. Nanaimo area, Vancouver Is., British Columbia Hornby Is. and Sucia Is., British Columbia Santa Ynez Mountains, Santa Barbara Co., California Martinez, Contra Costa Co., Northern California of a chemosynthetic macroinvertebrate fauna in cold-seep limestones from the Great Valley Group at Wilbur Springs (Hauterivian) and at Cold Fork of Cottonwood Creek (Albian), northern California. Xenomytilus fons, sp. nov. is locally abundant, and locs. 2b & 3 Maastrichtian |@ loc. 1 x4 @ loc. 2a fons Campanian Santonian on | Coniactent| : @loc.4 /. californica CAG Turonian Cenoman. 98.9 Figure 1. Index map showing locales of the new taxa. 1 = Garzas Creek. 2a = Cantinas Creek and north shore, Lake Na- cimiento area. 2b = Dip Creek. 3 = Warm Springs Mountain. 4 Santa Ana Mountains. n =) (e) o oO iar) vy o L O fe o Qa 1QO =) specimens are from moderately widespread locales in var- ious formations (Figure |). Although most of the speci- mens were collected by others and stored in museums, some of the specimens from Warm Springs Mountain and Lake Nacimiento area were collected by the authors. This new bivalve is unusual for a mytilid in that it possesses a strong tooth in either valve. It also possesses a sickle shape, a very sharp umbonal ridge that extends from the beak to the posterodorsal margin, abundant radial ribs, and a concave ventral margin. These external features are equivocal, and it is the hinge that is the most defining feature of this mytilid, which, so far, appears to be en- demic to California. Xenomytilus is unusual among my- tilids, so much so that it can be placed in its own sub- family, along with Lycettia Cox, 1937, an Old World Tethyan mytilid of Early Jurassic to Late Cretaceous age (see “‘Systematic Paleontology’). The occurrence of /canotia californica, sp. nov., is the first documented record of this genus from the Pacific slope of North America. A total of 25 specimens was detected in museum collections. As will be discussed later in this paper, /canotia is a warm-temperate and Tethyan R. L. Squires & L. R. Saul, 2005 bivalve, whose geologic range is nearly the entire Cre- taceous (Valanginian to Maastrichtian). It is predomi- nantly known from Europe and Asia and is a relatively uncommon bivalve (Casey, 1961). Most of its species are based on only a few specimens (Dhondt & Dieni, 1993). Prior to this present study, the only other report of this genus in North America was a single specimen from the “Coon Creek fauna” in the Coon Creek Tongue at the base of the Ripley Formation, McNairy County, Tennes- see (Wade, 1926:91—92). This fauna is of latest Campan- ian age (Cobban & Kennedy, 1995:2). Abbreviations used for catalog and locality numbers are: CSUE California State University, Fullerton; LAC- MIP, Natural History Museum of Los Angeles County, Invertebrate Paleontology Section; UCR, University of California, Riverside. STRATIGRAPHY AND DEPOSITIONAL ENVIRONMENTS OF THE NEW TAXA The following stratigraphic units are listed from oldest to youngest. Holz-Baker Transition of Ladd Formation Specimens of Icanotia californica were found in the upper part of the Baker Canyon Member and in the im- mediately overlying, lowermost part of the Holz Shale Member of the Ladd Formation, Santa Ana Mountains, Orange County, southern California (Figure 1, locale 4). These two members are gradational in lithology, and this gradational zone, which is commonly referred to as the ‘“‘Holz-Baker transition,’ constitutes the lowermost part of the Holz Shale. The Baker Canyon Member and the ““Holz-Baker transition”’ are late Turonian in age, based on gastropods, bivalves, and ammonites (Saul, 1982). Sundberg (1980:text-fig. 2) provided a generalized Cre- taceous stratigraphic column showing these two mem- bers. He also assigned fossils from this part of the Holz Shale Member to an Inoperna-Pterotrigonia association, which he reported as characteristic of a shallow-water, offshore environment that was above wave base. All but one of the specimens we were able to find of Icanotia californica are in siltstone; the one exception is in very fine sandstone. A few of the specimens found in siltstone are conjoined and one is “‘butterflied.”” We conclude that these specimens have undergone only minimal transport. Siltstone lithology is more consistent with an environ- ment below wave base and more in keeping with a shelf environment of moderate depth, like the one Saul (1982: fig. 2) envisioned for the mollusks in the Holz-Baker tran- sition. Garzas Sand Member of Moreno Formation The type locality of Xenomytilus fons is in the Garzas Sand Member of the Moreno Formation, Garzas Creek Page 123 area, Stanislaus County, central California (Figure 1, lo- cale 1). This member has a complex nomenclatural his- tory, but it has always had the term “‘Garzas”’ as part of its name. Regardless of how the term has been used, ““Garzas”’ consistently refers to the uppermost Cretaceous sandstone that crops out north of the San Luis Reservoir/ Pacheco Pass area, Stanislaus and western Mecerd coun- ties, along the western side of the northern San Joaquin Valley, central California (Bishop, 1970). This member contains an abundant molluscan fauna that is of late early Maastrichtian age (Saul, 1983:fig. 10), based primarily on turritellas (gastropods) and ammonites. The Garzas Sand Member was deposited in a nearshore shallow-water en- vironment in the Garzas and Quinto creeks area, where specimens of the new mytilid species were found. More conjoined valves of the new species were found in this member than in the other stratigraphic units where this species occurs. Beyond the Garzas and Quinto creeks area, the Garzas Sand Member was deposited in deeper water (Bishop, 1970), and some of its fauna was probably transported downslope and offshore (Saul, 1983). El] Piojo Formation Some specimens of Xenomytilus fons were found in this formation, which crops out at 1) Cantinas Canyon northwest of Lake Nacimiento in San Luis Obispo Coun- ty, west-central California and 2) nearby, along the north and south shores of Lake Nacimiento (Figure 1, locales 2a & 2b). This formation, which is of early late Maas- trichtian to earliest Paleocene age, based on turritellas (Saul, 1983, 1986; Squires & Saul, 1993), has not re- ceived detailed depositional-environment studies, but some of its mollusks are the same nearshore, warm-water ones found in other stratigraphic units that contain X. fons. Turritella chaneyi orienda Saul, 1983, is present in both the El Piojo Formation and the basal San Francis- quito Formation (see below). Turritella webbi paynei Saul, 1983, and the bivalve Glycymeris (Glycymerita?) banosensis Anderson, 1958, are present in both the El Piojo Formation and the Garzas Sand Member. The strata that comprise what is now referred to as the El Piojo Formation were deposited by debris flows associated with turbidity currents (Grove, 1986). Saul (1983, 1986) also reported that, most likely, the shallow-marine mollusks found in the El Piojo Formation underwent some down- slope movement into deeper water. No conjoined speci- mens of the new mytilid were found in the El Piojo For- mation, but this is in keeping with its turbidite-influenced sedimentation. Specimens of the new mytilid from this formation are very unusual because they occur in pebbly sandstone. Basal San Francisquito Formation Many specimens of Xenomytilus fons were found in the lowermost part of the basal San Francisquito For- Page 124 mation at Warm Springs Mountain, Los Angeles County, southern California (Figure 1, locale 3). Based on turri- tellas and ammonites (Saul, 1983), this formation ranges in age from late Maastrichtian to late Paleocene and rep- resents a transgressive sequence that consists of near- shore, shallow-marine deposits grading upsection into deeper submarine-fan deposits (Kirby & Saul, 1995). Kir- by & Saul (1995:text fig. 2) reported also that the Cre- taceous part of this formation 1) conformably overlies the granite-gneiss basement rock, 2) is approximately 110 m thick, 3) was deposited in a transition-zone environment, and 4) contains storm-lag concentrations of fossils. Field work by the senior author of this present report revealed that specimens of the new mytilid occur in small lenses interbedded with thicker intervals of fine sandstone in the lowermost 15 m of this 110 m-thick stratigraphic unit. The lenses consist of poorly sorted, very fine to coarse sandstone, with scattered very angular granules and small pebbles of granite. In addition to Xenomytilus fons, other fossils in these lenses are Turritella, thick-shelled oysters, Glycymeris, and plant debris. Some of the lenses consist almost entirely of oyster coquinas, and others consist of very coarse grains of decomposed granite. Although the fossils in the lenses have been transported, the distance of transport was not great, based on the presence of a few conjoined valves of the mytilid. Based on the poor sorting of the deposits, the small lateral extent of the lenses, and the presence of coquinas, we interpret that the lowermost part of the San Francisquito Formation at Warm Springs Mountains was deposited in a nearshore environment that was either in the shoreface environment or at the interface between lower shoreface and the proximal part of the transition zone. A few meters stratigraphically above these Xenomytilus-bearing beds, specimens of the the warm-water bivalve Roudairia Munier-Chalmas, 1881, were found. Roudairia is a predominantly Tethyan genus, and its presence is suggestive that the basal part of the San Francisquito Formation was deposited in warm wa- ters (Kirby & Saul, 1995). MODE OF LIFE Xenomytilus fons Adult mytilids are byssate (Coan et al., 2000). Appar- ently the new mytilid rested on its ventral surface because conjoined specimens are stable in that position, much like those of the malleid bivalve Nayadina (Exputens) Clark, 1934, known from Eocene strata of North America and Mexico (Squires, 1990). Nayadina (Exputens), however, was a nestler, based on the wide variability in its mor- phology. The new mytilid does not show as much vari- ability in its morphology. Icanotia californica The mode of life of /canotia is difficult to fully doc- ument because icanotiids are extinct. [canotia was un- The Veliger, Vol. 48, No. 3 doubtedly infaunal, because Casey (1961) stated that it has a deep pallial sinus. According to Casey (1961), Ican- otia does not have a posterior gape. Most of the conjoined specimens of /canotia californica also do not show a gape. One conjoined specimen of /. californica (Figure 23), however, shows what appears to be a gape, but it is, most likely, the result of post-burial processes. Sundberg (1980:text-fig. 8) depicted Jcantia [sic] as a deeply infaunal, suspension feeder bivalve with separate, long siphons, like those found on Recent tellinids. In spite of what he stated, no one can be sure whether or not Icanotia was a suspension feeder. Most tellinoids (e.g., tellinids) are actually deposit feeders, although some tel- linoids (e.g., psammobiids) are probably suspension feed- ers (Boss, 1982; Coan et al., 2000). Whereas modern tel- linoids have separate siphons (Boss, 1982; Coal et al., 2000), no one can be sure of the morphology of the si- phons of /canotia. PALEOBIOGEOGRAPHY OF THE STUDIED GENERA Xenomytilus Xenomytilus is known so far only from central and southern California. Although it is possible that it was endemic to this region, it seems likely that eventually it will be found elsewhere in the world. As mentioned under ““Systematic Paleontology,” there are similar age mytilids from central Asia and southern India, and careful cleaning of their hinges could reveal them to be the new genus. Icanotia Icanotia Stoliczka (1871) appears to be restricted to strata deposited in warm-temperate and tropical (Tethyan) seas in Europe, Africa, central Asia, and the Americas (Dhondt, 1987). The geologic occurrences of Jcanotia that we are aware of are listed in Table 2 and summarized graphically in Figure 2. As can be seen, this genus ranges from Early Cretaceous (Valanginian) to Late Cretaceous (Maastrichtian). References heavily utilized in our study were Pictet & Campiche (1865), Wade (1926), Olsson (1934, 1944), Casey (1961), Dhondt (1987), and Dhondt & Dieni (1993). Among these workers, only Wade (1926) and Olsson (1934, 1944) included any mention of Jcan- otia from California. Switzerland has the only known Valanginian and Hau- terivian occurrences of IJcanotia (Pictet & Campiche, 1865). No Barremian records are yet known, but the ge- nus is presumed to have been present in the vicinity of Switzerland at that time, because Aptian records are known from there (Pictet & Renevier, 1855). Also during the Aptian, /canotia migrated into England (Casey, 1961). The only Albian records are in upper Albian strata from England (Casey, 1961) and undifferentiated Albian strata in northern Peru (Olsson, 1934). This Peruvian occur- R. L. Squires & L. R. Saul, 2005 Page 125 Table 2 Stage and geographic distribution of species of [canotia. Stage Species Valanginian I. escheri (Pictet & Campiche, 1865) Hauterivian I. intermedia (Pictet & Campiche, 1865) Barremian None known Aptian I. pennula Casey, 1961 I. studeri (Pictet & Renevier, 1855) Albian I. peruviana Olsson, 1934 I. impar (Zittel, 1865) Cenomanian I. impar (Zittel, 1865) I. atlantica Dartevelle & Freneix, 1957 Turonian I. californica, sp. nov. I. discrepans (d’Orbigny, 1845) I. impar (Zittel, 1865) I. atlantica Dartevelle & Freneix, 1957 Coniacian I. discrepans (d’Orbigny, 1845) I. impar (Zittel, 1865) probably J. atlantica Dartevelle & Freneix (1957) Santonian I. discrepans (d’Orbigny, 1845) I. impar (Zittel, 1865) probably /. atlantica Dartevelle & Freneix (1957) Lowermost Camp. to upper- I. elicita Stoliczka, 1871 most Sant. Campanian I. impar (Zittel, 1865) Location Switzerland Switzerland England Switzerland Northern Peru England France, Austria, Germany, England Cameroon & Congo, Central Africa Southern California France France, Austria Cameroon & Congo, Central Africa France Austria, NE Italy Cameroon & Congo, Central Africa France Austria, NE Italy Cameroon & Congo, Central Africa Southern India Austria, Hungary, NE Italy, Central Asia possibly I. discrepans (d’Orbigny, 1845) France Uppermost Camp. Maastrichtian I. pulchra Wade, 1926 I. pacifica Olsson, 1944 Tennessee Peru rence is associated with the gastropod Sogdianella peru- viana (Olsson, 1934), which Olsson incorrectly placed in genus Actaeonella d’Orbigny, 1842, according to Sohl & Kollman (1985), who assigned this gastropod to the Al- bian. The arrival of [canotia in Peru probably coincided with both a global trend of rising sea level (Haq et al., 1987) and with warm and equable surface waters that occurred during the Albian (Frakes, 1999). This migration into the New World was, most likely, via westward-flow- ing equatorial currents. This Peruvian occurrence repre- sents the first known migration of this genus out of Eu- rope, but the migration was rather limited, because during the Cenomanian, the genus is only known from Europe (d’Orbigny, 1845; Geinitz, 1871-1875; Woods, 1904— 1913). With the exception of the late Turonian occurrence of I. californica in southern California, Europe continued to be the only area where this genus lived during the Tu- ronian. The exceptional warming during the early Turon- ian (Frakes, 1999) most likely induced the spread of Ican- otia into southern California. Additionally, there was a high-sea stand at this time, and the seaway across south- ern Mexico was apparently at its widest (Imlay, 1944; Alencaster, 1984). These conditions further enabled [can- otia to migrate. That this migration was via westward- flowing equatorial currents, rather than by way of the easternmost Tethys (Japan) and the northern Pacific gyre, is suggested by the lack of this genus in Japan, British Columbia, Oregon, or northern California. If Icanotia had arrived onto the Pacific slope of the Americas via a north- erly route, there should be some record of it in these higher latitude locales. The late Turonian New World oc- currence in California, like the Albian New World oc- currence in Peru, was relatively shortlived. During the Coniacian and Santonian, the genus is known to have lived only in Europe. During the Campanian, /canotia was again present in Europe, but the genus migrated south to Cameroon and the Congo regions of central Af- rica, and east to central Asia and southern India (Stoli- czka, 1871; Dartevelle & Freneix, 1957; Pojarkova, 1976). Again, these expansions into other regions did not persist very long. During the Maastrichtian, however, Icanotia was no longer present in the Old World, but was present only in the New World; namely, in Tennessee (Wade, 1926) and in Peru (Olsson, 1944). This Peruvian occurrence, which is based on Icanotia pacifica Olsson, 1944, is the youngest record for this genus. Olsson (1944: 217) assigned I. pacifica to his Baculites ammonite zone, which he believed to be of Maastrichtian age.. He reported that this zone contains Baculites lyelli (d’Orbigny, 1847). For an updated synonymy of d’Orbigny’s species, see Cobban & Kennedy (1995:26, 29), who stated that B. lyelli is a junior synonym of Eubaculites carinatus (Mor- Page 126 The Veliger, Vol. 48, No. 3 ton, 1834), which is a widely occurring baculitid of early to late Maastrichtian age. SYSTEMATIC PALEONTOLOGY Class BIVALVIA Linnaeus, 1758 Order MYTILOIDA Férussac, 1822 Superfamily MyYTILOIDEA Rafinesque, 1815 Family MYTILIDAE Rafinesque, 1815 Subfamily XENOMYTILINAE Squires & Saul, subfam. nov. Type genus: Xenomytilus, gen. nov.; Maastrichtian, cen- tral and southern California. Diagnosis: Small mytilid, smooth or ribbed, falcate (sick- le shape), single tooth in either valve with socket in cor- responding valve. Included genera: Lycettia Cox, 1937, and Xenomytilus. Discussion: Recognized subfamilies of Mytilidae in Soot- Ryen (1969) are: Mytilinae Rafinesque, 1815; Crenellinae Adams & Adams, 1857; Lithophaginae Adams & Adams, 1857; and Modiolinae Keen, 1958. Although the new subfamily has the pointed terminal beaks and falcate shape of some mytilines, the thickened anterior hinge margin of some crenellines, and the falcate shape of some modiolines, Xenomytilinae differs from all of these other subfamilies in having a single, strong tooth on either valve. The only other mytilid that has very similar den- tition is Lycettia Cox, 1937, a genus considered to be a mytiline by Soot-Ryen (1969) but regarded by us to be- long to the new subfamily. Genus Xenomytilus Squires & Saul, gen. nov. Type species: Xenomytilus fons, sp. nov.; Maastrichtian, central and southern California. Diagnosis: Small mytilid, falcate, sharply carinate ada- jacent to ventral margin, radially ribbed, ventral surface concave, prominent strong tooth on either valve, deep socket on other valve, and chevron-like pits commonly present on valve margins near hinge area. Discussion: The hinge of the new genus develops alter- natively in either valve, with the single prominent tooth located between two sockets, the ventral one being the larger one. The new genus does not have true transposi- tion, described by Popenoe & Findlay (1933), because the prominent tooth is variable in its location. The new genus is most similar in terms of dentition to Lycettia Cox, 1937, but the new genus differs by having ribs, more inflated valves, a sharp carina, a wider and more concave ventral surface, a more prominent tooth on the hinge, chevron-like pits posterior to the nymph, and a much Geographic Region Central Central rans Europe Peru | Tenn. |Calif. Figure 2. Global chronostratigraphic and geographic distribu- tion of Jcanotia. Dashed lines indicate presumed occurrence. 1 = France, Switzerland, Germany, Austria, Hungary, and Eng- land. 2 = Cameroon. 3 = Central Asia. 4 = Southern India. 5 = Northern Peru. 6 = Tennessee. 7 = Southern California. Geo- logic ages after Gradstein et al. (2004). shorter and less distinct nymph. Cox (1937) illustrated only left valves of Lycettia, but Tashiro (1992:pl. 14, fig. 12) illustrated a right valve. Cox’s (1937) description of the hinge of Lycettia is hard to make sense of, and his illustrations do not match his descriptions. His usage of the terms “‘above”’ and “‘below”’ are very confusing. He put too much emphasis on the ligamental ridge and did not provide enough information about the dentition. Lycettia is an Old World Tethyan bivalve that ranges from late Early Jurassic (Toarcian) to Late Cretaceous (late Campanian) (Tashiro, 1992). It is found in the upper Lower Jurassic of Spain (Gahr, 2002), the lower Middle R. L. Squires & L. R. Saul, 2005 Page 127 Jurassic (Aalenian to Bajocian) of England (Cox, 1937), the Middle Jurassic (Bathonian) of east Africa (Hallam, 1977), the Upper Jurassic of Portugal (Fiirsich & Werner, 1987), the Upper Jurassic (Oxfordian to Kimmeridgian) to Lower Cretaceous of western China (Yin & Fiirsich, 1991), and Lower Cretaceous (Berriasian to Valanginian) to Upper Cretaceous (upper Campanian) in southern Ja- pan (Tashiro, 1992). It is tentatively known from the Mid- dle Jurassic (Bathonian) of western China (Sha et al., 1998). One species, Lycettia tippana (Conrad, 1858) has been reported by various workers (see Akers & Akers, 2002) to be in upper Campanian and Maastrichtian strata of southeastern and eastern United States, but this so-called “Lycettia” is Cuneolus Stephenson, 1941. Soot-Ryen (1969) regarded Cuneolus to be a synonym of Lycettia, but Cuneolus is edentulous. In terms of the falcate shape and strong carina adjacent to the ventral margin, the new genus is similar to the Permian mytiline Coxesia Mendes, 1952; the Jurassic modioline Falcimytilus Cox, 1937, the Maastrichtian my- tiline Cuneolus, and the Recent mytiline Mytilaster Mon- terosato, 1884. The new genus differs from the first three of these by having ribs and not being edentulous. The new genus differs from Mytilaster by having a dentition- bearing plate in the umbonal cavity, terminal beaks, more concave ventral surface, and chevron-like pits on the valve margins near the hinge area. In terms of the overall shape and presence of ribs, Xe- nomytilus is similar to the Jurassic to Recent mytiline Brachidontes Swainson, 1840, but the new genus differs by having terminal rather than nearly always subterminal beaks, much more pointed beaks, an angulation rather than a rounded ridge between the beak and the postero- ventral margin, a concave ventral surface, and a strong tooth or deep socket in each valve rather than a variable number of very small dysodont teeth with intervening shallow sockets. In having chevrons along the dorso-posterior margin (see Olsson, 1961:pl. 12, fig. 5) and the possibility of only a single tooth on the hinge, Xenomytilus is also similar to Aeidimytilus Olsson, 1961, (type species: Mytilus adamsiana Dunker, 1857), a Recent mytilid from south- ern California to Ecuador (Coan et al., 2000). Aeidimy- tilus can have one to three central teeth. Xenomytilus dif- fers from Aeidimytilus by having larger size, terminal beaks, falcate shape, much weaker and flatter ribbing, usually non-bifurcating ribs, and a much more concave ventral surface. Olsson (1961) opined that Aeidimytilus is a subgenus of Scolimytilus Olsson, 1961. Keen (1971) and Coan et al. (2000) opted for making Aeidimytilus and Scolimytilus synonyms of Brachidontes, because the soft parts do not differ. Also, in terms of the general shape and presence of radial ribs, the new genus resembles the Triassic to Re- cent mytiline Septifer Récluz, 1848, sensu stricto, and the Recent mytiline Ischadium (Jukes-Brown, 1905). The new genus differs from Septifer by having dentition in- stead of a septum. The new genus differs from [schadium by having a narrower shell, an angulation between the disk and the concave ventral surface, and a single tooth in either valve. The hinge teeth of Ischadium are several in number (Soot-Ryen, 1955). Etymology: The genus name is a combination of xeno, Greek for strange, and mytilus. Xenomytilus fons Squires & Saul, sp. nov. (Figures 3-17) Brachidontes n. sp. Saul, 1986:26, 27. Brachidontes? n. sp. Kirby & Saul, 1995:24. Diagnosis: Same as for genus. Description: Shell small (most specimens approximately 30 mm in length and 15 mm in height, rare specimens up to 42.5 mm in length and 18 mm in height), longer than high, shell height/shell length ratio = usually 0.5 (largest specimen = 0.42). Falcate, inequilateral, equi- valved, valves inflated. Anterior end acutely pointed. Hinge margin long with portion posterior to ligament bent in low angle. Beaks pointed and terminal at anterior end. Umbones terminal, acutely pointed. Umbonal ridge pro- nounced and sharp (angulate), extending from beak to postero-ventral margin. Disk moderately broad. Ventral margin of both valves concave. Valves ornamented with abundant radial ribs, varying in strength on different ar- eas. Radial ribs of anterior area narrowest, slightly wider than interspaces but becoming approximately twice as wide as interspaces posteriorly. Radial ribs on disk flattish to lowly rounded. Radial ribs of anterior area diverging from ribs of flank along strong, arching beak-to-outer margin carina. Radial ribs on concave ventral margin nar- row, closely spaced, and approximately three times as wide as interspaces. Hinge consisting of single, prominent tooth on either valve; tooth located between two sockets with the ventral socket larger. Tooth long and narrow, usually dorsal of center of hinge, but can be slightly ven- tral of center or extending diagonally across central area. Tooth bears microscopic striae on both sides, especially on ventral side. Valve not bearing tooth has deep socket. Ventral area of hinge with relatively inconspicous tooth and subordinate socket (notch) at hinge margin. Lunular area can be infolded to form very narrow channel for corresponding valve margin of other valve. Nymph mod- erately long, with thin extension of nymph not quite reaching apex of valve. Dorsal margin of valve, including ligamental area, can be pitted with chevron-like crenula- tions. Dimensions of holotype: Conjoined valves, length 31.1 mm, height 16.5 mm, thickness 15.5 mm. Holotype: LACMIP 13290. Page 128 The Veliger, Vol. 48, No. 3 Type locality: LACMIP loc. 121°07'14"W. Paratypes: LACMIP 13291-13299. 10660, 37°13'41"N, Geologic age: Middle to latest Maastrichtian. Distribution: MIDDLE MAASTRICHTIAN: Moreno Formation, Garzas Sand Member, Garzas Creek, Stanis- laus County, north-central California (type locality). LOWER UPPER MAASTRICHTIAN: El Piojo Forma- tion, Cantinas Canyon northwest of Lake Nacimiento and north shore of Lake Nacimiento, San Luis Obispo Coun- ty, west-central California. UPPERMOST MAASTRICH- TIAN: El Piojo Formation, upper part, south shore of Lake Nacimiento, San Luis Obispo County, west-central California; San Francisquito Formation, basal part, Warm Springs Mountain, Los Angeles County, southern Cali- fornia. Discussion: The above description is based on 71 spec- imens: 35 left valves, 27 right valves, and 9 conjoined valves. Several valves were carefully cleaned to expose the hinge. The chevron-like crenulations on the nymph and posterior to it are not preserved on all specimens. Saul (1986:26, 27) mentioned Brachidontes, n. sp. from Cantinas Creek area just northwest of Lake Naci- miento and from the north shore of Lake Nacimiento, west-central California, and this bivalve is synonymous with Xenomytilus fons. She also mentioned in this same article that a similar form of Brachidontes, n. sp. is from the Moreno Formation, and this bivalve is also synony- mous with X. fons. Kirby & Saul (1995:24) mentioned Brachidontes? n. sp. from the Warm Springs Mountain, southern Califor- nia, and this bivalve is synonymous with X. fons. Xenomytilus fons somewhat resembles Brachidontes fulpensis Stephenson (1952:84—85, figs. 10-13) from the lower Cenomanian Woodbine Formation of northeastern Texas but differs from this Texas species by having a terminal beak, a sharper angulation between the beak and the postero-ventral margin, a concave ventral surface, a sloping rather than a long and straight antero-dorsal mar- gin, shorter nymph, and a single prominent tooth on either valve. The hinge of this Texas species has three or four, deep channels with intervening sharp teeth that are mod- ifications of crenulations along the anterior margin. The new bivalve is similar to Septifer? ahaaralensis Pojarkova (1976:92-93, pl. 47, figs. 8-9, pl. 48, figs. 6— 11) from lower Santonian to lower upper Campanian stra- ta in central Asia. Although the new bivalve differs by having a more concave ventral surface and a narrower beak, some illustrated specimens of the central Asian spe- cies do approach the shape of X. fons. The new bivalve is also similar to Modiola annectans Stoliczka (1871:380—381, pl. 23, figs. 8, 8a, 9, 9a) from the Ariyalur Group [= Arrialoor Group in Stoliczka] in the Vayalappadi [= Vylapauda in Stolickza] of the Cau- very Basin in southern India. Sastry et al. (1972) reported that these rocks at this locale are in the Sillakudi For- mation. According to Sundaram et al. (2001), this for- mation is of Santonian to Campanian age. The new bi- valve differs from Stoliczka’s species by having much stronger ribs and a more sloping dorsal-anterior shell margin. No information is known about the hinge of Sto- liczka’s species. Etymology: The new species is named for its occurrence on Warm Springs Mountain; Latin, fons, meaning of a spring. Order VENEROIDA H. & A. Adams, 1856 Superfamily TELLINOIDEA de Blainville, 1814 Family ICANOTIIDAE Casey, 1961 Genus /canotia Stoliczka, 1871 Type species: Psammobia impar Zittel, 1865, by original designation; Late Cretaceous (Cenomanian to Campani- an), southern England, western and southeast France, Germany, Austria (Gosau), Hungary, northeast Italy, cen- tral Asia, and southern India. For an updated synonymy of this species, whose nomenclatural history has been very confusing, see Dhondt & Dieni (1993). Discussion: Casey (1961) implied and Keen (1969) stated that Jcanotia has two cardinal teeth in either valve. Ican- otia pulchra Wade (1926:91, pl. 29, figs. 5, 6), from up- permost Campanian strata of Tennessee, however, has a single prominent cardinal tooth in the left valve, and this tooth is situated between a large socket posteriorly and a smaller socket anteriorly, just as in the new species de- scribed below. It is possible that J. pulchra and the new species constitute a new subgenus of Jcanotia that is char- acterized by having only a single cardinal tooth, but more specimens of both species are needed in order to docu- Explanation of Figures 3 to 17 Figures 3-17. Xenomytilus fons Squires & Saul, genus et sp. nov., specimens coated with ammonium chloride. Figure 3. Holotype LACMIP 13290, LACMIP loc. 10660, left valve, X 1.6. Figures 4—5. Paratype LACMIP 13291, LACMIP loc. 10660, left-valve interior. Figure 4. X 1.5. Figure 5. Close-up of hinge shown in previous figure, <4.6. Figure 6. Paratype LACMIP 13292, LACMIP loc. 9196, left-valve interior, <3. Figure 7. Paratype LACMIP 13293, LACMIP loc. 10660, left-valve interior, X1.8. Figure 8. Paratype LACMIP 13294, LACMIP loc. 10660, left-valve R. L. Squires & L. R. Saul, 2005 Page 129 interior, X3. Figure 9. Holotype LACMIP 13290, LACMIP loc. 10660, right valve, <1.6. Figure 10. Paratype LACMIP 13295, LACMIP loc. 26486, right valve, X 1.7. Figure 11. Paratype LACMIP 13296, LACMIP loc. 26526, right valve, X1.5. Figures 12—14. Paratype LACMIP 13297, LACMIP loc. 26352. Figure 12. Apical view of beaks, X1.6. Figure 13. Oblique ventral view, 1.4. Figure 14. Ventral view, 1.4. Figure 15. Paratype LACMIP 13298, LACMIP loc. 21193, rubber peel of right-valve interior, X3.5. Figures 16-17. Paratype LACMIP 13299, LACMIP loc. 10660. Figure 16. Right-valve interior, * 1.4. Figure 17. Close-up of hinge shown in previous figure, 3.7. Page 130 ment that this single tooth is a constant morphologic char- acter. The geologic range of [canotia is Valanginian to Maas- trichtian, although it has been variously referred to as Aptian to Maastrichtian by Casey (1961), Hauterivian to Maastrichtian by Soot-Ryen (1969), and Valanginian to Maastrichtian by Dartevelle & Freneix (1957). Stoliczka (1871:168) reported Jcanotia elicita Stoliczka, 1871, from the Aryialur Group [= Arrialoor Group in Stoliczka] north of Karappadi [= Karapaudy in Stoliczka] in the Cauvery Basin of southern India. Sastry et al. (1972) re- ported that the rocks at this locale are in the middle part of the Sillakkudi Formation. Sundaram et al. (2001) re- ported that this part of the formation is latest Santonian to earliest Campanian in age. Lundgren (1894:50, pl. 2, figs. 12, 13) described Ican- otia? grosseplicata Lundgren, 1894, of early late Cam- panian age from southernmost Sweden. We do not believe that Lundgren’s species belongs in /canotia, even tenta- tively, because the beak is located too far anteriorly, the ribs are much too strong, too widely spaced, too few, and the overall outline of the valves is too quadrate. Icanotia californica Saul & Squires, sp. nov. (Figures 18—29) Icantia [sic] sp. Sundberg, 1980:table 1, text-fig. 8. Diagnosis: A narrow, elongate /canotia with low but dis- tinct beaks and weak, closely spaced ribs confined to pos- terior slope. Description: Shell medium, thin, narrow and com- pressed. Soleniform, narrow and elongate (up to 46 mm in length, with height of 13.5 mm), shell height/shell length ratio = 0.29. Equivalve and inequilateral. Dorsal margin long, straight, ventral margin essentially parallel- ing dorsal margin. Anterior end narrowly rounded, pos- terior end subtruncate. Lunule poorly developed or want- ing. Escutcheon long and very narrow. Beaks low but distinct and prosogyrate. Beaks slightly less than one- third of distance fron anterior end, with maximum shell height at beaks. Shell sculpture confined to posterior slope, with ribs radiating from beak and covering sector about 20 to 25°, from beak to postero-ventral corner. Ribs 14 to 20 in number, narrow, closely spaced, and weak to moderately strong. Ribs strongest dorsally and posteri- orly, with noticeable weakening ventrally. Anterior mar- gin of shell with radial striae crossing incised growth lines and producing minute cancellate sulpture. Shell oth- erwise smoothish, with bands of growth lamellae. Hinge on left valve consisting of singular and prominent cardi- nal tooth perpendicular to hinge margin and situated be- tween large socket posteriorly and small socket anteriorly. Ligament external and opisthodetic, seated on weakly de- veloped nymph. Interior rib, very faint, extends from The Veliger, Vol. 48, No. 3 beak to half way toward postero-ventral corner. Muscle- scar pattern unknown. Dimensions of holotype: Right valve, length 39.5 mm, height 12 mm. Holotype: LACMIP 13300. Type locality: LACMIP 10883, 33°43'12’N, 117°37'26"W. Paratypes: LACMIP 13301-13306; UCR 4032/2. Geologic age: Late Turonian. Distribution: Ladd Formation, upper part of Baker Can- yon Member and Holz-Baker transition, Silverado Can- yon, Santa Ana Mountains, Orange County, southern Cal- ifornia. Discussion: The new species is based on 25 specimens: 9 left valves, 11 left valves, and 5 conjoined valves. The largest specimen is 71 mm in length and 20 mm in height. Most of the specimens are internal molds, and most spec- imens are from the Holz-Baker transition; only a single specimen is from the Baker Canyon Member. The new species is most similar to [canotia pacifica Olsson (1944:55—S6, pl. 5, fig. 6), known from a single specimen from Maastrichtian strata of northern Peru. The new species differs from /. pacifica by being smaller and having ribs that are less widely spaced and weaker throughout, ribs that do get weaker ventrally. In addition, the new species differs by having an umbo that is evident, rather than being nondistinct, and having the maximum height one-third from the anterior end, rather than at mid- length. The new species is the first fully documented report of Icanotia from the Pacific slope of North America. Sund- berg (1980) listed the genus in a faunal list but misspelled the name. He also did not provide any type numbers nor any illustrations of this bivalve. In addition to the new species, there are three other species of Jcanotia known from the Turonian. These three all have long geologic ranges that encompass the Turon- ian, and they are the following: Jcanotia impar (Zittel, 1865) of late Albian to Campanian age in Europe, central Asia, and southern India (Casey, 1961; Dhondt & Dieni, 1993); Icanotia discrepans (d’Orbigny, 1845) of Turonian to Campanian age in Europe (Dhondt & Dieni, 1993); and Icanotia atlantica (Dartevelle & Freneix, 1957) of Cenomanian to Senonian age (i.e., the latter term is un- differentiated as to Coniacian, Santonian, and Campani- an) in central Africa (Dartevelle & Freneix, 1957). The new species differs from both /. impar and I. discrepans by being more elongate and with weaker and more close- ly spaced ribs. The new species is more similar to /. at- lantica but differs from the central African species by being larger, having stronger and more widely spaced ribs on the posterior slope, absence of ribs along the ventral side and antero-ventral edge of the valves, and having commarginal bands on some specimens. R. L. Squires & L. R. Saul, 2005 Page 131 Explanation of Figures 18 to 29 Figures 18-29. JIcanotia californica Squires & Saul, sp. nov., specimens coated with ammonium chloride. Figure 18. Paratype UCR 4032/2, UCR loc. 4032, left valve, X1.3. Figure 19. Paratype LACMIP 13301, LACMIP loc. 10100, left valve, 3.5. Figure 20. Paratype LACMIP 13302, LACMIP loc. 10100, left valve, <1. Figure 21. Paratype LACMIP 13303, CSUF loc. 62-8, left valve, X1.6. Figures 22—23. Paratype LACMIP 13304, LACMIP loc. 8198, left valve. Figure 22. Hinge, 2.7. Figure 23. Dorsal view of hinge, X2.8. Figure 24. Paratype LACMIP 13302, LACMIP loc. 10100, dorsal view, <1. Figure 25. Paratype LACMIP 13305, LACMIP loc. 10100, right- valve interior, X 1.4. Figure 26. Paratype LACMIP 13302, LACMIP loc. 10100, right valve, <1. Figure 27. Paratype UCR 4032/2, UCR loc. 4032, right valve, <1.3. Figure 28. Holotype LACMIP 13300, LACMIP loc. 10883, right valve, X1.9. Figure 29. Paratype LACMIP 13306, CSUF loc. 44-24, right valve, 0.8. Etymology: The species is named for the state of Cali- tute two of the paratypes of Jcanotia californica: Annie Dhondt fornia. (Belgium) kindly provided us with valuable biostratigraphic in- formation about Old World /canotia. Alexandr Guzhov (Institute Acknowledgments. John Cooper (California State University, of Paleontology of RAS, Moscow, Russia) kindly provided us Fullerton) kindly allowed the donation of specimens that consti- with hard-to-find literature. 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Approximately 260 m N and 10 m W of SE corner of section 7, T. 5 S, R. 7 W, El Toro Quadrangle (7.5 minute, 1949), Silverado Can- yon, Santa Ana Mountains, Orange County, southern California. Ladd Formation, lower part of Holz Shale Member. Age: Late Turonian. Collector: E A Sund- berg, circa 1978. LACMIP 8198. Section 16, T. 5 S, R. 7 W, Santiago Peak Quadrangle (7.5 minute, 1954), Santa Ana Mountains, Orange County, California. Ladd Formation, Baker Canyon Member. Age: Late Turonian. Collector: W. P. Popenoe, March 14, 1934. LACMIP 9196. Conglomerate cropping out on E side of Cantinas Creek approximately 30 m N of S line of section 5, T. 25 S, R. 9 E, Bryson Quadrangle (7.5 minute, 1948), San Luis Obispo County, west-central California. El Piojo Formation. Age: Early late Maas- trichtian. Collectors: L. R. and R. B. Saul & V. M. Seiders, November 13, 1985. LACMIP 10100. Concretions in shale 30.5 m above stream bed and near fence on N side of canyon about 0.8 km N of road fork in Santiago Canyon, NW % of NW 4% of section 28, T. 5 S, R. 7 W, Santiago Peak Quadrangle (7.5 minute, 1973), Santa Ana Mountains, Orange County, California. Ladd Formation, Holz Shale Member. Age: Late Turonian. Collector: B. N. Moore, January 1, 1928. LACMIP 10660. In small gully, 59.5 m stratigraphically above base of Garzas Sand Member, 457 m E and 579 m S of NW corner of section 20, T. 8 S, R. 8 E, N side of Whitney Canyon, Howard Ranch Quadrangle (7.5. minute, 1953, photorevised, 1971), Stanislaus County, central California. Moreno Formation, Garzas Sand Member. Age: Early late Maastrichtian. Collec- tors: B. Adams & W. P. Popenoe, June 1, 1941. LACMIP 10883. Immediately above base of gray sand- stones overlying gray basal conglomerate, NW % of SW % of section 21, T. 5 S, R. 7 W, Santiago Peak Quadrangle (7.5 minute, 1954), Santa Ana Mountains, Orange County, southern California. Ladd Formation, Baker Canyon Member. Age: Late Turonian. Collector: W. P. Popenoe, April 21, 1932. LACMIP 21193. At contact with gneiss-basement com- plex in ravine just S of NE ridge of Warm Springs Mountain, approximately 228 m N and 287 m E of Warm Springs Mountains, T. 6 N, R. 16 W, Warm Springs Mountain Quadrangle (7.5 minute, 1958, pho- torevised 1974), NW San Gabriel Mountains, Los An- geles County, southern California. San Francisquito Formation, basal part. Age: Latest Maastrichtian. Col- lectors: J. Elam & R. Webb, June 5, 1942. LACMIP 21599. Light brown, dirty sandstone, in trail cut between exposures of gneiss basement on SE slope NW side of ravine along Warm Springs trail, approx- imately 305 m E of junction with firebreak S from Warm Springs Mountain, 122 m S and 269 m E of Warm Springs Mountain, T. 6 N, R. 16 W, Warm Springs Mountain Quadrangle (7.5 minute, 1958, pho- torevised, 1974), NW San Gabriel Mountains, Los An- R. L. Squires & L. R. Saul, 2005 Page 135 corner of section 30, T. 25 S, R. 10 E, Lime Mountain Quadrangle (7.5 minute, 1948), San Luis Obispo Coun- ty, California. Piojo Formation, upper part. Age: Latest Maastrichtian. Collectors: L. R. & R. B. Saul, Decem- ber 31, 1977. LACMIP 30141. Fossils in pebbly sandstone, about 1.6 km N of Nacimiento River on E side of road (?Bee geles County, southern California. San Francisquito Formation, basal part. Age: Latest Maastrichtian. Col- lectors: R. W. Webb & E. H. Quayle, June 26, 1941. LACMIP 26352. West side San Joaquin Valley, approx- imately 1.2 km S of Garzas Creek, 671 m S, 84m E of NW corner of section 20, T. 8 S, R. 8 E, Howard Ranch Quadrangle (7.5 minute, 1953, photorevised 1971), Stanislaus County, California. Moreno Forma- tion, Garzas Sand Member. Age: Early late Maastrich- tian. Collectors: R. B. Stewart & W. P. Popenoe, 1944. LACMIP 26486. North shore (at about water line) of Lake Nacimiento, 396 m N, 533 m E of SW corner of section 17, T. 25 S, R. 10 E, Bradley Quadrangle (15 minute, 1961), San Luis Obispo County, California. El Piojo Formation. Age: Early late Maastrichtian. Col- lectors: L. R. & R. B. Saul, November 25, 1977. LACMIP 26526. Poorly sorted conglomeratic sandstone, S side of Lake Nacimiento, E side of narrows of Dip Creek, elevation 225 m, 457 m S and 122 m W of NE Rock Road) near quarter corner of section 18, T. 25 S, R. 10 E, and about 1.6 km N of Nacimiento River, Tierra Redonda Mountain Quadrangle (7.5 minute, 1949), San Luis Obispo County, west-central Califor- nia. El Piojo Formation. Age: Early late Maastrichtian. Collector(s): unknown. UCR 4032. Approximately 260 m N and 10 m W of SE corner of section 7, T. 5 S, R. 7 W, El Toro Quadrangle (7.5 minute, 1949), Silverado Canyon, Santa Ana Mountains, Orange County, southern California. Ladd Canyon Formation, lower part of Holz Shale Member. Age: Late Turonian. Collector: unknown. The Veliger 48(3):136—142 (November 2, 2006) THE VELIGER © CMS, Inc., 2006 A New Genus of Indo-West Pacific Turridae (Gastropoda: Prosobranchia) ANTONIO BONFITTO Dipartimento di Biologia e. s., via Selmi, 3, 40126 Bologna, Italy (e-mail: bonfitto@alma.unibo. it) AND MAURO MORASSI via dei Musei, 17, 25121 Brescia, Italy (e-mail: vitmoras @tin.it) Abstract. Based on specimens collected in the Gulf of Aden at bathyal depths and stored primarily in Museo di Zoologia dell’ Universita di Bologna, Acanthodaphne sabellii, gen. & sp. nov. is here described and figured. The Recent Pleurotomella abbreviata Schepman, 1913, from Ceram Sea, Indonesia, and Puha pusula Laws, 1947, from the Early Miocene of Hokianga, New Zealand, are recognized as belonging to the new genus. SEM photographs of type specimens of both previously known Acanthodaphne species are provided. INTRODUCTION The examination of turrid samples collected by the Oceanographic ship MARION DUFRESNE, within the framework of the European Community project RED SED’ 92 (Red Sea and Gulf of Aden), in the Gulf of Aden led us to recognise the empty shells of an unknown, dis- tinctive raphitomine species. This species is undescribed and shares with the New Zealand Early Miocene (Otaian) Puha pusula Laws, 1947, and the Indonesian Recent Pleurotomella abbreviata Schepman, 1913, distinctive morphological features which can be used in recognition of a new supraspecific taxon here proposed as Acantho- daphne, gen. nov. Acanthodaphne sabellii, the new spe- cies from Gulf of Aden, is extremely similar to Puha pusula and examination of material relevant to this latter species proved to be necessary. The close resemblance between the new species and P. pusula indicates that shell characters in Acanthodaphne have remained remarkably stable over about 20 m.y. Remarks and SEM photos based on the examination of the paratype of Puha pusula stored in Institute of Geo- logical & Nuclear Sciences (New Zealand) are presented in order to define more precisely the differences between these two closely related species. Although none of its members is anatomically known, Acanthodaphne, gen. nov. is here assigned to the subfam- ily Raphitominae Bellardi, 1875. The protoconch mor- phology and anal sinus shape of all species here assigned to the group are consistent with such attribution. Under the cladistic classification of the superfamily Conoidea proposed by Taylor, Kantor & Sysoev (1993), the subfamily Raphitominae Bellardi, 1875 (synonym Daphnellinae Casey, 1904), previously included in the family Turridae, is transferred to the Conidae. However, this proposed classification has recently been critiqued by Rosenberg (1998) who demonstrated that their results cannot be reproduced and remarked that additional data are necessary before a more complete classification can be performed. As we consider the assignment of the Ra- phitominae to the family Conidae at present uncertain, the traditional arrangement will be followed in the present paper, as well as in papers in progress, referring to Tur- ridae sensu lato. Abbreviations used in the text are: a/] = ratio of ap- erture length to total shell length; b/l ratio of shell breadth to total length; IGNZ = Institute of Geological & Nuclear Sciences, New Zealand; MNHN = Muséum National d’Histoire Naturelle, Paris; MZB Museo di Zoologia-dell’ Universita di Bologna; ZMA = Zoologisch Museum Amsterdam. SYSTEMATIC DESCRIPTION Family TURRIDAE H. & A. Adams, 1853 Subfamily RAPHITOMINAE Bellardi, 1875 Acanthodaphne Bonfitto & Morassi, gen. nov. Type species by original designation: Acanthodaphne sabellii, sp. nov. Included species: The type species (Gulf of Aden); Pleu- rotomella abbreviata Schepman, 1913 (Indonesia); Puha pusula Laws, 1947 (Early Miocene of New Zealand). Description: Shell small (maximum length 8.5 mm), heavy and biconic. Whorls sharply angled below or near A. Bonfitto & M. Morassi, 2005 middle, sculptured by opisthocline axial folds extending from lower suture to periphery where they form sharp tubercles. Spiral sculpture consisting of a peripheral cord with 1—2 weaker cords below it on spire whorls. Suture bordered by a subsutural fold bearing tubercles more nu- merous than axial folds. Subsutural ramp wide and weak- ly concave. Inner lip wide. In’ the type species, median area of the inner lip sometimes obliquely thickened but not producing a pleat. Fasciole well developed. Anal si- nus moderately deep, reversed L-shaped. Protoconch multispiral with diagonally decussate riblets. Etymology: From Acanthus (Latin name for the plant) referring to the solid shell with sharp tubercles somewhat reminiscent of the bold plant with stately spikes. Gender feminine. Discussion: Laws (1947), in a work devoted to the study of the mollusca from Hokianga District, New Zealand, introduced the Early Miocene (Otaian) Puha pusula Laws, 1947. The species was subsequently listed by Pow- ell (1966:131) as characteristic to the genus Puha Mar- wick, 1931, a taxon erected with Puha fulgida Marwick, 1931, an early or middle Miocene (Altonian-Clifdenian) species from Gisborne District, New Zealand, designated as type species. Maxwell (1988) remarked that Puha pusula should be removed from the genus Puha and noted a close resem- blance between the former species and the Recent Indo- nesian Pleurotomella abbreviata Schepman, 1913. Powell (1966) and Sysoev (1997) regarded Pleurotomella ab- breviata as belonging to the genus Cryptodaphne Powell, 1942. Shuto (1971) transferred it with a query to genus Buccinaria Kittl, 1887. In the absence of a more appro- priate generic taxon, Maxwell (1988) also provisionally referred Puha pusula to genus Buccinaria. The new genus Acanthodaphne is here proposed for Pleurotomella abbreviata Schepman, 1913, Puha pusula Laws, 1947 and a third species, namely Acanthodaphne sabellii, sp. nov., from the Gulf of Aden. These three species constitute a peculiar species-group characterized by the small but heavy shell, biconic shape, tuberculate subsutural fold and sculpture of axial folds restricted to lower part of the whorl forming tubercles at periphery. MacNeil (1960:110, pl. 5, fig. 13) reported and figured a Pseudoinquisitor? cf P.? pulchra known from a single specimen from Yonobaru Clay, Miocene of Okinawa (Ja- pan). The author noted that the species “‘does not have any close relatives in the region of Japan.’ Judging from the figure provided, it closely resembles members of Acanthodaphne in type of sculpture. Unfortunately, the protoconch features are unknown and the position of the species therefore remains indeterminate. Acanthodaphne closely resembles the genus Puha Mar- wick, 1931, in some features such as the tuberculate sub- sutural fold and periphery. However, the former genus differs from the latter in shape (biconic compared to Page 137 ovate-elongate), smaller dimensions (5.6—8.5 mm com- pared to 9-12.4 mm in length), less gradate whorls with much broader subsutural ramp, and moderately deep re- versed L-shaped anal sinus compared to virtually absent. Powell (1966:130) noted that in Puha ‘‘there is no ante- rior fasciole’”” whereas in Acanthodaphne it is well de- veloped. Some specimens of Acanthodaphne sabellii, sp. nov. have an obliquely thickened inner lip. Marwick (1931:150) was probably referring to a similar feature when he described the inner lip of Puha fulgida as “grooved longitudinally near its raised outer edge.” The Upper Oligocene to Recent genus Cryptodaphne Powell, 1942, may resemble Acanthodaphne, gen. nov. in having a biconic shell shape and a similar type of anal sinus. However, Cryptodaphne pseudodrillia Powell, 1942, type species of the genus, totally lacks axial folds with peripheral tubercles, possessing spiral keels and nu- merous, somewhat ‘‘Tomopleura-like”’ sigmoid threads over entire whorl surface. Shuto (1971) proposed Acamp- todaphne Shuto, 1971, as a monotypic subgenus of Cryp- todaphne Powell, 1942, with Pleurotomella biconica Schepman, 1913, selected as type species. In its biconic shape and distinct fasciole, Cryptodaphne (Acamptodaph- ne) biconica (Schepman, 1913) is superficially similar to Acanthodaphne sabellii, but differs in lacking a strong, tuberculate subsutural fold and in the weaker axial ele- ments (Schepman, 1913, p. 444 referred to ‘“‘rather in- cospicuous axial ribs, nearly disappearing in last whorl’). Both Schepman (1913) and Shuto (1971) referred to very coarse growth lines (“nearly rib-like”’) occurring on the subsutural ramp of C. (A.) biconica. This feature is not noted in the species here assigned to Acanthodaphne. Also, Acanthodaphne species have fewer spiral elements. Shuto (1971) described and figured the anal sinus of C. (A.) biconica (Schepman, 1913) which, if correctly inter- preted on the basis of the growth lines, differs from that of Acanthodaphne being deeper and differently shaped. Finally, the measurements available in literature (Schep- man, 1913; Shuto, 1971) indicate that the type species of Acamptodaphne is narrower than any member of Acan- thodaphne (b/| 0.41 compared to 0.47—0.55). The new genus differs from the Miocene to Recent Mioawateria Vella, 1954, in possessing much stronger axial folds, less gradate whorls, broader subsutural ramp and in the moderately deep anal sinus compared to vir- tually absent (growth lines indicating at most a weak in- sinuation). Gymnobela Verrill, 1884, as construed by Sysoev (1996, 1997) and Sysoev & Bouchet (2001), includes the species previously assigned to Bathybela Kobelt, 1905, Speoides Kuroda & Habe in Habe, 1962, and Theta Clarke, 1959, taxa retained at most as subgenera. In ad- dition, Sysoev (1996) noted a certain resemblance be- tween Mioawateria Vella, 1954, and Gymnobela Verrill, 1884, and stated that the former “‘may be either a large and very widely distributed genus or a synonym of Gym- Page 138 The Veliger, Vol. 48, No. 3 nobela.”’ The same author (Sysoev, 1997) subsequently retained Mioawateria as a full genus and referred to it the West African Gymnobela rhomboidea Thiele, 1925. Acanthodaphne, gen. nov., Mioawateria Vella, 1954, and Puha Marwick, 1931, possess a somewhat Gymno- bela-like. These three genera are first reported in the early Miocene of New Zealand. Despite the large number of species included and the very wide distribution, we know of no Gymnobela species based on fossil material and the genus never occurred in the Cenozoic of New Zealand (Beu & Maxwell, 1990). Members of Acanthodaphne, Mioawateria and Puha are rather small compared to Gymnobela species, which usually exceed 20 mm and may reach 70 mm in length, and further differ from Gym- nobela in details of sculpture and/or anal sinus. Acantho- daphne, for example, is readily distinguished from Gym- nobela by the more biconic shell with well developed subsutural fold bearing tubercles, a feature not occurring in the latter genus which has at most thickened scars of the anal sinus. In addition, Acanthodaphne has a greatly reduced number of spiral sculptural elements on spire whorls, a developed fasciole and may possess an oblique- ly thickened inner lip. Given the differences in morphology, geological oc- currence and the most probably polyphyletic nature of Gymnobela as currently used in literature, we consider Acanthodaphne, Mioawateria and Puha worthy of full generic status. Acanthodaphne may be compared to the Tertiary to Recent genus Buccinaria Kittl, 1887, but the former dif- fers from the latter in dimensions (maximum length 8.5 mm vs. more than 20 mm of typical Buccinaria species), different shape (biconic shell vs. ovate-pyriform or buc- cinoid) and reduced spiral sculpture. Furthermore, in the new genus the anal sinus is moderately deep while in Buccinaria it is broader and shallower. The New Zealand genus Awateria Suter, 1917, super- ficially resembles Acanthodaphne in having a well de- veloped subsutural fold and axial sculpture. However, Awateria species have very different proportions (the spire is 1.5 times height of aperture and canal according to Powell, 1942, and Beu & Maxwell, 1990). Powell (1942) regarded Awateria “‘closely allied”’ to the raphitomine genus Gymnobela but Beu & Maxwell (1990) assigned it to subfamily Borsoniinae (= Clathu- rellinae). Acanthodaphne sabellii Bonfitto & Morassi, sp. nov. (Figures 1—10) Diagnosis: Acanthodaphne with subsutural fold sculp- tured by somewhat comma-shaped tubercles about twice as numerous as folds. Last whorl with 14—16 axial folds extending across base but not onto neck. Maximum length 6.6 mm. Description: Shell small (up to 6.6 mm in length), heavy and biconic. Teleoconch consisting of about 4.5 whorls which are sharply angled below middle on early whorls, near middle on later ones. Subsutural ramp wide and gently concave. Whorls separated by a weakly impressed suture margined by a subsutural fold. Sculpture consisting of short, narrow, opisthocline folds extending from shoul- der angle, where they form prominent tubercles, to the lower suture on spire whorls and across base evanescing at level of the neck on the last whorl. Axial folds of rounded-triangular cross-section with interspaces wider than them. There are 12—15 axial folds on penultimate whorl, 14—16 on last. Subsutural fold sculptured by tu- bercles about twice as numerous as folds on later whorls. Spiral sculpture consisting of a narrow peripheral cord doubled during growth by a weaker cord anteriorly. In larger specimens, additional weak cord, just above lower suture, developed on later whorls. Last whorl with 2-3 threads in interspace between two main cords; below line of suture 2 cords present with | thread in each interspace, 3 widely spaced cords, and fine threads on the neck (10 in the holotype). Under SEM magnification (Figure 1H), entire whorl surface seen to be covered by dense rows of granules. Aperture pyriform. Inner lip wide, columella forming a distinct angle with parietal region. In some specimens, median area of inner lip obliquely thickened but not producing a pleat. Under SEM surface of inner lip seen to be covered with prickly nodules (Figure 1J). Siphonal canal short and shallowly notched. Outer lip without labral varix. Anal sinus moderately deep, re- versed L-shaped. Color white with yellowish protoconch. Protoconch conical, of up to 3.5 whorls, first half whorl (protoconch I stage) covered with minute spiral threads crossed by even finer axial threads giving a reticulate ap- pearance; subsequent part (protoconch II stage) sculp- tured by opisthocyrt axial riblets extending from suture to suture decussate by oblique threads on anterior two- Figures 1-10. Acanthodaphne sabellii Bonfitto & Morassi, sp. nov. Figure 1. Holotype (MZB 40664); scale bar 1 mm. Figures 2—3. Paratype (MZB 40666) from RS92/1. Figures 4—6. Protoconch; scale bar 100 pm. (MZB 40666). Figure 7. Teleoconch; scale bar 100 tm. (MZB 40664). Figure 8. Microsculpture of the teleoconch; scale bar = 10 pm. (MZB 40666). Figure 9. Anal sinus; scale bar = 500 wm. (MZB 40664). Figure 10. Inner lip prickly nodules; scale bar = 10 ym. (MZB 40666). A. Bonfitto & M. Morassi, 2005 Page 140 The Veliger, Vol. 48, No. WW A. Bonfitto & M. Morassi, 2005 thirds of each whorl. Protoconch maximum diameter: 0.55—0.62 mm. Dimensions (in mm): Holotype 6.5 X 3.3 mm (b/I 0.51), aperture height 3.1 mm (a/I 0.47); largest paratype: 6.6 X< 3.3 mm (b/I 0.50), aperture height 3.2 mm (a/I 0.48). The available material was found at the following two localities: No. Station Co-ordinates Depth specimens RS92/1 From 11°55'95"N—44°22'70"E 810 m 3 To 11°55'82"N—44°22'53"E 795 m RS92/2 From 12°02'36"N—44°29'53"E 1400 m 6 To 12°02'46"N—44°30'82”"E 1395 m Type deposition: Holotype MZB 40664 and four para- types MZB 40665 from Gulf of Aden (RS92/2), 1 para- type MNHN, unnumbered, same data; | paratype MZB 40666 from Gulf of Aden (RS92/1), 1 paratype IGNS, unnumbered, same data, | paratype ZMA, unnumbered, same data. Type locality: Gulf of Aden, station RS92/2. Etymology: This species is named after Professor Bruno Sabelli of the University of Bologna in recognition of his support to our studies. Remarks: Acanthodaphne sabellii, sp. nov. most closely resembles the larger (up to 6.6 mm vs. 7.4—8.5 mm in length) A. pusula (Laws, 1947), these two species differ- ing in relatively minor characters. Acanthodaphne sabellii has the subsutural tubercles about twice as numerous as folds on later whorls while in the paratype of A. pusula the tubercles are fewer (on penultimate whorl there are 11 folds and 16 tubercles). With regard to the axial sculp- ture of the last whorl, A. sabellii has stronger folds with sharper crests extending across base (but not onto neck) while in A. pusula the axials are rapidly fading below the periphery. In addition, while in A. pusula the last whorl surface bears numerous fine axial threads (Figure 2E) this feature is not noted in the new species. Acanthodaphne pusula (Laws, 1947) (Figures 11—15) Puha pusula Laws, 1947:539, pl. 55, fig. 7. Puha pusula Laws. Powell, 1966:131. Page 141 Buccinaria (?) pusula (Laws). Maxwell, 1988:68. Buccinaria (?) pusula (Laws). Beu & Maxwell, 1990:420. Type locality: Sandstone with lenses of fine conglomer- ate and shell grit about %4 mile below junction of Taita Stream and Waimamaku River, Hokianga Harbour area, Northland, New Zealand. Occurrence: Known only from the early Miocene (Otai- an) of Hokianga, North Island, New Zealand. Material examined: The paratype stored in IGNS, n°. T™ 1803. Remarks: Laws (1947) described the holotype of this species, stored in Geology Department of University of Aukland, New Zealand (accession number not provided), as measuring 8.5 X 4 mm (b/] 0.47), with 14 axial folds on the penultimate whorl. The paratype is smaller 7.4 X 3.9 mm (b/l 0.53), aperture height 3.4 mm (a/I] 0.46) with 11 folds on penultimate whorl. From the original figure (pl. 55, fig. 7), it would also seem that the holotype has a somewhat more prominent subsutural fold than the paratype. However, these differences appear minor and presumably taxonomically not significant. The proto- conch (fig. 2B) is damaged in the paratype examined but clearly of planktotrophic type with raphitomine sculpture. Under SEM (fig. 2C), the whorl surface is seen to be covered by dense rows of granules as in A. sabellii. Fine axial threads are clearly visible on the last whorl (fig. 2E). Acanthodaphne abbreviata (Schepman, 1913) (Figures 16—21) Pleurotomella abbreviata Schepman, 1913:83 (447), pl. 30, fig. 6. Cryptodaphne abbreviata (Schepman). Powell, 1966:127. Cryptodaphne abbreviata (Schepman). Sysoev, 1997:348. Buccinaria? abbreviata (Schepman). Shuto, 1971:12, pl. 2, fig. 1-3. Type locality: 835 m, blue mud, Ceram Sea, Indonesia, 02°40'S, 128°37.5’E.. Material examined: The syntype stored in ZMA collec- tion number 3.13.088. Remarks: Schepman (1913) originally described this species on the basis of two shells. Shuto (1971) provided descriptive remarks based on the examination of the type — Figures 11-15. Acanthodaphne pusula (Laws, 1947) (SEM photos from specimen uncoated). Figure 11. Paratype of Puha pusula Laws, 1947 (IGNZ TM 1803); scale bar = 1 mm. Figure 12. Protoconch; scale bar = 100 wm. Figure 13. Microsculpture of the teleoconch; scale bar = 100 ppm. Figure 14. Teleoconch whorl; scale bar = 500 wm. Figure 15. Microsculpture of the body whorl; scale bar = 500 pm. Figures 16-21. Acanthodaphne abbreviata (Schepman, 1913) (SEM photos from specimen uncoated). Figures 16— 17. Syntype of Pleurotomella abbreviata Schepman, 1913 (ZMA 3.13.088); scale bar = 1 mm. Figures 18-19. Protoconch; scale bar = 100 pm. Figure 20. Teleoconch; scale bar = 100 ym. Figure 21. Inner lip prickly nodules; scale bar = 10 pm. Page 142 material. However, only one specimen of the original two is presently stored in Zoologisch Museum Amsterdam, the second (3.13.089) being lost. This specimen, which presumably is the one figured by Shuto (1971), measures 5.6 X 3.1 mm (b/1 0.55), aperture height 2.9 mm (a/] 0.52) and has 14 axial folds on penultimate whorl. The proto- conch (Figure 3C—D) is worn but clearly planktotrophic with traces of decussate riblets, and a maximum diameter of about 0.56 mm. The shell surface microgranulation occurring in both A. sabellii and A. pusula appears to be abraded in this specimen. However, the prickly nodules on the inner lip are evident (Figure 3F). As stated above, Powell (1966), followed by Sysoev (1997), assigned the species to genus Cryptodaphne Pow- ell, 1942 but did not provide elements supporting this attribution. We agree with Maxwell (1988:68) in consid- ering Pleurotomella abbreviata very similar in supra- specific features to Puha pusula, these two taxa are ac- cordingly here assigned to the new genus Acanthodaphne. However, A. abbreviata differs from its congeners in pos- sessing the peripheral tubercles just above lower suture (Figure 3A—E) on all teleoconch whorls while in the two other Acanthodaphne species the periphery is near middle on later whorls. Acknowledgments. We express our gratitude to Alan Beu (IGNS) and Robert Moolenbeek (ZMA) for the loan of type spec- imens. Bruno Sabelli (MZB) provided his valuable advice on the present manuscript. LITERATURE CITED Beu, A. G. & P. A. MAXwELL. 1990. Cenozoic Mollusca of New Zealand. New Zealand Geological Survey Paleontological Bulletin 58:1—518. Bouchet, P. & A. WaAREN. 1980. Revision of the north-east At- lantic bathyal and abyssal Turridae. Journal of Molluscan Studies, 8(Suppl.):1-119. Laws, C. R. 1947. Tertiary Mollusca from Hokianga district, North Auckland. Transactions of the Royal Society of New Zealand 76(4):537—-541. MACNEIL, FS. 1960. Tertiary and Quaternary Gastropoda of The Veliger, Vol. 48, No. 3 Okinawa. United States Geological Professional Paper 339: iv + 148 pp. Marwick, J. 1931. The Tertiary Mollusca of the Gisborne Dis- trict. New Zealand Geological Survey Paleontological Bul- letin 13:1—177. MAXWELL, P. A. 1988. Late Miocene deep-water Mollusca from the Stillwater Mudstone at Greymouth, Westland, New Zea- land: paleoecology and systematics. New Zealand Geologi- cal Survey Paleontological Bulletin 55:1—120. POWELL, A. W. B. 1942. The New Zealand Recent and fossil Mollusca of the family Turridae with general notes on turrid nomenclature and systematics. Bulletin of the Auckland In- stitute and Museum 2:1-188. PowELL, A. W. B. 1966. The molluscan families Speightiidae and Turridae an evaluation of the valid taxa, both Recent and fossil, with lists of characteristic species. Bulletin of the Aukland Institute and Museum 5:1—184. ROSENBERG, G. 1998. Reproducibility of results in phylogenetic analysis of mollusks: a reanalysis of the Taylor, Kantor, and Sysoev (1993) data set for conoidean gastropods. American Malacological Bulletin 14(2):219—228. SCHEPMAN, M. M. 1913. The Prosobranchia of the Siboga Ex- pedition. Part 5. Toxoglossa. Resultats Siboga-Expeditie, monograph, 49-1, (5)365—452, pls. 25-30. SHuTo, T. 1971. Taxonomical notes on the turrids of the Siboga- Collection originally described by M. N. Schepman, 1913 (part 3). Venus 30(1):5—22. SysoeEv, A. V. 1996. Deep-sea conoidean gastropods collected by the John Murray Expedition, 1933-34. Bulletin of the Nat- ural History Museum of London, Zoology 62(1):1—30. SysoEv, A. V. 1997. Mollusca Gastropoda: new deep-water turrid gastropods (Conoidea) from eastern Indonesia. Mémoires du Muséum National d’Histoire naturelle [Paris], série A, Zoologie 172:325—355. Sysoev, A. V. & P. BoucHeT. 2001. New and uncommon turri- form gastropods (Gastropoda: Conoidea) from the south- west Pacific. Pp. 271-320 in P. Bouchet & B. A. Marshall (eds.), Tropical Deep-sea Benthos. Vol. 22. Mémoires du Muséum National d’ Histoire naturelle [Paris], serie A, Zoo- logie 185. TayLor, J. D., Y. I. KANToR & A. V. Sysoev. 1993. Foregut anatomy, feeding mechanisms, relationships and classifica- tion of the Conoidea (= Toxoglossa) (Gastropoda). Bulletin of the Natural History Museum of London, Zoology 59(2): 125-170. THE VELIGER © CMS, Inc., 2006 The Veliger 48(3):143—150 (November 2, 2006) Growth and Activity Patterns in a Backyard Population of the Banana Slug, Ariolimax columbianus ANITA K. PEARSON, OLIVER P. PEARSON,* AND PETER L. RALPH Museum of Vertebrate Zoology, University of California, Berkeley, California 94720, USA (e-mail: opearson@ vdn.com) Abstract. The study was carried out on and around a redwood deck in a suburban area of central California, using bait-plates to attract nearby banana slugs. Over a period of two years 150 individual Ariolimax columbianus weighing between 0.4 and 75.4 g were identified using photographs of the patterns on their maculated bodies. In many subsequent sightings of these slugs, we recorded their weight, their appearance, the time and location, and their activity. In both 1999 and 2000, slugs weighing less than a gram appeared in January and, feeding both day and night, gained weight at an average rate of 0.11 g/day. The weight of individual slugs fluctuates widely, and only below 20 g does weight correlate with the age of a slug. Maximal weights are reached in summer. In the following autumn months, slugs may lose up to 50% of their maximal weight. Within one day, feeding slugs showed an average weight increase of 10.12%. Slugs over 20 grams appeared at the bait-plates during early morning hours and towards dusk; they did not feed during the night. Slugs frequented specific sites, usually appearing at the same or nearby bait-plates, or repeatedly selecting the same night roosting site, and little long distance movement was seen. Slugs recorded on the deck did not cross the grass to bait-plates 4.8 m away. They displayed a cyclic behavior, appearing at a bait-plate for one or two days followed by an absence of an equal or longer period before reappearing. Adult slugs did not feed at bait-trays with many young slugs. On a July day, 46 individual slugs weighing a total of 750 g were recorded in an area of approximately 60 square meters. Our evidence indicates that these slugs do not live much longer than two years. The number of slugs on the deck was highest with temperatures between 8° and 16°C and a relative humidity near 75%. On warm September days, slug numbers on the deck declined as temperatures rose and humidity fell. On cool, wet March days, their numbers increased with the temperature. INTRODUCTION b; Rollo, 1983a, b; Rollo & Wellington, 1981); others have described their role in forest dynamics (Richter, 1979; Gervais et al., 1998). Few publications are con- cerned with the behavior of unconfined banana slugs, however, and the most extensive study of this kind, in a dissertation, remains largely unpublished (Richter, 1976). In the small area of our study, most A. columbianus have melanistic areas on their dark, mustard-colored bod- ies. The pattern of these areas is distinct in young animals and remains as the slugs grow, thus providing a non- invasive means of identifying individuals. We have taken advantage of this fact to record the growth and activity of slugs in our local population, and to learn more of their biology. The term banana slug is applied to several large terrestrial gastropods of the genus Ariolimax occurring in western North America. The taxonomy within the genus has been complicated by the many color variations encountered: individuals within a population were found to vary from white, to yellow to brown, to almost black, and many may be monochromatic or maculated. In a revision of the genus, Mead (1943) identified species on the basis of gen- ital morphology. Ariolimax columbianus (Gould), a large slug well known in forested areas of coastal California from the Salinas Valley northward to Alaska, was rec- ognized as the only species of banana slug with both monochromatic and maculated individuals. The literature on slugs is greatly biased towards those that are agricultural pests, and there is little information about benign native slugs such as Ariolimax columbianus. Of the few existing studies, some have been concerned primarily with the reactions of these slugs in artificially controlled situations (i.e., Hamilton & Wellington, 1981a, METHODS The study site, in Orinda, Contra Costa County, Califor- nia, is located on the eastern side of the inner coast range. The climate is Mediterranean, with mild temperatures and with moderate rainfall between October and March. Al- though we have watched slugs on our property for many * Deceased March 4, 2003. years, the observations presented here were made be- Page 144 ae Bait- SOS Vi Geranium box o Bait-plate Figure 1. Diagram of the site where the study took place. The boards of the redwood deck are indicated. Containers with plants are shaded dark; grass and shrubbery are lightly shaded; deck furniture is unshaded. A brick wall lines the grassy area. tween May 1998 and May 2000, on and around a deck composed of 14 * 14 cm redwood planks separated by 6.5 mm spaces. The deck, 27 square meters in size, is at 37.9035°N;122.17508°W (datum WGS 85), and is a meter above the earth on the west side on the house (Figure 1). Observations were continuous over these two years, ex- cept for a period in 1998 from mid-October to the end of December. The house is on several uncultivated acres with California bay and coast live oak trees, shrubby ar- eas including coyote brush, blackberry, and poison oak. A few redwood trees planted 60 years ago are 6 meters from the western edge of the deck where our observations were made. In summer, plant containers on the deck are watered by a drip system, and the lawn between the deck and a brick retaining wall also is regularly watered. Over the two years of our study we placed on the deck a 27 X 35 cm aluminum tray, which, when we were in residence, was continuously supplied with lettuce, melon rind, cucumber, zucchini, apples, and occasional other food items. After the first three months, four 20 cm, round aluminum pie plates with similar food items as bait were added at 1.5 m intervals under the ferns and camellias which border the deck but are a meter below the level of the deck. For short periods (one month in 1998; one week in 1999), baited pie plates also were placed along the brick wall, a distance of 4.8 m from the deck. At each observation we recorded and identified slugs at bait-plates and at any other site on the deck, such as on a potted plant or on adjacent walls of the house. Ob- servations were made at irregular time intervals, both day and night, up to 27 observations in a 24-hour period. To identify individual slugs, we focused on the pig- mented pattern of the body just posterior to the mantle on the slug’s left side. Originally we sketched the pattern of dark spots in this area; later we photographed each slug facing left on a ruler (Figure 2). The photographs were valuable in identifying individuals, as slug appear- ance changes dramatically with posture, but patterns re- mained distinct and discernable. The slugs were trans- The Veliger, Vol. 48, No. 3 - eee G\ = OO; t wisi ca nid tnd iii ihaunatainl 2 oe GI fea) £\ oN S bili ilitihi } lili rhitili| lelitiriretil islileee height +| \r\i\adaeelalbrhahatehdiihihaly Figure 2. Photographs of slugs (#98-32, 3.2 g & #98-90, 31.1 g) showing areas posterior to the mantle that were used for iden- tification. ported inside the house on an index card to be photo- graphed, assigned an identifying number, and weighed on an electronic balance (Acculab Pocket Pro 150B, Newton, PA). Within five minutes they were returned to the site where they had been found. Date, time, location, activity and weights were recorded. Slugs were reweighed and rephotographed at irregular intervals thereafter; other data were recorded each time a slug was seen, without distur- bance to the slug. We identified and assigned numbers to 150 A. colum- bianus that weighed between 0.4 and 75.4 grams. We refer to those under 2 g as juveniles, from 2—20 g as sub- adult, from 21—50 g as adult, and above 50 g as large adult. Our handling methods apparently did not distress the slugs, for usually they continued their observed be- havior when returned to the site where they were found. We did not identify individually the numerous small slugs (i.e., <5 g), but ascertained their numbers in 1999 at ap- proximately two-week intervals by retaining for a day all small slugs that appeared on the bait-plates. They were then returned to the plates from which they had come. Over the two years of this study, the temperature and relative humidity were recorded with a Weather Monitor Il and accompanying software (Davis Instruments, Hay- ward, CA). The Monitor was situated nine feet above the deck, under the eaves on the side of the house. Data were processed using R (www.r1-project.org; pro- duced most of the figures), SPSS 11.0 and Quattro Pro 5.0 (Borland, Inc.). Robust regression using an iteratively reweighed least squares procedure with a Tukey bisquare weighing function (Huber, 1981) produced the initial growth rate estimates. Only the first year of data was used to capture the initial growth pattern for each cohort..A A. K. Pearson et al., 2005 robust regression was used because of the high variability of slug weight. RESULTS Appearance Ninety-five percent of the slugs in our study had a con- spicuous round black spot (about 3 mm diameter) in the center of the mantle and assorted melanistic spots on the body. Those without spots on the mantle lacked spots on the body, or had only a few faint spots. A few slugs could be recognized by superficial irregularities (possibly old wounds?) on their bodies. The smallest slugs that ap- peared on the bait-plates (<0.1 g) had markings similar to adults, with one conspicuous spot on the mantle and others on the body. Thirteen slugs had gouges and nicks on their bodies, including four slugs known to be over a year old. One slug was missing an eye stalk in addition to wounds on its back and mantle. A slug first identified at 3.3 g had a gash on its 38.9 g body 10 months later (May, 1999). After a further three months (August, 1999), the same slug (now 51.6 g) had a nick on the edge of its mantle. By September it had several more mantle nicks as well as more wounds on its body, and it was last seen in early October. This slug had been seen on 91 different days in the 15 months we had been recording it. We have no evidence concerning the source of the wounds, although slugs are known to have a variety of predators (Harper, 1988). Two dead slugs were seen. One, a squashed juvenile on the deck, was being eaten by an adult. The other was a dead adult slug, covered with mucous, in leaf litter near the deck, with no clues as to the cause of death. Reproduction In two years of observations we failed to see a mating pair or to find a cluster of Ariolimax eggs. However, on January 18, 1999 and on January 15, 2000, the first ju- venile Ariolimax appeared on bait-plates below the deck. The 1999 juvenile weighed 0.6 g; that in 2000 weighed 0.3 g, which is the average weight of an egg of this spe- cies (Rollo, 1983a). Subsequently, we recorded juvenile slugs on the bait-plates weighing as little as 0.1 g. Several slugs less than 2 g appeared on the bait-plates in February and March 1999, and a few appeared as late as July (Fig- ure 3). When the 1999 cohort of young first appeared on the deck during the day in March 1999, there were often as many as eight of the 1998 cohort (now adults, 21—50 g) feeding on the bait-tray together, apparently tolerating each other’s presence. As time passed, the young cohort of 1999 appeared in greater numbers during the day, and such aggregations of adults were no longer seen. By May Page 145 80 60 4 ° = 40 4 o oD 20 4 : 7 : ae f Pa a 04 ere | May 98 1 Sept 98 1 Jan99_ 1 May 99 1 Sept 99 1 Jan00 1 May 00 Figure 3. Weights of Ariolimax columbianus from May 1998 to June 2000. No observations were made from mid-October to late December, 1998. Regression lines for the 1998 and 1999 cohorts were calculated using robust regression on the first year of data for each cohort. The growth rate for 1999 (a weight gain of 0.11 + 0.003 g/day) seems more reliable because it includes data for the youngest slugs. 1999 it was usual to find only one adult slug with several sub-adults on the bait-plate, and sometimes only the latter. Weights of Slugs In the two years of our study, we identified and as- signed numbers to over 150 Ariolimax columbianus ini- tially weighing between 0.4 and 75.4 g. Subsequently, those that were seen again were reweighed at irregular intervals, some more than 80 times. These data, plus the weights of unidentified juveniles and sub-adults, also pre- sented in Figure 3, indicate that young slugs gain an av- erage of 0.11 + 0.003 g per day. A monthly analysis of the composition of the slug population by weight groups also supports a rapid weight increase, as numbers of ju- veniles rapidly become sub-adults (Figure 4). When initially identified, there were 12 slugs that weighed more than 50 g (i.e., large adults), and five of these were not seen again. Most large adults in the pop- ulation (Figures 3, 4) were slugs initially identified as 20 to 50 g adults (11 individuals) or as 2 to 20 g sub-adults (12 individuals). Weight records of four of the longest- recorded individuals show wide weight fluctuations (Fig- ure 5); maximal weight was recorded during the summer months and weight declined thereafter. One of these was first recorded as a sub-adult (12.8 g) in August, 1998. The regression lines of Figure 3 suggest an age of 5 months for a slug of this weight. A year later it weighed 54.0 g, and in subsequent months its weight fluctuated between 40 and 50 g in September and October, and be- tween 30 and 40 g after that. When last seen on April 22, 2000, it weighed 39.4 g and was approximately 26 months old. Over 20 months, we had recorded this slug i 40 bb na Ge 3 2 3 FI EI SIN May '98 Sept '98 Jan '99 May '99 Sept '99 Jan '00 May '00 Composition of Slug Population Figure 4. Age composition of the slug population in the area of our study during two years. The mean weight gain of a slug during each month was calculated to determine to which class it belonged. For the <2 g (juveniles) and 2—20 g (sub-adult) clas- ses, in which many slugs were not individually identified, a min- imal number was determined by keeping such slugs captive throughout the day they were weighed. Slugs 21—50O g are re- ferred to as adults; over 50 g are large adults. on 62 days. Using a similar approach with 16 slugs for which we have lengthy records, the estimated age when last seen varied from 13 to 26 months, with a mean of 18.3 months (SD * 3.10). Although weights of individual slugs fluctuate, some patterns are discernible. Many slugs first recorded in May and June, 1998, weighed between five and ten grams, indicating an age of two or three months (Figure 3). Dur- ing the following months they gradually gained weight, but in September and October this cohort of slugs—now ranging from 35 to 74 g—started to lose weight. Some lost a great deal of weight (53%, 43%, 41% of their max- imal weight), others lost a more moderate 10—25% of maximum. A slug that originally weighed 8.0 g in July 1998, increased to a maximal 74.1 g in early September of the following year. After an absence of three weeks, it weighed 54.5 g, and its weight fluctuated about this level until last seen at the end of October. We have 67 records of slugs that were weighed twice within one day, many weighed before and after a feeding bout. Five of the 67 lost weight (0.01 to 3.6 g); the weight increases of the remaining 62 ranged from 0.1 to 9.8 g, with an average of 3.48 g (SD + 2.35), an increase of 10.12% (SD + 7.48). Smaller slugs generally showed larger percentage weight gains. The largest recorded gain was a Slug that on a rainy February day, spent 105 min- utes on the bait-tray. At the beginning of the feeding bout it weighed 26.4 g; at the end it weighed 35.7 g, an in- crease of 35.2%. Although uptake of moisture may have contributed to this increase, similar percentage increases in weight were also recorded in the dry month of. August. There does not appear to be a correlation between the time of year and the amount of weight a slug may gain within one day. The Veliger, Vol. 48, No. 3 a @ 98.45 g 70 “al S ¢. Be : g = oe. = _ 605 i om 50 =» 40 4 y 2 3( Lene ees BEA 4 205 OS een we tere eee 4x 98.69 Oa) ige saree “4 Sa a a a T 1 June 98 1 Oct 98 1 Feb99 | June99 1 Oct 99 1 Feb00 1 June 00 Figure 5. Weight records of four slugs tracked for more than a year. The numbers (98.69, etc.) indicate the individuals. On several days we weighed once all the slugs that appeared on the bait-tray on the deck and the four bait- plates below it throughout the day, keeping unidentified young slugs in containers to avoid duplications. As many as 27 to 52 slugs, with a combined weight approaching a kilo, were living on and around the deck (an area of less than 100 square meters) on these days (Table 1). When feeding at night on bait-plates below the deck or at the brick wall, young Ariolimax were often found in the company of three species of small slugs, sometimes a snail, and occasionally, isopods. Movements Individual slugs showed a tendency to be found re- peatedly in the same area and at the same bait-plate. One sub-adult slug, first recorded at 12.3 g on an hydrangea plant in June, 1998, was subsequently observed on the same plant on six different days in the months that fol- lowed, and five more days in May and June of the fol- lowing year, when it weighed 65.4 g. Another slug, re- corded in a planter box with geraniums, was transferred on four successive days to the hydrangea 1.5 m away. Each time, it appeared back in the geranium box the day following the transfer. On several occasions we recorded the speed at which large adult slugs progressed across the boards of the deck. Values ranged from 11.7 to 19.2 cm/min. Slugs weighing 50 g were seen going through the 6.5 mm gap between the boards of the redwood deck, and a slug of similar size was seen to make its way through leaf litter and disappear into a one centimeter hole in the earth. Slugs climbed as far as three meters up the painted redwood siding of the house, leaving a slime trail that remained visible for several months. Slugs may remain in an extended position on the wall, but often formed a loop, frequently with the head down. With their mouth near the mucous pore at the rear of their body, they could be seen consuming the mucous and anything that has stuck to it. In the morning hours of all months, a few slugs were found on the wall and on the brick chimney, and it was usually the same individuals. At night, how- ever, wall-roosting had a different pattern. Adult slugs A. K. Pearson et al., 2005 Page 147 Table | Census of slugs on the deck and on 4 bait-plates below it, on selected days in 1999. Juveniles Sub-adults Adults Date (<2 gm) (2-20 gm) (20-50 gm) May 20 4 28 3 June 10 3 41 4 July 14 2 33 8 Aug. 19 1 19 4 Sept. 12 0) 27 4 Large adults Total (>50 gm) number Total weight 3 38 368.5 gm 52 628.0 gm 3 46 750.2 gm 3 Di, 531.3 gm 3 34 624.5 gm were on the wall at night only in the winter months of January to March; in August, only sub-adults (2—20 g) were found there. A slug may spend the entire night with eye stalks retracted, immobile on the wall, descending as morning approaches. Several slugs spent 12 to 19 hours in place on the wall, and one adult slug remained in situ over 24 hours. A coiled 10 cm string of excreta often remained behind when a large slug descended from a wall. Of slugs that were first identified on the deck, 71% of those that were seen again were also found on bait-plates below the deck. Slugs recorded first at the bait-plates be- low the deck more consistently remained in their shady original sites, only 32% appearing also on the deck. Of the 114 slugs originally identified on the deck or the area below it, none was found across the grass at the brick wall 4.8 m away. Six of the 36 slugs first recorded at the brick wall subsequently appeared on bait-plates below the deck. In September, a slug was discovered on the opposite side of the house, 28.4 meters from the site near the brick wall where it had been recorded two months earlier. This is the farthest an identified slug was found from its orig- inal location. 80 | 4 relative humidity (%) 60 4. 40 | temperature (C) 20 fen a heal #/\of slugs 0 ! 1 } i | rirybpe ry vf fy fo of © fo. hh rw 1 3 5 7 9 11 13 15 17 19 21 23 August Figure 6. Records of the first three weeks of August 1998, showing the number of slugs on the deck relative to temperature and relative humidity. The vertical shaded areas represent night- time. Numerous observations that found no slugs are shown on the zero line. Time of Activity No adult slugs were observed feeding during hours of darkness. When recorded on the deck at night, adult slugs were inactive, usually attached to the walls or doors of the house. They appeared at the bait-plates primarily dur- ing early morning hours; a few appeared towards dusk. In the dry summer of Orinda, when daytime temperatures may reach 30°C, slugs appeared on the deck during lows in the temperature cycle and highs in the cycle of relative humidity (Figures 6, 7): their numbers decreased as the temperature rose and humidity dropped. During the cool and humid winter conditions of March, slug numbers par- alleled the rise in temperature from lows of 7°C (Figure 8). Consistently, the number of slugs on the deck was highest with temperatures between 8° and 16°C. Few slugs appeared on mornings when there was frost on the deck, and usually it was the same one or two slugs that were present. In contrast to the diurnal feeding pattern of adults, ju- venile slugs, first appearing in January, fed both day and night on bait-plates below the deck and at the brick wall, sites which offered more cover than did the bait-tray on the deck. In late March, 1999, five young slugs (0.6, 1.2, 1.9, 2.4 and 2.4 g) were the first to make their way up through the cracks of the deck and were seen feeding on the bait-tray at 9:25 PM. Although in the following month they fed on the deck both day and night, sub-adult slugs were more often observed feeding on the deck at night. By the end of May some of this cohort of slugs weighed as much as 8 g and they no longer fed at night. Patterns of Activity Slugs over 50 g (adults) usually appeared alone on bait- plates, or with sub-adults (2—20 g), which they appeared to ignore. Although slugs occasionally visited a bait-plate and departed immediately, most remained between one and two hours. Longer bouts were not unusual. One slug remained on the deck bait-tray for ten hours, although it is not known how much of that time it was feeding. Analysis of the extensive records of five individuals shows a pronounced tendency for slugs to appear at the Page 148 The Veliger, Vol. 48, No. 3 August 1998 MB Mean Number of Slugs ES Temperature (C) [] Percent Relative Humidity 10 0:00 1:30 3:00 4:30 6:00 7:30 9:00 10:30 ma © Figure 7. 12:00 Average number of slugs on the deck and walls for each half-hour interval in August 1998 in relation to | | I ip i TORE Th | aT ah : i t i q | MAMET 13:30 © 15:00 18:00 1930 21-00 temperature and relative humidity. Shaded areas indicate sunrise and sunset on August 15, Pacific Standard Time. bait-plates for one or two consecutive days, and then to be absent for longer intervals. Consecutive days recorded at a bait-plate varied from one to five, but the great ma- jority of the 193 visits of these individuals were one day (70.5%) or two days (21.8%). Intervals between visits to the bait-plates were more spread out, ranging from one day (26.0%), two days (22.0%), to longer than 10 days (11.0%). The long absence intervals did not appear to be correlated with the seasons. DISCUSSION Approximately a third of the identified slugs, in all size categories, were seen only once (45 slugs), although there was no obvious reason for their disappearance. Others were seen repeatedly during the next two years. One slug was recorded numerous times on 89 different days during the two years of our observations. In 1998 we identified 86 slugs on and around the deck and within a few meters of our house. In January of the following year 25 of these slugs were still being recorded. By June 1999 we were seeing only 15 of the original slugs, and by January 2000, only one still survived. Meanwhile unidentified slugs (85% of them adults and large adults) were appearing: 33 in 1999 and 29 in the first months of 2000. When com- bined, these data indicate that within this 160 square me- ter area near our house, there was a large population of slugs with a limited life span and rapid turnover. There are striking differences between our observa- tions and the field observations of Richter (1976) on the same species in Washington State. In Orinda, the Arioli- max columbianus are remarkably larger than those noted in Richter’s study, which averaged 25 to 35 g and weighed a maximum of 45 g. Many slugs in our study weighed over 50 g. Additonally, in July 2003, three years after our supplemental feeding ended, an unidentified slug weighing 107 g appeared on the deck, demonstrating that large size in the local populations was not dependent on supplied food. The weight gain of young slugs in Richter’s popula- tions was estimated to be 0.2 to 0.3 g per month (Richter, 1976); the well-documented gain in weight of our young slugs was 10 times that. In Washington State, slugs were active and feeding both day and night from May to June, and were only seen between April and November, after which they disappeared into hibernation (Richter, 1976). In Orinda, adult slugs did not feed at night and appeared at the bait-plates throughout the year. In the Seattle area, the eggs of banana slugs hatch in late February and early March; in Orinda, recently hatched slugs appeared from January to as late as April. Climate differences may be partly responsible for these discrepancies. Average tem- peratures in Orinda are somewhat higher in winter, but in summer they are quite similar to those in Richter’s study areas. Another possible explanation may be that the spe- cies identifications of Mead (1943), based on genital mor- phology, are not valid. Recent studies indicate that the systematics of ariolimacine taxa are poorly resolved (Roth, 2004; Pearse et al., 2004). Up to 20 grams, the weight of a banana slug is a good indicator of the age group to which it belongs. Above this weight, however, slug weight fluctuates widely. Some of our monitored slugs continued to increase in weight; oth- ers remained hovering about 40 grams. Some weight fluc- tuations are probably due to the problems of maintaining hydration. Richter (1976) was concerned that egg pro- duction might mask normal increases in body weight. The A. K. Pearson et al., 2005 Page 149 March 1999 32 MB Mean Number of Slugs ISS Temperature (C) 24 [4 Percent Relative Humidity/10 16 a fl I] f MM A Al Ale ! | | TTT TA iit } WT i i] | ie | H | | | i H il i H i UAL ! A | 0:00 1:30 3:00 4:30 6:00 7:30 9:00 8610:30 Figure 8. | mt HY 13:30 18:00 §=$16:30 18:00 19:30 Ae 22:30 Average number of slugs on the deck and walls for each half-hour interval in March 1999 in relation to temperature and relative humidity. Shaded areas indicate sunrise and sunset on March 15, Pacific Standard Time. clutches of Ariolimax columbianus, are relatively small for a mollusk (an average 17.7 eggs), and eggs weigh an average 0.32 gm (Rollo, 1983a). A clutch of eggs, there- fore, would entail an increase in weight of approximately 6 grams, which is a modest weight change in the popu- lation we were observing. The weight fluctuations of slugs in our study probably can not be attributed to re- productive events. The dynamics of the slug population described sug- gested that habituation, a constant food supply, and a high reproductive potential contributed to the large numbers of slugs that appeared on and around the deck. The surge in numbers documented in 1999 could have been produced by a few reproductive individuals. Similar surges were occurring around the deck and at the brick wall 4.8m away, however, with little intermingling of the two pop- ulations. These observations, in addition to the noted homing tendency of slugs (Rollo & Wellington, 1981), suggest that what we were recording was the potential of this slug to expand in numbers under beneficial condi- tions - a potential reflected in the numerous clades exist- ing throughout its range. It has been stated repeatedly that Ariolimax columbi- anus has a life span between 4 and 6 years (Gordon, 1995; Richter, 1976; Rollo et al., 1983; and others). Our study finds no evidence that these slugs live more than a little over two years. Acknowledgments. The authors express their gratitude to Dr. Richard Sage for his advice and encouragement throughout the project, to Dr. James Patton, Dr. Carole Hickman, Dr. Barry Roth, Dr. Patricia Woolley and two unknown reviewers for helpful comments on the manuscript, and to Karen Klitz for her render- ing of Figure 1. LITERATURE CITED GERVAIS, J. A., A. TRAVESET & M. E WILLSON. 1998. The po- tential for seed dispersal by the banana slug (Ariolimax col- umbianus). American Midland Naturalist 140:103—110. Gorpon, D. G. 1995. Banana slugs: partners in slime. Pacific Discovery (Sept/Oct):42—43. HAMILTON, P. A. & W. G. WELLINGTON. 1981la. The effects of food and density on the movement of Arion ater and Ariol- imax columbianus (Pulmonata: Stylommatophora) between habitats. Researches on Population Ecology 23:299-308. HAMILTON, P. A. & W. G. WELLINGTON. 1981b. The effects of food supply and density on the noctural behaviour of Arion ater and Ariolimax columbianus (Pulmonata: Stylommato- phora). Researches on Population Ecology 23:309-317. Harper, A. B. 1988. The Banana Slug. Bay Leaves Press: Aptos, California. 32 pp. Huser, P. J. 1981. Robust Statistics. Wiley: New York. 308 pp. Meap, A. R. 1943. Revision of the giant west coast land slugs of the genus Ariolimax Moerch (Pulmonata: Arionidae). American Midland Naturalist 30:675—717. Pearse, J. S., J. L. LEONARD, K. BREUGELMANS & T. BACKELJAU. 2004. COI data reveal surprises in clades of banana slugs (Stylommatophora: Arionidae, Genus Ariolimax). Meetings of The Society for Integrative and Comparative Biology, New Orleans, Louisiana. (abstract). RICHTER, K. O. 1976. The foraging ecology of the banana slug Ariolimax columbianus, Gould (Arionidae). Ph.D. Disserta- tion, University of Washington, Seattle. 228 pp. RICHTER, K. O. 1979. Aspects of nutrient cycling by Ariolimax columbianus (Mollusca, Arionidae) in Pacific Northwest co- niferous forests. Pedobiologia 19:60—74. RoLLo, C. D. 1983a. Consequences of competition on the repro- duction and mortality of three species of terrestrial slugs. Researches on Population Ecology 25:20—43. RoLLo, C. D. 1983b. Consequences of competition on the time budgets, growth and distributions of three species of terres- trial slugs. Researches on Population Ecology 25:44—68. RoLio, C. D. & W. G. WELLINGTON. 1981. Environmental ori- entations by terrestrial Mollusca with particular reference to homing behavior. Canadian Journal of Zoology 59:225—239. Page 150 The Veliger, Vol. 48, No. 3 RoLLo, C. D., I. B. VERTINSKy, W. G. WELLINGTON & V. K. Rotn, B. 2004. Observations on the taxonomy and range of Hes- KANETKAR. 1983. Alternative risk-taking styles: the case of perarion Simroth, 1891 and the evidence for genital poly- time-budgeting strategies of terrestrial gastropods. Research- morphism in Ariolimax Morch, 1860 (Gastropoda: Pulmon- es on Population Ecology 25:321—335. ata: Arionidae: Ariolimacinae). The Veliger 47:38—46. The Veliger 48(3):151—169 (November 2, 2006) THE VELIGER © CMS, Inc., 2006 Lower Eocene Gastropods from the El Bosque Formation, Central Chiapas, Mexico MARIA DEL CARMEN PERRILLIAT,! JAVIER AVENDANO,? FRANCISCO J. VEGA,! AND JESUS SOLE! ' Instituto de Geologia, UNAM, Ciudad Universitaria, 04510, Mexico, D. FE, Mexico > Institudo de Historia Natural y Ecologia, Calzada de los Hombres Ilustres s/n, 29000—Tuxtla Gutiérrez, Chiapas, Mexico Abstract. Gastropods from the Lower Eocene El Bosque Formation in central Chiapas, Mexico, are reported in this contribution. Thirty-two species are described and figured, four of them are new species—Calliostoma granulata, Crom- mium globosa, Lyrischapa spinifera, and Volutocorbis minutus—and one new subspecies, Palmerella mortoni mexicana, is erected. The age for the sediments of the studied locality is redefined based on *’Sr/*°Sr analysis of well preserved Hercoglossa cf. H. ulrichi specimens as being Ypresian (52 Ma). Stratigraphic range of most of the species here described is concordant with the age inferred from isotopic data. Paleobiogeographic affinities with the Atlantic and Gulf Coastal Provinces are also confirmed, and some have affinities with the Tethyan Province. INTRODUCTION In recent years, a variety of fossil invertebrates that in- clude corals, crustaceans, and mollusks have been re- ported from locality IHN 1005, near Veinte de Noviem- bre, Depresion Central of Chiapas (Figure 1). Strata at this locality, originally thought to be part of the Middle Eocene San Juan Formation, have been dated as Early Eocene (Wilcox/Ypresian), using radiometric data from well-preserved nautiloid shells of Hercoglossa sp. cf. H. ulrichi (White, 1882). This species, described from the Midway Group of Arkansas, has a stratigraphic range of Paleocene-Lower Eocene (Miller, 1947). Several nauti- loid specimens were collected from the studied locality, most of them of relatively small size (Figure 2). *’Sr/*°Sr analysis made by one of us (Solé) yields a value of 0.707684 + 0.000008 (25) that, according to scales of Gradstein & Ogg (1996) and Odin (1994), is approxi- mately equivalent to 52 Ma (Figure 3). According to the stratigraphy and lithology described previously for the study area (Ferrusquia-Villafranca, 1996), we reassign lo- cality IHN 1005 to the lower part of the El Bosque For- mation. Paleobiogeographic affinities of the fauna described are consistant with Tethyan influence. STRATIGRAPHY AND PALEOENVIRONMENT Continental red beds exposed in Mesa Telestaquin, Soy- al6, and El Bosque were defined as the El Bosque For- mation (Allison, 1967). This unit was named as the Te- lestaquin Mesa Sandstone by Frost & Langenheim (1974), based on more complete and accessible outcrops in the area of Soyal6. However, Ferrusquia-Villafranca (1996) redefined this unit and suggested that the original name El Bosque Formation be preserved. Outcrops of the El Bosque Formation are scattered in the Ixtapa-Soyal6 area, and the formation has been recognized in the De- presion Central of Chiapas, where Frost & Langenheim (1974) suggested that this unit 1s present in the Copoya syncline. Based on a few molluscan species and benthic foraminifera, Frost & Langenheim (1974) and Ferrusquia- Villafranca (1996) suggested an Early Eocene age for the El Bosque Formation. Stratigraphic distribution of eight species here reported confirm an Early Eocene age for locality IHN 1005 (Table 1). This is the first report of the El Bosque Formation in Veinte de Noviembre area, where a thickness of less than 200 m is estimated. Thickness for this unit elsewhere was reported to be between 720 and 2000 m (Frost & Langenheim, 1974; Ferrusquia-Villa- franca, 1996). At Veinte de Noviembre, the El Bosque Formation rests conformably over siltstones of the Soyal6 Formation (Figure 4). The top of the sequence is marked by a coarse conglomerate. Some calcareous concretions near the top include coral remains, wood, and mollusks. The diversity of marine organisms is remarkable. Cypraeoidean gastro- pods and diverse crustaceans have been reported from this locality (Perrilliat et al., 2003; Vega et al., 2001). Annelid tubes, echinoids, fish remains, and rare seeds are also found. From the paleoenvironmental point of view, Frost & Langenheim (1974) and Ferrusquta-Villafranca (1996) defined this unit as being almost completely continental. The diversity of marine species at Veinte de Noviembre suggests a coastal environment, with some major conti- nental influence. Crustaceans represented only by cara- The Veliger, Vol. 48, No. 3 93°16" fo San Fernando Berriozabal to Mirador , to Soyalo Tuxtla =) Gutiérrez } = / Terdn San Antonio fo Ocozocuautla Cristobal San Vicente “—. ale / SS : : “Cres eC 3 ~ & N Cx AS vw~ S 6 to Villa Flores Tia @ Mexico City to Comitan Figure 1. Location map of study area at Depresion Central, Chiapas pace molts, with appendages or ventral parts not present, suggest at least some transport. However, delicate struc- tures such as spines are preserved, and the transport may not have been really significant, as most carapaces are complete (Vega et al., 2001). The studied fauna has its main affinities with gastro- pods reported from the Gulf Coastal Plains (Alabama, Texas, and Mississippi) and the west coast (California, Oregon, and Washington) of the United States. Only three species are common to France and England, and three species have also been reported to occur in the Mexican states of Nuevo Leon and Baja California Sur (Table 2). Specimens here described are deposited in the Instituto de Historia Natural y Ecologia, Chiapas, and in the Co- leccion de Paleontologia del Instituto de Geologia, Univ- ersidad Nacional Autonoma de México. Types are in- cluded in the Type Collection and classified under the acronyms IHN and IGM respectively. The type locality is registered in the locality catalog of the Instituto de Historia Natural y Ecologia, Chiapas. SYSTEMATIC PALEONTOLOGY Class Gastropoda Cuvier, 1797 Order Archaeogastropoda Thiele, 1925 Superfamily TROCHOIDEA Rafinesque, 1815 Family TROCHIDAE Rafinesque, 1815 Genus Calliostoma Swainson, 1840 Type species: Trochus conulus Linnaeus, 1758, by sub- sequent designation (Hermannsen, 1846:154); Recent, Mediterranean Sea. Figure 2. Hercoglossa sp. ct. H. ulrichi (White, 1882) IGM 8622 1.4, locality IHN 1005. Calliostoma granulata Perrilliat, Avendano & Vega, sp. nov. (Figures 5, 6) Diagnosis: Small Calliostoma; sculpture of four equally spaced, granulated spiral ribs and, from third whorl, three granulose threads, one between each rib. Description: Shell small sized, conical. Protoconch not preserved; teleoconch of six whorls that gradually in- crease in size; sculpture of four equally spaced, granulat- ed spiral ribs from suture and carina; from third whorl, three granulose threads in between these ribs. Entire sur- face covered by very thin radial threads. Edge of whorls finely granulated. Suture impressed. Base with spiral ribs and threads. Umbilicus deep and narrow. Aperture not preserved. Etymology: Name for the new species refers to sculpture of granulated spiral ribs. Types: Holotype IHN 5101, paratype IGM 8597. Holotype: IHN 5101, height 19.4 mm, diameter 16.6 mm; paratype IGM 8597, height 15.5 mm, diameter 16.9 mm. Discussion: The specimens from Chiapas differ from Calliostoma (Eutrochus) claibornianum Palmer (1944: 0.7080 0.7079 "Sr /*Sr 0.7076 Figure 3. *’Sr/8°Sr marine curve and 95% confidence interval for the upper Cretaceous—upper Eocene times (redrawn from Ho- warth and McArthur, 1997). The shaded area represents the range of *’Sr/8°Sr determined in Hercoglossa sp. cf. H. ulrichi (White, 1882), including analytical error. Most probable age is in the range of 50-55 Ma. M. del Carmen Perrilliat et al., 2005 Page 153 Table | Stratigraphic distribution of previously known species from locality IHN 1005. Paleocene Lower Eocene Middle Eocene Upper Eocene Oligocene Miocene Bittium (Bittium) estellensis »4 Potamides tricarinatus x xX x Turritella humerosa sanjuanensis x Mesalia alabamiensis x x x Calyptraea aperta x x x x Pachycrommium clarki x Xx x Galeodea koureos x Athleta petrosa petrosa x Architectonica alabamensis xX x Tornatellaea bella x 308, pl. 2, figs. 4-6), which are from the Gosport Sand, Claiborne, Alabama and very small with ornamentation of three spiral ribs and secondary spirals that increase in thickness until the last whorl, where they become five crenulated spiral ribs, and the base is covered also by spiral ribs. Calliostoma sp. (Clark & Martin, 1901:157) from the Aquia Formation, Upper Marlboro, two miles from Po- tomac Creek, Maryland, is not similar to the Mexican specimens, as only its last whorl presents spiral-crenulat- ed threads differing in thickness. Calliostoma sp. (Kellum, 1926:26, pl. 14, fig. 7), from the Eocene (Jackson) of Wilmington, New Hanover County, and Old Rocky Point, Pender County in North Carolina, is represented by a medium-sized internal cast with elevated spire. The sculpture is unknown, and there- fore it cannot be compared to the Chiapas specimens. Genus Trochus Linnaeus, 1758 Type species: Trochus maculatus Linnaeus, 1758, by original designation; Recent, Indo-Pacific. Trochus sp. (Figure 7) Description: Shell small sized, conical; protoconch of one and a half whorls, smooth; teleoconch of six slightly convex whorls, with sculpture of three thick, crenulate spiral ribs and a very fine thread between each rib; fine radial ribs in whole surface; suture impressed; base with same sculpture as rest of shell; aperture not preserved. Material examined: One specimen. IHN 5103, height 11.5 mm, diameter 8.6 mm. Discussion: One badly preserved specimen that cannot be assigned to any species. In the Eocene of North Amer- ica there are no known species assigned to this genus. Trochus (Tectus) bourdoti Cossmann & Pissarro (1902: 155, pl. 29, figs. 8, 9, 16, 17) from the Eocene of Fres- ville, France, is a large species with four prominent thick ribs, the anterior and posterior ones are thinner. The base presents seven thick ribs. The Chiapas specimen differs from the French species in being smaller in size (although the specimen illustrated in pl. 29, figs. 8, 9 is similar in size) and having ribs of equal size that are less crenulated. Order Caenogastropoda Cox, 1959 Superfamily CERITHIOIDEA Férussac, 1819 Family CERITHIDAE Fleming, 1822 Genus Cerithium Bruguiere, 1789 Type species: Cerithium adansonii Bruguiére, 1792, by tautonymy; Recent, Red Sea. “Cerithium” sp. (Figure 8) Description: Shell large, conical, slightly elongated. Pro- toconch and first whorls not preserved; rest of teleoconch with six flat whorls. Sculpture of rounded, closely spaced oblique broad axial ribs, straight in early whorls and curved at last whorl; ribs of last whorl are broad and thick in its medial part with interspaces deep and narrow; su- ture impressed; base rounded with one spiral rib; aperture not preserved. Material examined: One specimen. IHN 5200, height 61.5 mm, diameter 30.1 mm. Discussion: The specimen from Chiapas is incomplete, lacking apex and aperture. Weathering has removed axial ribs from abapertural side of three whorls; visible on ap- ertural side. Because of the incompleteness of this spec- imen, assignment to the genus Cerithium is uncertain. “Cerithium” sp. differs from Cerithium negritosense Woods (1922:87, pl. 11, figs. 1, 2) from the Eocene Ne- gritos Formation of Peru in having thicker and closely spaced axial ribs on every whorl; it does not present spi- ral sculpture. Page 154 Formation Gl-avi-brovre ne Eocene Juan Ae Reon middle ese Witcox 25 San 40 meters ‘eee Ri covered Paleocene * LOCALITY. THNAOOS Figure 4. Stratigraphic section at Veinte de Noviembre with position of locality IHN 1005. The Chiapas specimen is also different from Cerithium chaperi Bayan (1870:37, pl. 1, figs. 4, 5) from the Middle Eocene of Monte Postale, Verona, Italy, in that the latter species is more elongate, has more whorls, and lacks spi- ral sculpture. Subfamily CERITHINAE Fleming, 1822 Genus Bittium Leach in Gray, 1847 Type species: Murex reticulatus Montagu, 1803, by sub- sequent designation (Gray, 1847: 154); Recent, eastern North Atlantic Ocean. The Veliger, Vol. 48, No. 3 Table 2 Paleobiogeographic distribution of gastropod species from locality IHN 1005. Mexico USA Europe Bittium (Bittium) Alabama estellensis Texas Potamides tricarinatus Nuevo Leon Turritella humerosa Nuevo Leon sanjuanensis Mesalia alabamiensis Alabama Calyptraea aperta Mississippi France Texas England Alabama Pachycrommium Baja Cali- California clarki fornia Sur Oregon Washington Galeodea koureos Nuevo Leon Alabama Athleta petrosa petrosa Gulf Coast : Plain Architectonica Alabama alabamensis Tornatellaea bella Alabama Subgenus Bittium s. s. Bittium (Bittium) estellensis (Aldrich, 1921) (Figures 9, 10) Cerithiopsis estellensis Aldrich, 1921:15, pl. 2, fig. 11. Bittium (Bittium) estellensis (Aldrich). Gardner, 1933:274, pl. 24, figs. 12, 13. Bittium (Bittium) estellensis (Aldrich). Palmer & Brann, 1966:527. Description: Shell small sized, profile flat sided; tele- conch of five whorls; suture lineal; sculpture of two pri- mary broad nodulose spiral ribs, secondary spiral ribs be- tween the primary rib on anterior and posterior part of whorl; a nodulose very fine thread in anterior margin. Aperture not preserved. Material examined: Four specimens. IHN 5450, height 7.3 mm, diameter 3.8 mm; IGM 8598, height 5.4 mm, diameter 3.0 mm. Discussion: This species was described from the Paleo- cene Kincaid Formation of Alabama and Texas; speci- mens from there achieved a larger size than the specimens of Chiapas, Mexico. Genus /ddingsia Olsson, 1929 Type species: Cerithium laeviusculum Gabb, 1869, by original designation; Upper Eocene, Peru. ?Iddingsia sp. (Figure 11) Description: Shell medium sized, elongate; profile flat sided; protoconch and first teleoconch whorls not pre- M. del Carmen Perrilliat et al., 2005 Page 155 served; remainder three or four without sculpture; suture channeled; base not preserved. Material examined: Six specimens. IHN 6675, height 42.2 mm, diameter 28.4 mm; IGM 8599, height 43.1 mm, diameter 25.8 mm. Discussion: The channeled suture of the Mexican speci- mens suggests /ddingsia, but because they are incom- plete, they are tentatively assigned. ?/ddingsia sp. differs from Iddingsia laeviuscula saladoense (Olsson) (1929:82, pl. 4, figs. 2, 3) from the Middle Eocene Restin Formation of Quebrada Salado, Peru, in that the Mexican specimens are wider. Family POTAMIDIDAE Adams & Adams, 1854 Subfamily POTAMIDINAE Adams & Adams, 1854 Genus Potamides Brongniart, 1810 Type species: Potamides lamarckii Brongniart, 1810, by original designation; Oligocene, Paris Basin, France. Subgenus Potamidopsis Munier Chalmas, 1900 Type species: Cerithium tricarinatum Lamarck, 1804, by original designation; Middle Eocene, Paris Basin, France. Potamides (Potamidopsis) tricarinatus (Lamarck, 1804) (Figures 12, 13) Cerithium tricarinatum Lamarck, 1804:272. Cerithium mutabile Lamarck, 1804:344, vélin n° 14, fig. 13. Cerithium semicoronatum Lamarck, 1804:344, vélin n° 13, fig. 2. Potamides (Tympanotomus) semicoronatus (Lamarck). Cossmann & Pissarro, 1902:46, pl. 16, figs. 11, 13. Description: Shell medium sized; teleoconch with four whorls, profile flat sided; suture lineal; sculpture of three granulose spiral ribs, the posterior one being thicker than the other two; aperture not preserved. Material examined: Two specimens, IHN 6546, height 19.9 mm, diameter 12.1 mm; IGM 8600, height 16.7 mm, diameter 11.1 mm. Discussion: P. tricarinatus, P. mutabile, and P. semico- ronatus have been considered the same species as they present three spiral ribs, the posterior one being thicker than the other two. These species have been described from the Middle Eocene of the Paris Basin, France. Family TURRITELLIDAE Lovén, 1847 Genus Palmerella Allmon, 1996 Type species: Turritella mortoni Conrad, 1830, by orig- inal designation; Upper Paleocene, Maryland and Virgin- ia, USA. Palmerella mortoni mexicana Perrilliat, Avendano & Vega subsp. nov. (Figures 14—16) Diagnosis: Medium sized Palmerella. On last whorl, a rounded carina present below strong carina at spiral C. Description: Shell medium sized, four whorls preserved; whorls with convex profile; suture impressed; an angu- lose, strong peribasal carina present on each whorl, with spiral lines on rest of whorl; spiral lines granulated at intersection with growth lines; faint spirals below carina. On last whorl, a second rounded carina of similar thick- ness present below peribasal carina, four spiral threads present between these two carinae; edge of carina formed by a single spiral rib. Growth lines prosocline. Aperture unknown. Etymology: Subspecies name refers to Mexico. Types: Holotype IHN 5133, paratype IGM 8601. Holotype: IHN 5133, height 36.5 mm, diameter 21.3 mm; paratype IGM 8601, height 27.8 mm, diameter 19.2 mm. Discussion: All specimens incomplete. Palmerella mor- toni mexicana resembles P. m. postmortoni (Harris) (1894:302—304, in part, fig. 1, not fig. 2) from the upper Paleocene of Alabama in pronounced carina at C but dif- fers in that the carina of P. m. mexicana bears a single rib but that of P. m. postmortoni may be double ribbed (Allmon, 1996). ?Palmerella sp. (Figure 17) Description: Shell medium sized with five whorls, profile convex; rounded strong basal carina; thin spiral rib above carina on last two whorls, rest of whorl with fine spiral threads; and growth lines. Base and aperture not pre- served. Material examined: One specimen. IHN 5104, height 32.4 mm, diameter 17.2 mm. Discussion: One poorly preserved specimen, uncertainly assigned to Palmerella. Genus Haustator Montfort, 1810 Type species: Haustator gallious Montfort, 1810, by original designation; Upper Eocene, England. Haustator sp. cf. H. rivurbana (Cooke, 1926) (Figure 18) Description: Shell large sized; teleoconch of two and a half whorls; profile nearly flat sided; sutures impressed; two primary, raised, granulose ribs and a weaker spiral The Veliger, Vol. 48, No. 3 Page 156 M. del Carmen Perrilliat et al., 2005 Page 157 rib observed; weak spiral threads intercalate between ribs; interspace broad. Aperture unknown. Material examined: One specimen. IHN 5120, height 42.0 mm, diameter 21.6 mm. Discussion: Specimen incomplete and compared with Haustator rivurbana (Cooke, 1926) from the Eocene of Mississippi in having same sculpture. Turritella rivurbana Cooke (Harris & Palmer, 1947: 295, pl. 38, figs. 6, 7, 9, and Dockery, 1977:45—46, pl. 3, fig. 6) from the Eocene of Mississippi is represented by specimens of smaller size than the ones from Chiapas. Turritella rivurbana chiapasensis Allison, 1969 in AI- lison & Adegoke, 1969:1258, pl. 147, figs. 3, 5, 6, 8, 9— 12, 14; pl. 148, figs. 2, 11) and Turritella rivurbana mex- icana Allison, 1969 in Allison & Adegoke, 1969:1259— 1262, pl. 147, figs. 4, 13; pl. 148, figs. 1, 3, 4, 6, 7, 9, 10, 13, 14) from the Late Eocene of Simojovel, Chiapas, were considered synonym of 7. rivurbana (Allmon, 1996: 78). Genus? (“‘Turritella humerosa group’’) “Turritella” humerosa sanjuanensis Bowles, 1939 (Figures 19, 20) Turritella rivurbana sanjuanensis Bowles, 1939:314, pl. 34, fig. 18. Description: Shell large sized; three whorls preserved, whorl profile almost flat sided; last whorl with 15 spirals of similar width and separated by equal interspaces; growth lines present, forming small irregular nodes at in- tersection with spirals. Adapical carina rounded imme- diately below suture, slightly convex and with five spi- rals. Lateral sinus moderately deep. Material examined: Seven specimens. IHN 5770, height 50.7 mm, diameter 19.6 mm; IGM 8603, height 32.0 mm, diameter 14.5 mm. Discussion: Holotype of Turritella rivurbana sanjuanen- sis was described from the Lower Eocene of Rio San Juan, Nuevo Leén (Bowles, 1939:314). Specimens from Chiapas are larger than the ones described from Nuevo Leon. “Turritella”’ sp. (Figures 21, 22) Description: Shell large sized, four whorls of teleoconch preserved; profile flat sided; whorls with a rounded basal carina; spiral threads present between basal carina and suture; two spiral ribs on each whorl, one to two spiral threads between ribs; growth lines present. Material examined: Three specimens. [HN 5128, height 57.5 mm, diameter 20.6 mm; IGM 8604, height 25.2 mm, diameter 16.0 mm. Discussion: The most similar species to the Mexican specimens is Palmerella dutexata Harris (1895:82, pl. 9, fig. 8) from the middle Eocene of Texas. Palmerella du- texata Harris (illustrated in Stenzel & Turner, 1940b, pl. 46, fig. 5) is a small specimen and presents a basal carina similar to the specimens from Chiapas. The specimens from Chiapas differ from P. dutexata in the ornamenta- tion of the rest of the whorl. Genus Mesalia Gray, 1842 Type species: Turritella brevialis Lamarck, 1822, by subsequent designation (Gray, 1847:155); Recent, off the coasts of northern and western Africa. Mesalia alabamiensis (Whitfield, 1865) (Figures 23, 24) Potamides alabamiensis Whitfield, 1865:266, pl. 27, fig. 13; Dall, 1892:287; Whitfield, 1899:174. Turritella vittata abruta Conrad. De Gregorio, 1890:124, pl. 11, fig. 12. Mesalia alabamiensis Harris, 1897:25, 31; Cooke, 1926:264, pl. 94, fig. 9; Palmer, 1937:204. Figures 5-30. Specimens coated with ammonium chloride. All from IHN locality 1005. Figures 5, 6. Calliostoma granulata Perrilliat, Avendano & Vega, sp. nov. Figure 5. Holotype IHN 5101, 1.3. Figure 6. Paratype IGM 8597, 1.4. Figure 7. Trochus sp. Hypotype IHN 5103, <1.7. Figure 8. ““Cerithium” sp. Hypotype IHN 5200, 1. Figures 9, 10. Bittium (Bittium) estellensis (Aldrich, 1921). Figure 9. Hypotype IGM 8598, 4.4. Figure 10. Hypotype IHN 5450, <3.3. Figure 11. ?/ddingsia sp. Hypotype IHN 6675, X1. Figures 12, 13. Potamides tricarinatus (Lamarck, 1804). Figure 12. Hypotype IGM 8600, 1.5. Figure 13. Hypotype IHN 6546, 1.5. Figures 14—16. Palmerella mortoni mexicana Perrilliat, Avendano & Vega, subsp. nov. Figure 14. Paratype IGM 8601, <1.1. Figure 15. Holotype IHN 5133, X0.8. Figure 16. Paratype IGM 8602, <0.9. Figure 17. ?Palmerella sp. Hypotype IHN 5104, x1. Figure 18. Haustator sp. cf. H. rivurbana (Cooke, 1926). Hypotype IHN 5120, <1. Figures 19, 20. Turritella humerosa sanjuanensis Bowles, 1939. Figure 19. Hypotype IGM 8603, <1. Figure 20. Hypotype IHN 5770, x1. Figures 21, 22. Turritella sp. Figure 21. Hypotype IHN 5128, <1. Figure 22. Hypotype IGM 8604, x1. Figures 23, 24. Mesalia alabamiensis (Whitfield, 1865). Figure 23. Hypotype IGM 8605, 1.4. Figure 24. Hypotype IHN 5107, 1.5. Figures 25, 26. Calyptraea aperta (Solander, in Brander, 1766). Figure 25. Hypotype IGM 8606, 1.3. Figure 26. Hypotype IHN5201, <1.4. Figure 27. Terebellum (Seraphs) sp. Hypotype IHN 6684, <1. Figures 28, 29. Crommium globosa Perrilliat, Avendafio & Vega, sp. nov. Figure 28. Holotype IHN 5220, <1. Figure 29. Paratype IGM 8607, 1. Figure 30. Pachycrommium clarki (Stewart, 1927). Hypotype IHN 5145, x1. Page 158 The Veliger, Vol. 48, No. 3 Mesalia pumila var. alabamiensis Harris, 1899a:76, 77, pl. 10, fig. 9; Brann & Kent, 1960:550. Mesalia (Mesalia) alabamiensis (Whitfield). Cossmann, 1912:126. Mesalia alabamiensis (Whitfield). Bowles, 1939:327, pl. 34, fig. 10; Palmer & Brann, 1966:756. Description: Shell medium sized, four whorls preserved; whorl profile convex; whorls inflated, constricted at su- tures; sutures well defined; each formed by three fine spi- ral ribs; fine spiral threads in interspaces. First whorls with three spiral ribs; increasing to five on last whorl. Aperture unknown. Base with spiral threads. Growth lines weak. Material examined: 15 specimens. IHN 5107, height 25.7 mm, diameter 16.6 mm; IGM 8605, height 21.1 mm, diameter 15.3 mm. Discussion: Specimens from Chiapas are larger than the ones described from the Lower Eocene of Alabama. Superfamily CALYPTRAEOIDEA Lamarck, 1809 Family CALYPTRAEIDAE Lamarck, 1809 Genus Calyptraea Lamarck, 1799 Type species: Patella chinensis Linnaeus, 1758, by mon- otypy; Recent, Europe. Calyptraea aperta (Solander, in Brander, 1766) (Figures 25, 26) Trochus apertus Solander, 1766, in Brander:9, pl. 1, figs. 1, 2. Trochus opercularis Solander, 1766, in Brander:9, pl. 1, fig. 3. Calyptraea trochiformis Larmarck, 1802:385; Dall, 1892: 352° Calyptraea aperta (Solander). Harris, 1899a:84, pl. 11, figs. 13, 16; Maury, 1912:99, pl. 13, fig. 5; Palmer, 1937: 145, pl. 16, figs. 1-3, 5; Olsson, 1944:248, pl. 9, figs. 10-13; Harris & Palmer, 1947:260, pl. 31, figs. 2, 4— 12: Calyptraea (Trochatella) aperta (Solander). Olsson, 1928: 62. Calyptraea aperta (Solander in Brander). Palmer & Brann, 1966:547. Calyptraea (Trochita) aperta (Solander in Brander). Dock- ery, 1977:56, pl. 5, fig. 10. Description: Shell medium sized; protoconch of two whorls, smooth; first whorl of teleoconch smooth, follow- ing whorls with microscopic, irregular, and undulated ra- dial striae; nodes present on radial lines. Aperture and base not preserved. Material examined: 19 specimens. THN 5201, height 25.7 mm, diameter 27.1 mm; IGM 8606, height 24.1 mm, diameter 27.5 mm. Discussion: There is a wide gamut of variation in this species, from flat shells to those of considerable height. Mexican specimens are tall shells. This species has been described from the Lower Eocene to Miocene of Ala- bama, Mississippi, and Texas; Eocene of Peru; and Upper Eocene of England. The complete synonymy for this species can be found in Palmer (1937:145). Superfamily STROMBOIDEA Rafinesque, 1815 Family STROMBIDAE Rafinesque, 1815 Genus Terebellum R6ding, 1798 Type species: Terebellum nebulosum Roding, 1798, by subsequent designation (Winckworth, 1945:144); Recent, tropical western Pacific Ocean. Subgenus Seraphs Montfort, 1810 Type species: Seraphs convolutus (Lamarck, 1802), by original designation: Eocene, Europe. Terebellum (Seraphs) sp. (Figure 27) Description: Shell large sized, slender; first whorls in- volute; last whorl incomplete; aperture extends to almost 2.0 mm from apex of first whorls; aperture lineal except on its anterior third, where it is curved, widening from medial to posterior part. Surface smooth. Material examined: One specimen. IHN 6684, height 59.7 mm, diameter 29.2 mm. Discussion: Only one species has been described from the eastern part of America, Terebellum (Seraphs) belem- nitum Palmer (in Richards & Palmer, 1953:25, pl. 3, figs. 9, 12) from the Inglis Member of Moodys Branch For- mation of Florida, but the Chiapas specimen is larger and broader. Terebellum (Seraphs) belemnitum Palmer? (Woodring, 1959:192, pl. 25, fig. 6) from the Middle Eocene of Pan- ama is a smaller specimen than the one from Chiapas, and the apical end of shell thins rapidly. The specimen from Chiapas is similar to Terebellum (Seraphs) sopitum (Solander) Cossmann & Pissarro (1900:139, pl. 15, fig. 5) from the Middle and Upper Eo- cene of Fresville, France, in having similar width, but spire whorls are not involute. The following species from the Middle Eocene of France are smaller than the speci- men from Chiapas: 7. olivaceum Cossmann, 1889, T. chilophorum Cossmann, 1889, and T. isabellae Deshayes, 1865. More complete specimens are needed to support de- scription of a new species. M. del Carmen Perrilliat et al., 2005 Page 159 Superfamily NATICOIDEA Forbes, 1838 Family NATICIDAE Forbes, 1838 Subfamily AMPULLOSPIRINAE Cox, 1930 Genus Crommium Cossmann, 1888 Type species: Ampullaria willemetii Deshayes, 1825, by original designation; Eocene, France. Crommium globosa Perrilliat, Avendano & Vega, sp. Nov. (Figures 28, 29) Diagnosis: Large sized Crommium, globose, spire mod- erately elevated, whorls tabulate with faint spirals on tab- ulate portion and on last whorl. Description: Shell large sized, thick and globose; spire moderately elevated with four whorls; whorls evenly rounded but narrowly tabulate; suture deeply impressed; shell smooth except for faint spiral ribs on tabulate por- tion and persistant on last whorl; aperture broad; parietal callus not preserved; umbilicus narrowly open. Etymology: The name of the new species refers to its inflated shape. Types: Holotype IHN 5220, paratype IGM 8607. Holotype: IHN 5220, height 56.9 mm, diameter 50.6 mm; paratype IGM 8607, height 32.3 mm, diameter 31.9 mm. Discussion: The specimens from Chiapas differ from Crommium splendida (Deshayes) (1866, p. 61, pl. 67, figs. 8-10) from the Lower Eocene of Aisy, France, in having faint spirals on tabulate portion and last whorl with ribs. Genus Pachycrommium Woodring, 1928 Type species: “‘Natica phasianelloides d’Orbigny” of Guppy, 1866, by original designation; Miocene, Domin- ican Republic.: Pachycrommium clarki (Stewart, 1927) (Figure 30) “Amauropsis alveata (Conrad). Arnold, 1910:114, pl. 4, fig. 21; Arnold & Anderson, 1910:71, 286, pl. 26, fig. 21; Dickerson, 1915:86, pl. 5, fig. 9. Amaurellina (Euspirocrommium) clarki Stewart, 1927:336— 338, pl. 26, figs. 8, 9; Clark, 1929, pl. 11, fig. 10; Turn- er, 1938:86, pl. 20, fig. 3; Weaver, 1942:345, pl. 70, figs. 10, 18; Kenn & Bentson, 1944:127; Weaver & Klein- pell, 1963:188, pl. 27, fog. 15. _Amaurellina clarki Stewart. Gardner & Bowles, 1934:246. figs. 6, 8; Schenck & Keen, 1940:34, pl. 26, fig. 7; Keen & Bentson, 1944:127. Pachycrommium (?) clarki (Stewart). Vokes, 1939:26, 175, pl. 22, figs. 11, 30; Givens, 1974:73, pl. 8, figs. 6, 10. Amaurellina (?) multiangulata Vokes, 1939:26, 174, pl. 22, figs. 2, 8, 13. Pachycrommium clarki (Stewart, 1927). Marincovich, 1977: 238-241, pl. 20, figs. 4-10; Squires & Demetrion, 1992:32, fig. 84. Description: Shell medium sized, elongate; protoconch not preserved; spire elevated, with six whorls; last whorl moderately inflated; first whorls rounded; shoulder be- coming progressively more tabulate on last whorls; last whorl with strong tabulation; suture moderately im- pressed; surface smooth, except for growth lines; closed umbilicus; aperture not preserved. Material examined: 110 specimens. IHN 5145, height 34.9 mm, diameter 27.5 mm; IGM 8608, height 30.1 mm, diameter 21.6 mm. Discussion: This species is found in the Lower Eocene of California (Givens, 1974); the Middle Eocene of Oregon (Turner, 1938) and California (Stewart, 1927; Vo- kes, 1939); and the Upper Eocene of Washington (Weav- er, 1943) and California (Weaver & Kleinpell, 1963). It is also found in the Eocene Bateque Formation of Baja California Sur (Squires & Demetrion, 1992). It differs from Amauropsis jacksonensis Harris (1896b:474, pl. 19, fig. 3) of the Eocene of Jackson, Mississippi, in having a rounded shoulder and from A. perovata Conrad (1846:21, pl. 1, fig. 16) of the Middle Eocene of Alabama in being larger and having a well-defined shoulder. Superfamily TONNOIDEA Suter, 1913 Family CASSIDAE Latreille, 1825 Genus Galeodea Link, 1807 Type species: Buccinum echinophora Linnaeus, 1758, by monotypy; Recent, Mediterranean Sea. Galeodea koureos Gardner, 1939 (Figures 31, 32) Cassidaria brevidentata Aldrich ‘“‘var.” Harris, 1896b:479, pl. 22, fig. 10; Harris, 1899a:67, pl. 8, fig. 18; Harris 1899b:307. Galeodea (Mambrinia) koureos Gardner, 1939:23; LeBlanc, 1942:111, pl. 14, figs. 5, 6. Galeodea koureos Gardner. Palmer & Brann, 1966:695. Description: Shell medium sized; protoconch with three whorls, smooth; teleoconch with four whorls, first whorl with six spiral threads; following whorl with three pri- mary spiral ribs, with small nodes and two thin spiral threads in between each primary spiral rib; from next whorl, abapical spiral is more prominent and defines pe- riphery and external edge of shoulder, where 14 nodes are present. Two thinner abapical spirals with nodes on last whorl, with thin spiral threads between these spirals; whole surface with very fine growth lines; suture im- pressed; aperture not preserved. Page 160 The Veliger, Vol. 48, No. 3 M. del Carmen Perrilliat et al., 2005 Page 161 Material examined: 16 specimens. IHN 5242, height 24.8 mm, diameter 22.2 mm; IGM 8609, height 21.1 mm, diameter 18.3 mm. Discussion: This species has been described from the Lower Eocene of Alabama and west of Los Aldamas, Nuevo Leon. Although Chiapas specimens are smaller, they present the three major spirals on last whorl. Superfamily JANTHINOIDEA Lamarck, 1812 Family EPITONIDAE Suter, 1913 Genus Cirsotrema Morch, 1852 Type species: Scalaria varicosa Lamarck, 1822, by mon- otypy; Recent, Western Pacific. Cirsotrema sp. (Figure 33) Description: Shell small sized; turreted, five rounded whorls preserved; sculpture of eight prosocline axial la- mellae; aperture and base not preserved. Material examined: One specimen. IHN 5241, height 12.5 mm, diameter 7.2 m. Discussion: One poorly preserved specimen in which spi- ral lines are not observed on interspaces or in lamellae. It is similar to Cirsotrema (Coroniscala) newtonensis (Meyer & Aldrich, 1886) (Palmer, 1937:101, pl. 10, figs. 10, 11) from the Middle Eocene of Mississippi in size and number of lamellae. Cirsotrema (Coroniscala) clai- bornensis (Conrad, 1865) from the Middle Eocene of Al- abama, Cirsotrema (Coroniscala) octolineata (Conrad, 1860) from the Middle Eocene of Mississippi, and Cir- sotrema (Coroniscala) lintea (Conrad, 1860) from the Middle Eocene of Alabama all have more lamellae. Order Neogastropoda Thiele, 1929 Superfamily MurIcoIpEA Rafinesque, 1815 Family BUCCINIDAE Rafinesque, 1815 Genus Exilia Conrad, 1860 Type species: Exilia pergracilis Conrad, 1860, by origi- nal designation; Lower Eocene, Alabama, USA. Exilia sp. (Figure 34) Description: Shell medium sized, high spired, fusiform; teleoconch of three barely convex whorls; sculpture of numerous flexuous axial ribs and thin spiral ribs on in- terspaces; suture impressed; aperture narrow; outer lip not preserved; columella smooth. Material examined: Two specimens. IHN 5324, height 13.0 mm, diameter 4.3 mm; IGM 8610, height 8.9 mm, diameter 3.2 mm. Discussion: The specimens from Chiapas resemble Exilia pergracilis Conrad (1860:291, pl. 47, fig. 34) from the Lower Eocene of Alabama in having similar sculpture and a smooth columella, but they are smaller. The first whorls, aperture, and anterior siphon are not preserved. Family VOLUTIDAE Rafinesque, 1815 Subfamily FULGORARIINAE Pilsbry & Olsson, 1954 Genus Lyrischapa Aldrich, 1911 Type species: Lyrischapa harrisi Aldrich, 1911, by mon- otypy; Middle Eocene, Mississippi, USA. Figures 31-56. Specimens coated with ammonium chloride. All from IHN locality 1005. Figures 31, 32. Galeodea koureos Gardner, 1939. Figure 31. Hypotype IHN 5242, X1.5. Figure 32. Hypotype IGM 8609, 1.4. Figure 33. Cirsotrema sp. Hypotype IHN 5241, 2.1. Figure 34. Exilia sp. Hypotype IHN 5324, <2. Figures 35, 36. Lyrischapa spinifera Perrilliat, Avendafio & Vega, sp. nov. Figure 35. Holotype IHN 5282, <1. Figure 36. Paratype IGM 8611, x1. Figures 37, 38. Volutocorbis minutus Perrilliat, Avendafio & Vega, sp. nov. Figure 37. Holotype THN 5294, x1.5. Figure 38. Paratype IGM 8612, X2.5. Figures 39, 40. Athleta petrosa petrosa (Conrad, 1833). Figure 39. Hypotype IHN 5275, <1. Figure 40. Hypotype IGM 8613, X1.1. Figures 41, 42. Lapparia sp. cf. L. nuda Stenzel & Turner, 1940. Figure 41. Hypotype IGM 8614, X1.6. Figure 42. Hypotype IHN 5280, 1.1. Figures 43, 44. Volutilithes sp. cf. V. muricina (Lamarck, 1803). Figure 43. Hypotype IHN 5307, <1. Figure 44. Hypotype IGM 8615, X1. Figures 45, 46. Sulcobuccinum sp. cf. S. scalina Heilprin, 1880. Figure 45. Hypotype IGM 8616, x1. Figure 46. Hypotype IHN 5328, <1. Figure 47. Cornulina? sp. Hypotype IHN 5323, <1. Figure 48. Levifusus sp. Hypotype IHN 5326, <2. Figures 49, 50. Turricula sp. Figure 49. Hypotype IGM 8618, <1. Figure 50. Hypotype THN 5366, X1.3. Figures 51, 52. Architectonica alabamensis (Dall, 1892). Figure 51. Hypotype IHN 5386, 1.9. Figure 52. Hypotype IGM 8619, X2.5. Figure 53. Architectonica sp. Hypotype IHN 5388, <2. Figure 54. Archi- tectonica sp. cf. A. elaborata (Conrad, 1833). Hypotype IHN 5385, X2.1. Figures 55, 56. Tornatellaea bella Conrad, 1860. Figure 55. Hypotype THN 6550, 2.4. Figure 56. Hypotype IGM 8620, x3. Page 162 The Veliger, Vol. 48, No. 3 Lyrischapa spinifera Perrilliat, Avendano & Vega sp. Nov. (Figures 35, 36) Diagnosis: Medium sized Lyrischapa with low spire; blunt peripheral spines; columellar folds of same strength and spacing, strongest anteriorly. Description: Shell medium sized, conical; protoconch of two smooth whorls; spire low; teleoconch of four whorls, rapidly expanding, each succeeding whorl enveloping the preceding one; each whorl with nine prominent axial ribs, which gradually develop into erect, blunted peripheral spines; spiral sculpture not preserved; suture distinct, im- pressed, undulated around axial ribs; aperture narrow; on inner lip, only three columellar folds visible; parietal cal- lus thick. Etymology: Name of the new species refers to presence of blunted spines. Types: Holotype IHN 5282; paratype -IGM 8611. Holotype: IHN 5282, height 50.1 mm, diameter (with spines) 45.5 mm; IGM 8611, height 42.0 mm, diameter (with spines) 37.4 mm. Discussion: There are only three previously known spe- cies of Lyrischapa from the Middle Eocene of North America. Lyrischapa harrisi Aldrich (1911:11, pl. 4, fig. 8) from Mississippi differs from the Mexican specimens in having a larger protoconch, more rapidly expanding whorls, shorter peripheral spines, and a broader ramp. Lyrischapa lajollaensis (Hanna) (1927:320, pl. 52, figs. 1, 2) from the Llajas Formation, California, differs in be- ing slender and having more spines and a higher spire. Lyrischapa chiapasensis (Gardner & Bowles) (1934:248, figs. 10-12) from the Middle Eocene of Sayula, Chiapas, is smaller, more slender, and with upward pointed, larger, peripheral nodes. Genus Volutocorbis Dall, 1890 Type species: Volutocorbis (Volutilithes) limopsis (Con- rad, 1860), by original designation; Paleocene, Alabama, USA. Volutocorbis minutus Perrilliat, Avendano & Vega sp. nov. (Figures 37, 38) Diagnosis: Small Volutocorbis; fusiform; sculpture of ax- ial ribs and spiral ribs forming nodes. Description: Shell small sized, fusiform; protoconch smooth, with two and a half whorls; teleoconch of four whorls with slightly convex profile; first whorls with three spiral ribs, which form nodes at intersection with axial ribs; nearly 21 axial ribs on last whorl, with nine spiral ribs forming nodes and four smooth ribs on base; inter- spaces with fine spiral threads; suture deep; aperture nar- row, three folds on columella. Etymology: Species name refers to its relatively small size, compared with other species of this genus. Types: Holotype IHN 5294, paratype IGM 8612. Holotype: IHN 5294, height 16.8 mm, diameter 8.4 mm; paratype IGM 8612, height 11.2 mm, diameter 5.9 mm. Discussion: The specimens from Chiapas are most sim- ilar to Volutocorbis stenzeli (Plummer) (1932:813, pl. 9, figs. 12, 13) from the Middle Eocene of Texas, but this species has more axial ribs (40) and more spiral ribs. Volutocorbis limopsis (Conrad, 1860:292, pl. 47, fig. 24) from the Paleocene of Alabama differs from the Mexican specimen in being larger and having 27 axial ribs. Vol- utocorbis texana Gardner (1933:235, pl. 21, figs. 1, 2) from the Paleocene of Texas is larger and has fewer axial ribs; it has the same number of spiral ribs as the speci- mens from Mexico. Genus Athleta Conrad, 1853b Type species: Voluta rarispina Lamarck, 1811, by sub- sequent designation (Dall, 1890:75); Miocene, Aquitaine Basin, France. Athleta petrosa petrosa (Conrad, 1833) (Figures 39, 40) Voluta petrosa Conrad, 1833:29; Conrad, 1835:41, pl. 16, fig. 2; Lea, 1848:107; de Gregorio, 1890:63, pl. 4, figs. SOMSI 5375960! Volutilithes petrosa Conrad, 1854:31. Athleta petrosa (Conrad). Smith, 1907:230. Athleta petrosa (Conrad). Palmer, 1937:372, pl. 58, figs. 1— 14; pl. 88, figs. 1, 7, 11; Harris & Palmer, 1947:391— 393, pl. 53, figs. 1-4. Athleta petrosa (Conrad, 1833). Fisher, Rodda & Dietrich, 1964:40—43, pl. 8, figs. 1, 2; pl. 10, figs. 6-10. Description: Shell medium sized, fusiform; spire elevat- ed; whorl profile shouldered; protoconch not preserved; teleoconch of five whorls; early whorls with broad, rounded axial ribs, narrow interspaces; nodes on intersec- tion with axial ribs; nodes increase gradually in size, forming spines, and decrease in number on last whorl; axial and spiral lirae uniform on every whorl; lirae dis- appear on anterior part of last whorl; aperture narrow; two folds on columella. Material examined: Five specimens, IHN 5275, height 23.1 mm, diameter 14.6 mm; IGM 8614, height 17.1 mm, diameter 11.0 mm. Discussion: A broadly distributed species on the Atlantic and Gulf Coastal Plains. Complete synonymy in Fisher, Rodda & Dietrich, 1964:40—42. M. del Carmen Perrilliat et al., 2005 Genus Lapparia Conrad, 1855 Type species: Mitra dumosa Conrad, 1854, by monoty- py; Eocene, Mississippi, USA. Lapparia sp. cf. L. nuda Stenzel & Turner, 1940a (Figures 41, 42) Description: Shell large, heavy, subfusiform; only three whorls of teleoconch preserved; sculpture of axial ribs that persist to last whorl and nodes that gradually develop into spines; seven nodes per whorl; ribs narrow and in- terspaces broad; no spiral sculpture; a steeply sloping noded shoulder near middle of spire whorls; columella with three folds; aperture not preserved. Material examined: Two specimens. IHN 5280, height 54.3 mm, diameter 28.0 mm; IGM 8614, height 25.9 mm, diameter 20.7 mm. Discussion: The Mexican specimens are similar to Lap- paria nuda Stenzel & Turner (1940a:808, pl. 44, figs. 6, 9) from the Middle Eocene of Texas, but are larger in size, with persistent ribs on last whorl and fewer spines. Genus Volutilithes Swainson, 1831 Type species: Voluta muricina Lamarck, 1803, by sub- sequent designation (Dall, 1906:143); Middle Eocene, Paris Basin, France. Volutilithes sp. cf. V. muricinus (Lamarck, 1803) (Figures 43, 44) Description: Shell medium sized, fusiform; protoconch not preserved; teleoconch with five whorls; convex pro- file; first whorl with nine rounded axial ribs, interspaces twice the width of the ribs; following whorls with seven axial ribs; last whorl with acute periphery and spines on apex of axial ribs; no spiral sculpture; suture undulated; aperture not preserved; columella with three folds. Material examined: 16 specimens. IHN 5307, height 32.5 mm, diameter 16.0 mm; IGM 8615, height 31.4 mm, diameter 16.4 mm. Discussion: The specimens from Chiapas are similar to Volutilithes muricinus (Lamarck, 1803) from the Middle Eocene of France in their sculpture, but the Mexican specimens are smaller. Family PSEUDOLIVIDAE de Gregorio, 1880 Genus Sulcobuccinum d’Orbigny, 1850 Type species: Buccinum fissuratum Deshayes, 1835, by subsequent designation (Vermeij, 1998:60); Upper Paleo- cene, France. Page 163 Sulcobuccinum sp. cf. S. scalina Heilprin, 1880 (Figures 45, 46) Description: Shell medium sized, ovate; protoconch of two whorls, paucispiral; teleoconch of five whorls with shoulder; spiral sculpture of fine threads on first whorls of teleoconch; axial sculpture of 11 ribs, on last whorl reaching pseudolivid groove, forming nodes on shoulder; suture deeply channeled; base of last whorl below groove, smooth; aperture ovate. Material examined: 60 specimens. IHN 5328, height 26.9 mm, diameter 15.6 mm; IGM 8616, height 31.9 mm, diameter 17.2 mm. Discussion: The specimens from Chiapas are different from Sulcobuccinum scalina (Heilprin) (1880:371, pl. 20, fig. 12) from the Lower Eocene of Alabama in being smaller and lacking spiral threads below groove. Probably these specimens represent two different species. The classification suggested by Vermeij (1998:60) is followed in this work. Family MELONGENIDAE Gill, 1871 Genus Cornulina Conrad, 1853 Type species: Cornulina armigera Conrad, 1833, by sub- sequent designation (Fischer, 1884:621); Middle Eocene, Alabama, USA. Cornulina? sp. (Figure 47) Description: Shell large sized, heavy; two whorls pre- served; last whorl with two lines of short, equal sized spines; no spiral sculpture; suture undulated; aperture not preserved. Material examined: One specimen. IHN 5323, height 49.3 mm, diameter 40.8 mm. Discussion: Assignment to this genus is tentative because of the incompleteness of the specimen. Similarity to Cor- nulina is based on sculpture of last whorl, which has two lines of spines reminiscent of Cornulina, but the spire of this specimen is higher. Genus Levifusus Conrad, 1865 Type species: Fusus trabeatus Conrad, 1833, by subse- quent designation (Cossmann, 1901:14); Eocene, Ala- bama, USA. Levifusus sp. (Figure 48) Description: Shell small sized; protoconch not preserved; first whorl of teleoconch with eight widely spaced and rounded axial ribs, overrun by spiral ribs on whole whorl; following whorls with carina and 10 axial ribs; on last Page 164 whorl, a strong carina near mid whorl and a weaker one below it; spiral ribs and spiral threads between these two carinae; aperture not preserved. Material examined: Two specimens. IHN 5326, height 14.3 mm, diameter 12.0 mm; IGM 8617, height 16.0 mm, diameter 9.4 mm. Discussion: Two incomplete specimens are similar to Levifusus mortoniopsis carexus (Harris) (1895:72, pl. 7, fig. 5) from the Eocene of Texas. Although the Mexican specimens are smaller, they have the same number of ax- ial ribs, carina, and spiral ribs as the Texan specimens. Superfamily CONOIDEA Rafinesque, 1815 Family TURRIDAE Adams & Adams, 1858 Subfamily TURRICULINAE Powell, 1942 Genus Turricula Schumacher, 1817 Type species: Murex tornatus Dillwyn, 1817, by mono- typy; Recent, Indo-Pacific. Turricula sp. (Figures 49, 50) Description: Shell medium sized, slender; protoconch not preserved; teleoconch of five whorls; ramp slightly inclined and covers half of teleoconch whorl; surface bearing trace of labial sinus; shoulder prominent with 10 nodes; 18 spiral threads between shoulder and suture; threads disappear on base of siphonal neck. Material examined: Seven specimens. IHN 5366, height 28.6 mm, diameter 12.4 mm; IGM 8618, height 27.4 mm, diameter 12.3 mm. Discussion: The specimens from Chiapas are not similar to any species described from the Eocene of the Atlantic Coast of the United States. Pleurofusia raricosta (Gabb) (1864:93, 223, pl. 18, fig. 47) from the Eocene of California is a similar species with eight axial ribs terminating abruptly as slightly twist- ed nodes at shoulder. The Mexican specimens are similar to Turricula emerita Hickman (1976:36, pl. 1, figs. 14— 18) from the Lower Oligocene of Oregon in shape and sculpture. Order Heterostropha Fischer, 1885 Superfamily ARCHITECTONICOIDEA Gray, 1840 Family ARCHITECTONICIDAE Gray, 1850 Genus Architectonica Réding, 1798 Type species: Architectonica perspectiva Réding, 1798 (by subsequent designation, Gray, 1847:151); Recent, tropical western Pacific Ocean. The Veliger, Vol. 48, No. 3 Architectonica alabamensis (Dall, 1892) (Figures 51, 52) Solarium alabamense Dall, 1892:324, pl. 22, fig. 17; Harris, 1896a:114, pl. 12, fig. 4; Brann & Kent, 1960:803. Architectonica alabamensis (Dall). Palmer & Brann, 1966: 498. Description: Shell small sized, discoidal; protoconch of one and a half whorls, smooth; teleoconch with four whorls, first whorls with four spiral ribs increasing to eight on last whorl; growth lines not visible; flank of last whorl vertical with three strong, elevated ribs and two spiral threads between each rib; base with eight spiral ribs. Material examined: Two specimens. IHN 5386, height 5.7 mm, diameter 11.2 mm; IGM 8619, height 4.5 mm, diameter 8.9 mm. Discussion: The specimens from Chiapas do not clearly present growth lines and are larger than specimens de- scribed from Alabama. Architectonica sp. cf. A. elaborata (Conrad, 1833) (Figure 54) Description: Shell small sized, convex; protoconch of two whorls, smooth; teleoconch of three whorls, with sculpture of crenulated ribs alternating with spiral threads on adapical surface; base of shell with eight spiral ribs, one thickest at edge; aperture not preserved. Material examined: One specimen. IHN 5385, height 4.6 mm, diameter 10.1 mm. Discussion: One poorly preserved specimen with crenu- lated sculpture and small size similar to Architectonica elaborata (Conrad) (1833:344) from the Middle Eocene of Alabama. Architectonica sp. (Figure 53) Description: Shell small sized, discoidal; teleoconch of five whorls; sculpture of one strongly adapical rib to su- ture, rest of whorl smooth; margin with thick crenula- tions; umbilicus deep and wide. Material examined: One specimen. IHN 5388, height 6.8 mm, diameter 14.0 mm. Discussion: One poorly preserved specimen with a thick rib preserved; it differs from previously described speci- mens in having thick crenulations on the whorl periphery. Order Cephalaspidea Fischer, 1883 Superfamily ACTEONOIDEA Cernohorsky, 1972 Family ACTEONIDAE d’Orbigny, 1842 Genus Tornatellaea Conrad, 1860 Type species: Tornatellaea bella Conrad, 1860, by mon- otypy; Lower Eocene, Alabama, USA. M. del Carmen Perrilliat et al., 2005 Tornatellaea bella Conrad, 1860 (Figures 55, 56) Tornatellaea bella Conrad, 1860:294, pl. 47, fig. 23; Conrad, 1866:9; de Gregorio, 1890:166, pl. 16, fig. 19; Coss- mann, 1893:49; Cossmann, 1896:148, 150; Harris, 1899a:6, pl. 1, fig. 6; Harris & Palmer, 1947:460, pl. 64, fig. 10; Brann & Kent, 1960:869; Palmer & Brann, 1966:953; Toulmin, 1977:230, pl. 29, fig. 1. Tornatella (Tornatellaea) bella Conrad. Heilprin, 1879:22. Actaeon (Tornatellaea) bella Conrad. Harris, 1896a:74, in part. Description: Shell small sized, ovate; protoconch not preserved; teleoconch of six whorls; sculpture of promi- nent flat spiral ribs with narrow, punctuated interspaces; eight ribs on penultimate whorl and 23 ribs on last whorl; outer lip thick; columella with two prominent oblique folds. Material examined: 20 specimens. IHN 6550, height 12.0 mm, diameter 7.1 mm; IGM 8620, height 9.6 mm, diameter 6.0 mm. Discussion: Tornatellaea bella has been described from the Lower Eocene of Alabama. 7. lata (Conrad) (1834: 4) from the Middle Eocene of Alabama is more globose and has fewer spiral ribs. Acknowledgments. The authors wish to express their gratitude to CONACyT for the support provided to the project “‘Paleobi- ogeografia de moluscos y crustaceos del Eoceno y Oligoceno de Baja California Sur y Chiapas.”” We thank L. Saul and an anon- ymous reviewer for careful reviews of the manuscript. Thanks also are extended to José Manuel Padilla, who helped to elabo- rate the access and improvement of the graphics, and to Antonio Altamira and Fernando Vega for contributions to figures in this paper. PMERARURE CImED Apams, H. & A. ADAMS. 1853-1858. The genera or recent Mol- lusca arranged according to their organization. London. 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The Veliger 48(3):170-177 (November 2, 2006) THE VELIGER © CMS, Inc., 2006 Redescription of Two Antarctic Species of Cuspidaria: C. concentrica Thiele, 1912 and C. minima (Egorova, 1993) (Bivalvia: Cuspidariidae) DIEGO G. ZELAYA AND CRISTIAN ITUARTE Division of Invertebrate Zoology, Museo de La Plata, 1900 La Plata, Buenos Aires, Argentina; dzelaya@ museo.fcnym.unlp.edu.ar Abstract. Cuspidaria concentrica Thiele, 1912, frequently reported as a synonym of Cuspidaria kerguelensis Smith, 1885, is redescribed and revalidated following the study of new materials from the South Georgia Islands; this constitutes the first record of its presence in the Scotia Sea. The more elongated shell outline of C. concentrica and the smaller number of elevated lamellar concentric ridges clearly separate it from C. kerguelensis. Cuspidaria minima (Egorova, 1993), a species known from East Antarctica and the South Orkneys Islands, is redescribed based upon material from the Elephant Islands. C. minima is characterized by a small shell with a globose disk strikingly separated from the short rostrum and a shell surface with high, lamellated, upwardly bent commarginal ridges. Anatomical data, mainly from septal musculature of both species, are given. Type specimens of Cuspidaria tenella Smith, 1907, Cuspidaria plicata Thiele, 1912, and Cuspidaria infelix Thiele, 1912, are refigured and the reports on the presence of C. kerguelensis in the Scotia Arc Islands are discussed. INTRODUCTION The first record of Cuspidaria Nardo, 1840 in sub-Ant- arctic waters was provided by Smith (1885) who de- scribed C. kerguelensis from the Kerguelen Islands. Later, Smith (1907) described C. tenella from the Coulman Is- lands; Thiele (1912) described C. concentrica, C. plicata and C. infelix, all from Gauss Station, Antarctica (89°E). Egorova (1993) reviewed the Antarctic Cuspidariidae, and described Cuspidaria multicostata from off the South Sandwich Islands and Cuspidaria minima from the South Orkney Islands and East Antarctica, the latter, under Sub- cuspidaria, a genus proposed in the same paper (type species Neaera kerguelensis Smith, 1885). A number of recent contributions (Soot-Ryen, 1951; Dell, 1964, 1990; Miihlenhard-Siegel, 1989; Hain, 1990; Narchi et al., 2002) reported the above mentioned species as present also in West Antarctica and the Scotia Sea. However, rather poor original descriptions of all species (all based on a single valve and lacking in details) led to confusion and erroneous identifications and synonymies. In this regard, Soot-Ryen (1951) reported C. concentrica as synonymous with C. kerguelensis, a criterion also ac- cepted by Egorova (1993) but not by Dell (1990), who considered that C. concentrica differs from C. kerguelen- sis in having a fairly sparse sculpture. Dell (1964, 1990), Egorova (1993), and Poutiers & Bernard (1995) consid- ered C. plicata as synonymous with C. tenella. Nicol (1966), on the basis of similarities on shell sculpture, con- sidered C. plicaia and, surprisingly for the same reasons, C. concentrica as synonymous with C. tenella. Hain (1990) also included C. concentrica and C. plicata in the synonymy of C. tenella. Dell (1964) suggested that C. infelix could be regarded as an extreme variation of C. tenella; however, Dell (1990) considered the former as a valid species. As pointed out by Dell (1990), “‘elucidation of the Antarctic species of Cuspidaria is still difficult’. In the present paper Cuspidaria concentrica and C. minima are redescribed upon materials from the Scotia Arc Islands. Other species of Cuspidaria previously re- ported from the Scotia Arc Islands are also figured and compared with the species here studied in an attempt to clarify their identity. MATERIAL AND METHODS The material from the Scotia Arc Islands was collected during the 2002 Latin American Polarstern Studies (LAMPOS) aboard the R/V Polarstern and the 1996 sum- mer cruise to the South Georgia Islands by the R/V Ed- uardo Holmberg (voucher specimens in the collections of Museo Argentino de Ciencias Naturales “‘“Bernardino Ri- vadavia’” (MACN) and Museo de La Plata (MLP)). Spec- imens were fixed in a 10% formalin solution, sorted from the sediment under a stereoscopic microscope and pre- served in 80% ethanol. Shell morphology was studied and figured by scanning electron microscopy (SEM). Shell measurements were taken according to the following criteria: shell length (L), maximum anteroposterior distance; shell height (H), max- imum dorsoventral distance, perpendicular to length; and shell width (W), maximum distance across valves. Mor- phometric ratios H/L and W/H were calculated. The num- ber of specimens measured (n) and values of mean and D. G. Zelaya & C. Ituarte, 2005 standard deviation are given. Gross anatomy was studied under a stereomicroscope. Soft-part terminology of septal musculature follows Yonge (1928) and Allen & Morgan (1981). Several specimens for histological study were decal- cified by a 12-hr rinsing in a 10% formalin solution added with 2% acetic acid, embedded in Paraplast®, sectioned at 7 wm thickness, and stained with Meyer’s hematoxilin- eosin (Gabe, 1968). Type specimens of C. concentrica, C. plicata, and C. infelix (Museum fiir Naturkunde (ZMB), Berlin) and pho- tographs of the holotype of C. tenella (The Natural His- tory Museum (BMNH), London) were studied for com- parative purposes. The type of Neaera kerguelensis (BMNH) is presently badly damaged by Byne’s Disease (K. Way, in. lit.), and not photographable. Additional specimens of Cuspidaria plicata and C. infelix collected by the BANZARE Expedition from the South Australian Museum, Adelaide (SAM) were examined. Study of the types of Subcuspidaria minima (probably in the Zoolog- ical Institute, Russian Academy of Sciences, St. Peters- burg) was not possible. SYSTEMATICS Cuspidaria concentrica Thiele, 1912 (Figures 1—15) Cuspidaria concentrica Thiele, 1912:233, pl. 18, fig. 29 Diagnosis: Cuspidaria concentrica is characterized by its small size, the disk (i.e., the shell anterior to the rostrum) expanded anteriorly, and the shell surface with up to 16 lamellated commarginal ribs in larger specimens; a well- developed right-posterior lateral tooth, an inconspicuous left-posterior lateral tooth, and the anterior portion of the left-anterior septal muscle divided into three fascicles are also diagnostic. Description: Shell small (maximum L = 5.4 mm), ovate (H/L = 0.63 + 0.02, n = 15), somewhat globose (W/H = 0.75 + 0.03, n = 15), inequilateral, slightly inequivalve (in ventral view, left valve overlaps right one) (Figures 1-9). Anterior margin somewhat expanded into a wide curve, particularly evident in right valve; ventral margin evenly rounded at the anterior half, sinuous at posterior one. Anterior dorsal margin short, connected insensibly with anterior margin, which slopes gently; posterior dor- sal margin nearly straight in left valve, slightly curved in the right one (Figures 1—6, 8, 9). A weakly marked angle at the point of union of dorsal and anterior margins some- times present (Figures 3, 6). Rostrum moderately pro- jected, triangular in outline, dorsal and ventral margins not parallel, widely connected with disk. Transition be- tween disk and rostrum moderately demarcated. Two ros- tral ridges running from beaks to posterior end of rostrum delimit a triangular area where shell sculpture fades (Fig- ures 2, 4-6). Beaks wide, subcentral, directed posteriorly, Page 171 low but well visible above dorsal margin. Hinge: right valve has only a posterior lateral tooth present, being an elongated plate with low, central, or posteriorly displaced cusp (Figures 8, 10); left valve has only posterior lateral tooth represented by a thickening of dorsal margin, form- ing a weak articular relief (Figures 9, 11). Resilium elon- gated, posterior to beaks, inserted in a low, slender, re- silial pit (Figures 10, 11). Shell surface sculptured with a variable number of regularly spaced commarginal ribs: 9 to 12 in small specimens (3.5—4 mm L), 13-16 in larger ones (4.5—5.4 mm L). Ribs are lamellated, high in profile, anteriorly recurved, with rounded edges (Figure 13); in- terspaces much wider than ribs. Periostracum thin, form- ing weak, irregular, commarginal, and radial wrinkles par- ticularly noticeable between ribs. Prodissoconch well dis- cernible from dissoconch, about 195 jm in maximum diameter, finely granular (Figure 12). Dissoconch micro- sculpture formed by granules of two different sizes ran- domly distributed (Figure 14). Soft parts: Anterior half of left anterior septal muscle clearly divided into a few, usually three, bundles of fibers; inner septal muscle well developed, joining the anterior septal muscle slightly before the insertion; at this point, part of the fibers of both muscles merge in a single in- sertion area (Figure 15). Lateral septal muscle formed by a continuous series of fibers, not arranged in bundles. Incurrent and excurrent siphons surrounded by seven short tentacles: three in middorsal and dorsolateral posi- tions, and four in lateral and ventrolateral positions; each tentacle ending in an enlarged, rounded tip. Examined material: Holotype (ZMB 63120); 187 spec- imens, 54°18’S, 35°30’W, South Georgia Islands, 94 m (MLP 5655, MACN-In 36377); 32 specimens, 53°59’S, 37°38'W, 158-159 m, South Georgia (MLP 7352, MACN-In 36378), 7 specimens, 54°30'S, 56°08’W, Burd- wood Bank, 286 m (MLP 6873). Remarks: Cuspidaria concentrica resembles Cuspidaria minima (Figures 16-26) and Cuspidaria kerguelensis (Figure 28) in general shell shape, but consistently differs in having a disk more anteriorly expanded (particularly evident in the right valve), the rostrum less markedly dif- ferentiated from the disk, and a smaller number of higher lamellated ribs. C. concentrica also differs from C. min- ima in having a weaker right-posterior lateral tooth and an inconspicuous posterior lateral tooth in the left valve. Cuspidaria plicata (Figure 29) differs from C. concentri- ca in having a more elongated disc and nonlamellated commarginal ribs. The shell sculpture of Cuspidaria te- nella (Figure 30) and Cuspidaria infelix (Figure 31) with irregular and fine commarginal striae differs strikingly from that present in C. concentrica. Page 172 The Veliger, Vol. 48, No. 3 D. G. Zelaya & C. Ituarte, 2005 Figures 13-14, Cuspidaria concentrica from South Georgia Is- lands (MLP 5655): shell sculpture. Figure 13. Detail of shell sculpture. Figure 14. Detail of shell microsculpture. Scale bars: Fig. 13 = 140 um; Fig. 14 = 50 pm. Cuspidaria minima (Egorova, 1993) Figures 16—27 Subcuspidaria minima Egorova, 1993:164—165, pl. 3, figs. 2 de Diagnosis: Cuspidaria minima is characterized by its small size, short anterior end, and rounded, rather globose disk, strikingly demarcated from the rostrum. The shell surface with up to 15 lamellated commarginal ribs, the hinge with a robust right-posterior tooth, and a stout ma- melliform left-posterior lateral tooth, as well as the an- terior portion of the left-anterior septal muscle usually divided into several fascicles, are diagnostic features. Description: Shell solid, whitish, small (maximum L = Page 173 Figure 15. Cuspidaria concentrica: diagrammatic detail of an- terior septal musculature. Scale bar = 500 jm. (asm = anterior septal muscle, ilm = inner longitudinal muscle). 4 mm), ovate (H/L: 0.63 = 0.02, n = 13), inequilateral, slightly inequivalve (ventrally, left valve overlaps the right one) (Figures 16—19); disc and rostrum well-sep- arated. Disk globose (W/H: 0.81 + 0.04, n = 13), cir- cular in right valve, tending toward trigonal in left one; transition between disk and rostrum strikingly demar- cated by a depression of shell surface (particularly pro- nounced in left valve) (Figures 16-21); rostrum mod- erately projecting, handlike; a well-marked rostral ridge running from beaks to posterior end; a second, less pro- nounced rostral ridge running closer to posterodorsal margin (Figures 16-18). Beaks wide, subcentral, direct- ed posteriorly, low but well-visible above dorsal margin. Anterior margin short, evenly, and widely rounded in right valve, sloping markedly in left one; ventral margin evenly rounded anteriorly, steeply curved posteriorly. Anterior dorsal margin very short, insensibly connected with the anterior margin; posterior dorsal margin slightly curved (Figures 16—21). Shell surface sculptured with lamellated commarginal ribs (13 to 15 in specimens of 3.5—4 mm L), regularly separated; rib edge rounded; spaces between ribs wider Figures 1-12. Cuspidaria concentrica: Figure 1. Holotype (ZMB 63120). Figures 2, 4-12 Specimens from South Georgia Islands (MLP 5655). Figure 2. Left valve. Figure 3. Right valve of a specimen from Burdwood Bank (MLP 6873). Figures 4, 5. Right valve; Figure 6. Juvenile specimen; Figure 7. Ventral view; Figure 8. Inner view of right valve. Figure 9. Inner view of left valve; Figures 10, 11. Detail of hinge. Figure 10. Right valve. Figure 11. Left valve. Figure 12. Prodissococonch. Scale bars: Figs. 1-9 = 1 mm; Figs. 10, 11 = 250 wm; Fig. 12 = 100 pm. (plt = posterior lateral tooth, r = resilium) Page 174 The Veliger, Vol. 48, No. 3 Figures 16-26. Cuspidaria minima from Elephant Islands (MLP 6881). Figure 16. Left valve. Figure 17. Right valve. Figure 18. Juvenile. Figure 19. Ventral view. Figure 20. Inner view of right valve. Figure 21. Inner view of left valve. Figure 22. Right valve detail of hinge. Figure 23. Left valve detail of hinge. Figure 24. Detail of shell sculpture. Figure 25. Detail of shell microsculpture. Figure 26. Prodissoconch. Scale bars: Figs. 16-21 = 1 mm; Fig. 22 = 500 wm; Fig. 23 = 250 pm; Figs. 24, 26 = 100 um; Fig. 25 = 25 jum. (aar = anterior articular relief, plt = posterior lateral tooth, r = resilium). D. G. Zelaya & C. Ituarte, 2005 Page 175 Figure 27. Cuspidaria minima: diagrammatic detail of septal musculature. Scale bar = 500 pm. (asm = anterior septal muscle, ilm = inner longitudinal muscle, psm = posterior septal muscle). than ribs (Figure 24). Commarginal ribs not extending dorsal to ventralmost rostral ridge. Periostracum forming weak, irregular, commarginal, and radial wrinkles (Figure 24). Prodissoconch well-discernible from dissoconch, about 175 wm of maximum diameter, finely granulated (Figure 26). The entire surface of dissoconch covered by low, circular depressions, variable in size, and micro- scopic granules (Figure 25). Hinge: a triangular, recurved, robust posterior lateral tooth in right valve, cusp rounded, displaced posteriorly (Figures 20, 22); a blunt mamelli- form posterior-lateral tooth in left valve (Figures 21, 23). Articular reliefs anterior to beaks (anterior lateral teeth?), as shallow depressions and low protuberances, present in both right and left valves (Figures 20, 21). Resilium elon- gated, posterior to beaks, attached to a low and slender resilifer (Figures 22, 23). Soft parts: The anterior portion of left-anterior septal muscle divided into several (usually six or seven) well- discernible, slender bundles of fibers (Figure 27). The in- sertion area of the left anterior septal muscle is well-dif- ferentiated from the one that corresponds to the inner lon- gitudinal muscle. Incurrent and excurrent siphons sur- rounded by seven somewhat short tentacles: three in middorsal and dorsolateral positions and four in lateral and ventrolateral positions; tentacles ending in enlarged, rounded tips. Examined material: 69 specimens, 61°23’S, 55°26’W, Elephant Islands, 285 m (MLP 6881). Remarks: Cuspidaria minima is most similar to Cuspi- daria kerguelensis (Figure 28) in shell shape; judging from the figure in the original description of C. kergue- lensis, C. minima differs in having a smaller number of commarginal ribs; the spaces between the ribs are nearly equal to the rib width in C. kerguelensis, while in C. minima spaces are up to 3 times the rib width; in C. minima commarginal ribs fade above the rostral ridge while in the figure of C. kerguelensis, ribs extend onto the entire surface of the rostrum. The shorter anterior half of the disk in Cuspidaria min- ima, as well as the presence of a strong posterior-lateral tooth in the right valve and a larger number of muscular bundles in the anterior half of the left-anterior septal mus- cle, clearly separate C. minima from C. concentrica. The posterior curvature of the ventral margin in C. minima changes more abruptly than in C. concentrica, and as a consequence, the disk and rostrum appear strikingly dif- ferentiated. Also, C. minima has, within the same range of size, a larger number of commarginal ribs than C. con- centrica. Figures 28-31. Other Antarctic species of Cuspidaria. Figure 28. C. kerguelensis: figure from the original description. Figure 29. C. plicata: holotype (ZMB 63121). Figure 30. C. tenella: holotype (BMNN 1905.9.25.11). Figure 31. C. infelix: holotype (ZMB 63119). Scale bars: Fig. 28 = 1 mm; Fig. 29 = 5 mm; Figs. 30, 31 = 10 mm. Page 176 Cuspidaria minima differs from the Antarctic species C. tenella (Figure 30) and C. infelix (Figure 31) in shell sculpture: the two latter species do not have lamellated ribs but rather irregular commarginal striae and periostra- cal folds. C. tenella also differs in having a larger shell and a proportionally smaller rostrum (Figure 30); C. in- felix differs in being more expanded anteriorly, with the disk widely connected with the rostrum (Figure 31). Cuspidaria plicata (Figure 29), another Antarctic spe- cies, differs from C. minima in having a smaller number of low, nonlamellated commarginal ribs. Cuspidaria minima was originally described under Subcuspidaria, a genus proposed by Egorova (1993). Subcuspidaria was defined in the restricted context of a revision of Antarctic species of Cuspidaria, without con- sidering the wide variability known for the genus. Hence, the value of the set of characters proposed by Egorova (1993) as diagnostic at genus level needs to be reevalu- ated in a wider context. So far, we prefer to use Cuspi- daria for all the Antarctic species here studied. This is consistent with the catalogue of the Recent Anomalodes- mata by Poutiers & Bernard (1995). DISCUSSION The new data given in this paper represent a contribution towards a better definition of two of the most common Antarctic species of Cuspidaria. There is considerable confusion of the systematics of the sub-Antarctic and Antarctic species of Cuspidaria and, consequently, their actual geographical distributions are uncertain. Much of this confusion arises from the poor information on each taxon based on the small number of specimens studied; this has led to misinterpretations and erroneous synonymies. Such is the case, for example, for Dell (1990) who, based on ‘‘the small [shell] size, the strong but sparse commarginal sculpture and the whole aspect of Thiele’s original figure,’ considered that Cus- pidaria concentrica was described from a juvenile spec- imen (the type is 3.6 mm length). Our histological data showed that ripe eggs were present in females of C. con- centrica of about 4 mm length, a fact that clearly indi- cates the adulthood of these “small sized’ specimens. The same was observed in Cuspidaria minima, in which specimens of less than 4 mm length showed ripe ova and sperm. Another source of misleading information is the fact that authors often have not considered the differences in shell outline and general shell shape existing be- tween right and left valves: while the anterior half of the disk in the right valve is generally evenly rounded and more expanded anteriorly, in the left valve, the anterior end of the disk is shorter and slopes more abruptly. In this regard, it should be noted that a large number of the Antarctic species of Cuspidaria were described based on only one valve, and consequently, The Veliger, Vol. 48, No. 3 subsequent comparisons seeking similarities or differ- ences could be biased. Acknowledgments. The authors wish to acknowledge D. Na- habedian (University of Buenos Aires) for making available the specimens of C. concentrica collected by the R.V. E. L. Holm- berg from South Georgia Islands; W. Arntz (Alfred Wegener In- stitute) for allowing D. Z. to be a part of the “LAMPOS 2002” Expedition to Antarctica; M. Glaubrecht (ZMB) and R. Hamilton Bruce (SAM) for loaning specimens; K. Way and A. Campbell for kindly providing photographs and additional information on types lodged at BMNH, London; FE Scarabino for helping with the bibliography; and M. Szereszowiez for translating Egorova’s (1993) paper. This study was partially funded by student grants to D. Z. from the Western Society of Malacologists, the Santa Barbara Mala- cological Society, the Southwest Shell Club, the San Diego Shell Club, and the Northern California Malacological Club. The au- thors are members of the Consejo Nacional de Investigaciones Cientificas y Técnicas (CONICET), Argentina. LITERATURE CITED ALLEN, J. A. & R. E. MorGan. 1981. The functional morphology of Atlantic deep water species of the families Cuspidariidae and Poromyidae (Bivalvia): An analysis of the evolution of the septibranch condition. Philosophical Transactions of the Royal Society of London B, Biological Sciences 294(1073): 413-546. DELL, R. K. 1964. Antarctic and sub-Antarctic Mollusca: Am- phineura, Scaphopoda and Bivalvia. Discovery Reports 33: 93-250. Det, R. K. 1990. Antarctic Mollusca with special reference to the fauna of the Ross Sea. Bulletin of the Royal Society of New Zealand 27:1—311. Ecorova, E. N. 1993. Antarctic Cuspidaria (Cuspidariidae, Cus- pidariida, Bivalvia). [in Russian}. Antarktika 32:151—166. GaBE, M. 1968. Techniques Histologiques. Masson Ed.: Paris. 1113 pp. Hain, S. 1990. The benthic seashells (Gastropoda and Bivalvia) of the Weddell Sea, Antarctica. Reports on Polar Research 70:1—211+30 pls. MUHLENHARD-SIEGEL, U. 1989. Antarktische Bivalvia der Reisen des FS ‘‘Polarstern’’ und des FFS ‘Walther Herwig” aus den Jahren 1984 bis 1986. Mitteilungen aus dem Hambur- gischen Zoologischen Museum und Institut 86:153—178. Narculi, W., O. DOMANESCHI & FE. D. Passos. 2002. Bivalves Antarticos e Subantarticos coletados durante as expedicdes cintificas brasileiras a Antartica I a LX (1982-1991). Revista Brasilera de Zoologia 19:645—675. NIcot, N. 1966. Description, ecology and geographic distribution of some Antarctic pelecypods. Bulletin of American Pale- ontology 51:1—102. Poutiers, J. M. & E R. BERNARD. 1995. Carnivorous bivalve molluscs (Anomalodesmata) from the tropical western Pa- cific Ocean, with a proposed classification and a catalogue of Recent species. In P. Bouchet (ed.), Résultats de Cam- pagnes MUSORSTOM. Vol. 14. Mémoires du Muséum Na- tional d’ Histoire Naturelle 167:107—187. Situ, E. 1885. Report on the Lamellibranchiata collected during the voyage HMS ‘Challenger’. Challenger Expedition, Zo- ology 13:1—341. Situ, E. 1907. Mollusca. 5. Lamellibranchiata. National Ant- arctic Expedition 1901-1904. Natural History II, Zoology (Vertebrata, Mollusca, Crustacea): 1—6. D. G. Zelaya & C. Ituarte, 2005 Page 177, Soot-RYEN, T. 1951. Antarctic Pelecypods. Scientific results of the YONGE, C. M. 1928. Structure and function of the organs of Norwegian Antarctic Expeditions 1927-1928, 32:1—46 + 1 pl. feeding and digestion in the septibranchs, Cuspidaria and THIELE, J. 1912. Die Antarktischen Schnecken und Muscheln. Poromya. Philosophical Transactions of the Royal Society Deutsche Siidpolar Expedition 1901—1903, 13:183—285. of London B, Biological Sciences 216:221—261. THE VELIGER © CMS, Inc., 2006 The Veliger 48(3):178—205 (November 2, 2006) Shedding Light onto the Genera (Mollusca: Nudibranchia) Kaloplocamus and Plocamopherus with Description of New Species Belonging to These Unique Bioluminescent Dorids YVONNE VALLES ann TERRENCE M. GOSLINER University of California, Berkeley, Integrative Biology Department, Valley Life Sciences, Berkeley, California 94720, USA California Academy of Sciences, 875 Howard Street, San Francisco, California 94103, USA Abstract. Members of the Family Triophinae (Rudman, 1998), Gastropoda; Opisthobranchia, are characterized by having a reduced mantle, a narrow ridge along the sides of the body and a veil surrounding the head. This family is composed by seven genera of which only two have been observed to have bioluminescent species; Kaloplocamus Bergh, 1892 and Plocamopherus Riippell & Leuckart, 1831. In order to better understand the evolution of bioluminescence within these dorids a phylogenetic analysis based on morphological and molecular data comprising all genera pertaining to the subfamily Triophinae was undertaken. Because there is great taxonomic confusion regarding Kaloplocamus and Plocamopherus species, this study is an attempt to do a taxonomic and morphological revision of both genera. Further- more the description of new species for both Kaloplocamus and Plocamopherus are included. Five species of Kaloplo- camus were examined in this study, including three taxa described here: K. dokte n. sp.; K. peuldo n. sp. and K. maru n. sp. Seven species of Plocamopherus were examined including four new taxa: P. pecoso n. sp., P. maculapodium n. sp., P. margaretae n. sp. and P. lemur, n s.p. The review of the morphology of these species provides the basis for future phylogenetic studies. INTRODUCTION Dorids are marine hermaphroditic slugs that have lost the protective shell of most gastropods. As a result of the loss of this structure and the process of detorsion, dorids ex- hibit an incredible development of exuberant colors and forms. Color patterns are used as a defensive mechanism. Bright and attractive colors usually indicate that the man- tle contains chemicals that are generally toxic to predators (Avila, 1995). Subtler colors may be used as a camou- flage strategy (however most cryptic species have chem- ical defenses, as well). Other modes of protection are be- havioral (autotomy of the mantle outgrowths, swimming), presence of spicules and use of their prey defense system (storage of chemical compounds). Much less common within dorids, is the emission of bioluminescence to startle possible predators. Biolumi- nescence is the ability to create light as a result of a chemical reaction within a living organism. First noted by Aristotle as light of luminous wood (Johnson & Yata, 1966), early naturalists believed it to be a creation from god, just for the pleasure of the human eye, with no sig- nificant function. However, it is well known that the abil- ity to produce such light requires significant expense of energy and, depending on the organism, it has a well- defined function (i.e., communication, prey attraction, de- fense, etc.) (O’Day & Fernandez, 1974; Day et al., 2004). Although bioluminescence is known to occur in a di- versity of organisms such as bacteria, fungi, dinoflagel- lates, cnidarians, annelids, arthropods, echinoderms, tu- nicates, fishes and some mollusks (Cephalopoda), it has only been observed within few dorid species: Kaloplo- camus ramosus (Cantraine, 1835), Plocamopherus imper- ialis Angas, 1864, P. tilesii Bergh, 1877 and P. maderae (Lowe, 1842) of the two phanerobranch genera (Kaloplo- camus and Plocamopherus). In both cases, the lumines- cence is intrinsic which means that the production of light is dependent on the animal’s own biochemical processes (no symbiotic association with bacteria) (Wilbur & Yon- ge, 1966). However, in Kaloplocamus ramosus the chem- ical reaction takes place within luminescent cells or pho- tocytes being (intracellular luminescence) whereas in Plo- camopherus species, it results from the discharge of the luminous chemicals outside the cell where the chemical reaction occurs (extracellular luminescence). It is possible that since these organisms emit light only when disturbed, the role of bioluminescence could be to temporarily dis- tract the attention of potential predators (Wilbur & Yonge, 1966). Surprisingly, although both Kaloplocamus and Ploca- mopherus are the only dorids with this peculiar charac- teristic, they have been greatly understudied. In order to understand the origin and evolution of bioluminescence in these dorids, it is essential to understand the phylo- genetic relationships between them, which cannot be Y. Valles & T. M. Gosliner, 2005 Page 179 done unless the taxonomy of both genera is revised. The goal of this study is two-fold. First it includes a taxonom- ic and morphological revision of the genera Kaloploca- mus Bergh, 1879 and Plocamopherus Riippell and Leuck- art, 1831. Second we provide the descriptions of three new species of Kaloplocamus and four new species of Plocamopherus. MATERIALS AnD METHODS Twelve species have been anatomically examined. The specimens used for this study were obtained through sev- eral institutions: California Academy of Sciences, Mu- séum National d’ Histoire Naturelle, Paris, Zoologisk Mu- seum, Kgbenhavn University, Copenhagen and South Af- rican Museum, Cape Town. Most of the specimens were preserved in Bouin’s fixative and/or 95% ethanol. All type specimens that were available have been examined. For the newly described species, type material has been designated. Specimens were dissected with the aid of a Nikon SMZ 10 or a Nikon SMZ zoom 110 dissecting microscope and drawings of the different structures were made with the aid of a camera lucida. For all specimens, an initial de- scription of the external morphology was completed by microscopy to obtain details from the mantle and other external structures. A posteriori, for the internal exami- nation, an incision with a scalpel was made at the right side/ridge of the specimen from the head to the foot to expose the buccal mass and the reproductive system. Within the buccal mass, two structures, radula and jaws, were prepared for further examination. The radula was placed in a 10% sodium hydroxide solution for approxi- mately 48-72 hours, depending on the size and on the strength of the muscular tissue around it. The radula was then rinsed in distilled water and submerged in an ultra- sound bath (Branson, 1510) to eliminate any additional tissue. Finally, using fine pointed probes, the radula was mounted on a stub to air-dry. The jaws were opened and mounted on a stub immediately after their extraction from the buccal mass to dry. Stubs were then coated with gold for two minutes using a Hummer 5-sputter coater. Coated samples were viewed by using a LEO scanning electron microscope. All specimens were viewed with a current of 15 kv. A full external morphological description, SEM pic- tures of radula and jaws, and drawings of the reproductive system have been completed for all specimens available. The external morphological descriptions are based on photographs and notes made by the collector. Genus Kaloplocamus, Bergh, 1879 Kaloplocamus Bergh, 1879:634; 1884:56 Euplocamus Philippi, 1836:103; Alder, 1845:262; Alder and Hancock, 1855:xix; Bergh, 1879:623, Bergh, 1883:165, 1884:56; Vayssiere, 1901:65; Bergh, 1907:70; Vayssi- ere, 1913:341; O’ Donoghue, 1929:775 Idalia Leuckart, 1828; Philippi, 1844:76 Type species: Euplocamus croceus Philippi, 1836. The genus Kaloplocamus (Opisthobranchia: Doridoi- dea) is poorly known. All species were described between 1835 and 1955 with the exception of K. filosus (Cattaneo- Vietti & Sordi, 1988). Most of these species have only been reported when originally described. Because several generic names have been attributed to this genus a review of the taxonomic history of the generic name Kaloplo- camus is necessary. Cantraine (1835) briefly described the species Doris ramosa from the Adriatic Sea. It was characterized as an elongated specimen, with tubercles and bearing six ram- ified velar appendages and ramified branchial leaves. No illustrations of the animal were provided and because the information regarding the features exhibited by the spec- imen were scarce; a definitive identification of the species is controversial. A year later, Philippi (1836) described the genus Eu- plocamus based on a new species: Euplocamus croceus. The description was based on two specimens collected in 1832. The genus was characterized by specimens with a probosciform anterior part of the body and presence of branchial leaves of two types, ones shorter and positioned in circle surrounding the anus and the others longer and situated at the sides of the body (referring to the lateral appendages that are characteristic of the group). The ap- erture of the reproductive system was noted as situated at the right side of the body. The genus was described as intermediate between the dorids (branchial leaves sur- rounding the anus) and the tritonids (the latter includes genera possessing ramified appendages which function as respiratory systems). Because, until then, the specimens had just one kind of branchial leaves, either in circlet or as lateral appendages, Philippi considered the specimen as unique and therefore representative of a new genus, Euplocamus (Philippi, 1836). Forbes, stated that the genus Euplocamus should be synonymized with the genus Jriopa (Johnston, 1838), (Forbes, 1841). He based this judgement on his obser- vation of the lack of cilia on the lateral appendages. This would constitute proof of the misidentification of the function of the lateral appendages (respiration) provided by Philippi, since the genus Triopa was composed by specimens that had lateral appendages not involved in respiration as he noted for E. croceus. Cantraine (1841) considered two groups within Doris: Doris planes, composed of those with the capacity to de- tach some parts of the body, and Doris prismatiques, composed of those that do not have such a capacity. Since he did not consider the lateral appendages of Ewplocamus croceus as branchial leaves, and he did not observe the ability to lose body parts when disturbed, he included Page 180 Polycera and Euplocamus in the group of Doris prisma- tiques, and therefore the genus Doris (Cantraine, 1841). Cantraine did not refer to the comments that were made by Forbes pertaining to this matter. Following Forbe’s (1841) and Cantraine’s (1841) re- marks, Euplocamus was synonymized with /dalia (Phi- lippi, 1844). Even though a note was written on the va- lidity of the genus Euplocamus, explaining the differenc- es between the genera /dalia, Triopa and Euplocamus (Alder, 1845), Loven included Euplocamus as a synonym of Idalia, and reported that the generic name Euplocamus had already been given to a lepidopteran (Loven, 1846). In 1847, Gray created a list of genera that needed (ac- cording to him) to be synonymized including the genus Euplocamus. After examination of the different species that were described as part of the genus, Gray divided them into two groups. The species that had simple lateral appendages were synonymized with the genus /dalia and those that had ramified lateral appendages were synony- mized with the genus 7riopa (Gray, 1847). Several au- thors, (Alder & Hancock, 1854; Gray, 1857) provided again the necessary evidence to distinguish /dalia, Triopa and Euplocamus. Despite this, Abraham synonymized the latter once more with the genus 7riopa (Abraham, 1877), where he stated that “‘the differences between the two genera, however, appear to be more of degree than of kind, and serve better to distinguish sections of one genus It was not until much later that Bergh proposed, to use the generic name Kaloplocamus to replace Euplocamus to avoid any possible confusion with other taxa that had already the same generic name (Bergh, 1879a). The latter had already been given to both a butterfly and a bird. The name Kaloplocamus was also chosen because of its sim- ilarity with the initial name. However, from 1884 through 1925, even Bergh ignored this designation and published under the generic name Euplocamus. The first record of using the name of Kaloplocamus dates from 1925 (Thiele, 1925) for the description of Kaloplocamus orientalis. Surprisingly, after all the trial synonymies there is only one attempt to revise the genus, (Baba, 1989). However, this revision only involves two species that are commonly encountered in Japan, K. ramosus and K. acutus. Even though Cattaneo-Vietti & Sordi (1988) gave an outline of the species included in the genus, no real revision was attempted. Today the valid generic name considered is Kaloplo- camus being the type species Euplocamus croceus. Six- teen species have been described for this genus: K. ra- mosus (Cantraine, 1835); K. japonicus (Bergh, 1879b); K. principiswalliae (Collingwood, 1881); K. pacificus Bergh, 1884; K. atlanticus (Bergh, 1893); K. maculatus (Bergh, 1898); K. longicornis (Bergh, 1905); K. tristis (Bergh, 1905); K. capensis (Bergh, 1907); K. yatesi (An- gas, 1864); K. orientalis (Thiele, 1925); K. aureus The Veliger, Vol. 48, No. 3 (Odhner, 1932); K. acutus (Baba, 1955); K. gulo (Marcus, 1979); K. filosus (Cattaneo-Vietti & Sordi, 1988). Most of these species were described between 1836 and 1905. However, over time, many of them have been synonymized. Even though Bergh described K. maculatus in 1898 and K. capensis, K. tristis and K. longicornis in 1905 and 1907 as new species, in 1908 he states that the only valid species of the genus were K. croceus, (var., K. capensis and var. K. altanticus); K. japonicus and K. pa- cificus. In this article, he also doubted about the validity of the two latter species (Bergh, 1908). Kaloplocamus differs externally from all other genera included in the subfamily Triophinae, by several features. The lack of a globular structure on the apex of the ap- pendages and the absence of a penial sac in the repro- ductive system distinguish Kaloplocamus from Ploca- mopherus. The shape and length of the lateral append- ages, its ramifications, and several radular features (ab- sence of elongated and thin teeth as in Crimora or triangular rachidian plates as in Triopha) distinguish it from both genera Crimora and Triopha. The types of most described species are not available, since most have been lost through the years (misplace- ment of the specimens, or poor preservation). As a result of the lack of type material and the synonymies proposed over the years, three species are considered as valid in this study, K. pacificus, K. ramosus and K. acutus. Only K. ramosus is known to be bioluminescent. This work includes the description of three new spe- cies. Because it is difficult to test that these species are reproductively isolated, the description of new species is based on morphological features such as the external gen- eral shape, texture, color and shape of the mantle, lateral appendages and anatomical features such as those present in the radula and the reproductive system. TERMINAL TAXA Kaloplocamus ramosus (Cantraine, 1835) (Figures 1A, 2, 3) Doris ramosa Cantraine, 1835: 383 Type material: Doris ramosa Cantraine: the type mate- rial is lost. Material examined: CASIZ 072609, one spec- imen, dissected. Length (preserved): 9 mm. Ilha de Sao Miguel, Azores, Atlantic Ocean. Depth: 60 feet. 20 July 1988. Collector: T. M. Gosliner. CASIZ 087006, one specimen. Length (preserved): 9 mm. Faial, Azores, At- lantic Ocean. October 1992. Collector: J. Brun. Distribution: This species is known from the Mediter- ranean Sea, Japan, Australia, New Zealand, Hong Kong and the Atlantic Ocean including Angola and South Af- rica. Y. Valles & T. M. Gosliner, 2005 Figure 1. Living animals. A—Kaloplocamus ramosus (9 mm). B—Kaloplocamus acutus (12 mm). C—Kaloplocamus dokte (8 mm). D—Kaloplocamus peludo (7 mm). E—Kaloplocamus maru (8 mm). F—Plocamopherus cf. imperialis (13 mm). Photo by B. Bolland. G—Plocamopherus maculatus (19 mm). H—Plocamopherus tilesii (32 mm). Photo by B. Bolland. I—Plocamopherus maculapodium (13 mm). J—Plocamopherus pecoso (19 mm). K—Plocamopherus lemur (8 mm). L—Plocamopherus margaretae (40 mm). Photo by M. Strickland. All lengths were measured from preserved animals. All other photos by T. Gosliner. External Morphology: The living animal has a bright orange-red color (Figure 1A). The entire dorsum has small tubercles that are translucent orange. Kaloplocamus ramosus has eight velar ramified appendages, which are thin and have secondary sharp, elongate ramifications. The oral tentacles are flat and wide. The rhinophores are the- same color as the rest of the body. The clavus is slightly speckled with small white dots. The rhinophoral sheath is very short and speckled similar to the clavus of the rhinophore. The dorsum has four pairs of ramified lateral appendages. These appendages are not as thin as the ones on the veil, being somewhat flat and wide with sharp and elongate ramifications at the apex. The bran- chial leaves are of the same color as the rest of the body, tripinnate. Radula and Jaws on Buccal Armature: The radular for- mula of the specimen dissected is 17 X (17.6.0.6.17). Two well-differentiated types of teeth are observed in the radula (Figure 2A, C). The six inner lateral teeth are sim- Page 182 The Veliger, Vol. 48, No. 3 Figure 2. Kaloplocamus ramosus (CASIZ 072609) SEM photographs of the radula and jaws. A. Whole radula. B. Inner lateral teeth. C. Outer lateral teeth. D. Jaws. Figure 3. Reproductive system of Kaloplocamus ramosus (CASIZ 072609). Abbreviations: am, ampulla; bc, bursa copu- latrix; f, female gland mass; s, receptaculum seminis; u, uterine duct; v, vagina; vd, vas deferens. ilar in shape, sharply hook-shaped with a secondary cusp well developed (Figure 2B). This secondary cusp is more developed in the innermost tooth and becomes less evi- dent towards the outer lateral. There are seventeen outer lateral teeth, which are rectangular in shape and flat (Fig- ure 2C). The size of the outer lateral teeth decreases from the innermost to the outermost. The rachis is papillated and separated in different rectangular plates, which are wider in the upper part of the radula. Presence of jaws with thin, rounded, elongate and densely packed rodlets (Figure 2D). Reproductive System: The reproductive system (Figure 3) is triaulic. The vas deferens is not differentiated into a prostate. The female gland is oval somewhat elongate. The vagina is half as wide as the vas deferens and is two- thirds its length. It is connected to the bursa copulatrix. From the bursa copulatrix merges a thin and elongate duct that divides into two equally thin ducts. The first duct leads to the receptaculum seminis and second one, the uterine duct connects to the female gland mass. The latter is very thin but twice as long as the duct of the recep- Y. Vallés & T. M. Gosliner, 2005 taculum seminis. The ampulla, which is thinner than the vagina and almost as long, connects to the female gland mass near the connection of the vas deferens and the bur- sa copulatrix ducts. Remarks: The morphological features exhibited by the specimen dissected correspond exactly to those described by Cantraine (1835). Kaloplocamus ramosus and K. acu- tus differ primarly in their external morphology. Both of them have lateral appendages, but in the case of K. acutus the appendages are not as ramified at the apex, and are sharper. The white tips at the top of the clavus of the rhinophores, as well as the red-purple coloration on the lateral appendages present only in K. acutus, distinguish them. The length of the duct leading from the recepta- culum seminis to the female gland mass, which is twice as long as the one leading to the bursa copulatrix in K. ramosus and of the same length in K. acutus, separates these species. In a similar sized specimen, K. ramosus always has a larger number of both types of teeth. Also the hook shaped portion of the inner lateral teeth is much longer and sharper in K. ramosus than K. acutus. Kalo- plocamus ramosus differs from K. pacificus in the exter- nal morphology by not having a dorsal keel (present in K. pacificus). Anatomically they differ by the presence of rodlets in the jaws of K. ramosus and the absence in K. pacificus (Bergh, 1884). We have followed Marshall and Willan’s synonymy of K. yatesi to K. ramosus (Marshall & Willan, 1999). Kaloplocamus japonicus and K. ramosus were synony- mized (Eliot, 1913) since there were very few differences between them, based mostly on the number of appendages. Thiele (1925) described K. orientalis based on a different number of teeth. Since the variation in the number of teeth in the radula is highly variable within species in the genus it cannot be considered as a good character to establish new species. Therefore, K. orientalis should also be re- garded as a synonym of K. ramosus. Kaloplocamus prin- cipiswallie was synonymized with K. japonicus (Pruvot- Fol, 1935) based on the poor description given by Col- lingwood (1881). Nordsieck suggested that the form K. aureus of Odhner 1932 should be synonymized with K. ramosus (Nordsieck, 1972) and K. filosus was synony- mized with K. ramosus (Vallés et al., 2000). Kaloplocamus acutus Baba, 1955 (Figures 1B, 4, 5) Kaloplocamus acutus Baba, 1955: 45 Type material: Kaloplocaums acutus Baba. Material ex- amined: CASIZ 116905, two specimens, one dissected. Length (preserved): 12 mm. Depth: unknown. Puako in cave at night, Hawaii. 3 May 1981- Collector: Scott John- son. CASIZ 065370, one specimen, dissected. Length (pre- served): 8 mm. Madang, Papua New Guinea. Depth: 23 m. 22 May 1988. Collector: R. C. Willan. Page 183 Distribution: Kaloplocamus acutus was first described from Sagami Bay, Japan (Baba, 1955). It has been reported from the Indo—Pacific. Reported from Papua New Guinea and Hawaii (present study). External Morphology: The body shape of the living an- imal is elongate (Figure 1B). The anterior part of the body is somewhat wider than the rest of the body. The head bears six velar appendages, which are semi-translucent. Opaque white coloration is present on the basal third of the velar appendage, with the remainder being translucent. These appendages have acute, simple, and elongate rami- fications, each of which is bright red-carmine in color. The body has a background coloration of orange-red, with white dots over the entire dorsum. Both rhinophores are orange-red as the body, with a white line on the anterior part of the clavus. There are four lateral appendages along each lateral side of the body. The first three pairs of ap- pendages are situated between the rhinophores and the branchial leaves. The fourth pair is located just behind the branchial leaves. The lateral appendages are similar to the veil appendages in that they have the same coloration pat- tern. There are five bipinnate branchial leaves. The bran- chial leaves have an opaque white coloration throughout their whole length when observed form an anterior angle. However, when viewed from a posterior angle, they have the same coloration as the rest of the body. As with all the appendages, the branchial leaves have sharp and simple elongated ramifications that are red carmine in color. The posterior part of the foot is elongate, acutely pointed and red-orange in color, speckled with white dots. Radula and Jaws on Buccal Armature: The radular for- mula of the specimen dissected is 16 X (11.3.0.3.11). Two different types of teeth are observed in the radula (Figure 4A-C). The three innermost teeth are similar in shape and hook-shaped with a secondary cusp developed in a lower position of the tooth. This secondary cusp is more pro- nounced in the lateral innermost teeth and disappears by the third inner lateral tooth. All three inner lateral teeth are long and the hook portion is sharp. The eleven outer lateral teeth are flat and rectangular in shape (Figure 4A, C). Their size decreases towards the outer margins. The papillated rachis has a series of transverse plates that decrease in size towards the base of the radula. The jaws have rodlets that are thin, rounded, elongate and densely packed (Figure 4D). Reproductive System: The reproductive system (Figure 5) is triaulic. The vas deferens is not differentiated into a prostate. The vagina, although it is almost as wide as the vas deferens, is half as long. It is connected directly to the bursa copulatrix. The bursa copulatrix is similar in size to the receptaculum seminis to which is connected by a duct that enlarges and divides itself into two ducts. One duct opens into the receptaculum seminis and the other, is the uterine duct that opens into the female gland mass. The The Veliger, Vol. 48, No. 3 m7 io Yj [Pane ene Pan ; 3S 10jsm Figure 4. Kaloplocamus acutus (CASIZ 116905) SEM photographs of the radula and jaws. A. Whole radula. B. Inner lateral teeth. C. Outer lateral teeth. D. Jaws. Figure 5. Reproductive system of Kaloplocamus acutus (CAS- IZ 116905). Abbreviations: am, ampulla; bc, bursa copulatrix; f, female gland mass; s, receptaculum seminis; u, uterine duct; v, vagina; vd, vas deferens. ampulla, wider than the vas deferens, and as long as the vagina, penetrates into the female gland near the connec- tion of the vas deferens and the bursa copulatrix ducts. The connection between the ampulla and the vas deferens occurs within the female gland mass. Remarks: Kaloplocamus acutus was first described by Baba (1955). It was shortly described as having orange- yellow background coloration. Characterized by the pres- ence of six velar processes, four pairs of lateral appendages with a small number of sharp pointed branches that were always carmine red and a radula with two inner lateral teeth (Baba, 1955). The dorsum of the present specimen has four pairs of ramified appendages that are larger and longer than those of the veil. The branchial leaves have the same orange coloration as the rest of the body. All appendages have a pink-red-purple coloration at the apex of the ramifications and the rhinophores are red. Both, the red carmine coloration on the apex of the lateral and oral veil ramifications and the white lines on the anterior part of the clavus of the rhinophores are characteristic of the species. Also typical of the species is the presence of the Y. Vallés & T. M. Gosliner, 2005 Page 185 Figure 6. Kaloplocamus dokte (CASIZ 146066) SEM photographs of the radula and jaws. A. Whole radula. B. Inner lateral teeth. C. Jaws. simple and sparse ramifications on the veil and lateral ap- pendages. Kaloplocamus acutus differs from K. pacificus in its external morphology by not having a dorsal keel (present in K. pacificus) by having lateral and frontal ap- pendages that have simple ramifications. Kaloplocamus dokte n. sp. (Figures 1C, 6, 7) Type and examined material: Holotype, CASIZ 086448, one specimen. Length (preserved): 4 mm. Depth: 3 m. Bar- racuda Point, Madang, Papua New Guinea. 9 June 1992. Collector: T. M. Gosliner. Paratypes: CASIZ 146066, one specimen, dissected. Length (preserved): 8 mm. Depth: 8 m. Barracuda Point, Madang, Papua New Guinea. 30 Jan- uary 1988. Collector: T. M. Gosliner. CASIZ 078586, one specimen. Length (preserved): 3 mm. Depth: unknwon. Bwaken Island, Manado, Sulawesi, Indonesia. Date: un- known. Collector: P. Fiene. CASIZ 146060, one specimen. Length (preserved): 4 mm. Depth: 8 m. St 52, Barracuda Point, Madang, Papua New Guinea. 5 July 1989. Collector: T. M. Gosliner. CASIZ 146068, one specimen. Length (preserved): 4 mm. Depth: 8 m. Barracuda Point, Papua New Guinea. 16 July 1989. Collector: T. M. Gosliner. CASIZ 146057, one specimen. Length (preserved): 5 mm. Depth: 8 m. Barracuda Point, Madang, Papua New Guinea. 27 August 1989. Collector: T. M. Gosliner. CASIZ 075887, one specimen. Length (preserved): 6 mm. Depth: 6.6 m. Barracuda Point, Madang, Papua New Guinea. 19 Novem- ber 1990. Collector: T. M. Gosliner. CASIZ 146065, one specimen. Length (preserved): 2 mm. Depth: 8 m. Barra- cuda Point, Madang, Papua New Guinea. 14 August 1984. Collector: T. M. Gosliner. CASIZ 146067, three specimens. Length (preserved): 2—3 mm. Depth: 8 m. Barracuda Point, Madang, Papua New Guinea. 31 August 1989. Collector: T. M. Gosliner. CASIZ 146059, one specimen. Length (preserved): 5 mm. Depth: 8 m. Barracuda Point, Madang, Papua New Guinea. 22 July 1989. Collector: T. M. Gos- liner. CASIZ 146063, one specimen. Length (preserved): 3 mm. Depth: 8 m. Barracuda Point, Madang, Papua New Guinea. 14 August 1989. Collector: T. M. Gosliner. CASIZ 146064, one specimen. Length (preserved): 5 mm. Depth: 8 m. Barracuda Point, Madang, Papua New Guinea. 4 Feb- ruary 1988. Collector: unknown. CASIZ 146061, one Page 186 The Veliger, Vol. 48, No. 3 Figure 7. Reproductive system of Kaloplocamus dokte (CASIZ 146066). Abbreviations: am, ampulla; bc, bursa copulatrix; f, fe- male gland mass; s, receptaculum seminis; u, uterine duct: v, vagina; vd, vas deferens. specimen. Length (preserved): 5 mm. Depth: unknown. Rasch Passage, Madang, Papua New Guinea. 20 August 1989. Collector: T. M. Gosliner. CASIZ 083657, one spec- imen. Length (preserved): 6 mm. Depth: unknown. Luzon Island, Philippines. 19 February 1992. Collector: T. M. Gosliner. Distribution: Specimens of this species have been found in Papua New Guinea, Indonesia and the Philippines. External Morphology: When alive, this species appears to be ovoid in shape but upon a closer view, the posterior portion of the foot is elongated (Figure 1C). Most of the animal is opaque white except for the posterior end of the foot and the eight velar appendages, which are highly transparent. The dorsal and central portions of the body are translucent and the digestive gland is visible. The three tripinnate branchial leaves, are a translucent red-carmine. The rhinophores are of the same color as the branchial leaves, but more translucent with white dots on the apex. The long, large, lateral appendages are totally opaque white except at the apex, where they ramify and become more cream in tone. There are four pairs of lateral ap- pendages. They are smooth with ramifications in the apex only. Instead of being sharp and long these ramifications are rounded, almost globular and there are many of them resembling in shape to a bunch of grapes. These ramifi- cations only occur on the exterior portion of the lateral appendage, with the interior portion being smooth over its entire length. Radula and Jaws on Buccal Armature: The specimen dissected had a radula formula of 10 X (5.3.0.3.5). The radula is characterized by the presence of two different types of teeth (Figure 6A, B). The three inner lateral teeth, (Figure 6A, B) similar in shape, are elongated and hook- shaped at the apex. The hook-shaped portion of the teeth is short. The innermost lateral tooth has a smaller cusp than the two adjacent teeth and has a pronounced second- ary cusp near the primary one. The outer lateral teeth (Fig- ure 6A, B), five per side, are flat and rectangular in shape. The size of these rectangular teeth diminishes from the innermost to the outermost. The papillated rachis has a series of transverse plates that are regular in shape and size for the whole length of the radula. There are an equal number of rachidian plate rows, as rows of teeth. Presence of jaws with thin, elongate, rounded rodlets that are dense- ly packed (Figure 6C). Reproductive System: The reproductive system (Figure 7) is triaulic. The vas deferens is not differentiated in its distal portion into a prostate. The thin vagina is half the length of the vas deferens and before entering into the bursa copulatrix joins the duct from the receptaculum sem- inis. The duct connecting receptaculum seminis and bursa copulatrix, divides into two ducts, one leading to the oval receptaculum seminis and the other, the uterine duct, en- tering into the female gland mass. The ampulla is thin and short, and enters into the female gland mass near the con- nection of the vas deferens and the bursa copulatrix ducts. It presumably bifurcates into the vas deferens and oviduct within the female gland. Remarks: This species is distinguished by the opaque white coloration, never reported before for a Kaloplocamus species. The presence of four ramified appendages that are of the same length and are ramified are typical of the ge- nus. However the ramifications shape (similar to a bunch of grapes) makes this species unique. Another interesting feature that distinguishes K. dokte from other Kaloploca- mus is the small radula with a small number of teeth. Kal- oplocamus dokte differs from K. pacificus in the absence of a keel as well as differences on the lateral and frontal appendages that are not tripinnated. As in K. ramosus and K. acutus, four lateral appendages were observed as well as a total of eight veil appendages, which are shorter and smaller then the former. However, K. dokte has lateral ap- pendages that bear small secondary ramifications that in- stead of being sharp and elongated as in K. acutus or ram- ified as in K. ramosus, they are rounded. The radula is different from the typical structure of the K. ramosus and K. acutus specimens in that the three innermost lateral hook-shaped teeth are large relative to the size of the rad- ula. Etymology: This species is named “‘dokte” for the father of the senior author, whose nickname in Haitian kreol is Dokte. Y. Valles & T. M. Gosliner, 2005 Page 187 Figure 8. Kaloplocamus peludo (CASIZ 085981) SEM photographs of the radula. A. Whole radula. B. Inner lateral teeth. C. Outer lateral teeth. Figure 9. Reproductive system of Kaloplocamus peludo (CAS- IZ 085981). Abbreviations: am, ampulla; be, bursa copulatrix; f, female gland mass; s, receptaculum seminis; u, uterine duct; v, vagina; vd, vas deferens. Kaloplocamus peludo n. sp. (Figures 1D, 8, 9) Type and examined material: Holotype, CASIZ 083828, one specimen. Length (preserved): 6 mm. Depth: 20 m. Batangas, Luzon Island, Philippines. 23 February 1992. Collector: T. M. Gosliner. Paratypes: CASIZ 085981, four specimens, two dissected. Length (preserved): 8-4 mm. Depth: 33 m. Kirby’s Rock, NW side of the Maricaban Island, Batangas, Luzon Island, Philippines. 26 March 1993. Collector: T. M. Gosliner. CASIZ 118946, one spec- imen. Length (preserved): 11 mm. Depth: unknown. Ba- tangas, Luzon Island, Philippines. March 1995. Collector: Mike Miller. CASIZ 085934, two specimens. Length (pre- served): 9-11 mm. Depth: 23 m. Mindanao, Philippines. 31 March 1993. Collector: T. M. Gosliner. CASIZ 086294, two specimens. Length (preserved): 6-7 mm. Depth: 3 m. Madang, Barracuda Point, Papua New Guinea. 9 June 1992. Collector: T. M. Gosliner. CASIZ 115384, two spec- imens. Length (preserved): 7-10 mm. Depth: 55 m. Ser- agaki, Okinawa, Ryukyu Island, Japan. 127°52.60’E; Page 188 26°30.40'N. 26 March 1997. Collector: R. F Bolland. CASIZ 099222, one specimen. Length (preserved): 11 mm. Depth: unknown. Mtwara, Tanzania. 31 October 1994. Collector: T. M. Gosliner. CASIZ 121138, one spec- imen. Length (preserved): 6 mm. Depth: 13 m. Marshall Islands: Kwajalein Atoll. Ennubuj Island: Gea Island: ocean side of island. 24 October 1992. Collector: S. John- son. Distribution: Several specimens of this new species have been found in many different localities ranging from the Philippines, Papua New Guinea, Palau, Marshall Islands and Japan in the Western Pacific to Tanzania in the West- ern Indian Ocean. External Morphology: The living animal has a general orange coloration (Figure 1D). The whole dorsum has brown dots except at the apices of the lateral appendages, which are pale yellow. These brown dots are similar and size and regularly distributed. The dorsum has an opaque white, irregularly shaped line, which runs from the front of the head to the posterior end of the notum. This line bifurcates at the level of the branchial leaves, where it surrounds them and remerges again behind the branchial leaves. The posterior end of the foot is thin and acute and has a yellow color similar to that of the lateral appendage apices. Two types of appendages are observed in the spec- imen. The first type of appendages is the longest and wid- est. There are three pairs of these lateral appendages situ- ated along the margin of the sides. These appendages have a ramified apex. Each ramification of the apex has a round- ed base and a long, thin and sharp prolongation. The whole length of each of these appendages is covered by little and acute ramifications. The second type of appendages, are thin, long with many small sharp, thin and long simple ramifications. These appendages are present on the sides of the body, foot, dorsum and oral veil in number 15—17 per side with 6 on the oral veil. The rhinophores have a lamellate clavus and a peduncule with little, thin and sharp ramifications. Each rhinophoral sheath edge has three ram- ifications. The middle one is virtually identical to the lat- eral appendages. The other two ramifications are similar to the smaller tentacles of the rest of the body. The oral ten- tacles are simple folds of the mantle that have a cylindrical shape. There are three tripinnate branchial leaves. They present long, simple, and thin ramifications as the smaller appendages. Radula and Jaws on Buccal Armature: The radular for- mula of the specimen is 11 X (8.3.0.3.8). Two different types of teeth are present in the radula (Figure 8A—C). The three inner lateral teeth similar in shape have a blunt hook with a secondary cusp (blunt as well) developed in a lower position of the tooth. The outermost eight teeth are rect- angular in shape and their size decreases from the inner- most to the outermost. The papillated rachis is not well The Veliger, Vol. 48, No. 3 separated into different plates being just one whole plate for the radula. No rodlets were observed on the jaws. Reproductive System: The reproductive system (Figure 9) is triaulic. The vas deferens is differentiated in its distal portion into a wide, translucent prostate. A thin short ovi- duct emerges from the prostate joining the oval ampulla before entering into the female gland mass. The vagina is very thin and is a fourth of the length of the vas deferens. Before entering into the bursa copulatrix it joins the duct from the receptaculum seminis. The duct connecting re- ceptaculum seminis and bursa copulatrix, divides into two ducts, one leading to the oval receptaculum seminis and the other, the uterine duct, entering into the female gland mass. Remarks: This species is characterized by the several fea- tures, which make it very different from the other known species of Kaloplocamus. The presence of two kinds of lateral appendages which bear on the apex long, thin and simple filaments; the presence of tubercles with thin and long ramifications as well all over the body; and the par- ticular shape of the rhinophores sheaths disguising them- selves as lateral appendages are unique of K. peludo. Even though the general morphology of the radula is typical of the genus Kaloplocamus, several differences are found. The inner lateral teeth have a blunt apex instead of the typical sharp hook shaped. The presence of a granulated rachis characteristic of the genus, differs by not been trans- versally divided in rachidian plates rows. Other important differences are the absence of rodlets in the jaws and the presence of a distinctive wider prostate gland. Etymology: This species is named peludo for the long and thin filaments (hair-like) present on the apex of the lateral appendages. Peludo is a Spanish adjective to indicate that something or someone has a lot of hair. Kaloplocamus maru n. sp. (Figures 1E, 10, 11) Type and examined material: Holotype, CASIZ 170939. Length (preserved): 6 mm. Depth: 0-27 m. Badeldoab Is- land, Ngerduais Island, Palau. 21 September 1996. Collec- tor: T. M. Gosliner. Paratype: CASIZ 109685, two speci- mens, two dissected. Length (preserved): 6-10 mm. Depth: 0-27 m. Babeldoab Island, Ngerduais Island, Palau. 21 September 1996. Collector: T. M. Gosliner. Distribution: This species has been collected in Palau. External Morphology: The living animal has a general orange color (Figure 1E). The whole dorsum has brown dots except the tips or apex of the lateral appendages, which are pale yellow. The dorsum has a white diamond shaped spot that runs from the behind of the rhinophores to the tail (just behind the branchial leaves). The end of the foot is acute and has a similar color to the appendage apices. Two types of appendages are observed. The first Y. Vallés & T. M. Gosliner, 2005 20m Figure 10. Kaloplocamus maru (CASIZ 109685) SEM photographs of the radula and glands. A. Whole radula. B. Inner lateral teeth. C & D. Buccal glands. type of appendages are longer and wider and are posi- tioned at the margin of the sides of the animal. These ap- pendages have apical ramifications that are rounded at the base with a long, thin and sharp prolongation. Small acute ramifications cover the remaining parts of the appendage. The second type of appendage is thin, long with many small sharp, thin and long simple ramifications. These ap- pendages are present on the entire body, sides, foot, and dorsum. The oral veil has four appendages, two of which are longer, situated at the lateral edges of the oral veil and two that are shorter situated in the center. The rhinophores have a lamellate clavus and a peduncule with acute rami- fications. Each rhinophoral sheath edge has three ramifi- cations. The middle one is virtually identical to the lateral appendages. The other two ramifications are similar to the smaller tentacles of the rest of the body. The oral tentacles are.simple folds of the mantle with the appearance of cyl- inders. The three branchial leaves are tripinnate. They have the same simple and thin ramifications as the smaller ap- pendages. Radula and Jaws on Buccal Armature: The radula is enveloped in a cylindrical buccal mass, that has large glan- dular structures at both sides (Figure 10C, D). Their func- tion has not yet been identified. The radular formula of this species is 68 X (7.2.0.2.7). The radula of Kaloploca- mus maru is characterized by being a long ribbon of teeth (Figure 10A). There are two types of teeth. There are two inner lateral teeth with similar shape with the inner one being smaller than the outer one (Figure 10A, B). Both teeth are hook-shaped at the apex but the remaining tooth body seems to be folded forward, having each folded edge developed into a protuberance in the middle of their length. The outer lateral teeth are flat and have an irregular shape. The rachis is reduced to a line between both sides of the radula (Figure 10B). No jaws were observed. Reproductive System: The reproductive system is triaulic (Figure 11). The vas deferens is not differentiated into a prostate. It is as long and almost as wide as the vagina but becomes thinner in its proximal portion. Proximally the Page 190 vd Figure 11. Reproductive system of Kaloplocamus maru (CAS- IZ 109685). Abbreviations: am, ampulla; bc, bursa copulatrix; f, vagina; vd, vas deferens. vagina joins the duct coming from the receptaculum sem- inis, becoming a single duct. The latter splits into two other ducts, one of which enters into the bursa copulatrix and the other (uterine duct) connects with the female gland mass. The ampulla is wide and oval in shape and the upper part differentiates into a narrow and short duct that con- nects with the female gland mass near the connection of the vas deferens and the bursa copulatrix ducts. Remarks: Kaloplocamus maru is very similar to Kaloplo- camus peludo in the general external morphology. Kalo- plocamus maru is characterized by features such as the presence of long, thin and simple ramifications present in the apex of the lateral appendages; the presence of tuber- cles with thin and long ramifications as well all over the body (but many fewer then those present in K. peludo); and the particular shape of the rhinophoral sheaths dis- guising themselves as lateral appendages. Even though both species (K. peludo and K. maru) are externally very similar, anatomical differences clearly separate them. The radula of Kaloplocamus maru is atypical of the genus Kal- oplocamus. Two types of radular teeth are present in both species but differ markedly in their features. In K. maru the inner lateral teeth have a much shorter, blunt cusp than in K. peludo. The outer lateral teeth are irregular in K. maru but are rectangular in K. peludo. The absence of a wide granulate rachis is also atypical for the genus, in this The Veliger, Vol. 48, No. 3 case it is observed only as a very narrow line that runs longitudinally the radula. Usually, Kaloplocamus radular general morphology is wider than longer in contrast to that of K. maru, which is much longer than wider. Because of the atypical features of the radula two specimens were ex- amined having both the same radula. Other differences are the presence of a dinstinct prostate in K. peludo versus its apparent absence in K. maru. Etymology: This species is named “‘maru’”’ after the moth- er of the senior author, whom we all call Maruxa. GENUS PLOCAMOPHERUS RUPPELL AND LEUCKARGES3i Histiorophorus Pease, 1860:35 Peplidia Lowe, 1842:51 Plocamophoris Whitelegge, 1889 Plocamorphorus Fewkes, 1889 Plocamoceros J. E. Gray, 1857 Plocamophorus Okada and Baba, 1938:276 Type species: Plocamopherus ocellatus Rtippell & Leuck- art, 1831. The genus Plocamopherus was described based on a new species, Plocamopherus ocellatus characterized by having a shell-less body, elongate, doridiform, convex dor- sum, and armed at both sides with simple tentacles (Riip- pell & Leuckart, 1831). The anterior margin has ramified appendages, retractile rhinophores, a mid-dorsal anus sur- rounded by the branchial leaves, and a reproductive system opening on the right side. Plocamopherus ocellatus (type species) was characterized by a shaded color, dorsal oval spots of a sulfur color, and presence of conical papillae at both. This species was described from the Red Sea (Rtip- pell & Leuckart, 1831). Even though several changes of the spelling of the name Plocamopherus have been published throughout the his- tory of the genus (Lowe, 1842; Gray, 1857; Pease, 1860; Fewkes, 1889; Whitelegge, 1889; Okada & Baba, 1938), its validity has never been questioned, with the exception of Risbec, who synonimized it with Triopha and Triopa, because he found it impossible to distinguish the three gen- era (Risbec, 1928). Subsequent authors did not accept this synonymy. Plocamopherus has been considered as very closely re- lated to the genera Kaloplocamus and Triopha. Included as well in the subfamily Triophinae, Plocamopherus has several characteristics that allow its unequivocal identifi- cation. A limaciform body usually with up to three pairs of lateral appendages characterizes it. In most cases, the most posterior pair of appendages have a bioluminescent structure on their apices composed of columnar cells that produce a secretion that, when excreted produces light. This structure is sometimes also present in some species, on the other two pairs of appendages, but when present, it is usually much smaller. This structure is unique to Plo- Y. Valles & T. M. Gosliner, 2005 Page 191 Figure 12. Plocamopherus cf. imperialis (CASIZ 079194) SEM photographs of the radula and jaws. A. Whole radula. B. Inner lateral teeth. C. Outer lateral teeth. D. Jaws. camopherus species. However the production of biolumi- nescence is not limited to the bioluminescent structures. The head and the sides of the body as well as the crest of the tail emit a less intense light than the bioluminescent structures (as occurs Kaloplocamus ramosus). Another fea- ture of Plocamopherus is their ability to swim (Lowe, 1842, 1843). Most species of this genus have a vertically flattened postero-dorsal keel on the foot that allows them to swim by lateral undulation. There are seventeen described species: Plocamopherus amboinensis (Bergh, 1890); P. apheles (Barnard, 1927); P. ceylonicus Kelaart, 1858; P. flagellatus (Kruzenstern, 1813); P. fulgurans (Risbec, 1928); P. imperialis Angas, 1864; P. indicus (Bergh, 1890); P. insignis (Smith, 1884); P. levivarius (Abraham, 1876a); P. [ucayensis Hamann and Farmer, 1988; P. maculatus Pease, 1860; P. maderae Lowe, 1842; P. naeavatus (Abraham, 1876b); P. ocellatus Rtippell and Leuckart, 1831; P. pilatecta Hamann and Farmer, 1988; P. ramulosus (Stimpson, 1855); and P. ti- lesii Bergh, 1877. Plocamopherus naevatus was synonymized with Plo- camopherus imperialis (Bergh, 1883). Thompson in 1975 proposed P. amboinensis; P. apheles, P. flagellatus; P. ful- gurans, P. indicus, P. levivarius as dubious species. These descriptions were based on preserved material (Thompson, 1975). The descriptions not only lack data on the external coloration, but anatomical details as well, since most of them were not dissected. Thompson determined the fol- lowing valid species: Plocamopherus ocellatus, P. mad- erae, P. maculatus, P. ceylonicus, P. imperialis, and P. tilesii. TERMINAL TAXA Plocamopherus cf. imperialis Angas, 1864 (Figures 1F 12, 13) Plocamopherus imperialis Angas, 1864:59 Type material: Plocamopherus imperialis Angas, was lost. Material examined: CASIZ 079194, one specimen, dissected. Length (preserved): 13 mm. Depth: 1.5 m. Ser- agaki Beach, Ryukyu Islands, Okinawa. 25 March 1991. Collector: R. E Bolland. CASIZ 089062, one specimen. Length (preserved): 8 mm. Depth: 3 m. Seragaki Beach, Page 192 Figure 13. Reproductive system of Plocamopherus cf. imperi- alis (CASIZ 079194). Abbreviations: am, ampulla; bc, bursa co- pulatrix; f, female gland mass; p, prostate; s, receptaculum sem- inis; u, uterine duct; v, vagina; vd, vas deferens. Ryukyu Islands, Okinawa. 1 January 1993. Collector: R. FE Bolland. Distribution: Known from temperate Southeastern Aus- tralian waters and northern New Zealand (Rudman 1998) and from Okinawa. External Morphology: In life, its body shape is elongate, limaciform and anteriorly rounded (Figure 1F). The oral veil is wide, flattened and has eight velar processes. The presence of flat oral tentacles was observed. The present specimen has a yellowish background color. The entire body is highly speckled with brown dots that are more intense on the dorsum than the sides of the body. There are larger blotches of brown coloration surrounding the lateral appendage bases. The rhinophoral peduncles are pink with a brownish clavus. The rhinophoral sheaths are short with the margins minutely papillated. There are three pairs of lateral appendages, the first two being conical in shape and having small-branched papillae on the tip. Both tips are pink as is the globular structure on the third pair. The same structure and coloration is presented by the velar appendages. There are five tripinnate branchial leaves, which are very elongate and elevated. The posterior por- tion of the foot is elongated and thin and has a small keel that is slightly fringed on its upper margin. The posterior end of the foot is flattened laterally. Radula and Jaws on Buccal Armature: The specimen dissected had a radula formula of 15 x (12.5.0.5.12). Two types of teeth are present. The five inner lateral teeth (Fig- ure 12A—C) are similar in shape, slightly broad and hook- shaped at the apex. The apex is very short, flattened and The Veliger, Vol. 48, No. 3 almost blunt. The five innermost teeth have a small sec- ondary cusp. The outer lateral teeth (Figure 12A—C), twelve per side, are flat and rectangular in shape. The size of these rectangular teeth decreases from the innermost to the outermost. The rachis has a series of papillated trans- verse plates, whose size decreases towards the base of the radula. There is an equal number of rachidian plates rows as rows of teeth. The jaws have thin, elongate and flattened rodlets that are densely packed (Figure 12D). Reproductive System: The reproductive system is triaulic (Figure 13). The vas deferens is differentiated in its prox- imal portion into a wide penial sheath. On its distal portion the vas deferens becomes half as thick and differentiates into the prostate. The massive prostate is connected to the female gland by a thin and short duct. The vagina is almost as wide as the vas deferens but half as long. From the large and oval bursa copulatrix, emerges a short and thin duct that divides into two ducts, one of which is connected to the receptaculum seminis and the other is connected to the vagina. The uterine duct emerges from the receptacu- lum duct and connects with the female gland mass near the prostate duct. The ampulla is narrow, convoluted and long and enters into the female gland mass in a position distal relative to the penetration of both the vas deferens and the uterine ducts. Remarks: Even though Angas (1864) did not give any details of the anatomy P. imperialis in the original descrip- tion, this species has a rather unique external coloration characterized by the presence of the three lateral pink ap- pendages, a yellow-brown background color and the pres- ence of irregular brown dots all over the body. However some differences are found in the color pattern (brown blotches on the lateral appendages apices) and the geo- graphical distribution. In order to be certain that both spe- cies are the same, an analysis of the Australian species would be necessary. Anatomical details as a non-envel- oped bursa copulatrix (by the prostate) differentiates them from P. ceylonicus, P. tilesii, P. maculatus, where the bur- sa is totally enveloped by the prostate. Plocamopherus maculatus (Pease, 1860) (Figures 1G, 14, 15) Plocamopherus maculatus (Pease, 1860): 35 Type material: Plocamopherus maculatus Pease, was not available. Material examined: CASIZ 116824, three spec- imens, one dissected. Length (preserved): 19-20 mm. Depth: 10 m West side Pig Island, Madang, Papua New Guinea. 7 February 1988. Collector: T. M. Gosliner. CAS- IZ 073147, one specimen. Length (preserved): 8 mm. Depth: 20 m. Cement Mixer Reef, Madang, Papua New Guinea. 22 October 1986. Collector: T. M. Gosliner. CAS- IZ 071152, one specimen. Length (preserved): 15 mm. Depth: 9 m. Pig Island, Madang, Barracuda Point Papua Y. Vallés & T. M. Gosliner, 2005 Figure 14. Plocamopherus maculatus (CASIZ 116824) SEM photographs of the radula and jaws. A. Whole radula. B. Inner lateral teeth. C. Outer lateral teeth. D. Jaws. Figure 15. Reproductive system of Plocamopherus maculatus (CASIZ 116824). Abbreviations: am, ampulla; be, bursa copu- latrix; f, female gland mass; p, prostate; s, receptaculum seminis; u, uterine duct; v, vagina; vd, vas deferens. New Guinea. 7 February 1988. Collector: T; M. Gosliner. CASIZ 086317, three specimens. Length (preserved): 4— 12 mm. Depth: 10 m. South side Rasch Passage, Madang, Papua New Guinea. 16 June 1992. Collector: T. M. Gos- liner. South African Museum, Cape Town, A35560, one specimen. Length (preserved): 18 mm. Depth: unknown. K 212. 14 May 1981. Collector: unknown. Distribution: Plocamopherus maculatus has been reported from Hawaii (Kay & Young, 1969), Western Australia (Wells & Bryce, 1993), South Africa (Gosliner, 1987) and is here reported from Papua New Guinea. External Morphology: In life the body is elongate, li- maciform and anteriorly rounded (Figure 1G). The head has a wide flattened and fringed oral veil with numerous papillae. The oral tentacles are flat. Plocamopherus ma- culatus has translucent white background coloration. Typ- ical of the species are the red-orange dots on the dorsum between the rhinophores and in the central part of the dor- sum as well as crimson and white pigment surrounding the base of the branchial leaves. The whole body is slightly and regularly speckled with minute brown-orange dots. Page 194 The rhinophores are for their entire length translucent white and are speckled with minute crimson dots as is the rest of the body. The rhinophoral sheaths are short. There is only one pair of lateral appendages situated behind the branchial leaves that bears the white globular structure. There are two pairs of slightly ramified conical tubercles at the tip at each side of the body. There are five tripinnate branchial leaves, which are elongate. The posterior portion of the foot is elongate and thin, and has a small keel that is slightly fringed on its upper margin. Radula and Jaws on Buccal Armature: The specimen dissected had a broad radula with formula of 28 X (11.11.0.11.11). Two types of teeth are present. The eleven inner lateral teeth (Figure 14A—C), similar in shape, are elongate and slightly hook-shaped at the apex. The apex is very elongated, somewhat flattened and fairly sharp. All inner lateral (except for the two outermost) have a well- developed secondary cusp. The outer lateral teeth (Figure 14A, B), eleven per side, are flat and rectangular in shape. The size of these rectangular teeth decreases from the in- nermost to the outermost. The rachis has a series of pap- illated transverse plates that are very regular in shape and size through the whole length of the radula. There is an equal number of rachidian plates rows, as rows of teeth. The jaws have thin, elongate and flattened rodlets that are densely packed (Figure 14D). Reproductive System: The reproductive system is triaulic (Figure 15). The vas deferens is differentiated in its distal portion into a well-developed and glandular-looking pros- tate. The prostate is connected to the female gland by a thin and short duct. The vagina is almost as wide as the vas deferens in its proximal portion, and connects directly into the bursa copulatrix. The vagina is one-third the length of the vas deferens. From the rounded and small bursa copulatrix, emerges a short and thin duct that divides into two ducts, one of which is connected to the receptaculum seminis and the other penetrates into the female gland mass. Both ducts are of similar length and width. The am- pulla is wide, long and convoluted and enters into the fe- male gland mass in a position distal relative to the con- nection of both the vas deferens and the uterine ducts. Remarks: Described under the generic name Histiophorus (Pease, 1860), it was synonymized with the genus Ploca- mopherus by Bergh (1879b). Plocamopherus maculatus was characterized by the whitish background coloration with the orange speckles and the crimson dots at the base of the branchial leaves. The coloration of this species is rather unique among the Plocamopherus species. The fea- tures exhibited by the radula are also characteristic of the species where the number of innermost and outer lateral is identical (usually the former being more numerous). The Veliger, Vol. 48, No. 3 Plocamopherus tilesii Bergh, 1877 (Figures 1H, 16, 17) Plocamopherus tilesii Bergh, 1877:433 Type material: Plocamopherus tilesii Bergh, was not found. Material examined: CASIZ 079251, one specimen, dissected. Length (preserved): 32 mm. Depth: 50 m. Ser- agaki Beach, Ryukyu Islands, Okinawa. 7 July 1991. Col- lector: R. EK Bolland. CASIZ 97426, one specimen. Length (preserved): 11 mm. Depth: unknown. Oman Arabian Sea: Ra’s ad Duqm: tidepool. October-November 1993. Collec- tor: J. L. Earle. Distribution: Known from Japan, China, Arabian Sea. Also reported occasionally from New South Wales, al- though no published records (www.seaslugforum.net). External Morphology: In life its body shape is oval and anteriorly rounded (Figure 1H). The oral veil is wide, flat- tened and bears numerous short somewhat flattened ap- pendages that vary in length. The presence of flat oral tentacles was observed. Plocamopherus tilesii has a trans- lucent, yellowish background color and the whole body is homogeneously speckled with regular black and white spots. There are yellow patches that are irregular in size with distribution over the entire body. Also present are large brownish spots that are fewer in number. The oral veil margin is lined by a yellow coloration, however the oral appendages are black for their entire length. The rhin- ophores are translucent for their whole length, speckled extensively with brown dots. The rhinophoral sheaths are short and have large, irregular, black spots on the margin. There are three pairs of short lateral appendages, with the last two pairs having a prominent globular structure that is white in color. There are three tripinnate branchial leaves. The posterior portion of the foot is short and flattened forming a keel that has a small crest. Radula and Jaws on Buccal Armature: The dissected specimen. had a radular formula of 18 X (8.10.0.10.8). The radula of P. tilesii is characterized by the presence of two different types of teeth (Figure 16A—C). The inner lateral teeth, (Figure 16A—C) similar in shape, are hook-shaped. The primary cusp is elongated and sharp. Only the first three innermost teeth have a well-developed secondary cusp. The base of the innermost lateral teeth becomes less rectangular and more triangular towards the lower teeth rows of the radula. The outer lateral teeth (Figure 16A, C), eight per side, are flat and rectangular in shape. The size of these rectangular teeth decreases from the innermost to the outermost. The papillated rachis has a series of trans- verse plates that decrease in size towards the base of the radula. The jaws were lost. Reproductive System: The reproductive system is triaulic (Figure 17). The vas deferens is differentiated in its distal portion into a well-developed and glandular looking pros- Y. Valles & T. M. Gosliner, 2005 Page 195 Figure 16. Plocamopherus tilesii (CASIZ 079251) SEM photographs of the radula and jaws. A. Whole radula. B. Inner lateral teeth. C. Outer lateral teeth. tate that envelops the whole bursa copulatrix. The prostate is connected to the female gland by a very short and thin duct. The vagina is as half as wide as the vas deferens but is almost the same length. From the rounded bursa copu- latrix, emerges a short duct that divides into two ducts, one, which is connected to the vagina and the other to the receptaculum ‘seminis. This second duct before entering into the receptaculum seminis splits again into the uterine duct that penetrates the female gland mass. The ampulla is very elongate and convoluted and enters the female gland mass in a position distal relative to the penetration of both the vas deferens and the uterine ducts. Remarks: According to the original description (Bergh, 1877) this species has a light brown-gray background color speckled with dark purple and yellow spots all over the body. The yellow spots are larger and may be found at the margins of the mantle. Characteristic of the species is the black color present on the appendages of the oral veil whereas its margin is lined with yellow. The radula gof P. tilesii, as in P. maculatus, has more innermost teeth than typically found in other species of Plocamopherus. Plocamopherus maculapodium n. sp. (Figures II, 18, 19) Type and examined material: Holotype, CASIZ 099307, one specimen. Length (preserved): 25 mm. Depth: un- known. Ras Nungwi, Zanzibar, Tanzania. 6 November 1994. Collector: Don Pisor. Paratypes: CASIZ 070421, one specimen, dissected. Length (preserved): 13 mm. Depth: 3-6 m. Cement Mixer Reef, Madang, Papua New Guinea. 11 January 1988. Collector: T. M. Gosliner. CASIZ 073403, one specimen. Length (preserved): 11 mm. Depth: 33 m. Cement Mixer Reef, Madang, Papua New Guinea. 22 October 1986. Collector: T. M. Gosliner. Distribution: This new species has been collected from Papua New Guinea and Zanzibar, Tanzania. External Morphology: In life the body shape is elongate, limaciform and anteriorly rounded (Figure 11). The head bears a fringed oral veil without distinct appendages. The presence of flat oral tentacles was observed. Plocamo- pherus maculapodium has a red background color and the Page 196 Figure 17. Reproductive system of Plocamopherus tilesii (CASIZ 079251). Abbreviations: am, ampulla; bc, bursa copu- latrix; f, female gland mass; p, prostate; s, receptaculum seminis; u, uterine duct; v, vagina; vd, vas deferens. notum is slightly speckled with minute white dots. The white dots are usually situated on the notum along a line at the margin of both sides of the animal and joining be- hind the branchial leaves. They are also present along the margin of the fringed oral veil, on the tip of the posterior portion of the foot, tips of branchiae, clavus of the rhin- ophores, dorsal tubercles and oral veil appendages. There are a few black larger spots on the sides of the animal at the base of the foot. The rhinophores are elongate and at the peduncle have the same red background coloration. However, the clavus is red-brown with a white spot at the tip. The rhinophoral sheath is short. There are three pairs of short lateral appendages, with the last pair having a prominent rounded globular structure that is white in col- or. All lateral appendages are slightly ramified and whit- ish at the tip. At both sides of the body, four small tu- bercles are found in both sides situated at a regular dis- tance from each other. There are three tripinnate branchial leaves. The posterior portion of the foot is elongated and thin forming a keel that has a small crest tipped with white. The specimens examined were 11—25 mm long after being preserved. Radula and Jaws on Buccal Armature: The specimen dissected had a radular formula of 14 * (7.3.0.3.7). The radula of P. maculapodium is characterized by the pres- ence of two different types of teeth (Figure 18A—C). The inner lateral teeth, (Figure 18A, B) similar in shape, are broad and curve towards the rachis. There are three of them and the innermost tooth has a well-developed sec- ondary cusp. These hook-shaped teeth are blunt at the The Veliger, Vol. 48, No. 3 apex, not sharp. There are seven outer lateral teeth per side, (Figure 18A, C) which are flat and rectangular in shape. The size of these rectangular teeth diminishes from the innermost to the outermost. The papillated rachis has a series of transverse plates that decrease in size towards the base of the radula. The jaw rodlets are thin, elongate, flattened and densely packed (Figure 18D). Reproductive System: The reproductive system is triaul- ic (Figure 19). The vas deferens is differentiated in its distal portion into a well-developed and glandular-looking prostate that envelops the whole bursa copulatrix. The prostate is connected to the female gland by a very short and thin duct. The vagina is as wide as the vas deferens but half its length. From the oval bursa copulatrix merges a short duct that divides into two ducts, one that is con- nected to the vagina and the other to the receptaculum seminis. This second duct before entering into the recep- taculum seminis splits into the uterine duct that penetrates the female gland mass. The ampulla is very short and enters into the female gland mass in a position distal rel- ative to the penetration of both the vas deferens and the uterine ducts. Remarks: The most similar species in their external mor- phology to P maculapodium are P. lucayensis and P. pi- latecta. All three have red background coloration and are speckled with white small dots. However several char- acteristics distinguish them. First, in both P. maculapo- dium and P. pilatecta the white dots are situated at the margins of the body, along a line joining behind the bran- chial leaves whereas in P. lucayensis they are scarce and regularly distributed over the whole notum (Hamann & Farmer, 1988). These white dots are also present in P. maculapodium and P. pilatecta along the margin of the fringed oral veil on the margin of the keel, tips of bran- chiae, clavus of the rhinophores, dorsal tubercles and oral veil appendages as opposed to in P. lucayensis. Radular differences are found in the inner lateral teeth, where the curved portion of the teeth is much longer and wider in P. maculapodium than in P. pilatecta. Externally, P. ma- culapodium and P. pilatecta differ by the presence in the former of three lateral appendages whereas the latter has only one. The reproductive system of P. maculapodium differs from that of P. pilatecta by the presence of a con- voluted prostate that totally envelops the bursa copulatrix, as opposed to the naked bursa copulatrix in P. pilatecta and by the shape of the vagina which is flat in P. pila- tecta. Etymology: This species is named maculapodium refer- ring to the black spots present on the sides of the foot. Plocamopherus pecoso n. sp. (Figures 1J, 19, 20) Type and examined material: Holotype, CASIZ 110378, one specimen. Length (preserved): 20 mm. Y. Valles & T. M. Gosliner, 2005 100um Figure 18. Plocamopherus maculapodium (CASIZ 070421) SEM photographs of the radula and jaws. A. Whole radula. B. Inner lateral teeth. C. Outer lateral teeth. D. Jaws. Figure 19. Reproductive system of Plocamopherus maculapo- dium (CASIZ 070421). Abbreviations: am, ampulla; be, bursa copulatrix; f, female gland mass; p, prostate; s, receptaculum seminis; u, uterine duct; v, vagina; vd, vas deferens. Depth: unknown. Caban Island, Batangas, Luzon, Phil- ippines. 21 April 1997. Collector: T. M. Gosliner. Para- types: CASIZ 106548, one specimen, dissected. Length (preserved): 19 mm. Depth: O-10 m. Polonpoli, Marica- ban Island, Batangas, Luzon, Philippines. 19 April 1996. Collector: T. M. Gosliner. CASIZ 083738, one specimen. Length (preserved): 8 mm. Depth: unknown. Balayan Bay, Batangas, Luzon Island, Philippines. 19 February 1992. Collector: T. M. Gosliner. CASIZ 097401, one specimen. Length (preserved): 4 mm. Depth: 31 m. Ba- layan Bay, Batangas, Luzon Island, Philippines. 15 March 1994. Collector: T. M. Gosliner. CASIZ 088077, one specimen. Length (preserved): 11 mm. Depth: 10 m. Bus Stop Reef, Balayan Bay, Batangas, Luzon Island, Phil- ippines. 24 March 1993. Collector: T. M. Gosliner. Distribution: This species has been collected only in the Philippines. External Morphology: In life its body shape is elongate, limaciform and anteriorly rounded (Figure 1J). The head bears a fringed oral veil with eight short, small and ram- ified appendages. The oral tentacles are flat. Plocamo- Page 198 The Veliger, Vol. 48, No. 3 Figure 20. Plocamopherus pecoso (CASIZ 106548) SEM photographs of the radula and jaws. A. Whole radula. B. Inner lateral teeth. C. Outer lateral teeth. D. Jaws. pherus pecoso has a transparent white background color heavily speckled all over the body with small orange dots. These orange dots become much larger and fewer at the base of the foot. The presence of minute brown dots sur- rounding the lateral appendages base and along the keel was observed (these brown dots are present all over the body in other specimens of this species). There are some white dots that are usually situated on the notum along a non-continuous line at the margin of both sides of the animal and joining behind the branchial gills. They are also present along the margin of the fringed oral veil on the tip of the tail, tips of branchiae, clavus of the rhin- ophores, dorsal tubercles and oral veil appendages but they are slightly larger and scarce. The long rhinophores are translucent and speckled with brown at the peduncle and clavus. A white spot is present at the tip of the clavus. The rhinophoral sheath is long. There are three pairs of short lateral appendages; the two posterior pairs have a prominent, brown, rounded globular structure. Usually the posterior most pair has the larger globular structure, although exceptions have been observed. All lateral ap- pendages are slightly ramified and whitish at the tip. There are three principal tripinnate branchial leaves, which do not form a complete circle around the anus. The posterior portion of the foot forms a well-developed keel that has a small crest tipped with white. Radula and Jaws on Buccal Armature: The specimen dissected had a radular formula of 14 X (6.3.0.3.6). The radula of P. pecoso is characterized by the presence of two different types of teeth (Figure 20A—C). The inner lateral teeth, (Figure 20A, B) similar in shape, are thin, elongated and hook-shaped. There are three of them and the innermost tooth has a well-developed secondary cusp while in the other two it is prominent but somewhat less pronounced. These hook-shaped teeth are acute apically. The outer lateral teeth (Figure 20A, C), six per side, are flat and rectangular in shape. The size of these rectangular teeth decreases from the innermost to the outermost. The papillated rachis has a series of transverse plates that are regular in size along the whole length of the radula. The jaw rodlets are thin, elongate, flattened and densely packed (Figure 20D). Y. Vallés & T. M. Gosliner, 2005 Figure 21, Reproductive system of Plocamopherus pecoso (CASIZ 106548). Abbreviations: am, ampulla; bc, bursa copu- latrix; f, female gland mass; p, prostate; s, receptaculum seminis; u, uterine duct; v, vagina; vd, vas deferens. Reproductive System: The reproductive system is triaul- ic (Figure 21). The vas deferens is differentiated in its distal portion into a well-developed and glandular-looking prostate. The prostate is connected to the female gland by a short duct. The vagina, although almost as wide as the vas deferens in its proximal portion, becomes half as wide at its distal part and penetrates directly into the bur- sa copulatrix. The vagina is as long as the vas deferens. The bursa copulatrix is totally enveloped by the massive prostate. From the oval bursa copulatrix, emerges a some- what flattened duct that divides into two ducts, one which is connected to the receptaculum seminis and the other (the uterine duct) that penetrates into the female gland mass. Both ducts are of similar length and width. The ampulla is thin and elongated and enters into the female gland mass in a position distal relative to the entrance of both the vas deferens and the uterine ducts. Remarks: The most similar species to Plocamopherus pecoso are P. ceylonicus and P. maderae. Even though all of them are mottled with orange dots, Plocamopherus pecoso has a background color of translucent brown as opposed to the pale dull red of P. maderae. Plocamo- pherus pecoso does not have the brown pattern typical of P. ceylonicus nor the white pattern along margin edge of the body sides (Kelaart, 1858). The orange dots are slight- ly larger in Plocamopherus pecoso with smaller and more numerous ones in P. maderae and yellowish in P. cey- lonicus. No white dots are present on P. maderae. Plo- camopherus pecoso has two pairs of lateral appendages Page 199 with the globular structure whereas only one is found in P. maderae. Differences are found in the radula as well. In Plocamopherus pecoso the innermost teeth have a long and acutely pointed cusp, in P. madeare the innermost cusp is elongated wide and slightly blunt. Etymology: This species is named pecoso due to the presence of dots over its entire body appearing as freck- les. In Spanish the word pecoso means to have a lot of freckles. Plocamopherus lemur n. sp. (Figures 1K, 22, 23) Type and examined material: Holotype, CASIZ 146056, one specimen. Length (preserved): 9 mm. Depth: unknown. Les Cocotiers Hotel, [le Ste. Marie, Madagas- car. 17 April 1989. Collector: T. M. Gosliner. Paratype: CASIZ 074152, one specimen. Length (preserved): 12 mm. Depth: unknown. patch reef, Grand Passe, Aldabra Island, Seychelles. 21 March 1986. Collector: T. M. Gos- liner. CASIZ 146058, one specimen, dissected. Length (preserved): 9 mm. Depth: unknown. North side, Andi- lana Beach, Madagascar. 15 April 1989. Collector: T. M. Gosliner. CASIZ 146062, one specimen. Length (pre- served): 10 mm. Nosy Komba, Madagascar. 16 April 1989. Collector: T. M. Gosliner. Distribution: This species has been collected from both East and West coasts of Madagascar and the Seychelles. External Morphology: The body shape is elongate, li- maciform and anteriorly rounded (Figure 1K). The oral veil has 18 ramified appendages that differ in length. The oral tentacles are flat. Plocamopherus lemur has a brown- ish background color heavily speckled all over the body with minute brown dots, and minute orange dots that are clustered to form orange patches. White pigment is pre- sent in front of the branchial leaves, around the lateral appendages, at the base of the rhinophores and along the crest. This white coloration is situated on the notum along a non-continuous line at the margin of both sides of the animal and joining behind the branchial leaves. White pigment is also present along the margin of the sides of the body, the oral veil, tips of branchiae and clavus of the rhinophores. At the peduncle, the rhinophores are translucent and the clavus is translucent with a white apex. The rhinophoral sheath is short with clusters of or- ange dots. There are three pairs of lateral appendages, with only the last pair forming a rounded, brown, and prominent globular structure. All lateral appendages are conical having a short but highly ramified prolongation at the apex. The first two lateral appendages are very short and the last pair is very elongate. Six to eight small tubercles are present at each side of the animal. There are three principal tripinnate branchial leaves. The posterior Page 200 The Veliger, Vol. 48, No. 3 Figure 22. Plocamopherus lemur (CASIZ 146056) SEM photographs of the radula and jaws. A. Whole radula. B. Inner lateral teeth. C. Outer lateral teeth. D. Jaws. Figure 23. Reproductive system of Plocamopherus lemur (CASIZ 146056). Abbreviations: am, ampulla; be, bursa copu- latrix; f, female gland mass; p, prostate; s, receptaculum seminis; u, uterine duct; v, vagina; vd, vas deferens. portion of the foot forms a well-developed but short keel that is fringed on its upper margin. Radula and Jaws on Buccal Armature: The specimen dissected had a radular formula of 13 X (6—7.3.0.3.6—7). The radula of P. /emur is characterized by the presence of two different types of teeth (Figure 22A—C). The inner lateral teeth (Figure 22A, B), similar in shape, are thin, elongate and hook-shaped. There are three of them and only the innermost tooth has a well-developed secondary cusp. These hook-shaped teeth have an acute apex. The outer lateral teeth (Figure 22A, C), six to seven per side, are flat and rectangular in shape. The size of these rect- angular teeth decreases from the innermost to the outer- most. The papillated rachis has a series of wide transverse plates that are regular in size along the whole length of the radula. The jaws have rodlets that are thin, elongate, flattened and densely packed (Figure 22D). Reproductive System: The reproductive system 1s triaul- ic (Figure 23). The vas deferens is not differentiated into a prostate. The vagina is as wide as the vas deferens and penetrates directly into the bursa copulatrix. The vagina Y. Vallés & T. M. Gosliner, 2005 Page 201 Figure 24. Plocamopherus margaretae (CASIZ 162208) SEM photographs of radula and jaws. A. Whole radula. B. Inner lateral teeth. C. Outer lateral teeth. D. Jaws. is half as long as the vas deferens. From the oval bursa copulatrix, emerges a wide duct that splits into two ducts, one of which is connected to the small oval receptaculum seminis, and the other (the uterine duct) that penetrates into the female gland mass. The ampulla, is externally connected to .the vas deferens entering into the female gland mass in a position proximal relative to the penetra- tion of the uterine duct. Remarks: Plocamopherus lemur resembles P. ceylonicus and P. pecoso. All three species have a brownish trans- lucent background color with orange and brown dots on the notum. However, in P. /emur these dots are not dis- tributed regularly, but in patches. No branched papillae or yellow coloration are present on the notum of P. /emur, which are characteristic of P. ceylonicus. Only the last pair of lateral appendages, which is extremely long in P. lemur, has the globular structures while both P. pecoso - and P. ceylonicus have two pairs of globular structures. In both Plocamopherus pecoso and P. ceylonicus the prostate envelops the bursa copulatrix, while the prostate is not a discrete structure leaving the bursa copulatrix exposed in P. lemur. Etymology: This species is named lemur owing to sim- ilarity of coloration with the cryptic primates that are characteristic of its type locality, Madagascar. Plocamopherus margaretae Nn. sp. (Figures 1L, 24, 25) Type and examined material: Holotype, CASIZ 162208, one specimen. Length (preserved): 35 mm. Depth: unknown. Myanmar. Collector: Mark Strickland. Paratype: CASIZ 162207, one specimen, dissected. Length (preserved): 40 mm. Depth: unknown. Dubai, United Arab, Emirates. Collector: C. Harris. Distribution: This new species has been collected in Du- bai and Myanmar. External Morphology: In life, its body shape is elongate and anteriorly rounded (Figure 1L). The oral veil is wide, Figure 25. Reproductive system of Plocamopherus margaretae (CASIZ 162208). Abbreviations: am, ampulla; bc, bursa copu- latrix; f, female gland mass; p, prostate; s, receptaculum seminis; u, uterine duct; v, vagina; vd, vas deferens: flattened and fringed. The oral tentacles are flat. Ploca- mopherus margaretae has a pink or red background color heavily speckled all over the body with large white spots and black, almost oval spots. The white spots are of var- ious sizes and are regularly scattered on the notum. The black spots appear always surrounded by white, and are much fewer than the white spots. A yellow-orange line runs along the margin of the foot, the margin of the oral veil and the margin of the keel. The rhinophores are yel- low-pink for their whole length. At the tip of the clavus there is a short white line. The rhinophoral sheath is the same background color as the body and is short with its margin lined by white and yellow pigment. There are three pairs of lateral appendages, which are rounded and form prominent globular structures. These globular struc- tures are much larger in the posterior most pair of ap- pendages. The anterior pairs are very short and rounded. All are pink-yellowish in color. There are three tripinnate branchial leaves that each insert separately into the dor- sum. The posterior portion of the foot forms a well-de- veloped elongated keel that is slightly fringed in its upper margin. Radula and Jaws on Buccal Armature: The specimen dissected had a radula formula of 16 X (7.7.0.7.7). The radula of P. margaretae is characterized by the presence of two different types of teeth (Figure 24A—C). The inner lateral teeth, (Figure 24A, B) similar in shape, are wide and hook-shaped with a blunt apex. There are seven of them and the three innermost teeth have a well-developed secondary blunt cusp while in the other four is less pro- nounced. The outer lateral teeth (Figure 24A, C), seven per side, are flat and rectangular in shape. The size of these rectangular teeth diminishes from the innermost to The Veliger, Vol. 48, No. 3 the outermost. The papillated rachis has a series of trans- verse plates that are regular in size along the whole length of the radula. The jaws have rodlets that are thin, elon- gate, flattened and densely packed (Figure 24D). Reproductive System: The reproductive system is triaul- ic (Figure 25). The vas deferens is not differentiated into a prostate. The vagina, although almost as wide as the vas deferens, is a third of its length. The bursa copulatrix is large and oval, from which, emerges a duct that divides immediately into two ducts, one of which connects to the vagina and the other connects to the receptaculum sem- inis. From the latter also emerges the uterine duct that penetrates the female gland mass. The ampulla is thin convoluted and elongated entering into the female gland mass in a position proximal relative to the penetration of both the vas deferens and the uterine ducts. Remarks: The most similar species to P. margaretae is P. tilesii. However the differences between both are ob- vious. While P. margaretae has pink-red background col- oration, P. tilesii is whitish or yellowish. Both have black dots all over the notum, however they are more regularly distributed and scarcer in P. margaretae than in P. tilesii. Another external characteristic that differentiates both species is the number of lateral appendages and globular structures that are present, with three pairs in P. margar- etae and two in P. tilesii. Even though the number of teeth may be somewhat variable, in P. tilesii, the number of innermost teeth exceeds the number of outer lateral teeth, whereas in P. margaretae the outermost are more numerous than the inner lateral. In Plocamopherus mar- garetae the prostate is not differentiated and therefore the bursa copulatrix is exposed whereas in P. tilesii the pros- tate is massive and totally enveloping the bursa copula- trix. Etymology: This species is named for Carol Harris’ mother. Carol collected the first specimens of this species. CONCLUSION The ability of living organisms to produce light has mul- tiple functions depending on the organism. Fishes, like the Malacosteidae, produce red light to see their prey without alerting either prey or other possible predators (O’Day & Fernandez, 1974). Several beetles use light to communicate with their potential mates (Day et al., 2004), etc. In mollusks, light production has already been studied within cephalopods (Young, 1977; Johnsen et al., 1999). In dorids, bioluminescence has only been reported from the two genera Kaloplocamus and Plocamopherus. Both genera are phanerobranchs (having non-retractable branchial leaves) and have taxonomically been included in the same subfamily Triophinae. Both genera are almost exclusively Atlantic and Indo-Pacific and share several morphological and anatomical features: six to eight lateral processes, presence of a large rachis in the radula, similar Y. Vallés & T. M. Gosliner, 2005 outer and inner lateral teeth and the ability (of several species) to emit light among others. We have revised the taxonomy of both genera. As a result, we considered that there are six valid species for the genus Kaloplocamus (K. pacificus, K. ramosus, K. acutus, K. maru, K. peludo and K. dokte). However we believe that a comprehensive study of all K. ramosus specimens, of their external and internal morphology is necessary as well as a molecular phylogenetic analysis, allowing us to understand the distribution patterns shown by this spectacular species. Within the genus Plocamo- Dherus we suggest that the valid species are the follow- ing: P. ocellatus, P. maderae, P. maculatus, P. ceyloni- cus, P. imperialis, P. tilesii, P. lucayensis, P. pilatecta, P. pecoso, P. maculapodium, P. lemur and P. margare- tae. Interestingly not all species of both genera have been reported to be bioluminescent. Whether they are biolu- minescent or not remains to be observed both in the wild and by TEM studies of the body wall and the globular structure tissues of both genera. It is important to note that it may not have been observed in the wild because although a stimulation provokes the animal into swim- ming and emitting light, when the animal is stressed or receives a continuous mechanical stimulation it stops emitting light, suggesting that there is some kind of prod- uct that it stores, that is used for the light emission and therefore its availability is limited (Wilbur & Yonge, 1966). It is possible that since these animals emit light only when disturbed and for a short span of time, the role could be to temporally distract the attention of potential predators (Wilbur & Yonge, 1966). In order to better understand the evolution of biolu- minescence within these dorids a phylogenetic analysis based on morphological and molecular data comprising all genera pertaining to the subfamily Triophinae must be done. If Kaloplocamus and Plocamopherus were sister taxa, a plausible explanation could be that biolumensc- ence evolved just once. However, the information avail- able regarding the light production in such organisms is very limited. Even though Wilbur and Yonge (1966) es- tablished the different type of light emitting systems, there was no description of the cellular tissue of the man- tle (also capable of emitting light) of Plocamopherus spe- cies. Wilbur and Yonge stated that the light from the glob- ular processes is much more intense then that of the rest of the body wall, which raises the question whether it is, or not, the same type of cells that produce the light in the body wall and the globular structure. Acknowledgments. We would like to thank all the members of Nudibranch Central for their support. Special thanks to Benoit Dayrat for his valuable comments and encouragement, and to David Behrens, Carol Harris, Bob Bolland and Mark Strickland for making specimens and photos available. 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Western Australian Museum: Perth. 184 pp. WHITELEGGE, T. 1889. List of the marine and freshwater invertebrate fauna of Port Jackson and neighbourhood. Journal and Pro- ceedings of the Royal Society of New South Wales 23:769. WILBUR, K. M. & C. M. YONGE. 1966. Physiology of Mollusca, Vol. 2. Academic Press: New York. YounG, R. E. 1977. Ventral bioluminescent countershading in midwater cephalopods. Symposium of the Zoological Soci- ety of London 38:161—190. The Veliger 48(3):206—214 (November 2, 2006) THE VELIGER © CMS, Inc., 2006 A New Species of Lepidochitona (Mollusca: Polyplacophora) from El Salvador CEDAR I. GARCIA-RIOS Department of Biology, University of Puerto Rico at Humacao, Humacao, Puerto Rico 00792 (e-mail: cgarcia@www.uprh.edu) Abstract. Lepidochitona salvadorensis sp. nov. from El Salvador is the third species of the genus described for the Panamic Province. The species previously known were Lepidochitona beanii Carpenter, 1857 and L. corteziana Clark 2000. L. salvadorensis differs from L. beanii in having longer hyaline spicules, a smooth tegmentum, rounder lateral margins of the intermediate valves and smaller marginal spicules. L. corteziana differs from both in the lack of large hyaline spicules on the girdle. INTRODUCTION In July 2002 a group of zoologists from El Salvador and Puerto Rico studied the polyplacophoran fauna of the rocky coasts of El Salvador (Garcia-Rios et al., 2003). Eleven species of Polyplacophora from shallow water were collected, of which seven species were new reports, including Lepidochitona beanii Carpenter, 1857 and two specimens of an undescribed species of Lepidochitona. In December 2003 a new visit to the previous stations al- lowed the collection of additional material, enabling the description of the new species. The genus Lepidochitona Gray, 1821 is represented in the Panamic Province by two species, L. beanii and L. corteziana. This description adds a new species of this genus to the fauna of El Salvador. MATERIALS AND METHODS Thirty-three specimens of L. salvadorensis and four spec- imens of L. beanii were collected in the sublittoral, at depths of 0.5—2 m, at low tide. All animals were found under small rocks partially buried in the sand. They were preserved flat in 70% ethanol. Measurements of whole animals (including girdle) were made using a caliper under the dissection micro- scope. Four specimens selected for detailed studies of the valves, radula, and girdle were prepared according to the methods described by Bullock (1985). Soft parts of the chitons were dissolved in heated 2N KOH solution. The valves and radula were cleaned further using ultrasound for one or two minutes. After cleaning, these structures were rinsed with distilled water and then stored in 70% ethanol. Cleaned hard parts selected for study with scanning electron microscopy (SEM), were air dried, mounted on aluminum specimen stubs using carbon tape and coated with gold using a sputtering machine. SYSTEMATICS Higher classification follows Kaas & Van Belle (1985). Types are placed in the Zoological State Collection Mu- nich (ZSM), Munich, Germany; Museo de Historia Nat- ural de El Salvador (MUHNES), Central America; Acad- emy of Natural Sciences, Philadelphia (ANSP); United States National Museum of Natural History, Smithsonian Institution, (USNM). Class POLYPLACOPHORA Gray, 1821 Order NEOLORICATA Bergenhayn, 1955 Suborder ISCHNOCHITONINA Bergenhayn, 1930 Family ISCHNOCHITONIDAE Dall, 1889 Genus Lepidochitona Gray, 1821 Subgenus Lepidochitona s.s. Lepidochitona (L.) salvadorensis Garcia-Rios, sp. nov. (Figs. 1-10) Holotype: ZSM Moll 20040210: one complete specimen (Figure 1), preserved flat in alcohol, 4.6 mm long, 2.7 mm wide, Los Cébanos (13°31'40"N, 89°48’44”W), Son- sonate, El Salvador, sublittoral, 1 m depth at low tide, under stone partially buried in sand, 23 December 2003. Other material examined: Paratype MUHNES 90-416, 4.4 mm long, 2.5 mm wide, with the same locality and collection data of holotype (Figure 2); paratype MU- HNES 90-417, 6.1 mm long, 3.5 mm wide, Playa Maculis (13°09'30"N, 89°55'24"W), La Union, El Salvador, (Fig- ure 3); paratype ANSP A20089, 6.0 mm long, 3.8 mm wide, Playa Maculis, (same previous coordinates) (Figure 4); paratype USNM 1023580, 5.4 mm long, 3.0 mm wide, C. I. Garcia-Rios, 2005 Page Figure 1. Lepidochitona salvadorensis Garcia-Rios, sp. nov. Holotype, ZSM Moll 20040210, scale bar = 1 mm. Figure 2. Lepidochitona salvadorensis Garcia-Rios, sp. nov., Paratype, MUHNES 90-416, scale bar = | mm. Page 208 the Veliger, Voly43 Nowe (ott RT Figure 3. Lepidochitona salvadorensis Garcia-Rios, sp. nov., Paratype, MUHNES 90-417, scale bar = 1 mm. the previous locality (Figure 5); and others, not designat- ed as paratypes, in the author’s private collection. All the types and other material examined for this description were found under stones, 0.5—2 m depth, collected in De- cember 2003 and preserved flat in alcohol. All the ma- terial came from the two locations mentioned. Etymology: This species is named for the country, El Salvador, in which it was first discovered. Diagnosis: Recognized by its small size, oval shape, and reddish color. Head valve semicircular, posterior margin V-shaped. Intermediate valves with convex anterior mar- gin, rounded side margins, concave posterior margin at both sides of the protruding apex, lateral areas poorly defined. Tail valve is semicircular, smaller than the others, mucro at central position, concave postmucronal slope. The tegmentum is smooth. The girdle is wide, dorsally covered with minute bluntly pointed spicules and 20-21 tufts of long, curved hyaline needles. DESCRIPTION The species is small, up to 9.8 mm long, 6.5 mm broad, elongated oval, moderately elevated (dorsal elevation 0.37), carinated, girdle relatively broad. When alive, the color of the tegmentum and girdle is dark red, some with mottlings of white and brown marks. In alcohol the red color of the tegmentum changes from red to orange. The head valve always is reddish, sometimes with small white marks only in the posterior margin. The tail valve always has some white or brown marks in the postmucronal area (Figures 1-5). The head valve is semicircular, its posterior margin is V-shaped (Figure 6A), some notched in the middle. In- termediate valves have convex anterior margin, side mar- gins are rounded, their posterior margins are concave at both sides of the protruding apex, lateral areas are poorly defined or not defined at all (Figure 6B). Tail valve is semicircular, anterior margin almost straight in the center, with a 45° angle of both borders (Figure 6C), mucro at a central position, prominent, postmucronal slope is con- cave, stepped near the mucro (Figure 7). The tegmentum width size ratio of valve I:IV:VIII is 1:1.40:0.84 (N = 33). Frequently the tegmentum of the valve V is a little broader than on valve IV (mean of the differences is 0.04 mm). Tegmentum smooth, without any granules or sculpture under magnification. The megalopores are uniformly dis- tributed throughout the tegmentum, with a density of 1 per 930 ppm?. There are 9 to 10 micropores for each me- galopore. Articulamentum white, translucent, apophyses broad, subtriangular with rounded apex, sinus length is about 25% of the intermediate valves’ width and 15% of the tail valve tegmentum width. The insertion plates slit for- mula is: 8—9/1/7—11, slit rays present on all valves, teeth well defined, striated, eaves porous and narrow. Girdle wide (average is 32% of the total width), red- dish brown, dorsally covered with very small, juxtaposed, globulose, bluntly pointed spicules, up to 20-25 X 5-7 wm (Figure 8), and long, smooth and curved hyaline nee- dles mostly measuring 0.4—0.8 mm X 25-30 wm, (largest frequently broken). Different sizes of hyaline needles are C. I. Garcia-Rios, 2005 Page 209 Figure 4. Lepidochitona salvadorensis Garcia-Rios, sp. interspersed and bunched in tufts of 2—6, almost 20-21 tufts: 4 near the head, 2-3 post tail plate and 14 aligned with the sutures (Figure 1). Some smaller spicules (40— 250 X 15-20 wm) are dispersed in the dorsal surface of nov., Paratype, ANSP A20089, scale bar = 1 mm. the girdle (Figure 8). Marginal spicules are elongate, straight, amber color, dorsal surface vaulted and grooved in two directions on the distal half (Figures 8 and 9A), ventral surface flat, without any sculpture; up to 55—65 Figure 5. Lepidochitona salvadorensis Garcia-Rios, sp. nov., Paratype, USNM 1023580, scale bar = 1 mm. Page 210 The Veliger, Vol. 48, No. 3 Figure 6. Scanning electron micrograph of Lepidochitona salvadorensis Garcia-Rios, sp. nov. (whole specimen 8.1 mm long, 4.7 mm wide, including girdle, #CIGR 2049): dorsal view of A, head valve; B, valve IV; C, tail valve. x 10 wm. Undersurface paved with translucent, imbri- cating, pointed scales, 40-45 x 10-15 pm (Figure 9C); and a row of submarginal scales (between the marginal spicules and the ventral scales), less pointed, translucent, up to 80-100 pm xX 16um (Figure 9B). Central tooth of the radula is twice as long as wide, with a well defined blade; major lateral tooth with a tri- dentate cusp, central one a little wider and longer than the others (Figure 10). The arrangement of the gills is holobranchial, with 14 pairs of gills (in 7 and 9 mm long specimens). The gills are abanal, with the size of the gills increasing posteri- orly, with a free space between the two rows of gills and the anus, with a well defined pallial lappet. C. I. Garcia-Rios, 2005 Page 211 Figure 7. Scanning electron micrograph of Lepidochitona salvadorensis Garcia-Rios, sp. nov. (#CIGR 2049): lateral view of tail valve. lt” ae Paes dae >. Sas oat BA ong <5 a a. Figure 8. Scanning electron micrograph of Lepidochitona salvadorensis Garcia-Rios, sp. nov. (whole specimen 9.4 mm long, 5.7 mm wide, including girdle, #CIGR 2055): dorsal view of girdle. The Veliger, Vol. 48, No. 3 Figure 9. Scanning electron micrograph of Lepidochitona salvadorensis Garcia-Rios, sp. nov. (#CIGR 2055): ventral view of the girdle; A, marginal spicules; B, submarginal spicules; C, ventral spicules. DISTRIBUTION L. salvadorensis has been collected in two localities in El Salvador: Los Céobanos, Sonsonate (13°31'40"N, 89°48'44"W) and Playa Maculis, La Union (13°09'30'N, 89°55'24"W). REMARKS The new species always appear associated with crustose coralline algae (Corallinaceae: Neogoniolithon spp). The live chiton closely matches the color of the associated crustose alga, as illustrated in Littler & Littler (2000:71). The hyaline needles are frequently broken. The holotype has a fine calcareous overgrowth in the II, III] and VII plate. DISCUSSION Eernisse (2004) proposed three clades based on compar- ative biochemistry of the family Lepidochitonidae: one clade groups the Lepidochitona from northern Europe, which remain under Lepidochitona, another clade groups the species from the northern Pacific under the genus Cy- anoplax, and a third clade containing the Caribbean, Pan- amic and southern African species together with the ge- nus Nuttallina (Eernisse does not suggest a generic clas- sification for this clade). The genus designation for L. salvadorensis is based strictly on morphological charac- ters, following the criteria of Kaas and Van Velle (1998). It was included on this genus due to the presence of hy- aline spicules on the girdle. The genus Mopalia was not considered because the specimens have more than 8 slit in the head valve and the girdle is not hairy; neither does it belong to Dendrochiton because the girdle lacks cor- neous processes. L. salvadorensis differs consistently from L. beanii Carpenter, 1857, the only previously known Lepidochi- tona species from El Salvador, by its possession of con- siderably longer hyaline spicules, a smooth tegmentum, C. I. Garcia-Rios, 2005 Page 213 Figure 10. Scanning electron micrograph of Lepidochitona salvadorensis Garcia-Rios, sp. nov. (#CIGR 2049): radula. rounder lateral margins of the intermediate valves and smaller marginal spicules. The microgranular surface of the tegmentum in L. beanii (Figure 11) stands out both in the original brief description (Carpenter, 1857, illus- trated in Brann, 1966), as well as in the detailed and well illustrated description of Kaas & Van Belle (1985). Other differences between both species are the wider girdle and the amber color of the marginal spicules in L. salvadorensis. The examination of the radulae of three specimens of L. salvadorensis did not show the much longer central cusp in the major lateral tooth as illustrated for L. beanii in Kaas & Van Belle (1985). L. salvadorensis is similar in color and shape to L. corteziana, from the Gulf of California. The smaller size (to 3.5 mm) and lack of large spicules on the girdle (Clark, 2000) separate L. corteziana from the other two species. L. corteziana have shorter dorsal and ventral gir- dle elements, different slit formula (10-1-12) and differ- ent megapores to micropores ratio (1:2) in the tegmentum than L. salvadorensis. L. salvadorensis tegmentum surface and the ratio of 10 micropores for each megapore is similar to the Caribbean Lepidochitona liozonis (Dall & Simpson, 1901) (but L. liozonis has mega and micropores density three times greater). Both species have similar color patterns and are generally associated with crustose red algae (Garcia-Rios, 2003). Clark (2000) mentioned a similar resemblance of his L. corteziana with the South African L. turtoni (Ash- by, 1928) and the North Pacific L. (Spongioradsia) aleu- tica (Dall, 1878), all with coralline algal habitat. L. salvadorensis is the same species that appears illus- trated and described as Lepidochitona sp. in the guide to identification of the species of El Salvador (Garcia-Rios et al., 2003). Acknowledgments. For their help and assistance in the field work, appreciation is here expressed to Migdalia Alvarez-Ruiz (University of Puerto Rico at Ponce), Carolina and Carlos Rob- erto Hasbun (Fundacién Zoologica de El Salvador), Laura Mar- tinez, Ana M. Rivera (Museo de Historia Natural, San Salvador, El Salvador), José E. Barraza (Ministerio de Medio Ambiente y Recursos Naturales, El Salvador). I thank Enrico Schwabe, Bruno Anseeuw, Yves Terryn and Edgardo Ortiz for their valuable comments and for allowing me to use their collection of literature and specimens of L. beanii for comparison. The scanning electron microscope facilities were provided by the Department of Physics, UPR-Humacao and the Partnership to Integrate Research and Education in Materials (NSF-DMR-0353730). Facilities for obtaining microphotographs of the types were provided by the Advance Program (NSF SBE- 0123654). The review of Dr. Douglas J. Eernisse provided many useful ideas and suggestions. This research was partly supported by the Research Fund of the University of Puerto Rico at Humacao and by a contribution The Veliger, Vol. 48, No. 3 ~ AB brn Figure 11. Scanning electron micrograph of Lepidochitona (L.) beanii Carpenter, 1857 (whole specimen 3.6 mm long, 2.1 mm wide, including girdle, #CIGR 2076): valve IV. of the Puerto Rico Alliance for Minority Participation (PR- AMP). LITERATURE CITED BERGENHAYN, J. R. M. 1930. Kurze Bemerkungen zur Kenntnis der Schalenstruktur und Systematik der Loricaten Kungi. Svenska Vetensk. Hand] (3)9(3):3—54. BERGENHAYN, J. R. M. 1955. Die fossilen Schwedischen Lori- caten nebst einer vorlaufigen Revision der Systems der gan- zen Klasse Loricata. K. Fysiogr. Sallsk. Handl., Lund N.F 66(8):1—44. BRANN, D. C. 1966. Illustrations to “Catalogue of the Collection of Mazatlan Shells” by Philip P. Carpenter. Paleontological Research Institution: Ithaca, USA. 111 pp. BuLLock, R. C. 1985. The Stenoplax limaciformis (Sowerby, 1832) species complex in the New World (Mollusca: Poly- placophora: Ischnochitonidae). Veliger 27:291—307. CARPENTER, P. P. 1857. Pp. i-iv, ix—xvi, 1-552 in Catalogue of the Collection of Mazatlan Shells in the British Museum: Collected by Frederick Reigen. British Museum: London. CLARK, R. N. 2000. The chiton fauna of the Gulf of California rhodolith beds (with the description of four new species). Nemouria 43:1—18. DaLL, W. H. 1889. Preliminary catalogue of the shell-bearing marine mollusks and brachiopods of the southern coast of the United States, with illustration of many species. Bull. U.S. Natl. Mus. 37:3—271. EERNISSE, D. J. 2004. Revival of the genus Cyanoplax Pilsbry, 1892 for a clade of West Coast chitons [abstract]. Pp. 33- 35 in J. C. Martinez & R. V. Yeomans, (eds.), Program and Abstracts of the 37th Annual Meeting of the Western Society of Malacologists, Ensenada, Baja California, México. GarciA-Rios, C. I. 2003. Los Quitones de Puerto Rico. Editorial Isla Negra: San Juan/Santo Domingo. 208 pp. Garcia-Rios, C. L, M. ALvarez-Ruiz, J. E. BARRAZA, A. M. RIVERA & C. R. HAsBUN. 2003. Los Quitones (Mollusca: Polyplacophora) de El Salvador: Una Guia para la Identifi- cacion de las Especies. UPRSG-H-85. Impresos Sea Grant: Puerto Rico. 32 pp. Gray, J. E. 1821. A natural arrangement of Mollusca, according to their internal structure. London Med. Repos. 15:221—239. Kaas, P. & R. A. VAN BELLE. 1985. Monograph of Living Chi- tons (Mollusca: Polyplacophora). Vol. 2, Suborden Ischno- chitonina: Ischnochitonidae: Schizoplacinae, Callochitoni- nae and Lepidochitoninae. E. J. Brill Publ.: Leiden, Neth- erlands. 198 pp. Litter, D. S. & M. M. Litter. 2000. Caribbean Reef Plants: An Identification Guide to the Reef Plants of the Caribbean, Bahamas, Florida and Gulf of Mexico. Offshore Graphic, Inc.P Washington, U.S.A. 542 pp. The Veliger 48(3):215—219 (November 2, 2006) THE VELIGER © CMS, Inc., 2006 On the Occurrence of Rhomboidella prideaux (Leach, 1815) (Mollusca: Bivalvia: Mytilidae) in the Eastern Mediterranean BILAL OZTURK,* JEAN-MAURICE POUTIERS,** MESUT ONEN,* aANp ALPER DOGAN* *Ege University, Faculty of Fisheries, Department of Hydrobiology, 35100 Bornova, Izmir, Turkey (e-mail: bilal.ozturk @ege.edu.tr, mesut.onen @ege.edu.tr, alper.dogan @ege.edu.tr) **Muséum National d’ Histoire Naturelle, Département Systématique et Evolution, Unité Taxonomie et Collections (Mollusques), Case Postale N°51, 55 rue de Buffon, F-75231 Paris cedex 05, France (e-mail: malaco@mnhn.fr, jampoutiers @ free.fr) Abstract. na study performed in 2000 on the benthic fauna inhabiting the Turkish coasts of the Aegean Sea, a small less known Mytilidae species, Rhomboidella prideaux (Leach, 1815), which has not been reported from the eastern Mediterranean so far, was found. In the present study, the taxonomic, ecological and distributional aspects of this species, based on the material collected from the Aegean Sea and the eastern coast of Sicily, are provided. INTRODUCTION Rhomboidella Monterosato (1884:13) is one of seventeen genera of Mytilidae having representatives in the Medi- terranean Sea (Sabelli et al., 1990). The genus includes only one Mediterranean species, which is an epibenthic form, living attached to the substrate with its byssal threads. Although some Mediterranean species of Mytil- idae (for example some species of Mytilaster) are distrib- uted in environments showing greatly reduced levels of salinity (Oztiirk et al., 2002) Rhomboidella prideaux (Leach, 1815) appears to be a strictly marine species. This tiny species is found mainly along the European and Af- rican coasts of the Atlantic Ocean (Berkeley, 1827; Forbes & Hanley, 1849; Tebble, 1966; Segers, 2002) and in the western Mediterranean Sea (Van Aartsen et al., 1984; Salas, 1996; Giannuzzi-Savelli et al., 2001). Here, we report the species from the eastern Mediterranean ba- sin for the first time. The present study provides new locality records for R. prideaux and gives additional information about its ecol- ogy and general distribution. MATERIALS AnD METHODS In the framework of a research realized in 2000 to deter- mine the benthic fauna distributed along the Aegean coasts of Turkey, samplings were carried out at 314 sta- tions in depths ranging from 5 to 200 m. Samples were collected by the R/V Hippocampus, using a dredge cap- turing a volume of almost 30 dm?. Material was sieved ~ with 0.5 mm mesh and fixed in 4% formaldehyde. In the laboratory, samples were sorted under a stereomicroscope and preserved in 70% ethanol. During the research, spe- ciemens of R. prideaux were found at only one station located in the southern part of the Aegean Sea (Figure 1). Dimensions were measured using an ocular microme- ter. Species description and morphological terminology follow Cox (1969), amended by Ockelmann (1962, 1983) for early stages of development. As usual for Mytilidae, dimensions were defined as follows: when ventral margin is placed horizonally, length is the maximum size of valves measured according to that direction, and height along a perpendicular direction. The specimens (one specimen and a valve) are depos- ited in the Department of Hydrobiology (Ege University, Turkey). Additionally, series housed in the Muséum Na- tional d’Histoire Naturelle (MNHN) Paris, were studied by one of us (J.M.P.) for comparision. This additional material was recently collected by diving and dredging during a sampling campaign conducted by MNHN in the Gulf of Catania, eastern Sicily. RESULTS anp DISCUSSION Rhomboidella prideaux (Leach, 1815) Modiola prideaux Leach, 1815:35. Synonyms: Modiola rhombea Berkeley, 1827:229, pl. 18, fig. 1. Modiola asperula Wood, S., 1840:252 (nomen nul- lum); 1851:64, pl. 8, fig. 8. Other references: Modiola prideauxiana, Brown, 1827: 3, pl. 39, fig. 17; Crenella rhombea, Forbes & Hanley, 1849:208-209, pl. 45, fig. 3; Rhomboidella rhombea, Monterosato, 1884:13; Modiolaria (Rhomboidella) rhom- bea, Dall, 1898:805; Winchworth, 1932:240; Crenella (Rhomboidella) prideaux, Bowden & Heppell, 1966:103, 115; Crenella prideauxi, Tebble, 1966:48—49, fig. 19b & 23; Solamen (Rhomboidella) pridaux(i), Soot-Ryen, 1969:276; Solamen (Rhomboidella) prideauxi, Bruschi et Page 216 Cc ~ 40° ae A E G E 39° A N 38° 37° 26° 2m 28° Figure 1. Map of the study area with location of the sampling site where Rhomboidella prideaux was found. al., 1985:48; Rhomboidella prideaux, Sabelli et al., 1990: 283. Material examined: West of Bodrum Peninsula, Turkey (see Figure 1), 36°56'45”N—27°16'32"E, 31 m, 20th Sep- tember 2000, temperature 22.0°C, salinity 39.1 ppt, dis- solved oxygen concentration 4.97 mg/L, sandy substrata with algae fragments, one specimen (1.4 mm long, 1.2 mm high), and a left valve (3.0 mm long, 1.9 mm high) (Figures 2, 3). Gulf of Catania, eastern Sicily (M.N.H.N., 1990), from North to South: Aci Trezza, Capo Mulini, 37°34.5'N— 15°11.9'E, 60-90 m, 5 left valves, 4 right valves; same region, 37°34.5’N—15°20.0’E, 90-120 m, 2 right valves. Brucoli, Capo Campolato, 37°17.8'N-15°12.8'’E, 40— 60 m, | right valve; same region, Cozzo dei Turchi, 37°17.7'N-15°09.8'E, 15-20 m, 2 left valves; same re- gion, Brucoli Bay, 37°17.3'N—15°11.7'E, 3-9 m, sand bottom with Posidonia, | left valve, | right valve; same region, off Monte Amara, 37°16.5'N—15°13.5'E, 22-25 m, 2 left valves, 2 right valves. Description: Shell very small (up to about 3.5 mm long), moderately thin for its size, equivalve, well inflated and strongly inequilateral; rounded triangular to obliquely The Veliger, Vol. 48, No. 3 Figure 2. Rhomboidella prideaux (Leach, 1815), left valve: A) from the outside, B) from the inside. rhomboidal in outline. Shape very convex, becoming more or less laterally compressed posterodorsally. Byssal gape poorly developed, a narrow to obsolete slit just an- terior to mid-ventral margin of the valves. Umbones pro- sogyrate, prominent and nearly terminal. Anterior margin short, strongly convex, not extending beyond the umbo- nes. Ventral margin long and straight or nearly so (weakly convex in juvenile stages, tending to be slightly depressed in the middle in mature specimens). Posterior margin large and widely rounded, becoming less convex towards the dorsum. Posterodorsal margin long and_ slightly arched, oblique in relation to ventral margin, forming a rounded obtuse angle with posterior margin. Outer surface of valves covered with numerous, dense- ly set radial riblets, slightly diverging on posterodorsal and posteroventral areas, and crossed over by concentric ridges. Radial riblets rounded, about as wide as their in- tervals, increasing in number with growth by branching or intercalation. Concentric ridges more widely spaced Figure 3. Rhomboidella prideaux, living specimen. B. Oztiirk et al., 2005 than radial riblets, raised and sharp, most prominent in later growth stages, giving the shell periphery a decussate aspect. Umbonal area with distinct nepioconch and prod- issoconch. Nepioconch large (about 680 to 700 wm long), rounded-oblique in outline, sculptured with numerous, conspicuous, incised concentric lines. Prodissoconch large-sized (about 240 ym long), rather convex, D-shaped in outline, slightly more rounded anteriorly than posteri- orly, with a narrow and somewhat flattened marginal area (corresponding to prodissoconch II); surface of prodis- soconch dull white, roughened by minute dense granu- lations. Periostracum very thin, adherant, smooth and transparent, resulting in a somewhat glossy and iridescent aspect to the outer shell surface. Shell translucent greyish white in colour, under the pale straw-coloured periostra- cum. Ligament inframarginal, not strong, brownish yellow in colour, extending along posterodorsal margin to about two-thirds the distance from umbo to posterodorsal angle; resilial ridge deeply inset, very thin, not pitted. Hinge with a thickened process forming two poorly separated denticles under the beak with tiny, irregular transverse ridges. Shell margin smooth for a short distance just an- terior to the beak. Posterior part of hinge line bearing a series of small granules along and behind the otherwise smooth ligamental margin; size of these granulations in- creasing progressively from umbonal area to posterodor- sal angle, where normal crenulations of the valve margins begin. Interior of shell somewhat glossy and subnacreous, nearly smooth, with the outer sculpture showing through. Muscle scars and pallial line indistinct. Internal margins finely crenulate in accordance with the outer sculpture. Taxonomic status: Romboidella prideaux was first re- ported from southern England by Leach (1815:35), who briefly described it as Modiola prideaux but failed to il- lustrate it. A figure accompanying this short description was provided much later (Leach & Gray, 1852:333, pl. 12, fig. 9). In the intervening years, the species in ques- tion, still included in the genus Modiola, has been rede- scribed and figured by Brown (1827:3, pl. 39, fig. 17) under the emended name of M. prideauxiana, or as a different species as M. rhombea by Berkeley (1827:229, pl. 18, fig. 1). Wood (1851:64, pl. 8, fig. 8) created M. asperula for a Pliocene species from the Crag Formation of England, and this was generally considered as a syn- onym of R. prideaux later on (Glibert & Van de Poel, 1965:79). The species has been placed in other genera, such as Crenella (Forbes & Hanley, 1849:208—209; Winckworth, 1932:240; Bowden & Heppell, 1966:103, 115; Tebble, 1966:48—49; Seaward, 1990:69), Modiolaria {a junior synonym of Musculus (Dall, 1898:805; Lamy, 1937:7)], or Solamen (Soot-Ryen, 1969:276). Though Monterosato (1884:13) pointed out the struc- tural differences between the genus Modiola and the shell Page 217 characters of this species and introduced the new genus Rhomboidella for it, true identity and diagnostic features of R. prideaux have been poorly understood, during the following 100 years, as stated by Van Aartsen et al. (1984:58). As a consequence, the systematic position and nomenclatural status of R. prideaux has much varied in literature up to recent years. On the assumption that the species name prideaux was incorrect (as non latinized), Leach’s name has long been either variously emended (as prideauxi, prideauxii, or prideauxiana), or even rejected by different authors. The opinion of major authors of the XIXth century English school (for example Forbes & Hanley, 1849; Jeffreys, 1863) in the use of Berkeley’s name rhombea, seemed a determining factor in that sit- uation, until Bowden & Heppell (1996:103, 115) gave good reasons to reinstate prideaux for the species. How- ever, many authors still use rhombea or prideauxi (e.g., Poppe & Goto, 1993; Salas, 1996). Although the shell form and hinge of R. prideaux differ completely from those of the genus Crenella Brown, 1827, Rhomboidella has been often placed in Crenella as a subgenus. In addition, Rhomboidella was included in the Solamen as a subgenus by some other authors as well (Soot-Ryen, 1969; Bruschi et al., 1985). This varying ge- neric allocation reflects the poor understanding of the spe- cies and of its genus, as well as its frequent confusion with other mytilid species. For instance, authors dealing with Mediterranean fauna (Parenzan, 1974:55) have fol- lowed Nordsieck (1969:30—31), who added to the con- fusion in considering R. rhombea and R. prideaux as dis- tinct species, and in distinguishing Crenella arenaria Monterosato, 1875 at subspecies level within R. prideaux. This situation may have partly arisen from the difficulty of detailed examination of these tiny shells, since their maximum size does not exceed a few millimeters. The outer sculpture of the shell is clearly distinct in specimens of Crenella arenaria, owing to the absence of radial rib- lets in this species (Giannuzzi-Savelli et al., 2001:fig. 189, p. 115). A correct appraisal of the variability of R. prideaux is also necessary for its unambiguous distinction from re- lated species. This species varies 1n shell form and sculp- ture. Shape may be variable in length, the posterodorsal angle well-marked to rounded and more or less obtuse. Furthermore, the mid-ventral margin may be straight to somewhat depressed in adult stages. The comprehensive information and drawings provided by Tebble (1966:48— 49, fig. 19B & 23) illustrate the variability of shell shape in this species. The shell shape of the specimens from the Aegean Sea and eastern Sicily are similar to shells from Algesiras Bay, Spain (Van Aartsen et al., 1984:126, fig. 292). Some variation in shape and sculpture may be related to ontog- eny. Concentric sculpture tends to be less developed in juveniles, and there is an allometry of growth with a pro- gressive reduction of the height to length ratio. This can The Veliger, Vol. 48, No. 3 be correlated with a fast growth of the posterior part of shell, the umbones being shifted from a submedian po- sition on the dorsal margin at nepioconch stage to a mark- edly anterior one in the adult. Ecology and distribution: The Aegean Sea is an ecolog- ically distinct part of the Mediterranean Sea due to its peculiar hydrographic characteristics. It is an area where the brackish waters of the Black Sea (17%c) communicate with the saline waters of the Eastern Mediterranean Sea (39%c). According to Kocatas & Bilecik (1992), great fluctuations exist among the ecological features such as temperature, salinity and nutrients, in the southern Ae- gean Sea influenced by the Eastern Mediterranean Sea and in the northern Aegean Sea affected by the Black Sea. This ecological variation affects the flora and the fauna. There are tropical as well as boreal molluscs, but most of the more than 300 bivalve species have an At- lantic-Mediterranean distribution similar to that of R. pri- deaux. : Only scant information on the ecological requirements of R. prideaux exists, and most deals with its depth range and preferred substrate. Previously, the species was only reported from the Atlantic Ocean and from the Western Mediterranean region with a mean salinity of 35—36%o. However, the fact that the species has been found in the southern Aegean Sea with a salinity value of 39%o, leads to the conclusion that this species tolerates environmental conditions showing different levels of salinity, like many other species of Mytilidae. Specimens of R. prideaux live attached to the substrate with their byssal threads, like most species of Mytilidae. Berkeley (1827:229) reported a specimen from Wey- mouth adhering to a large mass of slate by its byssal threads. Nevertheless, living specimens of the species have been reported from different types of substrates, in- cluding “‘coarse bioclasts, mud, rocks and gravel, sand, laminaria, algae’? (Salas, 1996), and it is not excluded that it can lead an infaunal, superficially buried mode of life, as it has been hypothesized for the West African R. obesa Ockelmann (1983:102). The large size of the prod- issoconch and the reduction of prodissoconch I, suggests that development in R. prideaux is lecithotrophic and pos- sibly direct. This small sized Mytilid species may live in a range of depths on the continental shelf and possibly the upper slope. Although it is found at a depth of 31 m in the southern Aegean Sea, it is reported from much deeper zones in the Western Mediterranean Sea and the Atlantic Ocean. For example, in the scope of her work on the Bivalve fauna collected during the “‘Balgim”’ and “‘Fauna 1” expeditions in the Alboran Sea, the Gibraltar Straits and the Ibero-Moroccan Gulf, Salas (1996) noted that the species was found living at depths of 12—118 m. In the same work, R. prideaux was also found at the maximum depth of 332 m, but only as empty valves. In the past, R. prideaux was reported from the Western Mediterranean and in the Eastern Atlantic, from the Brit- ish Isles and Ireland (Marshall, 1897) to West Africa. Re- cent data (Seaward, 1990; Smith & Heppell, 1991) show that the northern limit of the species is in southern Eng- land (from Margate, Kent, in the East, to the Sicily Is- lands and the Bristol Channel in the West) and western Ireland (northwestern Connaught). In the South, it has been reported from Morocco to Santa Helena Island (Go- mez Rodriguez & Perez Sanchez, 1997), but its exact distribution is much uncertain because of recurrent con- fusion with similar Rhomboidella species. In that area, it certainly occurs in northern Morocco (Salas, 1996) and probably also in the northern oceanic islands of Madeira and the Canaries, where it co-occurs with R. canariensis (Odhner, 1931). It seems to be absent from the tropical coasts of Africa, where R. canariensis and R. obesa Ock- elmann, 1983 are rare and patchily distributed (Rolan & Ryall, 1999; R. von Cosel, personal communication). Easternmost records are the Gulf of Tunis, Tunisia (Wein- kauff, 1867:213, 299), Maltese Islands (Cachia et al., 1993) and the western Ionian Sea, in southern Italy (Par- enzan, 1961:44) and Sicily (Gulf of Catania, new data). In recent years, researches conducted in the eastern Ae- gean Sea and its coastal lakes have contributed to the knowledge of the faunal diversity of that area. Thus, shortly after Mytilaster marioni (Locard, 1889) was re- ported from Bafa Lake by Oztiirk et al. (2002), the dis- covery of R. prideaux significantly increased the diversity of the Mytilid fauna in this area. LITERATURE CITED BERKELEY, M. J. 1827. A short account of a new species of Mo- diola, and of the animal inhabitants of two British Serpulae. Zoological Journal 3:229—231. BowpDeN, J. & D. HEPPELL. 1966. Revised list of British Mollus- ca. 1. Introduction; Nuculacea-Ostracea. Journal of Con- chology 26:99—124. Brown, T. 1827. Illustrations of the Recent Conchology of Great Britain and Ireland. W. 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Woes ase © CMS, Inc., 2006 The Veliger 48(3):220—227 (November 2, 2006) Seasonality, Habitat Preference and Life History of Some Willamette Valley Wet Prairie Terrestrial Molluscs in Western Oregon, USA PAUL M. SEVERNS U.S. Army Corps of Engineers, Willamette Valley Projects, RO. Box 429, Lowell, Oregon 97452, USA; present address: Department of Botany and Plant Pathology, Oregon State University, Corvallis, Oregon 97331, USA (e-mail: severnsp @science.oregonstate.edu) Abstract. Willamette Valley wet prairie of western Oregon, USA is a seasonally inundated habitat that currently exists in small, highly fragmented reserves. Permanent traps in one wet prairie parcel were repeatedly sampled every five to ten days over three years in order to describe the terrestrial mollusc community. Ten species comprised the community: three exotic slugs (Arion ater s.1., Arion hortensis s.1., Deroceras reticulatum), five native snails (Catinella rehderi, Cochlichopa lubrica, Monadenia fidelis, Vertigo modesta, Verpericola cf. depressa), and two native slugs (Deroceras laeve, Prophysaon andersoni). The wet prairie mollusc community became more active with increased precipitation, with the snails having a longer active season than the slugs. There were also differences among species in the habitat they were trapped in (wet vs. dry) and the time of year they were sampled, indicating that the environment partially structures the terrestrial mollusc community. The presence of standing water in the wet prairie likely explained why many of the native species actively foraged diurnally under sunny conditions and the overlap of generations in reportedly ‘“‘annual’”’ species. The unique seasonal flooding of the wet prairie is a harsh condition for terrestrial mollusc existence and the habitat should be investigated further for new species and adaptive mollusc behaviors. INTRODUCTION Little is known about the terrestrial molluscs inhabiting Willamette Valley wetland prairie of western Oregon, USA, perhaps due to the rarity of the habitat. Once a dominant ecosystem along the 200 km long Willamette River floodplain, wetland prairie, like other North Amer- ican prairies and wetlands has experienced drastic reduc- tions in size. Before the 1850s, anthropogenic fires likely maintained the Willamette Valley wetland prairie as a grassland (Boyd, 1986), without which it succeeds to a wet Oregon ash (Fraxinus latifolia Benth. 1844) forest. The loss of habitat through urbanization, agricultural de- velopment, succession, and invasion of exotic species has left Willamette Valley wetland prairie highly fragmented. Currently, Willamette Valley wetland prairie estimates are circa 600 ha, which is considerably less than 1% of the historical breadth of the habitat not more than 150 years before the present (Christy & Alverson, 1994). Due to the high percentage of habitat loss, Willamette Valley wet prairie 1s considered to be one of the most endangered ecosystems in the United States (Noss et al., 1995). Wetland prairie is a seasonally inundated grassland that has standing water from November through May. The perched water table is caused by a semi-impervious al- luvial clay layer that is approximately | to 3 m below the soil surface (Finley, 1995). As precipitation increases, the water table level rises to exposure surrounding tussocks of the dominant native bunchgrass, Deschampsia cespi- tosa (L.) Beauv. 1753, creating small islands of terrestrial habitat (Figure 1). Eventually, the accumulated surface water evaporates by June and by July the soils are hard and occasionally cracked. A perennially dry mound hab- itat, which contains plant species characteristic of Wil- lamette Valley upland prairie, persists due to a slight in- crease in elevation.of less than a meter from the surround- ing inundated habitat. This perennially dry habitat is like- ly a refuge from terrestrial molluscs when the lower regions of the prairie are flooded. A history of repeated seasonal flooding in the wetland prairie has likely led to the evolution of endemic plant species adapted to the soil conditions, eight of which are listed as either endangered, threatened, or species of con- cern (Oregon Natural Heritage Program, 2001). Some en- demic or wetland obligate faunal taxa are also known to occur in the Willamette Valley wet prairie but little is known about their distribution and life history (Lattin & Schwartz, 1986; Oregon Natural Heritage Program, 2001; Severns, 2003; Severns & Villegas, 2005). One of these species is informally named the “Bald Hesperion” (Ves- pericola cf. depressa) and appears to be limited to the confines of remnant wet prairie parcels in the southern Willamette Valley and nearby upland grasslands (Oregon Natural Heritage Program, 2001). The Willamette Valley wet prairie terrestrial mollusc community has not been formally studied and has only been sporadically collected from in the past. Seasonal in- undation and rapid evaporation of standing water in the P. M. Severns, 2005 Page 221 Figure 1. Photograph of the Horkelia Prairie Management Unit study site. The inset photo shows the “‘small islands” of terrestrial habitat in the inundated portion of the prairie. wet prairie creates a dynamic and rapidly fluctuating hab- itat for terrestrial molluscs compared with the more en- vironmentally stable forest floors of the Pacific North- west. Given that there are endemic plants adapted to the seasonal flooding of Willamette Valley wet prairie there may similarly be unique terrestrial mollusc species other than Vespericola cf. depressa that have not yet been col- lected from this ecosystem. A multiyear repeated sam- pling of terrestrial molluscs in the southern Willamette Valley wet prairie parcel was initiated in 1999 to generate a species list of terrestrial molluscs for the site and to determine thé effects of prescribed fire on the mollusc community (Severns, 2005). In this paper I describe a Willamette Valley wet prairie terrestrial mollusc com- munity, temporal patterns in species’ relative abundance, species’ preference for microhabitats, and contribute life history observations from the three years of field study. MATERIALS AnD METHODS Study site: I sampled a parcel of wet prairie managed by the US Army Corps of Engineers, located approximately 10 km west of Eugene, Oregon, USA (44°20'38’N, 123°10'03"W). The study site, Horkelia Prairie Manage- ment Unit (circa 4 ha and 114 m elevation) is typical tufted hairgrass (Deschampsia cespitosa (L.) Beauv.)- dominated Willamette Valley wet prairie habitat that has shallow, remnant furrows historically used (unsuccessful- ly) to drain the standing water for the creation of pasture. There is no evidence suggesting that the study site was used for crop cultivation. Horkelia prairie is primarily dominated by gaminoids, with occasional ash trees (Frax- inus latifolia) and serviceberry (Amelanchier alnifolia Nutt. 1834) that dot the landscape (Figure |). Dominant wet prairie plant species at the study site other than Des- champsia_ cespistosa are: Anthoxanthum odoratum L. 1753 (exotic species), Rosa nutkana Presl. 1851, Rubus armeniacus L. 1874 (exotic species), Horkelia congesta var. congesta Dougl. 1829, Camassia quamash (Pursh) Greene 1814, Aster hallii Gray 1943, and Grindelia in- tegrifolia Nutt 1836. A semi-rural residential area with interspersed culti- vated hay fields borders the east edge of the study site and Rose Prairie Research Natural Area unit is to the north. Horkelia Prairie is bounded on the south by a canal constructed in the 1950s that separates the study site from the Amazon unit of the wet prairie Research Natural Area. To the west the study site is confined by Fern Ridge Reservoir and to the northeast the wet prairie habitat tran- sitions into a small, degraded white oak (Quercus gar- ryana Doug. ex Hook. 1840) forest that was not part of the surveyed habitat. Page 222 The Veliger, Vol. 48, No. 3 The climate of western Oregon and the study site is Mediterranean. A weather station approximately 6 km northeast of the study site receives approximately 104 cm of mean annual rainfall, with 86% of the annual precip- itation occurring from November through June. The mean annual temperature is mild, 11.2°C, with the coldest month having a mean temperature of 3.9°C (January), and the warmest month (July) averaging 19.1°C (National Weather Service, 2002). Mollusc sampling: I used coverboards, 0.6 m X 0.6 m pieces of 1 cm thick plywood, to sample terrestrial mol- luscs at the study site because trapping overcomes some sampling bias (Boag, 1982) and it may be more easily replicated by novices compared to hand searching (Ward- Booth & Dussart, 2001). The coverboards sampled a gra- dient of wet and dry habitats within the study site and their position was permanent. The distance between traps (n = 21) ranged from approximately 8 to 40 meters. I estimated the cover of standing water, beneath the cov- erboard to the nearest 1% for 1—10% covers and in 5% increments thereafter. Coverboards were also placed in an adjacent burned area, but I chose to present only the cov- erboard data from the unburned portion of the study (n = 21 coverboards) because some species were not de- tected in the burned area (Severns, 2005). I sampled all coverboards every 5 to 10 days throughout three terres- trial mollusc active seasons, October—July of 1999-2002. In year | (1999/2000) I checked the coverboard arrays on 42 occasions, while in year 2 (2000/2001) the site was visited 41 times, and in year 3 (2001/2002) on 29 occa- sions. Traps were checked before 1300 hr on days fol- lowing nights that were favorable for mollusc activity, generally mild temperatures and high relative humidity. I recorded the number and species of terrestrial molluscs encountered on top of and adhered to the underside of the coverboard as well as the ground below the cover- board. When encountered, dead snails were counted and then discarded from beneath coverboards so that resam- pling of dead individuals was avoided. Live molluscs cen- sused using the coverboards were left in situ except for a small number of voucher specimens of each species when required for a positive identification. These voucher specimens were deposited in the Oregon State Arthropod Collection (OSAC) with individual bar codes for six spe- cies (See Table 1) and the remainder deposited under ac- cession #00133. I used trap efficiency, the number of molluscs captured per trap unit, as an index for relative mollusc abundance. For monthly estimates of mollusc abundance, the total number of captures was divided by the number of traps multiplied by the number of days sampled in the month. I considered a trap having less than 3% standing water during the time interval that the wet prairie maintains standing surface water to be perennially dry habitat. Molluscs were identified using Pilsbry (1939, 1940, 1946, 1948) and Burch and Pearce (1990). Arion Table 1 Total number of species captures and the percent of cap- tures occurring under dry mound coverboards. Total # of % captures on dry Species captures mounds (<3% water) Vespericola ct depressa 1784 46.4 Deroceras reticulatum 424 60.4 Miiller 1774 Deroceras laeve Miller 372 20.7 1774 Catinella rehderi Pils- 148 23.6 bry 1948 Arion hortensis s.l. Fér- oy) O23 ussac 1819 Monadenia fidelis Gray 29 75.9 1834 Vertigo modesta Say 20 85.0 1824 Prophysaon andersoni 19 Wael Cooper 1872 Arion ater s.l. L. 1758 16 81.3 Cochlicopa lubrica 5 80.0 Miiller 1774 * OSAC specimen numbers: 44544, 44546, 44547, 44551— 44563, 44583-44588. ater and Arion hortensis were identified sensu lato al- though segregates of each species complex are recog- nized (e.g., Rollo & Wellington, 1975; Davies, 1977; Pearce & Bayne, 2003). RESULTS Community: There was a total of 2869 mollusc obser- vations from checking the coverboard array 112 times in the three years of sampling (Table 1). I detected ten spe- cies in the study site. Vespericola ct. depressa was the dominant species, followed by Deroceras reticulatum Miiller 1774, Deroceras laeve Miller 1774, Catinella rehderi Pilsbry 1948, Arion hortensis s.1. Férussac 1819, Mondenia fidelis Gray 1834, Vertigo modesta Say 1824, Prophysaon andersoni Cooper 1872, Arion ater s.l. L. 1758, and Cochlicopa lubrica Miller 1774 (Table 1). Nine species were trapped in the first year of sampling and the last species, Cochlicopa lubrica, was captured during December of the second year. Total site diversity calculated using the Shannon Index (Magurran, 1988) was H’ = 1.22 for the pooled three years of data. Accumulation of precipitation and seasonality were consistent indicators of community activity and relative abundance during the sampling period. Plotting mean dai- ly rainfall by month against monthly coverboard trap ef- ficiency showed that the wet prairie mollusc community as a whole became more active with increasing precipi- tation (Figure 2). Snails were more abundant than slugs in the early fall and late spring/early summer, extending P. M. Severns, 2005 1.44 -- @-- Mean Precip/Day/Month —o— Snail Trap Efficiency — a-— Slug Trap Efficiency trap efficiency Dec Feb | Apr Jun .?) a a c oa o ® ao 5 = (<< Aug & Oct-00 | Figure 2. Mean daily precipation by month and monthly trap efficiency for slugs and snails over three active seasons. their growing season past that of the slugs which peaked in late winter (Figure 2). The native species Vespericola cf. depressa, Catinella rehderi, and Deroceras laeve de- creased in abundance in January, the coldest month of the year, while the exotic slug species Deroceras reticu- latum and Arion hortensis increased in relative abundance during the same time (Figure 3). Species accounts: Vespericola cf. depressa was encoun- tered throughout the year from October through June (Figure 3) and was also active in the summer months of July, August, and September on humid days during and following thunderstorms. Although the relative abun- dance of Vespericola generally decreased during the win- ter (Figure 3), some individuals were active beneath coy- erboards on days that were below freezing and snowing. In February and March, Vespericola were often found covered with mud, suggesting that they spent time below ground during the winter. This species foraged diurnally on mild (<20°C), overcast and partly sunny days when the relative humidity was high. Vespericola consumed the biofilm that accumulated beneath the coverboards, fungal mycelia, green algal mats that remained after the inun- dated part of the wet prairie dried in May, and unidenti- fied plant seedlings. Vespericola were evenly distributed between the wet coverboards (>3% mean standing water) and the peren- nially dry traps (Table 1). Adults and subadults were found in both dry mound and wet habitats, but neonate snails (up to 3—5 individuals/coverboard) were encoun- tered under the dry mound traps 80% of the time (n = 25 observations) suggesting that Vespericola eggs were laid near the mound coverboards. Crushed Vespericola shells were found in the nests of shrews (Sorex vagrans Baird) and deer mice (Peromiscus maniculatus Wagner) under dry mound coverboards. Oc- casionally Pterostichus sp. (Carabidae) and rove beetles (Staphylinidae) were present under the coverboards but Page 223 only a handful of intact shells were found (27 individu- als), suggesting that snail predation by beetles is rare. None of the Vespericola shells at the study site had holes near their aperture indicative of Ancotrema sp. predation. Deroceras reticulatum, an exotic slug and the second most abundant species, slightly preferred dry mound hab- itat over the wetter intermound areas (Table 1) but was rarely found under coverboards that had more than 20% standing water present (4 of 424 wet coverboard obser- vations). D. reticulatum became increasingly common in the late fall and declined markedly in abundance begin- ning in May (Figure 3). It seemed to prefer a cool, moist climate and appeared to be greatly stressed when air tem- peratures exceeded 21°C. More than 90% of the individ- uals observed had wounds or scars on the dorsal and lat- eral tail regions but I did not directly observe antagonistic behavior in this species. Eggs of this species were also encountered under dry coverboards in the months of Jan- uary, February, March, and April. Adults readily ate Ro- maine lettuce in captivity but I had no direct observations of herbivory in the field. Deroceras laeve, the dominant native slug, preferred wet areas with standing water over dry mound habitat (Table 1) and was most abundant during the rainiest por- tion of the wet season, from November—April (Figure 3). Throughout the winter (December—March) D. laeve com- monly foraged in full daylight on rainy, overcast, and even sunny days above freezing but less than 10°C. Der- oceras laeve was associated with the ant mounds con- structed by Formica occulta Francoeur. In the winter, when the ants were not at their peak activity, D. laeve were on the outside and inside of the ant mounds and consumed the middens from the colony. D. laeve also consumed the biofilm on the underside of the coverboard and unidentified plant germinants, but the species may be discriminating herbivores as they refused Romaine lettuce in captivity. In March of 2002, I observed three D. laeve consuming a wounded earthworm (Lubricus terrestris L.) that had partially drowned in shallow water, suggesting carnivory in this species. Catinella rehderi, like Deroceras laeve, had a strong affinity for wet microhabitats (Table 1) and the two spe- cies were often found beneath the same traps. C. rehderi consumed cotyledons of two leguminous plants, Lupinus sulphureus ssp. kincaidii (Smith) Phillips and Lotus for- mossissimus Greene, and also grazed on biofilm that coat- ed the coverboards and decomposing vegetation. In late April and early May (all three years), Catinella rehderi rested atop coverboards on sunny days that exceeded 22°C. These individuals died in the subsequent days (ap- parent desiccation) and were adhered to the exposed sur- face of the coverboard. I encountered live C. rehderi be- neath the same coverboard that dead snails were adhered to. Monadenia fidelis, Arion ater s.1., Arion hortensis s.1., Prophysaon andersoni, Cochlicopa lubrica, and Vertigo Page 224 Arion hortensis —O— Yr 1 trap —#— Yr 2 trap 0.1 —a— Yr 3 trap trap efficiency oO fo) a Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sept The Veliger, Vol. 48, No. 3 Catinella rehderi —o— Yr1 trap —a-— Yr 2 trap —a— Yr 3 trap Oct Nov Dec Jan Feb Mer Apr May Jun Jul Aug Sept —o— Yr 1 trap —a— Yr 2 trap —a— Yr 3 trap eee Deroceras laeve 0.35 4 Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sept Deroceras reticulatum —o— Yr 1 trap —a— Yr 2 trap —a— Yr 3 trap Vespericola cf depressa —o— Yr 1 trap —a_ Yr 2 trap 14,5 —a— Yr 3 trap > (2) c £4 2 b= o 2 s Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sept Vertigo modesta —o— Yr 1 trap —a— Yr 2 trap —a— Yr 3 trap > oO i= ao 2 = o Qa £ Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sept Figure 3. modesta were uncommon or rare species at the study site. All six species were more frequently encountered under dry coverboards than semi-inundated traps (Table 1). Monadenia fidelis, Prophysaon andersoni, Cochlicopa lu- brica, and Arion ater were active throughout the year, whereas Vertigo modesta and Arion hortensis displayed strong seasonality trends. Arion hortensis was most abun- dant in the winter months January and February (Figure 3). Vertigo modesta activity patterns were seasonally bi- modal, appearing during the mid-fall after the rains began but before there was standing water on the prairie and also in the late spring as the prairie dried out (Figure 3). V. modesta grazed on the algal and microbial layer grow- 0 UE T T T T Oct Nov Dec Jan Feb Mar Apr [ae May Jun Jul Aug Sept Monthly trap efficiency for six species of wet prairie molluscs over three active seasons. ing on the underside margins of the coverboards, and oc- casionally foraged on dead but prostrate grass stems. Monadenia fidelis and Prophysaon andersoni were in- frequently encountered but appeared to be actively mov- ing about the study area. Both species were frequently captured once under a coverboard and were not generally recaptured at the same trap during the same year, sug- gesting that both the slug and the snail may be wide rang- ing. Monadenia fidelis found in a wet prairie parcel less than 0.3 km to the south of the study site were common and appeared to be tightly associated with small mammal holes, often found inside the mouth of the small mammal burrow. Furthermore, in the winter and early spring, M. P. M. Severns, 2005 Page 225 fidelis foraged diurnally and consumed the thallus of rein- deer lichen Cladina mitis (Sandst.) Hustich 1918, unin- dentified plant cotyledons and young, tender foliar plant tissue. Multiple overlapping generations of the reportedly “annual’’ species Deroceras reticulatum, Deroceras lae- ve, Catinella rehderi, Arion ater, and Cochlicopa lubrica were found beneath coverboards during the fall, winter, and spring, indicating that these species are likely not “‘annual”’ in the wet prairie study habitat. DISCUSSION Regionally, the wet prairie parcel I surveyed has low spe- cies richness compared to the richest forested Pacific Northwest habitats but has comparable species numbers to slightly disturbed sites and forests with lower plant diversity (Branson, 1977; Cameron, 1986). A diversity of ten species is either intermediate or on the lower end of Pacific Northwest mollusc study site diversity, but three of the ten Willamette Valley wet prairie species were ex- otic slugs, Arion ater, Arion hortensis, and Deroceras re- ticulatum, suggesting the native mollusc richness is low. I considered Cochlicopa lubrica to be native to the wet prairie despite the presence of human housing develop- ments, which have been suggested by Roth and Pearce (1984) to be sources for non-native C. /ubrica genotypes. With only seven native species found in the three years of sampling, the wet prairie study site has a comparably low native species component to other Pacific Northwest ecotypes (Cameron, 1986). Low species richness at the wetland prairie study site is not unexpected because grassland and old-field terrestrial mollusc diversity is usu- ally lower than that of nearby forested habitats in North American temperate climates (Karlin, 1961; Gleich & Gilbert, 1976; Cameron, 1986; Ports, 1996; Nekola, 1999) and the potentially harsh environmental conditions in the wetland prairie are unlikely to support a rich ter- restrial mollusc community. Moreover, a history of at least semi-annual anthropogenic fires set by native Amer- icans that burned entire portions of the Willamette Valley (Boyd, 1986) may have resulted in the loss of historical mollusc diversity as prescribed burning was associated with lower wet prairie mollusc richness and abundance (Severns, 2005). Similar reductions in terrestrial mollusc richness and abundance following fires in dry grasslands and forests have also been documented (Nekola, 2002; Kiss & Magnin, 2003). However poor, the wetland prairie mollusc diversity may be compared to nearby coniferous forests in the Coast Range and Cascade Mountains. The wetland prairie mollusc community is unique because there are few mol- lusc species found in the wetland prairie that are char- acteristically associated with Pacific Northwest Pseudot- suga menziesii (Mirbel) Franco 1825 (Douglas-fir) for- ests. Monadenia fidelis is found in both fir-forests (Bran- son, 1977; Cameron, 1986) and the Willamette Valley wetland prairie, however the wetland prairie race is di- minutive (about half the size of Monadenia fidelis found in fir-forests) suggesting that it may be distinct from the typical Douglas-fir forest race. Vespericola depressa, likely the closest related species to the wet prairie Ves- pericola (2002, B. Roth, personal communication) occurs in tallus slopes near the Columbia River gorge, 280 km to the north of the study site (Pilsbry, 1940). Adult wet- land prairie Vespericola lack shell setae, similar to Ves- pericola depressa, and both species occupy non-forested habitats. Another Vespericola species, one with shell se- tae, exists sympatrically with the wetland prairie Vesper- icola, except that it lives in the nearby Quercus garryana woods bordering Willamette Valley upland prairie. Nei- ther of the above described Vespericola species are the typical types found in Douglas-fir forests, but Douglas- fir is slowly invading the Willamette Valley floor and the study site, primarily through human efforts. These Doug- las-fir forests are relatively young, circa 100 years old, but they may create a corridor for the dispersal of typical Douglas-fir forest mollusc species into the wetland prai- rie. Evidence for recent mollusc dispersal coinciding with human modification of forests may reside in the presence of Prophysaon andersoni in the wet prairie. P. andersoni is frequently encountered on disturbed edges of young Douglas-fir forests (personal observation). Seasonality and the presence of standing water ap- peared to give some structure to the wet prairie mollusc community. In general, mollusc activity on the wet prairie appears dependent on accumulated precipitation. Once the wet prairie fills with standing water, the soils remain saturated through May despite a lack of substantial pre- cipitation, perhaps extending the season of mollusc activ- ity relative to nearby drier upland prairie habitats. Fur- thermore, some species like Catinella rehderi and Dero- ceras laeve were tightly associated with flooded micro- habitats whereas the other species appeared to prefer drier microhabitats. Snails emerged earlier than slugs and also remained active longer in the wet prairie (Figure 2). Pre- sumably, having a shell allows snails to remain active under semi-dry conditions that slugs in the wet prairie do not tolerate, however within the group of slug and snail species there appeared to be an effect of seasonality on patterns of abundance (Figure 3). These patterns of sea- sonality indicate that sampling and monitoring the wet prairie terrestrial mollusc community should span the whole active season as some species peaked in abundance during the summer and others were present primarily in the fall and spring (Figure 3). A severe drought occurred during the active mollusc seasons of year 2000/2001 (year 2). Precipitation in the drought year was nearly half of the normal mean yearly rainfall (National Weather Service, 2002) and as a result the study site did not have the amount of standing water that is normally present and soils became dry earlier in Page 226 the year. This lack of standing water appeared to affect snail abundance during the drought year, but slug abun- dance did not decrease until the following year (Figure 2). Vespericola ct. depressa and Monadenia fidelis snails were associated with small mammal burrows and tunnels, often being covered in mud even during the wettest months of the year. In the drought year, snails may have moved to small mammal burrows to seek refuge from the warm temperatures and lack of precipitation, resulting in low trap efficiency during the drought. Contrary to the snails, slugs were either encountered sheltering beneath coverboards, under tussocks of the dominant bunchgrass Deschampsia cespitosa, or beneath scattered patches of thatch that were approximately 2 cm in thickness. During the drought year, coverboards likely provided temporary shelter for slugs which were sampled efficiently by traps. However, the drought year’s slug cohort likely had low survival explaining the decline in the slug population the year following the drought while snails were trapped at relatively comparable levels to the normal precipitation year (Figure 2). Moreover, both the native slug (Dero- ceras laeve) and the two exotic slugs (Deroceras reticu- latum, Arion hortensis) had similar patterns of drought and post-drought year patterns (Figure 3), suggesting that the drought affected all slugs, not just a subset of them. Seasonal flooding and the fluctuation of standing water in the wetland prairie is likely related to some of the mollusc behavior observed in the three years of field study. Some species, Deroceras laeve, Vespericola cf. de- pressa, and Monadenia fidelis commonly foraged diur- nally at the study site despite being reported as primarily nocturnal species (Getz, 1963) or species whose conspe- cifics are nocturnal (Roth & Pressley, 1986; Szlavecz, 1986). It seems plausible that the high relative humidity created by the standing water in the wet season may al- lows for diurnal foraging without a high risk of dehydra- tion. Additionally, the presence of standing water and rel- atively mild temperatures likely explains the overlap of generations in some ‘‘annual’’ species that are reported to have a single synchronous lifestage cohort (Boag & Wishart, 1982). Either the high humidity and mild tem- peratures extend the lifespan of “‘annual’’ species like Deroceras laeve, or the active season is long enough to allow multiple cohorts to be produced. Deroceras laeve is often collected from “‘wet’’ habitats (DeWitt, 1955; Rollo & Wellington, 1979; Boag & Wis- hart, 1982; Ports, 1996; Frank, 1998) and at the study site it preferred wet coverboards over the dry ones (Table 1), demonstrating a consistent affinity for wet habitats across the range of the species. Curiously, Pearl (1902) noted that D. laeve moved into standing water with decreasing temperature and left the water with increasing tempera- ture. I observed no behavior suggesting that D. laeve chose to be submerged at any time of the year, but the slugs did appear to be content in shallow water conditions when their pneumostome was not submerged. The Veliger, Vol. 48, No. 3 The other wetland prairie mollusc species preferring an inundated habitat is Catinella rehderi and it was often found in cohabitation with Deroceras laeve. Catinella rehderi, however, had a conspicuous and unexplained be- havior at the study site. Some individuals of C. rehderi appeared to “commit suicide”’ by resting atop of cover- boards on warm, sunny days in April/May and exposing themselves to direct sun for the entire day. Not all Cati- nella rehderi had this behavior and I noticed no ocular tentacle swelling that would suggest a nematode parasite in the individuals crawling atop the coverboards on warm, sunny days. It is unclear if the coverboard was an attrac- tive resting surface for the snails and they inadvertently died from choosing an exposed resting site or that C. rehderi is a host to a parasite that alters resting behavior. Acknowledgments. I thank Kat and Jim Beal for access to the study site and support of this project. I am indebted to John S. Applegarth for his constant encouragement and aid with the iden- tification of molluscs. I also thank both T. J. Frest and B. Roth for their verification of mollusc identifications and an anonymous reviewer for helpful comments on this manuscript. LITERATURE CITED Boaa, D. A. 1982. Overcoming sampling bias in studies of ter- restrial gastropods. Canadian Journal of Zoology 60:1289— 1292) Boac, D. A. & W. D. WisHart. 1982. Distribution and abun- dance of terrestrial gastropods on a winter range of bighorn sheep in southwestern Alberta. Canadian Journal of Zoology 60:2633—2640. Boyp, R. 1986. Strategies of Indian burning in the Willamette Valley. 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Butterflies hanging on to existence in the Willamette Valley: a relict population of the great copper (Lycaena xanthoides Boisduval). Northwest Science 79:77—80. SZLAVECZ, K. 1986. Food selection and nocturnal behavior of the land snail Monadenia hillebrandi mariposa A.G. Smith (Pul- monata: Helminthoglyptidae). The Veliger 29:183—190. WarD-Booth, J. E & G. B. J. Dussartr. 2001. Consistancy in hand-searching for snails. Journal of Molluscan Studies 67: 502-506. The Veliger 48(3):228—229 (November 2, 2006) THE VERIGER © CMS, Inc., 2006 Exallocorbula (Bivalvia: Corbulidae), a New Name for the Amazonian Molluscan Fossil Pebasia Nuttall ANDRE NEMESIO Departamento de Zoologia, Instituto de Ciéncias Biold6gicas, Universidade Federal de Minas Gerais, Caixa Postal 486, Belo Horizonte, MG. 30.123-970 Brazil (e-mail: andre @nemesio.com.br) AUDREY ARONOWSKY AND LAURIE C. ANDERSON* Department of Geology and Geophysics, Louisiana State University, E235 Howe-Russell Geoscience Complex, Baton Rouge, Louisiana 70803 Abstract. Exallocorbula nom. nov. is proposed for Pebasia Nuttall, 1990 (Bivalvia: Myoida: Corbulidae), a homonym of Pebasia Roewer, 1947 (Arachnida: Opiliones: Cosmetidae). Both morphology and recent phylogenetic analyses sup- port the distinctness of Pebasia as a lineage within the corbulid bivalves, thus a new generic name is necessary to replace Pebasia Nuttall. The new name, Exallocorbula, refers to the extreme morphologic difference of the left and right valves of the type species. Exallocorbula nom. nov. Pebasia Nuttall, 1990: 315 [type-species: Pachydon (An- isorhyncus?) dispar Conrad, 1874, by original designa- tion (Nuttall, 1990:315)], nec Pebasia Roewer, 1947 [(type-species: Pebasia singularis Roewer, by monotypy (Roewer, 1947:25)]. Nuttall (1990) erected the genus Pebasia (Bivalvia: Myoida: Corbulidae) for P. dispar Conrad, 1874, a mid- dle Miocene species of lacustrine mollusk from the Pebas Formation, Loreto, Peru. Nuttall (1990:315) erected this monospecific genus after noting many morphologic dif- ferences between P. dispar and the type species of Pach- ydon Gabb, 1869 (Pachydon obliquus Gabb, 1869), a ge- nus in which it had been previously placed. Recent phy- logenetic analyses conducted by Anderson et al. (2006) support Nuttall’s separation of Pebasia dispar from Pach- ydon because Pebasia is consistently reconstructed out- side of the Pachydon crown group (see Anderson et al., 2006:figs. 6, 7). Pebasia Nuttall, however, is preoccupied by Pebasia Roewer, 1947, a genus of extant arachnid (Opiliones, Cosmetidae), also with a type locality at Pebas, Peru. Roewer (1947) erected the genus with the type species Pebasia singularis Roewer, 1947, a Peruvian Amazon en- demic whose name is valid and in use (Kury & Pinto da Rocha, 2002). According to the principle of priority, an unavailable or invalid name must be replaced by its next oldest avail- able synonym (Article 23.3.5, International Commission * Corresponding author, e-mail: laurie @ geol.lsu.edu on Zoological Nomenclature, 1999). Pebasia dispar, however, is markedly distinct from Pachydon (as outlined above) and from Anisorhnycus Conrad (a misspelling of Anisorhynchus Conrad in Meek, 1871), the subgenus in which it was originally placed. Further, Anisorhynchus is itself a junior homonym of the coleopterid Anisorhynchus Schoenherr 1842, and Vokes (1945) proposed the replace- ment name Ursirivus for the bivalve genus. The type spe- cies for Ursirivus is Corbula (Potamomya?) pyriformis Meek 1860 from the Cretaceous, Bear River Group of Wyoming. Pebasia dispar is morphologically distinct from Ursirivus pyriformis, possessing a concavoconvex shell, a bilobed right valve, and a relatively thin left valve whose cardinal tooth and chondrophore are highly re- duced. In contrast, U. pyriformis is nearly equivalved, with both valves convex, a right valve that is not bilobed, and a left-valve cardinal tooth and chondrophore that are well developed. Therefore, because Pebasia Nuttall lacks a current syn- onym, Exallocorbula nom. nov. is here established to re- place it (article 60, recommendation 60A: International Commission on Zoological Nomenclature, 1999). The name is a combination of exallo (Greek for quite differ- ent), to denote the profound shape asymmetry of the left and right valves, and corbula (Latin for little basket), to describe the inflated nature of the right valve of many corbulid taxa. As a result, the following new combination is introduced: Exallocorbula dispar (Conrad) comb. nov. Acknowledgments. We are indebted to Dr. Adriano B. Kury and Dr. Ricardo Pinto da Rocha for providing a copy of the original description of Pebasia singularis C. FE Roewer (A. B. A. Nemésio et al., 2005 Kury), and for information on the status of this genus (A. B. Kury and R. Pinto da Rocha). LITERATURE CITED ANDERSON, L. C., J. H. HARTMANN & E WESSELINGH. 2006. Close evolutionary affinities between freshwater corbulid bivalves from the Neogene of western Amazonia and Paleogene of the northern Great Plains, USA. Journal of South American Earth Sciences 21:28—48. ConrabD, T. A. 1874. Remarks on the Tertiary clay of the Upper Amazon with descriptions of new shells. Proceedings of the Academy of Natural Sciences Philadelphia, 1874:25—32, pl. 1. Gasp, W. M. 1869. Descriptions of fossils from the clay deposits of the Upper Amazon. American Journal of Conchology, 4: 197-200, pl. 16. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE. 1999. International Code of Zoological Nomenclature. 4th ed. The International Trust for Zoological Nomenclature: Pa- dova. 306 pp. Kury, A. B. & R. PINTO DA Rocua. 2002. Chapter 4.4, Opiliones. Pp. 345-362 in J. Adis (ed.), Amazonian Arachnida and Myriapoda, Pensoft Series Faunistica 24. Pensoft Publishers: Moscow. Page 229 MEEK, E B. 1860. Descriptions of new fossil remains collected in Nebraska and Utah, by the exploring expeditions under the command of Capt. J. H. Simpson, of the U.S. Topograph- ical Engineers, [extracted from that officer’s forthcoming re- port]. Proceedings of the Academy of Natural Sciences Phil- adelphia, 12:308—-315. MEEK, FE B. 1871. Part I, Palaeontology. In C. King, Report of the Geological Exploration of the Fortieth Parallel made by order of the Secretary of War According to Acts of Congress of March 2, 1867, and March 3, 1869, Under the Direction of Brig. and Bvt. Major General A. A. Humphreys, Chief of Engineers. Vol. 4:669 pp., 24 pls. NuTTALL, C. P. 1990. A review of the Tertiary non-marine mol- luscan faunas of the Pebasian and other inland basins of north-western South America. Bulletin of the British Mu- seum of Natural History, Geology 45(2):165—371. Roewer, C. Fr. 1947. Diagnosen neuer Gattungen and Arten der Opiliones-Laniatores (Arach.). Senckenbergiana 28(1—3):1— 58, 12 pls. SCHOENHERR, C. J. 1842. Genera et species curculionidum cum synonymia hujus familiae. Vol. 6, pt. 2. Roret: Paris. 495 Pp. VoKEs, H. E. 1945. Supraspecific groups of the pelecypod family Corbulidae. Bulletin of the American Museum of Natural History 86:1—32. The Veliger 48(3):230—233 (November 2, 2006) THE VELIGER © CMS, Inc., 2006 Cryptodaphne kilburni, a New Species of Bathyal Turrid (Gastropoda: Prosobranchia) from the Gulf of Aden (Northwestern Indian Ocean) MAURO MORASSI Via dei Musei 17, 25121 Brescia, Italy (e-mail: vitmoras @tin.it) AND ANTONIO BONFITTO Dipartimento di Biologia e.s., Via Selmi 3, 40126 Bologna, Italy (e-mail: bonfitto@alma.unibo.it) INTRODUCTION In September 1992 a series of dredgings was performed in the southern Red Sea and Gulf of Aden by the French research vessel Marion Dufresne in order to contribute to the knowledge of the bathyal thanatocoenoses and bio- cenosis of that area. Among the abundant material of pe- lagic mollusks (thecosomate pteropods and heteropods) numerous empty shells of a distinctive turrid species were found. This species has conchological characters similar to those of the early Miocene Cryptodaphne pseudodrillia Powell, 1942, from New Zealand, type species of Cryp- todaphne Powell, 1942, a genus comprising species usu- ally occurring at upper bathyal depths and reported from the Indo-Pacific area. The species from Aden differs dis- tinctly from all its previously described congeners and is accordingly here proposed as Cryptodaphne kilburni, sp. nov. Abbreviations used in the text are: a/l] = ratio of ap- erture length to total shell length; b/1 ratio of shell breadth to total length; MZB = Museo di Zoologia dell’ Universita di Bologna; NM = Natal Museum, Pietermar- itzburg. SYSTEMATIC DESCRIPTION Family TURRIDAE H. & A. Adams, 1853 Subfamily RAPHITOMINAE Bellardi, 1875 Genus Cryptodaphne Powell, 1942 Type species by original designation: Cryptodaphne pseudodrillia Powell, 1942 Description: Shell small to medium-sized (5.1—18.5 mm in length), claviform or biconic, with high, sometimes pagodiform, spire and excavated last whorl forming rel- atively short to moderately long neck. Sutural ramp wide, remaining part of whorl bearing spiral cords or keels. Ax- ial folds typically absent, reduced to vestigial nodes on periphery of early whorls, or relatively developed and extending from periphery to lower suture on spire whorls (in potential subgenus Acamptodaphne). Whorl surface covered by numerous collabral threads (particularly evi- dent on the sutural ramp). Anal sinus reversed L-shaped. Protoconch multispiral with decussately sculptured whorls. Included species: The type species (Early Miocene of New Zealand); Cryptodaphne kilburni, sp. nov. (Gulf of Aden); Cryptodaphne rugosa Sysoev, 1997 (Indonesia); Cosmasyrinx semilirata Powell, 1942 (Upper Oligocene of New Zealand); Pleurotomella affinis Schepman, 1913 (Ceram Sea, Indonesia); Pleurotomella biconica Schep- man, 1913 (near Waigeu Island, Indonesia) and Pleuro- tomella gradata Schepman, 1913 (Halmahera Sea, Indo- nesia). Discussion: Powell (1942) introduced Cryptodaphne for Cryptodaphne pseudodrillia Powell, 1942, from the Early Miocene (Otaian) of Kaipara, New Zealand. The same author (Powell, 1966) subsequently recognized four Re- cent Indonesian species as members of the genus. How- ever, one of the species listed by Powell (1966), Pleu- rotomella abbreviata Schepman, 1913, does not belong to Cryptodaphne (Morassi & Bonfitto, submitted). Shuto (1971) proposed Acamptodaphne Shuto, 1971, as a monotypic subgenus for Pleurotomella biconica Schep- man, 1913. That species has axial ornamentation on all teleoconch whorls (but fading below periphery on the last whorl) and a deep, narrow anal sinus differing in shape from that of Cryptodaphne pseudodrillia. Beu & Maxwell (1990) transferred the Upper Oligocene New Zealand Cosmasyrinx semilirata Powell, 1942, to Cryptodaphne on the basis of the protoconch sculpture. More recently, Sysoev (1996, 1997) reported Cryptodaphne gradata (Schepman, 1913) from the Gulf of Aden and described Cryptodaphne rugosa Sysoev, 1997, from Indonesia. The specimen figured by Sysoev (1996) as Crypto- M. Morassi & A. Bonfitto, 2005 daphne gradata has axial folds on early whorls, a feature not reported in the descriptions provided by Schepman (1913) and Shuto (1971). If this identification is correct then the occurrence and relative development of axial folds is a feature variable within the same species and need not necessarily represent a character valid in su- praspecific segregation. Axial sculpture confined to spire whorls also occurs in Cosmasyrinx semilirata (see Pow- ell, 1942). Schepman (1913) reported P. biconica from a bottom of “‘coarse sand”’ while P. affinis, which is a typical Cryp- todaphne species, was found on “‘blue mud.”’ The pres- ence of axial sculpture may simply represent an adapta- tion against predation by molluscivorous crabs on a sand bottom. Acamptodaphne should perhaps be retained for those Cryptodaphne species, including P. biconica, P. gradata, and C. semilirata, which possess a deep, narrow anal si- nus with the lower arm far extending forwardly. In C. pseudodrillia, as well as in C. kilburni, sp. nov., the sinus is moderately deep and broadly reversed L-shaped. However, the morphological limits of Acamptodaphne need to be redefined and in the absence of additional ma- terial we refrain from making a final judgement on the status of this taxon. Members of Cryptodaphne are characterized by spiral- ly keeled whorls and sinuous axial threads giving the shell a somewhat “‘tomopleurid”’ appearance. These fea- tures, as well as the microscopic granules reported in most species (but most probably normal for the genus), are usually encounted in the subfamily Clathurellinae H. & A. Adams, 1858. Among the “‘tomopleurid”’ group of genera, Cryptodaphne most resembles the widely distrib- uted Eocene to Recent Microdrillia Casey, 1903, but the diagonally decussate protoconch occurring in the former genus clearly indicates a different subfamilial allocation. The specimen of Pleurotomella sp. from the Pliocene of Tuscany figured by Chirli (1997:39—40, pl. 11, fig. 1— 2) is almost certainly a Cryptodaphne. Similarly, from the photograph of the holotype provided by Bouchet & Wa- rén (1980:37), it would seem that the Recent Pleurotoma anceyi Dautzenberg & Fischer, 1897, from the Azores closely resembles Cryptodaphne and may prove to be an Atlantic member of the genus. Cryptodaphne kilburni Morassi & Bonfitto, sp. nov. (Figures 1—7) Description: Shell small, thin but solid, biconic-clavi- form (b/l 0.33—0.39; a/] 0.41—0.46) with high spire and broadly excavated last whorl with relatively short neck. Teleoconch of up to 5.5 whorls separated by deep suture. Whorls with wide and shallowly concave sutural ramp occupying slightly more than half of whorl height; re- maining part of whorl gently convex and sculptured by Page 231 narrow cords. First teleoconch whorl with subsutural cord, weakly projecting peripheral keel well below mid- dle and one cord on either side of it. On subsequent whorls peripheral keel lying at one-third of whorl height and 1-2 cords occurring below it increasing to 3—4 on last whorl. Last whorl with 12-14 cords on base and neck, relatively uniform in strength and well spaced, lack- ing interstitial threads. In some specimens feeble spiral threads present on sutural ramp. Axial sculpture consist- ing of numerous sinuous collabral threads particularly ev- ident on sutural ramp. Under SEM (Figure 6), entire whorl surface seen to be covered by rows of granules rendering somewhat crisp axial threads on sutural ramp. Aperture narrowly pyriform, acute posteriorly; columella almost straight; parietal region weakly convex. Inner lip thinly callused; fasciole absent. Siphonal canal short, wide and shallowly notched. Outer lip rather thin, smooth within, lacking labral varix. Anal sinus (Figure 7) mod- erately deep, broadly reversed L-shaped, with its apex below middle of sutural ramp. Protoconch tall and narrowly conical of 3,25 decus- sately sculptured whorls. Color of teleoconch yellowish- white; protoconch reddish-brown. Dimensions: Holotype 6.5 X 2.3 mm, aperture height 2.7 mm; largest paratype: 6.7 X 2.2 mm, aperture height 2.7 mm; smallest paratype: 5.1 * 2 mm, aperture height 2.3 mm. Type locality: Gulf of Aden, station RS 92/1. No. Station Co-ordinates Depth specimens RS92/1 From 11°55'95”N—44°22'70"E 810 m 18 To 11°55'82"N—44°22'53"E 795 m Type material: Holotype and 17 paratypes off Gulf of Aden (RS92/1). Additional samples of the new species here not con- sidered paratypes occur in other lots of material dredged at the same station (RS92/1). Type repository: Holotype MZB 42988 and 16 paratypes MZB 42989; 1 paratype NM L6153/T1982. Etymology: Named after Richard Neil Kilburn (NM) in recognition of his continuous assistance to our work on Turridae s.1. Remarks: Judging from the available material, Crypto- daphne kilburni is the smallest Recent species in the ge- nus and is abundant at station RS92/1. Smaller paratypes have a more biconic shape, lower and slightly more prominent peripheral keel. Cryptodaphne kilburni super- ficially resembles C. affinis (Schepman, 1913) but the lat- ter species is larger (9 mm compared to 5.1—6.7 mm in length) and somewhat broader (b/l 0.41 compared to 0.33—0.39), has a wider sutural ramp and two prominent spiral keels just above lower suture. The new species has The Veliger, Vol. 48, No. 3 Figures 1-7. Holotype of Cryptodaphne kilburni, new species (MZB 42988). Figure 1. Apertural view; scale bar = 1 mm. Figures 2—3. Protoconch; scale bar = 100 pm. Figure 4. Protoconch; scale bar = 50 wm. Figure 5. Teleoconch whorl; scale bar = 500 pm. Figure 6. Microsculpture of teleoconch; scale bar = 100 wm. Figure 7. Anal sinus; scale bar = 500 pm. only one weakly projecting peripheral keel which is high- er on whorl height. Cryptodaphne kilburni is similar to Cryptodaphne rugosa Sysoev, 1997, but it is much small- er than the latter (5.1—6.7 mm compared to 7.7—12.8 mm in length) with different sculptural features. The new spe- cies has spiral elements, other than the peripheral keel, of relatively uniform strength and lacking secondary threads while C. rugosa has spiral cords of primary and secondary magnitude. Furthermore, in C. rugosa the col- umella is markedly twisted; in the new species it is almost straight. Actually, the closest morphological similarity of Cryptodaphne kilburni appears to be with the early Mio- cene Cryptodaphne pseudodrillia Powell, 1942. These two species have similar shape, the same dimensions (5.1-—6.7 mm and 6 mm in length respectively) and type of anal sinus. However, C. pseudodrillia has a strongly projecting peripheral keel at the lower fourth of the whorl (Powell, 1942) while in the Recent species the peripheral M. Morassi & A. Bonfitto, 2005 keel is weakly projecting, barely stronger than other cords, and lies at one-third of the whorl height. Crypto- daphne pseudodrillia has 1—2 interstitial threads between cords which are absent in the new species. Finally, the protoconch of C. kilburni has fewer whorls (3,25 com- pared to 5). Powell (1942) referred to the presence of “‘close- spaced spiral threads producing reticulation” covering the entire shell surface of C. pseudodrillia but this statement is presumably erroneous as the threads are axially, not spirally, oriented. Acknowledgments. John K. Tucker of the Illinois Natural His- tory Survey (USA) and Richard N. Kilburn (NM) provided help with literature. LITERATURE CITED Beu, A. G. & P. A. MAXwELL. 1990. Cenozoic Mollusca of New Zealand. New Zealand Geological Survey Paleontological Bulletin 58:1—518. BOucHET, P. & A. WAREN. 1980. Revision of the north-east At- Page 233 lantic bathyal and abyssal Turridae. Journal of Molluscan Studies 8(Suppl.):1—119. Cur, C. 1997. Malacofauna Pliocenica Toscana. Vol. 1. Su- perfamiglia Conoidea. C. Chirli: Firenze, Italy. xi + 129 pp. POWELL, A. W. B. 1942. The New Zealand Recent and fossil Mollusca of the family Turridae with general notes on turrid nomenclature and systematics. Bulletin of the Auckland In- stitute and Museum 2:1—188. PowELL, A. W. B. 1966. The molluscan families Speightiidae and Turridae an evaluation of the valid taxa, both Recent and fossil, with lists of characteristic species. Bulletin of the Aukland Institute and Museum 5:1—184. SCHEPMAN, M. M. 1913. The Prosobranchia of the Siboga Ex- pedition. Part 5. Toxoglossa. Resultats Siboga-Expeditie, Monograph 49-1, (5)365—452, pls. 25-30. SHuTo, T. 1971. Taxonomical notes on the turrids of the Siboga- Collection originally described by M. N. Schepman, 1913 (Part 3). Venus 30(1):5—22. SysoEv, A. V. 1996. Deep-sea conoidean gastropods collected by the John Murray Expedition, 1933-34. Bulletin of the Nat- ural History Museum of London, Zoology 62(1):1—30. SysoeEv, A. V. 1997. Mollusca Gastropoda: new deep-water turrid gastropods (Conoidea) from eastern Indonesia. Mémoires du Muséum National d’ Histoire Naturelle [Paris], série A, Zoo- logie, 172:325—355. i Instructions to Authors The Veliger publishes original papers on any aspect of malacology. All authors bear full responsibility for the accuracy and originality of their papers. Presentation Papers should include an abstract (approximately 5% of the length of the manuscript), Introduction, Materials and Methods, Results, and Discussion. Short notes should include a one- sentence abstract. 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ITHSONIAN INSTITUTION LI mene’ 3 90 595 N! 88 01256 0 Seasonality, Habitat Preference and Life History of some Willamette Velley Wet Prairie Ter- restrial Molluscs in Western Oregon, USA PAWIL Mi. SEVERINS heiiiccd pavsne ecto elieke ole cogs has saat c aa etree ae ot ES 220 Exallocorbula (Bivalvia: Corbulidae), a New Name for the Amazonian Molluscan Fossil Peba- sia Nutall ANDRE NEMESIO, AUDREY ARONOWSKY, AND LAURIE C. ANDERSON...........0--0005- 228 Cryptodaphne kilburni, a New Species of Bathyal Turrid (Gastropoda: Prosobranchia) from the Gulf of Aden (Northwestern Indian Ocean) MAURO MORASSI_AND ANTONIO: BONEITIO? sane cee Gos eee eee ee 230