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MALACOLOGIA, 1993, 35(2); 261-313

PHYLOGENETIC RELATIONSHIPS AND GENERIC REVIEW OF THE BITTIINAE (PROSOBRANCHIA: GERITHIOIDEA)

Richard S. Houbrick

Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A.

ABSTRACT The anatomy of seven members of the Bittium group is described, clarifying the status of the genus-level taxa comprising it. Bittium reticulatum, the type species of Bittium Gray, is described in depth, thereby establishing criteria for comparisons with other taxa of Bitliinae. The type species of Stylidium Dell and LirobiWum Bartsch, and representatives of Bittiolum Cossmann and Cacozeliana Strand are examined and compared with Bittium, s.s. Results of anatomical studies and a phylogenetic analysis using the Hennig86 and CLADOS programs, with Cehtt)ium as an outgroup, establish monophyly for Bitliinae Cossmann and reveal six different genus-level taxa. A new genus, ittibittium, from the Indo-Pacific, is proposed. Synonymies of each genus- level taxon and representative species examined are presented. Brief accounts of the ecology and zoogeography of each taxon are given. Two taxa formerly attributed to the 6/ff/um-group are herein excluded from it and referred to Cerithium Bruguière. These are Cerithium zebrum Kiener, 1841, and Cerithium boeticum Pease, 1861. The subfamily Bittiinae Cossmann, 1906, is thought to comprise nine genera (four of which were not included in phylogenetic analyses) : Bittium Gray, 1847; Bittiolum Cossmann, 1906; Ittibittium gen. n., Stylidium Dalí, 1907; Lirobit- tium Bartsch, 1911 ; Cacozeliana Strand, 1928; Argyropeza Melvill & Standen, 1901 ; Varicopeza Gründel, 1976; Zebittium Finlay, 1927. The genus Cassiella Gofas, 1987, of uncertain place- ment, is included as a possible member of the group. Key words: Bitliinae, Bittium, Cerithioidea, anatomy, taxonomy, phylogenetic analysis.

INTRODUCTION anatomical characters, most of the small- sized cerithioideans were left out of my anal- Shells of most small-sized cerithilds are no- ysis of cerithioidean phylogeny (Houbrick, tably difficult to classify, even to familial and 1988). generic levels. There has been much confu- The most recent revision of the Bittium sion and disagreement among malacologists group was published by Gründel (1976), who as to the limits and subdivisions of genus- based his taxonomy and phylogeny of the level taxa, because most genera have been group on sculptural characters of the proto- defined or based upon convergent shell fea- conch (embryonic spiral formation), ontoge- tures alone. Reflective of this unstable taxon- netic sculptural development of the teleo- omy, unreliable curatorial systems exist in conch, and overall shell form. Gründel (1976) most museums, where many lots of small- included many fossil and extinct taxa in his sized cerlthild taxa are randomly intermixed revision, but did not consider radular, opercu- with each other and with immature specimens lar, and anatomical characters of Recent of larger-shelled genera, such as Cerithium. taxa. Although he noted the similarities of Bit- These mixed lots frequently are assigned to tium and Cerithium Bruguière, 1789, he indi- the convenient "trash basket" category Bit- cated that Cerithium differs considerably from tium. Bittium in shell form, sculpture, aperture, and The genus Bittium Gray, 1847, sensu lato, especially in ontogenetic sculptural develop- comprises many poorly understood species ment. On the basis of the ontogeny of early placed in the family Cerithiidae Bruguière, spiral shell sculpture, Gründel (1976; 38) be- 1789. The concept of Bittium has been gen- lieved that genera in the Bittium group (Bit- erally broad, encompassing many other di- tium, Lirobittium, Bittiolum, Semibittlum) were verse genera, and opinions on the relation- descendents of the Jurassic genus Procerith- ships of the genus with other small-shelled ium Cossmann, 1902, of the family Procerithi- cerithiid groups have also been varied. For idae Cossmann, 1906. Indeed, he remarked these reasons and due to the lack of good that Bittium and Procerithium shared greater 261 262 HOUBRICK TABLE 1. Bittium-gmup genera and species used for anatomical studies (asterisk indicates type species). Genus Species Geographic Region Bittium 'reticulatum (DaCosta, 1778) Sao Miguel, Azores BiWum /mpendens (Hedley, 1899) Honolulu, Hawaii Bittiolum varium (Pfeiffer, 1840) Ft. Pierce, Florida BiWolum alternatum (Say, 1822) Provincetown, Massachusetts Ittibittium parcum (Gould, 1861) Honolulu, Hawaii Lirobittium subplanatum Bartsch, 1911 Palos Verdes, California Lirobittium attenuatum (Carpenter, 1864) Catalina Id., California Stylidium "esc/ir/c/ii//(Ivliddendorf, 1849) Carmel, California Cacozeliana 'granaría (Kiener, 1842) Albany, Western Australia

similarities in ontogenetlc sculptural develop- published: Dahlaida (Houbrick, 1978), Argyro- ment and overall shell morphology than did peza (Houbrick, 1980a), Varicopeza (Hou- BiWum and Cerithium. Grundel (1976; 40) brick, 1980b, 1987a), Glyptozaria (Houbrick, noted that the genera Argyropeza Melvill & 1981a), Alaba and Litiopa (Kosuge, 1964; Standen, 1901, and Varicopeza Grundel, Houbrick, 1987b; Luque et al., 1988), Courra 1976, usually placed near Bittium, were strik- (Houbrick, 1990a), Plesiotrochus (Houbrick, ingly similar In their ontogenetlc sculptural de- 1990b), and Diala (Ponder, 1991). Many of velopment and morphologies to species of these papers include anatomical data that the Jurassic genus Cryptaulax Täte, 1869 have helped partially to untangle the confus- (Procerithiidae), and stated that he consid- ing mixture of cehthlid genera of similar small- ered Argyropeza and Varicopeza to be shelled morphology. Recent members of Procerithiidae. Under The relationships of small-shelled species Procerithiidae, he assigned the Argyropeza- of the family Obtortlonidae Thiele, 1925, Cryptaulax group to the subfamily Cryptaul- which are very similar to those of members of axlnae Grundel, 1976, which he believed the BIttiinae, remain uncertain because ana- showed many of the "ancient characteristics" tomical characters are unknown. It is unclear of the family, and the Bittium-Procerithium If Obtortlonidae constitutes a separate family group to the subfamily Procerithlinae Coss- or should be included under BIttiinae. mann, 1902. Gründe! (1976) considered both subfamilies to have developed independently of one another and to have been separate MATERIALS AND METHODS since the Dogger (Middle Jurassic). Houbrick (1977) discussed the status of Bit- The goals of this study are threefold: first, to tium Gray, 1847, and included a historical re- examine the anatomy of Bittium reticulatum view, extensive synonymy, and a concholog- (DaCosta, 1778), the type species of the ge- ical redescription of the genus. This paper nus, thus setting the limits of the genus with a noted that most of the supraspecific taxa as- description of distinctive anatomical charac- sociated with the Bittium group are parochial ters; second, to study the anatomy of a num- In conception and scope, based on specific ber of other "Bittium" species, thereby estab- rather than generic characters, and convey lishing the validity or artificiality of other little or misleading phylogenetic information component groups or closely related higher about the group. In the Interest of pragmatism taxa; and third, to make a phylogenetic anal- and taxonomic parsimony, it was suggested ysis of the group based on a morphological that many of the generic and subgeneric data set that Includes more than shell char- names be abandoned or synonymized with acters. Bittium, sensu lato, until the entire group was This revision is based primarily on collec- properly evaluated on the basis of more than tions of preserved material in the USNM and shell characters. on living material studied In the field. Fossils Since Gründers (1976) work and my paper representing extinct genera and species were on Bittium (Houbrick, 1977), studies on a not considered, although a brief survey of ex- number of Bittium-l'\ke genera and other tinct forms and their possible relationships to small-shelled cerlthioldean taxa have been living members of the Bittium-group is in- GENERIC REVIEW OF BITTIINAE 263 eluded. The great number of species and robittium, and a new genus described herein, higher category groups traditionally included are less detailed and emphasize the anatom- under Bitiium, sensu lato, and the difficulties ical differences from Bittium reticulatum. of obtaining good anatomical material pre- The anatomy of the genera Argyropeza and cluded an exhaustive, comprehensive ana- Varicopeza is only superficially understood. tomical study of all members the group. In- Anatomical knowledge about Zebittium Fin- stead, I decided to look at representative taxa lay, 1927, and Cassiella remains unknown, of genera assigned to the S;Wum-group com- because I was unable to obtain preserved prising species having diverse shells from material of species representing them; conse- widely different geographic regions. A total of quently, only the shells are considered in this seven ß/fi/um-group species representing five review. higher taxa (genera) from different localities were examined by dissecting live-collected Phylogenetic Analysis material and by studying living populations in situ, where possible. These species are listed The guiding principles of this study are those below in Table 1 and include the type species of phylogenetic systematics (Hennig, 1966; of Bittium Gray, 1847, Stylidium Dall, 1907, Wiley, 1981). The Hennig86 computer pack- and Cacozeiiana Strand, 1928, and represen- age, version 1.5, ie and bb options (copyright tative species of Bittioium Cossmann, 1906, James S. Farris, 1988) and CLADOS, version Lirobittium Bartsch, 1911, and a new genus, 1.2 program (copyright Kevin C. Nixon, 1988, described herein. Two other species, each 1991, 1992) were used to analyse data and having a distinctive shell morphology, and construct trees. considered as putative genera formerly attrib- Phylogenetic analysis of six genus-group uted to "Bitiium," S.I., were also studied in the taxa of the Bittiinae (Bittium, Ittibittium, Bitti- field: "Bittium" zebrum (Kiener, 1841) from oium, Lirobittium, Stylidium, and Cacozeii- Pago Bay, Guam, and Enewetak Atoll, Mar- ana) was undertaken using 21 morphological shall Islands; and "Bittium" boeticum (Pease, characters comprising 51 character states de- 1861), from Honolulu, Hawaii. When the soft rived from the shell, operculum, radula, and parts of these two species were examined, soft anatomy of the taxa listed in Table 1. Ini- they were found to lack an epipodial skirt, and tially, there were 30 characters, but these the ciliated ridge tract and spermatophore were reduced to 21. Seven of the 21 charac- bursa in the lateral lamina of the palliai ovi- ters were multi-state characters. Autapomor- duct, characters distinctive of members of the phies defining terminal branches, which were Bittium-gmup. Therefore, both species were not part of multistate series, were not included excluded from the ß;Yf/um-group and as- in the analysis, but were retained for the di- signed to Cerithium Bruguière. Due to the cur- agnosis of each genus-group taxon. Multi- rent alpha-level taxonomic disarray of the Bit- state characters were unordered. f/um-group, I have attempted to present a comprehensive, annotated synonymy and Genus-Group Taxa Analysed have illustrated the shells of the species stud- ied in this review. I hope that this will give Six genus-group taxa were included: Caco- other workers an unequivocal idea about the zeiiana, Lirobittium, Stylidium, Bittium, Ittibit- species and genera they represent. tium, and Bittioium (Table 1). The phyloge- All specimens were dissected under water netic analysis excluded poorly known genera in wax-filled petri dishes using a Wild M-5 dis- that have been assigned without justification secting microscope. Méthylène blue was to Bittiinae, such as Zebittium and Cassiella. used to enhance anatomical features during Although the shell morphologies, opercular dissection. Sections were made at 5 tJ.m and and radular characters of Argyropeza and stained with Hematoxolin and Eosin. Photo- Varicopeza have been well studied (Houbrick, micrography was done using a Zeiss Photo- 1980a, 1980b), these genera also were left microscope III. out of the analysis because of lack of anatom- The emphasis of this study is on the anat- ical data. omy of Bittium reticulatum, the type species of Bittium, s.S., which is the criterion against Outgroup Selection which other Bittium-group genera are de- scribed and compared in this paper. Descrip- The genus Cerithium Bruguière, family Cer- tions of Bittioium, Cacozeiiana, Stylidium, Li- ithiidae Férussac, 1819, was selected as the 264 HOUBRICK TABLE 2. Comparison of dentition of radular teeth among genera (C central or main cusp; numbers signify no. of denticles). Taxon Rachidian Lateral Inner Ivlarginal Outer Marginal Bittium 2-3+C+2-3 1+C+3-6 3-4+C+4 3-4+C+O Bittiolum 3 + C + 3 2+C+3-4 3-4+C+2-3 6 + C + O IttibiWum 2 + C + 2 1+C+3-4 2 + C + 3 5 + C + O LirobiWum 6 + C + 6 6 + C + 15-17 15-19 + C + 5-6 15-19 + C + O Stylidium 2 + C + 2 1+C+3-4 4-5+C+3 4 + C + O Cacozeliana 2 + C + 2 1+C + 3-4 5-6+C+3-4 4 + C + O Argyropeza 2-3+C+2-3 1+C+5-6 5-6+C+4-5 5-6+C+O Varicopeza 3-4+C+3-4 1+C+5-6 3-4 + C + 3 3 + C + O

outgroup to root the trees generated by the among cerithioideans. Ponder's (1991) phy- analyses. The BiWum-group traditionally has logenetic analysis showed that dialids were been considered as a subfamily (Bittiinae) of closely related to litiopids and far removed Cerithiidae by various authors (see below, for from Cerithiidae (Ponder, 1991; 514). Diaia history). Cerithium, subfamily Cerithiinae, is was therefore rejected as an outgroup. the most appropriate group to use for out- group comparison, because it is the closest Characters sister group that is well known anatomically. The anatomy of Cerithium species has been The characters listed below comprise three described by Houbrick (1971, 1978, 1992) categories; shell characters (1-5), anatomical and is very similar to that of Bittiinae mem- characters (6-19), reproductive characters bers. However, Cerithium species have more (20-21). Radular characters were eliminated generalized and less complex external fea- from the final analysis because of their au- tures. Several external anatomical features of tapomorphic condition. Nevertheless, radular members of the S/tt/um-group, such as a characters are important diagnostic charac- metapodial mucus gland, and the epipodial ters of genera and are summarized in Table 2. skirt and associated papillae, are lacking in Because the polarities of multistate charac- Cerithium. The anatomy of such small-sized ters were largely speculative, all character snails as Bittium may be highly derived and/or states were left unordered; i.e., the integer modified due to their reduction in size. Cerith- assignment was arbitrary. The coding of ium species are generally much larger ani- these characters and their states are pre- mals than "Bittium" species, but a number of sented in Table 3. An annotated list of the species are very small and often are confused morphological characters and character with "Bittium" species. states used in the phylogenetic analysis is Among small-shelled cerithioideans, Litiopa presented below; and Alaba, family Litiopidae, were considered as possible outgroup candidates. These small Shell Characters: 1. Shell sculpture•0 = snails have external features, such as an spiral; 1 = cancellate. Most members of the epipodial skirt and epipodial tentacles, similar subfamily are characterized by a markedly to those seen among members of the Bittii- cancellate shell sculpture, in contrast to Cer- nae, and are well known anatomically; how- ithium species where spiral elements domi- ever, they differ from bittiid species in internal nate sculptural patterns (Houbrick, 1992). Ex- anatomy (Kosuge, 1964; Houbrick, 1987b; ceptions are species of the genera Stylidium Luque et al., 1988). Phylogenetically, Litiop- and Ittibittium, where spiral sculpture domi- idae is far removed from the family Cerithiidae nates and axial ribs are either lacking or (Houbrick, 1988; 114), and is therefore re- poorly developed. jected as a suitable outgroup. 2. Anal canal•0 = well developed; 1 = Another group of small-shelled species, the weakly developed or missing. A well-devel- Dialidae, was also considered as a possible oped anal canal is present in Cerithium mem- outgroup. However, only one species is bers (the outgroup), but occurs only in two known anatomically (Ponder, 1991), and genera of the S/fi/um-group, Cacozeliana and Healy (1986) has shown that the parasperma- Varicopeza, and is exceptionally well devel- tozoa of Diala are unique and highly derived oped in the latter genus (Houbrick, 1980b). GENERIC REVIEW OF BITTIINAE 265

TABLE 3. Data matrix derived from morphological characters of species representing six genus-group taxa of Bittiinae. Cerithium is the outgroup.

Character Taxon 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 Outgroup 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 Bittium 1 1 0 1 0 1 1 0 0 0 2 0 1 2 1 1 1 1 1 0 0 IttibiWum 0 1 0 1 0 0 1 0 0 1 2 1 0 1 1 0 0 0 1 1 1 Stylidium 0 1 1 1 2 0 0 1 1 0 1 0 2 1 1 1 1 1 1 0 1 Cacozeliana 1 0 1 0 2 2 2 0 0 0 1 0 0 0 0 1 1 1 0 0 Ü BiWolum 1 1 0 1 1 0 1 0 0 1 3 0 1 1 1 0 1 1 1 0 0 LirobiWum 1 1 1 1 0 1 0 1 1 0 1 1 2 1 1 0 1 1 1 2 1

3. Varices•0 = present; 1 = absent. Va- 8. Cephalic tentacle length•0 = elongate; rices, thickened, former growth lines, are a 1 = short. Among cerithiids and the Bittiinae, common feature of most cerithiids and occur cephalic tentacles are usually elongate and among members of Bittiinae with the excep- much longer than the snout, but in the eastern tion of Lirobittium and Stylidium. Pacific genera Lirobittium and Stylidium, the 4. Anterior canal•0 = well developed; 1 tentacles are much shorter than the length of = weakly developed. The anterior siphonal the snout. canal is a strong feature on most cerithiids, 9. Eye size•0 = normal; 1 = small; 2 = but in smaller-shelled taxa frequently is poorly large. Most cerithiids have eyes of normal size, developed (most Bittiinae) or absent (Cass- but in such deep-water species as Argyropeza iella, Cerithidium). and Varicopeza, the eyes are very large, pos- 5. Protoconch sculpture•0 = two spiral sibly an adaptation to water depth and poor lirae; 1 = one spiral lira; 2 = entirely smooth. light. In contrast, the eyes of Styliodium and Most outgroup species have strong spiral Lirobittium species are exceptionally small. sculptural elements on their protoconchs 10. Metapodial mucus gland•0 = absent; (Houbrick, 1992). Bittiinae genera range from 1 = present. Although this structure is absent species with spiral sculpture to those having in the outgroup, it does occur among a few only one weak spiral lira or no sculpture, but other cerithioidean groups (Litiopidae [Alaba, this is probably reflective of the type of devel- Litiopa], Cerithiidae [Colina]; Houbrick, opment. 1987b, 1990a, respectively). This gland may be an adaptation to an algal and/or high en- Anatomical Characters: 6. Opercular mor- ergy habitats. Species having a metapodial phology•0 = ovate shape; 1 = round, cir- gland are known to use the mucus thread se- cular shape; 2 = round shape with fringed creted by the gland to anchor themselves spiral edges. Cerithium species have oper- while they climb about the algal fronds cula with an ovate shape (Houbrick, 1992), (Houbrick, 1987b, 1990a), and it is thought herein that the more circular 11. Epipodial skirt•0 = rudimentary; 1 = shape observed among several S/ff/um-group well developed, smooth; 2 = well developed, taxa are modifications due to size reduction, papillate along edges; 3 = well developed, although this is not always the case (excep- scalloped. Cerithium species have a weak tions in Ittibittium and Bittiolum, both small operculigerous lobe on the rear of the foot, shelled genera). The spirally fringed condition which is here Interpreted as a rudimentary seen in Cacozeliana departs from the norm posterior epipodial skirt. In Bittiinae species, and is probably derived. the skirt extends forward along the sides of 7. Snout shape•0 = wide; 1 = narrow, the foot to form a fully developed epipodial elongate; 2 = short, narrow. This character is skirt. An epipodial skirt occurs also among a variable feature among cerithiids. Cerithium small-shelled members of the Litiopidae (Ko- species usually have large, wide, muscular suge, 1964; Houbrick, 1987b; Luque et al., snouts (Houbrick, 1992), whereas they tend 1988) and the Dialidae (Ponder, 1991). Al- to be narrow and elongate in members of the though this character is homoplastic among Bittiinae, especially among taxa of the Bittium cerithioideans, an epipodial skirt is character- clade {Bittium, s.S., Ittibittium, Bittiolum). istic of Bittiinae. 266 HOUBRICK TABLE 4. Comparison of developmental features among Bittiinae genera and species.

Max. Shell Protoconch Developmental Taxon Length Sculpture Type Egg Size Bittium reticulatum 15 mm 2 spirals planktonic 0.1 mm IttibiWum parcum 6 mm 2 spirals direct 0.2 mm Bittiolum varium 7 mm 1 spiral planktonic 0.1 mm Lirobittium subplanatum 10 mm 2 spirals direct 0.5 mm Stylidium eschhchtii 17.5 mm smooth direct 0.2 mm Cacozeliana granaría 24 mm smooth planktonic 0,1 mm Argyropeza divina 7.6 mm 2 spirals planktonic ? Varicopeza vahcopeza 10 mm 1 spiral planktonic ?

12. Ovipositor•0 = present; 1 = absent. bittium, the openings have a wide separation, This gland, although common among cerithio- whereas in Bittium they are not as far apart. In ideans, is absent in some taxa, such as those other bittiids and in most other cerithiids, the having internal brooding (Houbrick, 1987c). openings are close together. The absence of an ovipositor in females may 17. Spermatophore bursa location•0 = be falsely scored, as it is thought that its pres- located in medial lamina; 1 = located in lat- ence can be easily ascertained only during eral lamina. The spermatophore bursa is breeding season; moreover, this gland is also found in the lateral lamina in most members difficult to detect in some preserved speci- of the Bittium-group, but in Ittibittium and in all mens. Among Bittiinae, the ovipositor is ab- other known cerithiids, it occurs in the medial sent only in Ittibittium and Lirobittium. lamina (Houbrick, 1988). 13. Osphradial morphology•0 = bipecti- 18. Ciliated ridge tract•0 = absent; 1 = nate; 1 = monopectinate; 2 = vermiform. present. This structure, one of the synapo- This character varies greatly among Bittiinae morphies defining Bittiinae, is lacking in Ittibit- genera. Although the osphradium in Cerith- tium members and in most other cerithiids. ium species is bipectinate, it is vermiform 19. Seminal receptacle with grape-like mor- among most other cerithioidean families, phology•0 = present; 1 = absent. This such as the estuarine Potamldidae and fresh- grape-like configuration may not represent a water families Thiaridae and Pachychilidae distinct morphology, but may be due to the (Houbrick, 1988, 1991). highly filled condition of the receptacle. This 14. Osphradial length•0 = equal to condition occurs only in Cacozeliana. ctenidial length; 1 = a little less than ctenidial length; 2 = one-half the ctenidial length. This Reproductive Characters: 20. Spawn mor- is a highly variable character, but often diag- phology•0 = formed into gelatinous string nostic of some taxa. No overlap among char- wound into mass; 1 = short gelatinous tube; acter states was detected in the species stud- 2 = balloon-like cluster. A gelatinous string ied. mass is the common spawn morphology seen 15. Zygoneurous nervous system•0 = among cerithioidean taxa and within Bittiinae. absent; 1 = present. Bouvier (1887) docu- The balloon-like cluster of eggs in members mented a zygoneurous condition among of Lirobittium is unique, whereas a short ge- some cerithiids, and this was summarized by latinous tube morphology is seen only in It- Houbrick (1988). Zygoneury is absent in Cer- tibittium: both taxa have few, large eggs and ithium, and in all Bittiinae except for Bittiolum. undergo direct development (Table 4). 16. Common opening to sperm pouch and 21. Type of development•0 = planktonic; seminal receptacle openings•0 = close to- 1 = lecithotrophic (demersal/direct). Most gether; 1 = far apart. In Stylidium and Liro- members of the outgroup have a planktonic GENERIC REVIEW OF BITHINAE 267

Cacozeliana Lirobittium Stylidium Bittium Ittibittium Bittiolum

FIG. 1. Cladogram showing relationships among six genera of Bittiinae, using Cerithium as the outgroup (L = 41 ; C! = 70; Rl = 53; trees two. Numbers to left of black bars indicate characters: those to right of bars represent character states. Only characters with a CI of 100 are shown). larval phase In their development. It is thought strongly support the recognition of six genus- that planktotrophy can evolve to lecilhotrophy level taxa. The monophyly of Bittiinae is es- but not vice-versa (Strathmann, 1978). Direct tablished by three synapomorphies (11[1], developers have larger, fewer eggs per 18[1], 20[0]) and one homoplastic character spawn mass (Table 4). (17[1]). The layout of the palliai oviduct, dis- cussed in greater detail below, is the source of two good synapomorphous characters: a RESULTS ciliated ridge tract and a spermatophore bursa in the medial lamina. An epipodial skirt, Phylogenetic analysis resulted in two while distinctive of the Bittium-group, is plesi- equally parsimonious trees, each with a omorphic, because it occurs also in other cer- length of 41 steps, a consistency index of 70, ithioidean groups. and a retention index of 53 (Fig. 1). The num- Cacozeliana stands apart at the base of the ber of steps and the consistency indices of cladogram from the other taxa and is closest each character used in the construction of the to Cerithium, the outgroup. Cacozeliana is de- cladogram are shown in Table 5. The support- fined by two autapomorphous characters ing branches of both cladograms had identi- (6[2], 7[2]) and by two homoplastic characters cal tree topologies except for the clade sup- (5[2], 16[1]). Cacozeliana is well separated porting Bittium, Ittibittium, and Bittiolum. In the from all other genera of Bittiinae higher on the first tree, illustrated herein (Fig. 1), Ittibittium tree by five synapomorphies (2[1], 4[1], 14[1], and Bittiolum are sister groups of Bittium, 15[1], 19[1]) and with one homoplastic char- while in the second tree, Bittium and Bittiolum acter (13[1]). are sister groups of Ittibittium. Both analyses The Lirobittium-Stylidium clade, which is 268 HOUBRICK

TABLE 5. List of steps and consistency indices of characters used in construction of cladogram.

Character 1 2 3 4 5 6 7 8 9 10 11 Steps 3 1 2 1 3 3 2 1 1 1 3 C.I. 33 100 50 100 66 66 100 100 100 100 100 Character 12 13 14 15 16 17 18 19 20 21 Steps 2 3 2 1 3 2 2 1 2 2 C.I. 50 66 100 100 33 50 50 100 100 50 geographically confined to the west coast of characters. If the cladogram shown in Figure North America, is supported by two synapo- 1 is interpreted strictly, Ittibittium and Bittiolum morphies (8[1], 9[1]), and two homoplastic may be regarded as subgenera of Bittium; characters (13[2], 21 [1]) In this clade, Stylid- however, because this is a preliminary revi- ium is poorly defined by three homoplastic sion of the Bittium-group, based on only a few characters (1[0], 5[2], 16[1]), whereas Lirobit- representatives of each genus, and not in- tium is better founded on one autapomorphy cluding other poorly known taxa, it is best not (20[2]) and three homoplastic characters to assign differential rank to genus-group taxa (6[1], 12[1], 16[0]). at this stage. Therefore, I have decided to The Bittium clade is supported by one sy- treat all terminal nomina as full genera. napomorphy (7[1]) and two homoplastic char- As noted in an earlier paper (Houbrick, acters (3[0], 13[1]). Bittium, s.S., is defined by 1977), other genus-level taxa have been pro- one autapomorphy (14[2]) and three ho- posed under the Bittium-group or are thought moplastic characters (2[0], 12[1], 18[1]). It- to be linked closely to it. Many of these taxa tibittium and Bitiiolum, the sister taxa to Bit- are synonyms of Bittium-gmup genera de- tium, are separated from it by one scribed herein or have been proposed for fos- synapomorphy 10[1]). Bittiolum is supported sils. The subfamily Bittiinae, as understood in by two autapomorphies (5[1], 11 [3]) and two this paper, is thought herein to comprise nine, homoplastic characters (11 [3], 16[0]). A sin- possibly ten, Recent genus-group taxa: Bit- gle autapomorphy (20[1]) and six homoplastic tium Gray, 1847; Bittiolum Cossmann, 1906; characters (1[0], 12[1], 13[0], 16[0], 17[0], Ittibittium gen. n.; Stylidium Dall, 1907; Liro- 18[0], 21 [1]) define Ittibittium. The characters bittium Bartsch, 1911; Cacozeliana Strand, listed above are those derived only from the 1928; Argyropeza Melvill & Standen, 1901; data matrix (Table 3) used in the construction and Varicopeza Gründel, 1976. The genera of the cladogram (Fig. 1). Other autapomor- Zeblttium Finlay, 1927, and Casslella Gofas, phies defining terminal branches but not pari 1987, are provisionally referred to Bittiinae of multiState series were not included in the until more information is available. data matrix. These characters are given un- Argyropeza and Varicopeza have been der the diagnosis of each genus in the sys- treated previously by Houbrick (1980a, tematic portion of this paper. 1980b, 1987a), but their anatomy remains poorly known and they are not described in great detail here. An epipodial skirt has been DISCUSSION recorded in Varicopeza crystallina (Houbrick, 1987a: 80), but due to poorly preserved ana- The phylogenetic analysis of morphological tomical material, this structure could not be characters of the species in Table 1 resulted ascertained in Argyropeza species; however, in recognition of six different morphological the radula of Argyropeza species (Houbrick, groups (Fig. 1), which are herein interpreted 1980a) is similar to those of members of the as genus-group taxa under the stJbfamily Bit- BIttlum-gmup. tiinae Cossmann, 1906. Generic names al- Anatomical knowledge about potential Blt- ready exist for five of these groups: Bittium tlum-group species as yet unstudied, such as Gray, 1847; Bittiolum Cossmann, 1906; Ca- Casslella from the eastern Atlantic, Zeblttium cozeliana Strand, 1928; Stylidium Dall, 1907; from New Zealand, and the many species of and Lirobittium Bartsch, 1911. A new genus, small-shelled, Bittium-Wke cerithioideans from from the Indo-Pacific, is described herein. All the Indo-Pacific, may reveal even more new of the above genera, with the exception of genus-level taxa to be included under Bittii- Stylidium, are defined by autapomorphous nae. GENERIC REVIEW OF BITTIINAE 269 SYSTEMATIC TREATMENT OF BITTIINAE taxa Cosmocerithium Cossmann, 1906, In- fracerithium Gründel, 1974, and Rhabdocol- The species studied have been placed into pus Cossmann, 1906, were treated as sub- groups (genera) according to the above phy- genera of Procerithium. Gründel (1976) also logenetic analysis. The type- or representative included Argyropeza Melvill & Standen, 1901, species of each genus is described, and notes Varicopeza Gründel, 1976, and the extinct on reproductive biology and ecology are in- genus Cryptaulax Gründel, 1976, with sub- cluded, when possible. Shell-length measure- genera Pseudocerithium Cossmann, 1884, ments for each species represent the largest and Xystrella Cossmann, 1906, in the Bittium specimen obsen/ed. Representatives of other group under the subfamily Cryptaulaxinae genera for which anatomical material was Gründel, 1976. Excluding the Jurassic taxa, lacking are described from shell morphology the Recent genera Argyropeza and Varico- and radular morphology, if available. peza should probably be included in the Bit- tiinae, because the few morphological and BITTIINAE COSSMANN, 1906 anatomical characters known about these taxa strongly suggest affinity to this subfamily. Bittiinae Cossmann, 1906: 61. The other extinct genus-group taxa and Pro- Procerithiinae Cossmann, 1906, sensu Grün- cerithium should be excluded from Bittiinae, del, 1976 (in part). because the evidence supporting a relation- ship of these taxa with the S/rt/um-group is Diagnosis based solely on the ontogenesis of spiral sculpture as seen on the early shell spire, a Shell small, turreted, narrowly elongate to character which is, at best, tenuous: more pupate, with moderate spiral and axial sculp- characters are needed to lend credence for ture frequently cancellate and/or beaded. Ap- such a relationship. While Gründel's (1976) erture with short but distinct anterior canal. hypothesis poses interesting questions, it is Spiral sculpture usually 4-5 spiral cords per founded mostly on shell sculpture, which is whorl. Animal with epipodial skirt, opercular taxonomically informative but potentially phy- lobe, and palliai oviducts comprising large logenetically misleading. Considering the Ju- sperm bursa and seminal receptacle in pos- rassic age of the Procerithium group and the terior part of medial lamina, and spermato- great likelihood of homoplasy in shell mor- phore bursa and ciliated ridge tract in poste- phology, the belief that the Bittium- and Pro- rior lateral lamina. Ciliated gutter leading from cerithium- groups are of the same lineage is oviduct down right side of foot in females. largely speculative, cannot be falsified, and Glandular ovipositor at base of right side of should not be accepted as evidence for a phy- foot in most species. Nervous system dialy- togeny (Houbrick, 1988). neurous. Spawn consisting of gelatinous, The name Elassum Woodring, Bramlette & winding strings. Kew, 1946, has been traditionally associated with the Bittium-group in the literature, and Taxonomic Remarks was proposed by Woodring et al. (1946; 68) for Pleistocene and Recent material from The S/fi/um-group (Bittiinae Cossmann, southern California previously named Bittium 1906) has been placed under Cerithiidae by californicum Dall & Bartsch, 1901, and origi- nearly all authors (Cossmann, 1906; Thiele, nally assigned to the subgenus Elachista Dall 1929; Wenz, 1938; Golikov & Starabogatov, & Bartsch, 1901. Bittium californicum is the 1975; Ponder & Waren, 1988), except Grün- type species of Elachista by monotypy. How- del (1976), who assigned the group to the Ju- ever, as Elachista is preoccupied, a new rassic family Procerithiidae Cossmann, 1906 name, Alabina Dall, 1902, was proposed to (erroneously cited by Cossmann as 1905). He replace it. Woodring et al. (1946) did not be- allocated 12 genus-group taxa to the subfam- lieve the taxon californicum Dall & Bartsch, ily Procerithiinae (= Bittiinae). Of these, Bit- 1901, was an Alabina and thus proposed tlum, Bittiolum, SemibiWum and Procerithium Elassum to accomodate it, noting that the were treated as full genera; Cerithidium Mon- species was more Bittium-Wke than Alabina- terosato, 1884, ñasb/Wum Gründe!, 1976, Li- like. Because Elachista, Elassum, and Alab- robittium Bartsch, 1911, Cacozeliana Strand, ina have the same type species, Elassum be- 1928, and Stylidium Dall, 1907, were consid- comes a junior synonym of Alabina. The shell ered to be subgenera of Bitiium. The extinct of the type species somewhat resembles 270 HOUBRICK those of members of the Bitlium-group, and I nus, Cerithlum submamillatum Rayneval & concur with Woodring et al. (1946) that it pos- Ponzi, 1854, which he considered a synonym sibly should be included as a component ge- of Turritelia pusllla Jeffreys, 1860. As Gofas nus of the Bittium-group; however, as there is (1987: 110) remarked, the former name was no preserved material of living animals of this originally given to a Pleistocene fossil which is taxon to confirm this supposition, Alabina [ = not conspecific with the Recent species. Go- Elassium] is not further treated herein. fas (1987) remarked that the designation of Houbrick (1977: 103) initially placed 13 Cerithlum submamillatum as the type species nomina into the synonymy of Bittium, sensu of Cerithidium by Cossmann (1906) should lato. Subsequent studies on the BiWum-group prevail over that of Turritelia pusllla by Wenz and evidence derived from anatomical char- (1940). I agree with Gofas (1987: 109-110) acters presented herein now allow exclusion that both species are congeneric and have of six genera originally included in that syn- sculpture similar to Bittium reticulatum; how- onymy and a more focused diagnosis of Bit- ever, in a Cerithidium species examined by tium, s.S. An annotated list of taxa previously Ponder (Ponder, in litt.), the female palliai ovi- included in the Bittium-group, but now ex- duct was closed, which is very different from cluded, is presented below (Jurassic genera the open systems known in all other members not included): of Bittiinae. A closed palliai oviduct has not 1. Bittinella Dall, 1924 (type species: Bittium yet been demonstrated in the type species of hiloense Pilsbry & Vanatta, 1908). The type Cerithidium, but on the basis of the closed species of this genus is a rissoid of the genus system noted by Ponder, Cerithidium is ex- /sse//e//a Weinkauff, 1881, subfamily Rissoin- cluded provisionally from Bittiinae. inae (Ponder, 1985: 95; Kay, 1979: 80). Bit- 5. Dahiakia Biggs, 1971 (type species: tium parcum Gould, 1861, has been errone- Dahiakia lellae Biggs, 1971). The type spe- ously assigned to Bittineila (see below). cies is a junior synonym of Cerithlum proteum 2. Bittiscalia Finlay & Manwick, 1937 (type Jousseaume, 1930 (Houbrick, 1978), and I species: Bittium simplex Marshall, 1917). It is believe both names are probable synonyms unclear to which group this extinct species of Cerithlum scabridum Philippi, 1848. should be assigned. Although Finlay & Mar- 6. EubiWum Cotton, 1937 (type species: wick (1937: 44) placed it under Cerithiidae, Bittium lawleyanum Crosse, 1863) [not Eublt- they noted its similarity to Zeacumantus Fin- tium Cossmann, 1902). The syntypes of the lay, a batillariid (Houbrick, pers. obser.). Their type species of this genus (MNHN, Paris) are drawing of the type species (Finlay & Marwick, Batlllariella estuarlna (Täte, 1893), which is a 1937: pi. 5, fig. 20) shows a shell with an an- batillariid (family Batillariidae), and not closely terior canal that is a wide shallow notch, similar related to Cerithiidae. In any case, the name to poorly developed anterior canals seen in Eubittium Cotton is a secondary homonym. some Bittium and Alabina species. Because 7. Paracerlthium Cotton, 1932 (type spe- this is a fossil, we may never know with cer- cies: Bittium lawleyanum Crosse, 1863) [not tainty the correct family assignment. Although Paracerlthium Cossmann, 1902]. This taxon the authors placed it under Cerithiidae, they is a secondary homonym and has the same were obviously equivocal about this assign- type species as the previous taxon, which is a ment. It is best to leave Bittiscalia under the batillariid. broader category of Cerithiidae and to exclude 8. Sundablttium Shuto, 1978 (type species: it from the more narrow assignment of Bittinae. Cerithlum fritschi Boettger, 1883). It is highly 3. Brachyblttium Weisbord, 1962 (type spe- unlikely that this fossil genus is related to the cies: Bittium [Brachyblttium) caraboboense Bittium group. Shuto himself (1978: 152) was Weisbord, 1962). The type species, a fossil, equivocal in assigning it to Bittium. The fig- appears to be an immature or fragmentary ures of C. fr/iscA)/depicted by Martin (1914: pi. Cerithlum species, judging from its illustration 5, figs. 132-134) suggest an Abyssochrysos (Weisbord, 1962: pi. 15, figs. 5-6). species, but this assignment needs confirma- 4. Cerithidium Monterosato, 1884 (type tion by examination of the type material. species: Cerithlum submamillatum Rayneval & Ponzi, in Rayneval etal., 1854). Cerithidium Discussion was introduced by Monterosato (1884) who noted that it was characterized by a rounded The subfamily Bittiinae is characterized by aperture and lack of an anterior canal. Mon- small-shelled species generally having can- terosato listed a single species under the ge- cellate sculpture and short canals. Monophyly GENERIC REVIEW OF BITTIINAE 271 for Bittiinae is tentatively established by the mucus gland, but differ considerably from synapomorphous layout of the palliai oviduct members of the Bittium-group in other ana- (see description under Bittium reticulatum; tomical characters (Houbrick, 1990b: 247- Fig. 6C); i.e., the presence of three sperm 248), and are an unusual family. chambers: a large bursa (1), and smaller The relationship of the ß/fi/um-group to seminal receptacle (2) in the posterior half of other small-shelled cerlthioidean genera such the medial lamina, and a spermatophore as Scaliola A. Adams, 1860, and Finella A. bursa (3) in the posterior lateral lamina. The Adams, 1860, remains unclear because the position of the spermatophore bursa in the anatomy of these taxa is still unknown. Ponder lateral lamina appears to be a unique synapo- (1991) recently described the anatomy of a morphy defining Bittiinae, but this needs to be species of Diala A. Adams, 1861, which re- confirmed by observation of spermatophores sulted in his recognition of a separate family, in the bursa in other members of the subfam- Dialidae Ludbrook, 1941. According to Ponder ily. This character does not occur in Ittibittium, (1991: 504-506), Diala species have a weak a new genus described herein; thus, it had a epipodial fold (epipodial skirt), a pair of lateral CI of 50 in the analysis. The ciliated ridge tract opercular lobes, and a posterior opercular (Fig. 6B, C, ctr) on the lateral lamina epithe- flap, which appear to be homologous with the lium leading into the spermatophore bursa is epipodial skirt and opercular lobe described in also a synapomorphy defining Bittiinae. This the Bittiinae members above. However, unlike is an uncommon feature among cerithioide- the situation in Bittiinae, Diala species lack the ans, and is unusually long. Some plesiomor- metapodial mucus gland and the glandular phic characters, such as the well-developed ovipositor on the right side of the foot in fe- epipodial skirt and epipodial tentacles, occur males. Additionally in Diala species, the lateral in other cerlthioidean groups, but in combina- lamina of the palliai oviduct does not have a tion with the above synapomorphous fea- sperm pouch and the paraspermatozoa are tures, are characteristic of the Bittiinae. Ittibit- unique among Cerithioidea (Healy, 1986). tium, new genus, deviates from other The rachidian radular tooth of most mem- members of the subfamily in having the albu- bers of the S/ff/um-group is characterized by men gland protrude beyond the posterior being wider than tall and usually has a basal mantle cavity into the visceral coil. In other plate with concave sides. This differs from the respects, it generally agrees with the remain- hour-glass shape of the rachidian tooth found ing genera of the Bittiinae. in small-sized species of Diala, Litiopa, Alaba, The Recent genera treated herein are each and Varicopeza (Ponder, 1991: fig. 3F, G; characterized by external anatomical charac- Houbrick, 1987a: figs. 14, 19; 1987b: figs. 9, ters (Fig. 2), which allow easy classification of 10), taxa frequently confused with Bittium- living animals. Two genera of the subfamily group members. For dental cusp patterns (Bittlolum and Ittibittium, gen. n., have a large among Bittiinae taxa, see Table 2. metapodial mucus gland marked by an elon- Although members of Bittiinae are primarily gate slit in the middle of the sole (Fig. 2), lead- grazers of epiphytic microalgae, many species ing deep into the center of the foot. While the appear to feed on particulate matter gathered epipodial skirt and opercular lobe are charac- by cilia and mucus on the antehor ctenidial teristic of Bittiinae, these characters and the filaments when the animal is stationary. metapodial mucus gland also occur in spe- The ultrastructure of the sensory epithelium cies of Alaba H. Adams & A. Adams, 1854, of the osphradia of members of the Bittium- and Litiopa Rang, 1829 (Litiopidae Fischer, group is typical of Cerithioidea, and Haszpru- 1885), in members of Colina H. Adams & A. nar (1985:479) has shown that the osphradial Adams, 1854 (Cerithiidae Férussac, 1819), cells bear paddle cilia. The osphradial classi- and in species of Plesiotrochus Fischer, 1878 fication of Bittiinae species falls under Hasz- (Plesiotrochidae Houbrick, 1990b) (Kosuge, prunar's (1985) group "S¡2." Haszprunar 1964; Houbrick, 1987b; Luque et al., 1988; (1985) repeated the Fretter & Graham (1962: Houbrick, 1990a, 1990b, respectively). I have 367) statement that the osphradium is a "sim- previously pointed out the anatomical fea- ple brown ridge," but this is not concordant tures shared by Colina with members of the with my observations of the pectinate condi- Bittiinae (Houbrick, 1990a: 50-51). Species tion in many taxa of the group. of Plesiotrochus Fischer, 1878, also have a The phylogeny and relationship of mem- papillate epipodial skirt and an elongate bers of the Bittium-group will remain unclear metapodial slit leading into a large metapodial until the anatomy of other cerlthioidean taxa is 272 HOUBRICK

BITTIUM

ITTIBITTIUM

BITTIOLUM

STYLIDIUM

LIROBITTIUM

CACOZELIANA

FIG. 2. External anatomical characters of five genera of the BiWum-group. Figures to left represent right lateral views of headfoot, showing mantle edge, ciliated gutter, ovipositor and epipodial skirt configuration; figures to left show sole of foot, anterior mucus gland, metapodiai mucus gland (when present) and con- figuration of epipodial skirt. GENERIC REVIEW OF BITTIINAE 273 better understood and a phylogenetic analy- era, Triphora Blainville, 1828, and Cerithiop- sis can be accomplished. sis Forbes & Hanley, 1851. Neither a descrip- tion of Bittium nor a type species were given. Three months later, Gray (1847b) cited only BITTIUM GfíAY. 1847 Bittium reticulatum (Da Costa, 1778) under Bittium, and this citation is a subsequent des- Bittium Gray, 1847a (Oct.): 270 (Type species ignation. (Gray's system is explained in his by subsequent designation, Gray, 1847b: introduction, pp. 129-130, and the species so Strombiformis reticulatus DaCosta, listed are to be taken as type designations). 1778). Thiele, 1929: 211; Wenz, 1940: The earliest diagnosis of Bittium is that of H. 755; Nordsieck, 1968: 68; Houbrick, Adams & A. Adams (1854) who besides de- 1977: 103. scribing shell characters, noted the opercu- Cerithiolum TIberi, 1869: 263 (Type species lum, epipodial skirt, and opercular lobe. by original designation, Strombiformis re- My original paper on Bittium (Houbrick, ticulatus DaCosXa, 1778). 1977) reviewed the nomenclatural history of Manobittium Monterosato, 1917: 20 (Type the genus, and should be consulted for de- species by monotypy, Cerithium latreillei tailed information about the confusion and Payraudeau, 1826, = S. reticulatus). taxonomic problems between Bittium and Thiele, 1929: 212. other taxa of small-shelled cerithioideans. Inobittium Monterosato, 1917: 20 (Type spe- Subsequent to that review, there have been cies by monotypy, Cerithium lacteum many changes and the synonymy of Bittium Philippi, 1836, = S. reticulatus). Thiele, originally published (Houbrick (1977: 103) 1929; 212; Wenz, 1940: 757. has been modified herein: some taxa have Rasbittium Gründel, 1976: 53 (Type species been excluded, and genera not originally in- by original designation, Cerithium /aire/V- cluded have been added. A commentary on te/Payraudeau, 1826, = S. reticulatus). the present synonymy follows: Cerithiolum is an objective junior synonym of Bittium: both Diagnosis genera share the same type species, Bittium reticulatum. Gründel (1976) regarded Cer- Shell small, elongate, with short anterior ca- ithidium and Rasbittium Gründel, 1976, as nal and sculptured with 4-5 spiral cords with subgenera under Bittium, s.S., but as shown many aligned small beads formed where axial before, Cerithidium is excluded from Bittiinae. riblets are crossed by spirals. Operculum cir- Rasbittium is a primary objective synonym of cular, paucispiral with subcentral nucleus. Epi- Manobittium as seen in the synonymy above. podial skirt with many small, short papillae. Manobittium and Rasbittium are considered Opercular lobe with small pointed papillae. subjective junior synonyms of Bittium be- Well-developed ovipositor comprising parallel cause both share the same type species, Cer- glandular ridges and bisected by egg-laying ithium latreillei, which is considered by me gutter on right side of foot near edge of epi- and a number of authors to be conspecific or podial skirt. Osphradium ridge-like, weakly subspecific with Bittium reticulatum (see Ver- monopectinate, one-half the ctenidial length. duin, 1976). The eastern Atlantic species, Openings to sperm bursa well separated from Cerithium lacteum, which is the type species opening to seminal receptacle. of Inobittium, also is considered herein to be conspecific with Bittium reticulatum. Wenz Remarks (1940: 757) regarded Inobittium as a syn- onym of Lirobittium, but I see no close resem- Bittium Gray, 1847a, was first proposed in blance between the shells of the two. Should manuscript by Leach in 1818 for a classifica- Cerithium lacteum be a distinct species, as tion of British IVIollusca, and it was subse- thought by Verduin (1976), the differences quently made available by Gray (1847a). are certainly not of generic weight; conse- Leach's list referred Bittium and several other quently, Inobittium is regarded as a subjective diverse genera to Purpuridae and under the junior synonym. 65th entry listed Murex retlculatum, M. tuber- Discussion culare, M. adversum, M. elegantlssimum, and M. spencerl, consecutively, under Bit- The genus Bittium is characterized by a can- tium. Besides Bittium reticulatum, the other cellate, beaded shell sculpture formed by 4-5 species listed by Leach represent two gen- dominant spiral cords and numerous axial rib- 274 HOUBRICK lets (Fig. 3A-E), a circular operculum with sub- All living, observed members of the Bittii- centric nucleus (Fig. 3F), and by the small nae appear to be feeders of epiphytic microal- papillae along the edge of the epipodial skirt gae, such as diatoms, which occur commonly and opercular lobe (Fig. 2). The ovipositor in on sea grasses. Most species occur in large females is a highly developed, raised glandu- populations and are highly gregarious. lar lump at the base of the foot near the sole Species of the genus Bittium appear to be edge, forming a series of parallel, glandular primarily concentrated in the eastern Atlantic; ridges bisected by the deep ciliated egg-laying the Bittium reticulatum complex and species groove (Fig. 4B, ovp). The ridge-like monopec- closely related to it are commonly found tinate osphradium is unusual in having the throughout the Mediterranean, north African, pectins on its right side. It is half the length of and western European regions, and appear the ctenidium. The openings to the sperm to be adapted to temperate and cold waters. bursa and seminal receptacle in the lateral Bittium impendens from the Indo-Pacific, lamina of the palliai oviduct (Fig. 68, C, osr, which differs from the Atlantic Bittium species osp) are well separated from each other in only in lacking a monopectinate osphradium, contrast to most other members of the Bittium- is herein included under the genus Bittium. If group. this species truly belongs in Bittium s.S., and if The shells of small-sized Cerithium species other anatomically unknown Indo-Pacific spe- frequently are erroneously misclassified as cies are examined, the geographic distribu- BiWum species. Griindel (1976) presented tion of the genus Bittium may be far wider several conchological features that he be- than is now thought. lieved separated the two genera. He stated that Cerithium differs from Bittium in having a Bittium reticulatum (Da Costa, 1778) more complex aperture, but this is only true (Figs. 3-6) for larger Cerithium species; some small spe- cies, such as Cerithium atromarginatum, Cer- Strombiformis reticulatus Da Costa, 1778: ithium egenum, and Cerithium zebrum, have 117, pl. 8, fig. 13. apertures like those of Bittium (Houbrick, Murex reticulatus (Da Costa). Montagu, 1803: 1978). Gründe! (1976) further indicated that 272. ontogenetic sculptural development in Cerith- Cerithium latreillei Payraudeau, 1826: 143. ium begins with a single primary spiral cord Cerithium lacteum Philippi, 1836: 195. that becomes stronger and more prominent, Cerithium reticulatum, Risso, 1826; 157; G. B. forming a keel that is not integrated with the Sowerby, 1855: pl. 15, fig. 8; Jeffreys, weaker axial riblets; moreover, there are 1867; 258; 1869; pl. 80, fig. 4; 1885; 57. many fine spiral threads of varying strength. Bittium reticulatum, Watson, 1886; 540; Buc- In Bittium, whorl sculpture begins with two quoy et al., 1884: 212-215, pl. 25, figs. spirals that quickly become four primary spiral 3-9; Tryon, 1887: 15Ô-151, pl. 29, figs. cords forming a network with sharply defined 78-83; Dautzenberg, 1889; 40-41. axial riblets. The so called "definitive" shell characters proposed by Griindel (1976) are Description unreliable, because the more species that are examined, the more exceptions and ambigu- Shell (Fig. 3A-H): Shell elongate, reaching ities one encounters. 15 mm in length, comprising 9-10 moderately Marcus & Marcus (1963) cited the pres- inflated whorls. Protoconch (Fig. 3G) com- ence of a metapodial mucus gland in Bittium prising two weakly sculptured whorls. Early reticulatum, crediting this information to Fret- whorls beginning with two spiral cords and ter (1948). However, no such gland was ob- broad subsutural ramp (Fig. 3H). Adult whorls served in living or preserved, sectioned spec- sculptured with 4-5 spiral cords beaded imens from the Azores; furthermore. Ponder where many small axial riblets cross over (in litt.) did not note this structure on speci- them, creating cancellate sculpture. Suture mens of Bittium reticulatum from the western deeply impressed. Body whorl a little under coast of Sweden. Fretter's (1948; 628) paper one-third shell length, having weak basal con- merely cites the presence of this gland in striction and small anterior canal weakly re- such small gastropods as Bittium, Cerithiop- flexed to left. Body whorl sculptured with five sis, and Triphora, but as she mentioned only major spiral cords and 5-6 weaker cords on generic names, it is unclear what "Bittium" its base. Aperture ovate, a little over one-third species she actually observed. shell length, with concave columella having GENERIC REVIEW OF BITTIINAE 275

FIG. 3. Representatives of genus Bittium: A-H, B. reticulatum; l-N, S. impendens. A-C, SEM micrographs of B. reticulatum from Sao Miguel, Azores (USNM 878030), 6 mm length; D, E, S. reticulatum from Tunisia (USNM 754051), 11 mm length; F, SEM micrograph of operculum of B. reticulatum, bar = 0.5 mm; H, SEM micrograph of immature shell of B. reticulatum, bar = 0.5 mm; l-L, SEM micrographs of shell of B. impendens from Honolulu, Hawaii (USNM 857098), 5 mm length; M, SEM micrograph of operculum of B. impendens, bar = 0.5 mm; N, SEM micrograph of protoconch of B. impendens, bar = 150 fim. 276 HOUBRICK

slight columellar callus; anterior canal short, white spot. Mantle edge thickened at inhalant shallow; anal canal very small; outer lip (Fig. 4C, inh) and exhalant siphons. rounded, weakly crenulate. Periostracum thin, light tan. Palliai Cavity: Palliai cavity deep, comprising about two whorls. Osphradium olive colored, Animal (Figs. 4-6): Head-foot of animal pig- ridge-like, pectinate on right side only, bor- mented light yellowish-brown overlain by dered on each side by narrow ciliated strip. large dark brown blotches and small white Osphradium wide, about one-half ctenidial spots. Visceral mass with 8 visceral whorls, length, beginning close behind inhalant si- comprising mostly digestive gland and over- phon and extending length of ctenidium. lying gonads. Ovary white; testis dirty yellow. Ctenidium bluish-gray, comprising numerous Stomach about one whorl in length. Kidney finger-like, triangular filaments with narrow large, light tan, about two-thirds whorl in bases. Hypobranchial gland narrow, glandu- length. Columellar muscle white, broad, short, lar comprising several kinds of large gland about one-half length of palliai cavity. Head cells that stain dark blue. Rectal tube dis- (Fig. 4A) with elongate, narrow snout (Fig. 4B, tended, filled with elongate, ovoid-shaped fe- sn), flattened dorso-ventrally, expanded at bi- cal pellets. Palliai gonoducts open, beginning bbed tip, with bright yellow, oval-shaped oral behind mantle edge and extending posteriorly pad at anlero-ventral end (Fig. 4A, C, 1). as far as kidney. Cephalic tentacles (Fig. 4A, t) elongate, nar- Reno-pericardial System: Kidney large, about row, with broad peduncular bases each with two-thirds whort in length, beginning at ante- large dark eye. Foot narrow, elongate, cres- rior end of style sac, extending anteriorly well cent shaped anteriorly. Deep transverse slit into mantle cavity roof, lying over one-third of (Fig. 4C, amg) between epipodial lips marks posterior palliai gonoduct. Kidney with simple entrance to large ovate anterior mucus gland kidney opening, but no renopericardial duct. extending via central duct deep into anterior Pericardium typically monotocardian, lying ad- foot. Epipodium separated from lower foot jacent to posterior wall of mantle cavity. and densely ciliated sole by deep, laterally placed groove (Fig. 4B, epg) forming broad Alimentary System: Mouth (Fig. 4A, m) lying epipodial skirt (Fig. 4B, C, eps) extending antero-ventrally on snout, opening into oral posteriorly on each side of foot from corners cavity between two semicircular lips (Fig. 4A, of anterior epipodial lips of anterior mucus C, 1). Buccal mass (Fig. 4D, bm) relatively gland around entire foot base, joining behind small, about one-third snout length, loosely and below opercular lobe. Lateral epipodial attached to snout wall by numerous thin mus- skirt scalloped along edges of each side of cle strands. Jaw tan, semicircular, comprised median and posterior parts of epipodium, of cuticular cones and lying on either side of having small papillae (Fig. 48, C, ep); epipo- entrance to anterior buccal cavity. Radular dial skirt forming long opercular lobe (Fig. 4B, ribbon (Fig. 5A; Table 2) folded beneath buc- C, opi). Sole of foot (Fig. 4C, s) indistinctly cal mass and radula sac emerging behind it. divided into two parallel axial parts, forming Rachidian tooth (Fig. 5C) with dorso-ventrally anterior longitudinal fold. No metapodial mu- compressed basal plate with concave sides cus gland. Operculum (Fig. 3F) corneous, rounded base and with V-shaped base but- tan, circular, paucispiral with subcentral nu- tressed on each side with a basal lateral ex- cleus and with thin, transparant border. Cili- tension; rachidian broader above than below, ated gutter (Fig. 4B, C, eg) emerging from having cutting edge with slightly concave top, right side of mantle cavity (Fig. 4C, ex) and and comprising large, spade-shaped central running down right side of foot; ciliated gutter cusp flanked on each side by 2-3 small, leads to large glandular ovipositor (Fig. 4B, C, pointed denticles. Lateral tooth (Fig. 5B) with ovp) and egg-laying pit at base of epipodium broad basal plate comprising long, ventrally in females. Ovipositor oval-shaped, com- extending, central pillar having small pustule prised of glandular, transparant white tissue on its face, and with moderately long lateral formed into many parallel pleats divided extension; cutting edge comprising very large transversely by deep central slit. Mantle bi- spade-shaped cusp with one inner denticle lobed at edge, having smooth outer lobe and and 3-6 outer denticles. Marginal teeth (Fig. inner lobe with many small papillae, becom- 5A) curved, elongate, with broad, swollen ing smooth ventrally. Mantle papillae (Fig. 4B, shafts, narrowing and becoming spatulate at C, mp) slender, darkly pigmented, each with tips; inner marginal tooth with tip having long GENERIC REVIEW OF BITTIINAE 277

FIG. 4. Anatomical representations of Bittium reticulatum. A, head and snout; B, lateral view of headfoot; C, fiead and sole of foot; D, anterior alimentary system exposed by dorsal longitudinal cut through wall of buccal cavity, aes = anterior esophagus; amg = anterior mucus gland; beg = subesophageal gland; bg = buccal ganglion; bm = buccal mass; c = ciliated strip; eg = ciliated gutter; eg = esophageal gland; ep = epipodial papilla; epg = epipodial groove; eps = epipodial skirl; ex = exhalant siphon; inh = inhalant siphon; I = lip; leg = left cerebral ganglion; Ipg = left pleural ganglion; Isg = left salivary gland; m = mouth; mp = mantle papilla; op = operculum; opi = opercular lobe; ovp = ovipositor; pes = posterior esophagus; reg = right cerebral ganglion; rpg = right pleural ganglion; rsg = right salivary gland; s = sole; seg = supraesophageal ganglion; sn = snout; t = tentacle. 278 HOUBRICK

FIG. 5. Scanning electron micrographs of radula of Bittium reticulatum from Sao Miguel, Azores (USNM 878030). A, half row with marginal teeth folded back, bar = 19 ^m\ B, rachidian and lateral teeth, bar = 15 |i,m; C, detail of rachidian teeth, bar = 4 jxm.

central cusp, 3-4 inner denticles, 4 outer folded, forming large, olive-brown esophageal denticles; outer marginal tooth same, but gland (Fig. 4D, eg). Internal epithelium of lacking outer denticles. Salivary glands (Fig. esophageal gland (Fig. 7A, B, eg) forming nu- 4D, rsg, Isg) comprising pair of narrow, un- merous transverse folds or lamellae, staining coiled, shiny tubes, beginning behind nerve dark blue with f\/lethylene blue. Posterior ring, extending through it anteriorly, opening esophagus (Fig. 4D, pes) narrow and straight, into far anterior portion of buccal cavity. Buc- running on top of columeilar muscle, entering cal cavity opening and enlarging immediately into left side of stomach. Stomach large, com- behind nerve ring, having pair of prominent prising about one whorl of visceral mass, in- dorsal folds and smaller pair of smaller ventral cluding style sac. Esophageal opening into folds. Interior mid-esophageal w/alls highly median ventral part of stomach floor. Large GENERIC REVIEW OF BITTIINAE 279 sorting field with many fine folds adjacent to glandular, comprising prostate gland (Fig. 6A, right side of esophageal opening. Minor pg). Anterior half of male gonoduct glandular, typhlosole bordering right side of esophageal not as thick, putative spermatophore-forming opening. Large central elevated pad in center organ (Fig. 6A, so). of stonnach adjacent to single duct to diges- Ovary opaque white, thin-walled, overlying tive gland lying short distance below esoph- digestive gland, extending anteriorly, ending ageal opening. Digestive gland comprising about one-half whorl before stomach. Coelo- single brown lobe consisting of digestive cells mic oviduct (Fig. 68, C, cod) short tube, highly and secretory cells with dark brown granules. ciliated within, beginning anterior to stomach Gastric shield on right side of stomach having with duct wall lying against pericardium (no cuticular lining with protruding, toothed edge. connection), ending at posterior mantle cavity Depressed epithelial pocket on floor of stom- where circular sphincter muscle separates it ach adjacent to posterior part of gastric from palliai oviduct. Female palliai oviduct shield. Style sac short, about one-third the (Fig. 6B, C) large, comprising two laminae, stomach length, nearly spherical, and con- enlarged and glandular at their bases, at- taining crystalline style. Style sac adjacent to tached basally to each other and to mantle but separate from intestine opening, except floor, forming ciliated oviductal groove (Fig. for limited connection where both enter stom- 68, C, ovg). Posterior end of palliai oviduct ach. Anterior part of stomach with many par- closed. Medial, free lamina with wide anterior allel ciliated folds and closed off from style ciliated sperm gutter (Fig. 68, C, sg) along its sac by major typhlosole. Internal intestinal edge leading to two, well-separated, pocket- walls with many fine folds where exiting stom- like openings. First opening (Fig. 68, C, osp) ach. Intestine curves around style sac, turns leading into large, deep bursa having smooth to right, and runs straight forward. Rectum inner epithelium and containing large num- with thin muscular wall, terminating in anal- bers of non-directed spermatozoa (Fig. 7C, D, bearing papilla. sp); ciliated gutter continuing posteriorly to open (Fig. 7C, osr) into pouch-like, muscular Nervous System: Nervous system epiath- seminal receptacle (Fig. 6C, B sr; 8C, D, sr) roid, dialyneurous. Nerve ring comprised of containing oriented euspermatozoa with large ganglia. Pleural ganglia (Fig. 4D, rpg, heads embedded in receptacle walls. Lateral Ipg) close to cerebral ganglia (Fig. 4D, rcg, lamina attached to palliai wall, having anterior leg). Cerebral connective equalling length of ciliated tract comprising many parallel elon- cerebral ganglion. Buccal ganglia (Fig. 4D, gate, fine ciliated folds (Fig. 68, C, ctr; 7A, B, bg) small, lying at posterior edge of buccal ctr) running posterior to open into thin-walled mass. Subesophageal ganglion (Fig. 40, tube leading into posterior pouch-like bursa beg) very close to left pleural ganglion (Fig. having highly vacuolated epithelium and func- 4D, Ipg). Supraesophageal connective mod- tioning as spermatophore bursa (Fig. 6B, C, erately long, about twice length of right pleural sb). Ciliated tract and folds opening to semi- ganglion; dialyneury between left palliai nerve nal receptacle on lateral lamina located oppo- and nerve emerging from supraesophageal site sperm gutter and opening to seminal re- ganglion (Fig. 4D, seg). Visceral ganglion lo- ceptacle of medial lamina, both edges cated in floor of posterior mantle cavity. interdigitating to form closed system. Poste- Reproductive System: Testis creamy yellow, rior half of glandular portion of both laminae overlying dark brown digestive gland, extend- opaque white color, comprising albumen ing anteriorly about five whorls, ending one- gland (Fig. 68, C, ag; 7C, D, ag); anterior half half whorl before stomach. Testicular ducts dirty white, comprising capsule gland (Fig. on inner side of visceral coil, joining to form 68, C, eg; 7A, 8, eg). spermatic duct, enlarging anteriorly, becom- Spawn comprising thin gelatinous string ing seminal vesicle and containing two kinds (about 25 mm length, uncoiled) tightly coiled of spermatozoa; euspermatozoan with single clockwise or irregularly folded on itself and long flagellum and paraspermatozoan with attached to substrate. Jelly string containing [four ?] flagellae. Males aphallate. Male palliai many small opaque eggs (0.65 |a,m diameter) gonoduct (Fig. 6A) open, comprising two thin each within thin, transparent hyaline capsule walled laminae (Fig. 6A, 11, ml) with thicker (110 ^.m diameter). Entire spawn mass con- transverse glandular folds at their attached tains about 800 eggs. Free swimming bilobed bases bordering gonaductal groove (Fig. 6A, planktotrophic veliger larval stage present. gd). Posterior half of male gonoduct thick. Larval shell ranging from 170-330 ^im, de- 280 HOUBRICK

cod

osr

°^'XJv^ Ictr ovg II . osb

ml

0.2mm so II B 0.2 mm ml osp

cod

FIG. 6. Representation of palliai gonoducis of Bittium reticulatum. A, male palliai gonoduct, showing section through mid-duct beneath, represented by dotted line; B, palliai oviduct showing three cross sections of duct represented by dotted arrows and sections to right; C, reconstruction of palliai oviduct showing configuration of ducts and glands (anterior to right), ag = albumen gland; ant = anterior; eg = capsule gland; cod = coelomic oviduct; ctr = ciliated ridge tract; gd = gonaductal groove; II = lateral lamina; ml = medial lamina; osb = opening to spermatophore bursa; osp = opening to sperm bursa; osr = opening to seminal receptacle; ovg = oviductal groove; po = closed portion of palliai oviduct; sb = spermatophore bursa; sg = sperm gutter; sp = sperm bursa; sr = seminal receptacle; so = spermatophore-forming organ. pending upon age. Larval shell wWh rounded, Discussion nearly smooth whorls having thin spiral thread forming weak keel and with deep sinusigeral The status of the many specific and sub- notch (Thorson, 1946; 192, fig. 109). specific names comprising the Bittium reticu- GENERIC REVIEW OF BITTIINAE 281

FIG. 7. Successive sections, anterior to posterior, through palliai oviduct of Bittium reticulatum. A, anterior of palliai oviduct showing relationship of mantle cavity organs to oviduct, bar = 0.25 mm; B, mid-section shov^ing ciliated ridge tract and opening to sperm bursa, bar = 0.25 mm; C, section through enlarged sperm bursa in posterior palliai oviduct, bar = 0.25 mm; D, section through closed posterior of palliai oviduct, bar = 0.25 mm. ag = albumin gland; eg = capsule gland; ct = ctenidium; ctr = ciliated ridge tract; eg = esophageal gland; hg = hypobranchial gland; os = osphradium; osp = opening to sperm bursa; ovg = oviductal groove; r = rectum; sb = spermatophore bursa; sg = sperm gutter; sp = sperm bursa; sr = seminal receptacle. latum complex is controversial (Verduin, species or a closely related species of the 1976). It is not my intention to address alpha- Bittium reticulatum complex. Bittium reticula- level problems in this generic review, but the tum is exceedingly variable in shell sculpture Azorean population used for the anatomical throughout its range (compare Figs. 2A, C, study herein Is considered by some as a sub- D), but this is not unusual among cerithioide- 282 HOUBRICK ans. The palliai oviduct described by Johans- functions as a spermatophore bursa in Bittium son (1947) and notes and sketches made by reticulatum and probably in most other mem- Ponder (Ponder, in litt.) on the anatomy of bers of the Bittium-group, because Marcus & specimens from western Sweden agree sub- Marcus (1963) found spermatophores in this stantially with my observations of Azorian structure in the western Atlantic Bittiolum var- specimens. For the purposes of this study, ium. I was unsuccessful in finding spermato- the Bittium reticulatum complex is regarded in phores in either structure in specimens of the broad sense {sensu lato), as a single spe- Bittiolum varium from Florida. A new genus cies. from the Indo-Pacific, Ittibittium, described The epipodial skirt, characteristic of mem- herein, deviates from the typical palliai ovi- bers of the ß/fi/um-group, forms a highly cili- duct layout in lacking the spermatophore ated lateral groove where it overhangs the bursa in the lateral lamina and in having the foot, and carries detrital particles posteriorly albumen gland protrude posteriorly beyond to the back of the foot where they are dis- the back of the palliai cavity Into the visceral carded. coil. The posterior roof of the palliai cavity is The spawn of Bittium reticulatum was first covered by the anterior extension of the renal deschbed and figured by Meyer & Möbius organ, which overlays the posterior palliai (1872), and the spawn and larvae described gonoduct. The renal organ opens via a mus- by Lebour (1937) and Graham (1988). cular sphincter, the renal opening, into the Spawn, larvae, veliger, protoconchs, and ju- posterior palliai cavity. venile shells of this species were described The ridge-like osphradium of Bittium retic- and well illustrated by Thorson (1946: 192, ulatum is unusual in being pectinate on its fig. 109). Other depictions of the larval shell of right side. Although these pectins are small, this species are those of Fretter & Pilkington they are clearly visible and very unlike simple (1970: 10-11, fig. 6) and Richter & Thorson nonpectinate osphradia of closely related (1975: pi. 3, figs. 16-17). According to Gra- taxa. ham (1988), British Bittium reticulatum is a The rachidian tooth of the radula of Bittium summer breeder and attaches its spawn to reticulatum is similar to those of members of shells, stones or weeds. Spawn comprises a other genera in the group, but unlike that of cylindrical ribbon about 3 mm in diameter, Cacozeliana (see below). Table 2 gives the having a total length of 25 mm, and coiled in comparative dentition of the radular teeth. tight spirals. A spawn mass contains about Bittium reticulatum has three sperm stor- 1000 eggs, which develop to veliger larvae. age spaces, two connected to the ciliated The geographic range of the Bittium reticu- groove of the non-glandular portion of the me- latum complex is broad, comprising western dial free lamina, and one in the posterior part Europe, the Azores, North Africa, and the of the non-glandular attached lateral lamina Mediterranean. (Fig. 6B, 11 ). It is not entirely clear how these three bursae function. Of the two bursae in Bittium impendens (Hedley, 1899) the medial lamina, the smaller one is clearly (Fig. 3, l-N) the seminal receptacle, because oriented eu- Cerithium impendens Hedley, 1899: 434- spermatozoa are found in it, exclusively (Fig. 435, fig. 23 (Holotype: AMS 05944; type 7C, D, sr). The larger bursa (Fig. 6B, sp) con- locality: Funafuti Atoll, Ellice Islands); tains considerable numbers of unoriented Kay, 1979: 118, 120, fig. 45A. sperm, and much nondescript material (pre- sumably disintegrating paraspermatozoa and Description degenerating spermatophores), although some euspermatozoa occur with heads ori- Shell: (Fig. 31-N). Shell short, stout, with ented on the inner wall epithelium, especially wide base, reaching 7 mm length and com- near the opening to the sperm gutter (Fig. prising 8-9 convex whorls. Protoconch (Fig. 7D). Although this large bursa in the medial 3N) comprising 2.5 whorls; protoconch 1 lamina contains spermatophores in most cer- smooth; protoconch 2 sculptured with thin ithiids, this is not the case in members of the central, spiral keel and weak presutural spiral Bittium-group, where it appears to function as thread; lower part of each whorl with micro- a sperm storage and ingesting area. It is in- scopic pustules. Whorls slightly pendant ferred that the pouch in the posterior of the abapically, constricted at suture. Adult shell lateral lamina (Fig. 6C, sb, Fig. 7C, D, sb) sculptured with 3-4 major spiral cords inter- GENERIC REVIEW OF BITTIINAE 283 spersed with spiral threads. Spiral cords dens is very similar to that of Bittium reticula- weakly beaded and beads aligned to form ax- tum except that the marginal teeth have fewer ial riblets. Suture well defined. Weak varices outer and inner denticles. Aside from these randomly distributed. Body whorl very broad, differences, the animal shares most of the an- about one-half the shell length, with promi- atomical features of Bittium reticulatum. Al- nent wide, dorsal varix (Fig. 3J, L); body whorl though an argument could be made that this sculptured with about 14 spiral cords and species represents yet another new genus, I strongly constricted at base. Aperture a little have conservatively placed Bittium impen- over twice shell length, broadly ovate, with dens under Bittium, s.s, with a query, be- short, wide, shallow anterior canal and cause it does have many characters in com- smooth outer lip extending widely at shell mon with the type species of Bittium. base (Fig. 31). The shell of Bittium impendens differs from other SMum-group genera by its fir-tree out- Animal: Headfoot pinkish white, blotched line and wide body whorl with prominent dor- with brown, covered with white spots and with sal varix (Fig. 31-L). The protoconch (Fig. 3N) chestnut stripes. Kidney bright pink. Right is smooth except for a thin spiral thread and a side of foot in females with ciliated gutter end- deep sinusigeral notch, indicative of a plank- ing in small ovipositor at edge of lateral tonic larval phase. Judging from specimens groove. Epipodial skirl having very small pus- from other regions that appear to be concho- tules or protuberances along lateral edges on logically conspecific, this species has a wide each side of foot; opercular lobe scalloped Indo-Pacific distribution, occurring from cen- and pointed at end. Sole of foot pink, without tral Pacific islands throughout the Indo-West- metapodial mucus gland. Mantle edge fringed Pacific to east Africa. dorsally with papillae; underside of inhalant siphon with three large papillae. Marginal miBITTIUM, New Genus teeth of radula having three inner denticles. Osphradium a thin brown ridge, non-pecti- Diagnosis nate. Openings to sperm pouch and seminal receptacle in medial lamina close to each other, situated within common aperture at end Shell small, reaching 6 mm length, with in- of sperm gutter in edge of anterior third of flated whorls and dominant spiral sculpture of medial lamina adjacent to opening of sper- 4-5 cords. Protoconch with depressed, con- matophore bursa of lateral lamina. No ciliated cave apex, broad sutural ramp, sculptured tract leading to spermatophore bursa. with minute axial striae and two strong spiral cords. Operculum ovate, paucispiral with ec- centric nucleus. Each side of propodium with Discussion elongate papilla. Epipodial skirt laterally fringed with slender papillae. Large opercular Examination of the type lot (holotype and 7 lobe having elongate papillae. No ovipositor paratypes) of Cerithium impendens confirms in females. Sole of foot with long, central lon- that the Hawaiian specimens studied herein gitudinal slit marking entrance into large are conspecific with this taxon. This species metapodial mucus gland. Osphradium weakly has not been cited frequently in the literature. bipectinate. Albumen gland extending past The assignment herein of Bittium impen- posterior of palliai cavity into visceral coil. No dens to the genus Bittium is made with some spermatophore bursa in lateral lamina of pal- doubt. The shell morphology of this wide- liai oviduct. Spawn compnsing short gelati- spread Indo-Pacific species is quite different nous tube. from that of the type species of Bittium, Bit- tium reticulatum (compare Fig. 3A-E and Type Species: Bittium parcum Gould, 1861. 31-L), and unlike the shells of other eastern Atlantic Bittium species. In addition, the os- Etymology: A compound of "itti," American phradium is ridge-like rather than mono- vernacular prefex for very small, and Bittium. pectinate, and there does not appear to be a ciliated tract associated with the spermato- Remarks phore bursa on the lateral lamina. Instead, the opening to the spermatophore bursa is adja- This genus is perhaps one of the most dis- cent to the two openings of the bursae in the tinctive of the Bittium group, in terms of its medial lamina. The radula of Bittium Impen- unusual protoconch and anatomical features. 284 HOUBRICK The proloconch with depressed apex and keel. Adult whorls angulate, sculptured with broad sutural ramp (Fig. 81) is unique among keel-like median cord, 7-8 minor spiral cords, the Bittium-group. The distinctive propodial each cord abapically overlapped by succes- and epipodial papillae, well-developed epipo- sive one. Eight to nine weak to strong axial dial skirt, and long metapodial mucus gland ribs occasionally on whorls, especially on up- are conspicuous autapomorphiic characters per ones (Fig. 8J). Varices randomly placed. in living specimens (Fig. 2). The lack of a Suture moderately impressed. Body whorl spermatophore bursa in the lateral lamina of (Fig. 8L) slightly constricted at base, compris- the palliai oviduct and the protrusion of the ing a little less than half shell length, sculp- albumen gland through the posterior palliai tured with 15-19 weak flattened spiral cords, cavity into the visceral coil are highly unusual occasional weak axial ribs and with broad autapomorphies, and set Ittibittium, gen. n., varix. Aperture about one-third shell length, apart from the rest of the Bittiinae. The place- ovate with smooth outer lip and short broad ment of the spermatophore bursa in the lat- anterior canal. Slight columellar callus eral lamina is one of the synapomorphous present. Periostracum thin, nearly transpar- character used in this review to define the ent. subfamily Bittiinae; therefore, it is noteworthy Animal: Animal pigmentation highly variable, that Ittibittium, gen. n., has lost this feature. ranging from greenish-yellow to pink and The spawn mass of Ittibittium, gen. n., is also brown and covered with white blotches. unusual in being a simple, short tube. Cephalic tentacles wide at bases, elongate, In some museum collections, Bittium par- twice snout length. Snout elongate, narrow, cum and species similar to it are incorrectly bilobed at tip. Operculum (Fig. 8K) thin, cor- assigned to Bittinella Dall, 1924, a genus neous, tan, circular-ovate, paucispiral with based on Bittium hiloense Pilsbry & Vanatta, subcentral nucleus. Anterior part of foot cres- 1908, which has been shown to be a rissoid of cent-shaped, cowl-like, having single long pa- the genus Isselia (Ponder, 1985: 95; Kay, pilla on each side (Fig. 2). Narrow transverse 1979:80). slit at edge of propodium leading into large, spherical anterior mucus gland, staining deep ittibittium parcum (Gould, 1861) purple in toluidine blue. Lateral epipodial skirt (Figs. 8-11) with about 10 small, slender papillae along edges (Fig. 2) on each side of foot, extending Bittium parcum Gould, 1861: 387 (Lectotype, posteriorly to large opercular lobe having long R. Johnson, 1964, USNN/I 2040; type lo- papillae along its edges; papillae show cality Okinawa, Ryukyu Islands); G. B. through edges of opercular border. Sole of Sowerby, 1866: pi. 18, fig. 125; Tryon, elongate, narrow foot having deep, centrally 1887: 155, pi. 30, fig. 20; R. Johnson, placed, narrow longitudinal slit (Fig. 2) begin- 1964: 122, pi. 12, fig 14; Kay, 1979: 120, ning behind anterior mucus gland slit (Fig. 2) figs. 22D, 45D, E. and extending posteriorly to back of foot; slit Cerittiium tiawaiensis Pilsbry & Vanatta, leading by way of ciliated duct into deep, mas- 1905: 576 (Holotype ANSP; type locality: sive, metapodial mucus gland, staining deep Hilo, Hawaii). purple in toluidine blue. Males with ciliated sthp on right side of foot, emerging from right Description side of mantle cavity and extending down to edge of sole. Ciliated gutter on right side of Shell (Fig. 8): Shell small, pupale-elongate, foot in females deep, running down side of comprising about 8 inflated, angulate whorls foot and extending through lateral epipodial and reaching 5.8 mm length. Protoconch (Fig. groove (Fig. 2). No ovipositor present. Mantle 8F-I) compnsing two concave whorls, con- edge dorsally fringed with many small papil- cavely flattened apex, very broad sutural lae. ramp sculptured with minute axial striae (Fig. 8F); protoconch whorls sculptured with two Palliai Cavity: Osphradium a little less long strong, keel-like spiral cords, with central spi- than ctenidium, broad, about one-third ctenid- ral cord becoming dominant one. Early whorls ial width, dark brown, weakly bipectinate with sharply angulate (Fig. 81); first post-larval small pectins on each side but unconnected whorl with keel-like median spiral cord; sec- dorsally; osphradium becoming monopecti- ond whorl with another spiral cord above keel nate at inhalant siphon. Ctenidium narrow, and third whorl having 3 spiral cords above extending length of palliai cavity, comprising GENERIC REVIEW OF BITTIINAE 285

FIG. 8. SEM micrographs of IWbiWum parcum from Honolulu, Hawaii (USNM 857100). A, B, apertural and lateral views of shell, 3.6 mm length; C-E, apertural, lateral and dorsal views of shell, 3.6 mm length; F, newly hatched larval shell showing protoconch and details of whorl sculpture, bar = 63 (j,m; G, H, embryonic shells removed from eg capsule, bar = 23 ^jLm; I, larval and early whorls of shell, bar = 0.4 mm; J, shell with strong axial ribs, 5.3 mm length; K, operculum, bar = 0.2 mm; L, detail of penultimate and body whorl, showing details of sculpture and aperture, bar = 0.6 mm; M, apertural view of shell, 3.6 mm length. 286 HOUBRICK

FIG. 9. SEM micrographs of radula of IWbittium parcum from Honolulu, Hawaii (USNM 857100). A, middle of radular ribbon with right marginal teeth folded back, bar = 30 n,m; B, detail of rachidian and lateral teeth, bar = 8 ^.m.

long, finger-like, triangular filaments. Hypo- about two-thirds the stomach length. Intestine branchial gland partially overlaying rectunn, leaving stomach looping dorsally and across well developed, composed of several large, anterior style sac, turning sharply, running an- dark-staining glandular cells. teriorly, adjacent to right side of kidney and albumen gland. Rectum slightly wavy, wide, Reno-pericardial System: Pericardium lying containing large ovoid fecal pellets. adjacent to posterior palliai wall. Kidney large, extending from anterior of style sac forward, Nervous System: Cerebral ganglia very into roof of posterior palliai cavity. large, twice size of pleural ganglia. Sube- Alimentary System: Snout tip and lips of sophageal ganglion very close to left pleural mouth yellow. Buccal mass large, about two- ganglion. Supraesophageal ganglion sepa- thirds snout length. Radula (Fig. 9A) short, rated from right pleural ganglion by connec- about one-tenth shell length. Rachidian tooth tive two-thirds ganglion length. having weak hour-glass shape and cutting edge with large central cusp flanked by 2 den- Reproductive System: Testis white, overlay- ticles on each side. Lateral tooth (Fig. 98) ing brown digestive gland. Males aphallate having cutting edge with large pointed cusp, with open palliai gonoducts. Palliai oviduct one inner denticle, 3-4 outer denticles. Inner open, with large albumen gland extending marginal tooth with 2 inner denticles, large through posterior of mantle cavity mantle cav- elongate major cusp and 3 outer denticles; ity, protruding into visceral coil. Albumen outer marginal tooth with 5 inner denticles. gland staining cream-green in toluidine blue. Salivary glands paired, comprising tangled Capsule gland very large, swollen, staining mass behind nerve ring, extending through it dark blue In toluidine blue. Large spermato- anteriorly as slender tubes. Esophagus be- phore bursa in posterior medial lamina. No coming wide behind nerve ring, developing ciliated ridge tract or seminal receptacle in lat- lateral glandular pouches with many small eral lamina. Spawn mass comprising wide ge- transverse internal folds, comprising short latinous tube covered with thin membrane esophageal gland. Stomach large, about one forming compact, short tube about 2 mm long, whorl in length, having single opening to di- and 1.2 mm wide, containing large opaque, gestive gland, central raised pad, gastric compacted eggs each 0.2 mm in diameter. shield, short crystalline style and style sac, Eggs arranged in short jelly tube about 3-4 GENERIC REVIEW OF BITTIINAE 287 deep. Development direct with young snails ter throughout the Hawaiian chain, and also tiatching from eggs. occurs in French Polynesia (Naim, 1982) where it is very abundant in some localities. Discussion Naim (1982) found that this species repre- sented 89% of the molluscan fauna associ- "Bittium" parcum has not been cited com- ated with algae in Tiahura Lagoon in French monly in the literature, and due to great inter- Polynesia. specific variability in shell sculpture and color, A species from Western Australia, very is frequently misclassified or unidentified in similar to the type species, recently has been museum collections. Shell shape can vary described in great detail (Ponder, in press), from slender, elongate (Fig. 8J) to shorter, and appears to be closely related to Ittibittium more inflated (Fig. 8C-E), and shell sculpture parcum. is highly variable: the axial ribs seen in some specimens may be entirely lacking in others. The protoconch with its flattened apex, broad BITTIOLUM COSSMANN, 1906 sutural ramp and concave whorls is highly distinctive and unusual (Fig. 8F-H). However, Bittiolum Cossmann, 1906: 139. (Type spe- Ittibittium parcum is readily distinguished from cies by original designation: Bittium pod- by several external anatomical features: (1) agrinum Dall, 1892). Wenz, 1940: 755; the epipodial skirt and opercular lobe are Olsson & Harbison, 1953: 289-290. fringed with well-developed papillae; (2) a pair of long epithelial extensions (papillae) of the Diagnosis front of the foot (propodium); (3) the longitu- dinal slit marking the entrance to the metapo- Shell small, turreted, stout, sculptured with dial mucus gland is very long. Ittibittium par- 4 spiral cords and many axial ribs, and occa- cum has an unusual palliai oviduct in that the sional weak varices. Protoconch with one spi- albumen gland projects posteriorly past the ral lira. Whorls presuturally constricted, body posterior end of the mantle cavity into the vis- whorl elongate, narrow at aperture and con- ceral coil, and there is no seminal receptacle stricted at base, having less width than pen- in the lateral lamina of the pallia! oviduct. ultimate whorl. Operculum ovoid-circular, Living snails are quick, active crawlers, and paucispiral and with subcentral nucleus. An- even when removed from their shells showed terior canal weakly defined, short. Mantle a great deal of movement. edge smooth, epipodial skirt scalloped. Foot The operculum in this species tends to be elongated anteriorly and having median lon- more ovate than circular; in most other spe- gitudinal slit in posterior part of sole, leading cies of the Bittium-group, the operculum is cir- into large metapodial mucus gland. Ovipositor cular. The opercular lobe papillae show small. Osphradium bipectinate, wide, one- through the transparent edges of the opercu- third ctenidial length. Nervous system with lum. right zygoneury and with short supraesoph- This species undergoes direct develop- ageal connective. ment. The embryos pass through a veliger stage and hatch out as juvenile snails after Remarks losing the velar lobes. Direct development, while also occurring in Stylidium, is not the Bittiolum species have small shells (Table common mode of development among mem- 3) and are distinctive in having the body whorl bers of the Bittium-group. The comparatively elongated and constricted basally so that the large eggs of Ittibittium parcum are each en- aperture width is less than that of the penul- closed within individual hyaline capsules timate whorl. The smooth mantle edge, nar- about 0.2 mm diameter, and the egg capsules row elongate anterior foot, right zygoneury are stacked within a short, wide gelatinous and short supraesophageal connective are tube and deposited on the substrate in an ir- autoapomorphous characters of this genus. regular mass. Here they undergo develop- The type species of this genus is a Neo- ment, passing through a modified veliger gene fossil from Florida that has a shell mor- stage and producing a well-developed embry- phology very similar to that of living Bittiolum onic shell (Fig. 8F-H), after which they varium and Bittiolum alternatum. As the fossil emerge as small snails. species occurs in mid- to late-Neogene strata, Ittibittium parcum is common in shallow wa- and in the same geographic area as Recent 288 HOUBRICK Bittiolum varium, it is not unreasonable to in- Diastoma varium (Pfeiffer). Abbott, 1974: fer that the two species belong to the same 107, fig. 1037. clade, and the living species is considered to be congeneric with Bittium podagrinum. Description Cossmann (1906: 140) pointed out that Bitti- Shell (Fig. 10): Shell turreted, pendent- olum varium (Pfeiffer) (cited as Ceritttium) oc- shaped, comprising about 10 flat-sided curred from the Pleistocene of Florida and ex- whorls and reaching 7 mm length. Protoconch tended into the Recent. He further noted the (Fig. 101) comprising 2.5 whorls; protoconch 1 superficial resemblance of Bittiolum varium to smooth, protoconch 2 with central keel-like some fossils of Aneurychilus Cossmann, spiral lira and microscopic pustules on abapi- 1889, which he placed in the Diastomatidae cal part of whorl. Early whorls (Fig. 10H) with (as Diastomidae, Cossmann, 1906: 174). two weak spiral lirae, and sculptured with Dall (1889) was the first author to confuse dominant suprasutural spiral cord and two American members of Bittiolum with the ge- weaker spiral cords above it, and with weak nus Diastoma Deshayes, 1850, when he re- axial hbs. Adult whorls sculptured with 4 spiral ferred Bittiolum varium to that genus. Abbott cords and 14 strong axial ribs forming small (1974), probably following this cue, later re- beads at crossover points and producing can- ferred western Atlantic species of Bittium, s.l., cellate pattern. Body whorl elongate, more to Diastoma Deshayes, 1850, but this subse- than one-third shell length, constricted at ap- quently has been shown to be incorrect erture and more at siphon; body whorl sculp- (Houbnck, 1977: 102, 1981b), as the latter tured with about 10 flattened spiral cords and genus belongs to the Diastomatidae Coss- 14 weak axial ribs. Aperture ovate, con- mann, 1894, a totally different lineage repre- stricted, not as wide as width of body whorl, sented by individuals of much larger size and narrowing posteriorly and having short, dis- different anatomy that are not closely related tinct siphonal canal. Columella concave with to the Bittium-group (Houbrick, 1981b). slight callus. Outer lip of aperture smooth, The anatomy of "Bittium" alternatum, from rounded, thin and pendant, extending beyond the northeastern coast of North America, is siphonal canal. Periostracum thin, tan. identical to that of its southeastern, Carib- bean Province congener, Bittiolum varium. Animal: Snout, cephalic tentacles, and neck Thus, these two species and probably all slender, extremely long and extensible. Snout other American western Atlantic species be- bilobed at tip. Foot narrow, extremely elon- long in the genus Bittiolum, which is also rep- gate anteriorly, three times snout length, and resented by several eastern Pacific species, with crescent-shaped propodium (Fig. 2). such as Bittiolum fastigiatum (Carpenter, Deep crescentic transverse slit formed by two 1864). lips in anterior foot and leading via a central Because the two Bittiolum representatives duct into large anterior mucus gland (Fig. studied, B. varium and S. alternatum, are so 11 A, amg). Corners of anterior pedal lips ex- alike, they are treated jointly in the section tending laterally and posteriorly forming uncil- below. iated undulating epipodial skirt (Fig. 11A-B, es) delineating lateral groove between epipo- Bittiolum varium (Pfeiffer, 1840) dium and sole; epipodial skirt weakly scal- (Figs. 10-11) loped posteriorly (Fig. 2), forming lanceolate opercular lobe, scalloped around edges. Cili- Cerithium varium Pfeiffer, 1840: 256. ated gutter (Fig. 11B, eg) in both sexes Cerithium columellare Orbigny, 1842: pi. 23, emerging from floor of right side of palliai cav- figs. 13-15; 1845: 244 (In part; syntypes ity, running down right side of foot leading into Bf^NH). epipodial groove. Ciliated gutter terminafing Cerithium gibberulum C. B. Adams, 1845; 5 in small glandular ovipositor (Fig. 11B, ovp) at (Lectotype MCZ 186078, type locality Ja- edge of foot in females. Posterior third of sole maica). with median longitudinal slit leading into mas- Bittium varium (Pfeiffer). Tryon, 1887:152, pi. sive mesopodial mucus gland (Fig. 11 A, 29, fig. 86; Perry, 1940: 134, pi. 28, fig. mmg), extending deeply into head foot up to 202. nerve ring and cephalic hemocoel. Opercu- Cerithium (Bittium) gibberulum (C. B. Ad- lum (Fig. 10F, G) corneous, light tan, circular- ams). Kobelt, 1898: 245-246, pi. 43, ovate, paucispiral with subcentric nucleus. fig. 1. Mantle edge (Fig. 11B, me) bilobed, smooth. GENERIC REVIEW OF BITTIINAE 289

FIG. 10. SEM micrographs of BiWolum variumUom Ft. Pierce, Florida (USNM 77639). A, B, D, E, two shells showing sculptural variation and shell shape; length 3.2 nnm; C, immature shell, length 2.8 mm; F, G, operculum, bar = 0.2 mm; H, sculpture of early whorls, bar = 0.3 mm; I, protoconch, bar = 88 (xm. without papillae, slightly scalloped, iridescent marginal tooth with 6 small inner denticles. at edges. Midesophagus with wide ciliated dorsal food groove; posterior esophagus narrow. Palliai Cavity: Osphradiunn wide, one-third ctenidial length, weakly monopectinate, com- Nervous System: Cerebral ganglia slightly prising snnall, dorsally placed pectins, flanked larger than pedal ganglia and with short con- on each side by weak ciliated strip. Ctenidium nective (about one-third cerebral ganglion comprising long, triangular filaments with soft length). Pedal ganglia nearly fused at connec- rods and mucus glands. tive, each with posterior statocyst; two pairs of accessory pedal ganglia present: pair of small Alimentary System: Radula (Fig. 11C) short. propodial ganglia, and larger pair of metapo- Rachidian tooth (Fig. 11D) with cutting edge dial ganglia. Subesophageal connective be- of 3 small denticles on each side of central tween subesophageal ganglion and left pleu- cusp. Lateral tooth (Fig. 11D) with two outer ral ganglion equal in length to left pleural and 3-4 inner denticles. Inner marginal tooth ganglion; supraesophageal connective about with 3-4 inner and 2-3 outer denticles. Outer equal in length to subesophageal connective. 290 HOUBRICK

FIG. 11. SEM micrographs of BiWolum varium from Ft. Pierce, Florida (USNM 776639). A, B, critical point dried specimens showing external anatomical features of headfoot, bar = 0.2 mm; C, mid-section of radula, bar = 21 |i,m; D, detail of rachldian and lateral teeth, bar = 7 \¡.m. amg = anterior mucus gland; eg = ciliated groove; eps = epipodial skirt; I = lip of mouth; mmg = metapodial mucus gland; op = operculum; ovp = ovipositor.

Right zygoneury between subesophageal anterior as seminal vesicle, containing dimor- and right pleura! ganglion. phic sperm. Males producing crescent- shaped spermatophore with flared bifurcate Reproductive System: Ducts of testicular fol- end and pointed, filamentous tip. Spermato- licles joining to form spermatic duct, moving phores containing both eu- and parasperma- GENERIC REVIEW OF BITTIINAE 291 tozoa. Ovary cream colored, overlying brown (Fig. 11B, ovp) is barely visible only during the digestive gland, extending forward to stom- breeding season, but is basically the same as ach. Palliai oviduct open, but closed in far that observed in Bittium. The massive posterior portion. Common aperture to open- metapodial mucus gland located in the pos- ing of spermatophore bursa in lateral lamina terior part of the sole differs from that seen in anterior to opening of sperm pouch and open- tttibittium species, in which the slit is much ing of seminal receptacle located on edge of longer. This gland secretes a string of mucus medial lamina one-third from posterior of lam- by which the animal can suspend itself in the ina. Opening to spermatophore bursa not ad- algae, but the thread does not have the ten- jacent to opening on medial lamina, but lo- sile strength of the mucous threads produced cated one-third back from anterior of lateral by members of the Litiopidae (Houbrick, lamina. Spermatophore bursa comprising cil- 1987b). Except for major differences in exter- iated and high vacuolated epithelial cells. nal features, the radula and internal anatomy Spawn mass composed of spirally wound thin of Bittiolum varium is quite similar to that of jelly string containing many small eggs 100- Bittium reticulatum. The radula differs only mi- 120 [i.m in diameter, hatching as veliger lar- nor details (Table 2). Although Bittiolum var- vae, becoming planktotrophic. ium primarily is a grazer of epiphytic microal- gae, Marcus & Marcus (1963: 79) have shown that the snail can use its anterior Bittiolum alternatum (Say, 1822) ctenidial filaments for particle feeding while Turrltella alternats Say, 1822: 243. stationary. Pasithea nigra Totten, 1834; 369, figs. 7a, b. Marcus & Marcus (1963: 88-89) found four Bittium nigrum (Totten), Gould, 1870: 321, fig. spindle-shaped spermatophores, each 1 mm long and 0.06 mm wide, in the bursa of the 590. Bittium alternatum (Say), C. W. Johnson, lateral lamina in Bittiolum varium, and noted that the spermatophores dissolve in this 1915: 127. Diastoma alternata (Say), Abbott, 1974: 107, bursa. The location of the spermatophore bursa in the lateral lamina is a unique feature fig. 1037. among cerithioidean taxa, and this layout is probably the same among other members of Description the Bittium-group, in which the bursa in the lateral lamina has been confirmed. However, This species is essentially the same as spermatophores have not been observed in Bittiolum varium, described above, although this bursa in any other species. the shell differs slightly in being more pupoid Bittiolum varium lays its eggs mostly on and less narrowly elongate. seagrasses. In the Indian River, Florida, I ob- served numerous irregular egg masses com- prising strands of eggs embedded in a loose Remarks jelly matrix deposited on Halodule grass blades and on ramose algae. In the spring, Marcus & Marcus (1963) thoroughly de- nearly all adults were ripe and egg laying con- scribed the anatomy of Bittiolum varium in tinued through the summer months tapering Brazil. My work on populations of this species off in September. from Florida basically confirms their detailed Bittiolum varium has been the subject of a observations. In addition, the basic anatomy number of ecological investigations. Virnstein of the Brazilian and Florida specimens is very & Curran (1986) measured the colonization similar to that of Bittiolum alternatum from the time of this species in seagrasses in the In- American northeastern coast, suggesting that dian River, Florida. Hardison & Kitting (1985) the latter is probably a sister taxon of Bittiolum found that Bittiolum varium fed primahly on varium. diatoms and coralline algae in seagrass Bittiolum is the only genus studied in which meadows of the northwest Gulf of Mexico. the mantle edge is smooth, with no trace of Despite the high population densities of this papillae, a character noted by Marcus & Mar- snail (3,000/m^), little impact on its food could cus (1963). A wavy epipodial skirt and nar- be detected. In Chesapeake Bay, Van Mont- rowly elongate anterior foot are also distinc- frans et al. (1982) found that the grazing ac- tive external features (Fig, 2) of both tivities of Bittiolum varium, which selectively examined Bittiolum species. The ovipositor eats diatoms from blades of marine grasses. 292 HOUBRICK could have important implications for the Institute, St. Petersburg; type locality, abundance and distribution of Zostera. Sitka, Alaska). Bittiolum varium has a wide range in the Bittium (Stylidium) eschrichtii icelum Bartsch, western Atlantic, occurring from Chespa- 1907:178 (Holotype USNM 15209a; type peake Bay south to Florida and the Gulf of locality, Neah Bay, Washington); 1911: Mexico, throughout the Caribbean, and south 388, pi. 57, fig. 3; Ruhoff, 1973: 81. to Brazil. Bittium eschrichtii (Middendort). Oldroyd, 1927: 18-19, pi. 79, fig. 4. STYLIDIUM DALL, 1907 Bittium (Stylidium) eschrichtii (Middendort). Stylidium Dall, 1907: 178 (Type species by Abbott, 1974: 106, fig. 1010. original designation: Bittium eschrichtii Middendort, 1849). Thiele, 1929: 211; Description Wenz, 1940; 757; Abbott, 1974; 106. Shell (Fig. 12): Shell large, turreted, reaching Diagnosis 17.5 mm in length, comprising 9-11 convex Shell relatively large, dirty chalky white, whorls. Protoconch (Fig. 12G) has two smooth, weakly sculptured with four broad smooth whorls. Early whorts (Fig. 12E-G) spiral cords defined by incised lines. Proto- sculptured with three spiral bands. Adult conch unsculptured. Snout twice length of whorls sculptured with 4 weak, widely flat- cephalic tentacles. Epipodial skirt poorly de- tened spiral bands separated from one an- veloped, smooth along edges, but opercular other by deep incised spiral grooves. Penul- lobe with small, pointed papillae. No metapo- timate whorls with 5 wide, spiral, weak bands. dial mucus gland. Osphradium non-pectinate. Suture well defined, slightly counter-sunk into Common aperture to sperm bursa and semi- each abapical whorl. Body whorl a little less nal receptacle in edge of anterior third of me- than one-third shell length, sculptured with dial lamina of palliai oviduct. Openings to about 8 broad spiral cords and incised lines. sperm bursa and seminal receptacle well- Shell base weakly constricted at base; ante- separated. Long ciliated ridge tract in lateral rior siphon broad and shallow. Aperture ovate lamina of palliai oviduct. Development direct. having concave columella with weak callus; outer lip of aperture circular, crimped where Remarks spiral grooves end. Shell color chalky white- gray, covered by thin tan periostracum. This genus is represented by species living in cold-water habitats from California north to Animal: Base color dirty white with trans- Alaska. The shell is dull and chalky under the verse black stripes on snout, head, and epi- periostracum. Shell length can be quite large podium (Fig. 14A). Ciliated epithelial strip run- (Table 3) for a member of the Bittiinae, and ning from mantle cavity floor on each side of the large smooth protoconch, without sinusig- headfoot and ending beneath peduncle of eral notch, is indicative of direct development. each cephalic tentacle. Ciliated gutter on right At first glance, the shell of Stylidium does side of foot in females ending in small pink, not appear to fit the Bittium-group mold. How- glandular ovipositor at foot edge. Snout very ever, anatomical features, such as the epipo- long, twice length of cephalic tentacles, wide, dial skirt, large opercular lobe (Fig. 2) and pal- bibbed at tip. Eyes very small. Lateral epipo- liai gonoduct configuration unmistakably dial skirt with minute pointed papillae along place it Into the Bittiinae. The common aper- edge of posterior third of foot; opercular lobe ture to sperm pouch and seminal receptacle long, pointed posteriorly, darkly pigmented is unusual in being located in the far anterior and with small pointed papillae along edge edge of the medial lamina of the palial ovi- (Fig. 2). Anterior foot crescent-shaped with duct, and not adjacent to the opening of the long slit along edge leading into centrally spermatophore bursa of the lateral lamina. placed, ovate mucus gland deep within propo- The length of the ciliated ridge' tract of the dium. No metapodial mucus gland. Opercu- lateral lamina is also atypical. lum (Fig. 12H, I) thick, ovate, paucispiral, with eccentric nucleus. Mantle edge bilobed, with Stylidium eschrichtii (Middendortf, 1849) small papillae, and with slightly elongate ex- (Figs. 12-14) halant siphon. Mantle roof folded longitudi- Turriteila eschrichtii Middendort, 1849: 396- nally over exhalant siphon forming dorsal, 397, pi. 11, fig. 1 (Holotype, Zoological posteriorly extending ridge. GENERIC REVIEW OF BITTIINAE 293

FIG. 12. Stylidium eschrichtii \rom Carmel, California. A-D, two shells showing sculptural variation (USNM 804376), 22.4 and 20.2 mm length, respectively; E, F, SEM micrographs of immature shells showing early sculptural patterns, bar = 0.5 mm; G, SEM micrograph of protoconch and early whorls, bar = 0.3 mm; H, I, SEM micrographs of operculum, showing eccentric nucleus and attachment scar, 2.4 mm length.

Palliai Cavity: Osphradiunn tan, vermiform, 3 inner denticles, and 3-5 outer denticles; in- non-pectinate, extending length of palliai cav- ner marginal tooth with 4-5 inner and 3 outer ity, but slightly shorter than ctenidium. Ctenid- denticles. Paired salivary glands vermiform, ium pink, comprising long, finger-like fila- loosely compacted, lying mostly anterior to ments twice length of their attached bases. nerve ring, but beginning behind it as thick swellings, and passing through as thin tubes. Stomach large, about one whorl in length; in- Alimentary System: Radular ribbon (Fig. ternally with large sorling area and roundish 13A) short. Lateral tooth (Fig. 13B) w/ith long central pad; single opening to digestive gland lateral basal extension and cutting edge with on right side of pad; 6-7 large transverse ribs 294 HOUBRICK

FIG. 13. SEM micrographs of ra(iit\a oi Stylidium eschrichtii{USNM 804376); A, section of mid-raduiar ribbon with marginal teeth folded back, bar = 38 |j,m; B, detail of rachidian and lateral teeth, bar = 12 y.m. on left side of pad, posterior to cuticular gas- medial lamina. Very long ciliated ridge tract tric shield; short, wide style sac one-half stom- beginning in anterior part of lateral lamina, ach length, separate from intestinal opening. leading into posterior spermatophore bursa. Intestine opening separated from lumen of Spawn comprising thin gelatinous string style sac by typhlosole ridge. wound into irregular mass. Eggs 0.2 mm in diameter. Development direct. Nervous System (Fig. 14); Nerve ring large with thick commissure connecting cerebral Remarks ganglia. Dialyneury (Fig. 14B, d) between left palliai nerve and nerve arising from supra- esophageal ganglion. Supraesophageal con- Several subspecific taxa have been de- nective (Fig. 14A, sec) twice length of right scribed, but it is debatable if all of these nom- pleural ganglion. Subesophageal ganglion inal taxa are good subspecies or merely cli- (Fig. 14A, sbe) closely adjacent to left pleural nal/ecophenotypic varieties of Stylidium ganglion. eschrichtii. Abbott (1974) synonymized the subspecies icelum Bartsch with S. eschrichtii. Reproductive System Posterior half of palliai Stylidium eschrichtii is characterized by its oviduct with thick, white, opaque albumen chalky gray, smooth shell sculptured with gland comprising flocculant transverse glan- broad flattened spiral cords. The protoconch dular ridges; mid-section of palliai oviduct with is large, unsculptured, and lacks a sinusigeral thin, weak glandular transparent walls; very notch (Fig. 12G). The ovate operculum (Fig. thick, opaque transverse glandular ridges 12H, I) with eccentric nucleus is a departure present in anterior third of palliai oviduct, from a more circular operculum with subcen- comprising capsule gland. Sperm gutter in tral nucleus, as seen in other bittiid species. anterior edge of medial lamina having elon- Shell length seems to vary greatly among gate common aperture to spermatophore populations, but some individuals can be very bursa and seminal receptacle. Openings to large, approaching 18 mm length (Table 3). sperm pouch and seminal receptacle within Large shell size appears to be more common common aperture well separated. Long tube in northern populations. within edge of medial lamina leading to pos- This species lives on intertidal to subtidal teriorly placed pouch-like seminal receptacle. rubble in cool waters of the northeastern Pa- Large sperm pouch with internal transverse cific. I observed a large intertidal population epithelial folds, occupying posterior third of living among the intertices of gravel and algae GENERIC REVIEW OF BITTIINAE 295 Diagnosis

Shell turreted, elongate, sculptured with ax- ial riblets and spiral beaded cords. Proto- conch with two spiral lirae. Varices not present on adult whorls. Operculum circular. Radular ribbon very small; radular teeth with many small denticles. Snout long; head with small cephalic tentacles and small eyes. Ovi- positor and ciliated groove on right side of foot absent. Mantle edge with long papillae. Epi- podial skirt very weakly developed. Osphra- sec dium vermiform, wide. Spawn comprising ses sbe 'sg large egg capsules, each attached to long B stalk and anchored together. Development di- rect.

Remarks

Bartsch (1911) divided ß/rt/um-group spe- cies from the American west coast into four genera: Bittium, Lirobittium, Semibittium, and Stylidium. His groups were defined only on superficial shell characters, such as the pres- ence or absence of varices, protoconch sculpture, and axial and spiral sculpture. Many of the species Bartsch (1911) included under his generic scheme have been ignored or referred by subsequent authors to different generic taxa. The genus Lirobittium Bartsch, from the FIG. 14. Anatomical features of Stylidium eschrich- temperate eastern Pacific, was based on mi- tii. A, head and anterior foot, sfiowing pigment pat- tern; B, position of salivary glands relative to nerve nor shell sculptural characters: Bartsch ring, d = left dialaneury; leg = left cerebral gan- (1911: 384) noted that the defining characters glion; Ipg = left pleural ganglion; Isg = left salivary of Lirobittium were a protoconch with two spi- gland; rcg = rigfit cerebral ganglion; rsg = right ral lirae and the absence of varices from the salivary gland; rpg = right pleural ganglion; sbe = adult whorls. These features were also men- subesophageal ganglion; sec = supraesophageal tioned by Gründel (1976: 54), who addition- connective; sag = supraesophageal ganglion. ally noted that of the two primary spiral cords, the abapical one was inserted a little later. at Carmel, California. According to Slrath- Gründel (1976: 54-56) assigned Cacozeli- mann (1987), Stylidium eschriciitii has direct ana and Stylidium (with a query) as subgen- developnnent. Spawn is deposited on the sub- era of Lirobittium. He indicated that Cacoze- strate in gelatinous masses (presumably liana differed from Lirobittium by the formation comprising coiled strings) containing egg cap- of varices, and Stylidium by the suppression sules measuring 0.2 ^j^m diameter in which or complete absence of axial ribs. It has been the embryos undergo direct development, shown herein that the Cacozeliana is sepa- passing through the veliger stage and hatch- rated from Lirobittium by many significant ing as small snails. characters. The above history of Lirobittium shows that Z./flOß/TT/L'/W BARTSCH, 1911 much of the confusion regarding the place- ment of the numerous California species Lirobittium Bartsch, 1911: 384 (Type species stems from the original superficial generic de- by original designation, Bittium catalinen- scriptions based solely on shell morphology. sis Bartsch, 1907). Thiele, 1929: 211; It is obvious that the characters derived by Wenz, 1940: 757; Abbott, 1974: 106; these authors from minor sculptural details Gründel, 1976: 54. hardly seem to be of generic weight and have 296 HOUBRICK resulted in poorly defined, ambiguous genera the inhalant siphon has large paddle-shaped with broad or discordant limits, and that have papillae. The buccal mass is small, and the been used in varying combinations. Although radula minute, about one-thirteenth the shell shell sculpture may have some value at the length. The rachidian tooth has a triangular specific level, it is generally not useful at the basal plate with a long glabrella and is as generic level, especially in cerithiids. Not a wide as tall; there is a deep concave inden- single author has included radular or opercu- tation and a cutting edge with a long pointed lar characters and no mention is made of an- central cusp flanked on each side by 4-5 atomical features in the definition of genera. small denticles. The lateral teeth are deeply Abbott (1974: 106) considered both Bittium concave on the top, have a wide basal plate catalinense and B. subplanatum to be syn- with a large central buttress, and have numer- onyms of Lirobittium attenuatum Bartsch, ous small denticles. The marginal teeth are 1911, but gave no reasons for this decision. slender, and serrated along their tips with Hertz (1981: 40) showed that Lirobittium sub- many small pointed denticles (Fig. 15). planatum (cited as Bittium) was a valid spe- cies. I have examined two species of Lirobit- Lirobittium subplanatum (Bartsch, 1911) tium: L. catalinense.(one dried specimen) and (Figs. 15-17) well-preserved material of L. subplanatum. Observations on the poorly preserved, dried Bittium (Semibittium) subplanatum Bartsch, animal of L. catalinense are included because 1911: 395-396, pi. 57, fig. 5 (Holotype, it is the type species of the genus, but the bulk USNM 160076; type locality, Catalina Id., of the descriptive anatomical characters of Li- California); Oldroyd, 1927: 23: Ruhoff, robittium are derived from study of L. sub- 1973: 130. planatum. The two species are anatomically Bittium subplanatum Bartsch. Dali, 1921:146; very similar, have similar radulae, and are un- Hertz, 1981:40, figs. 23-27. doubtedly congeneric. The above diagnosis Bittium subplanatum Bartsch. Oldroyd, 1927: and following specific descriptions represent 23. an integrated analysis of generic characters, Bittium (Lirobittium) subplanatum (Batsch). based on these two species. Abbott, 1974: 106.

Lirobittium catalinense Bartsch, 1907 Description Bittium catalinensis Bartsch, 1907; 28, pi. 57, Shell (Fig. 15): Shell elongate, turreted, com- fig. 13 (Holotype: USNM 165232, type lo- prising 8-9 moderately inflated whorls. Pro- cality: Santa Barbara, California [Pleis- toconch (Fig. 15) about 1.5 whorls, well tocene]); Abbott, 1974: 106, fig. 1013. rounded, smooth. Early whorls sculptured Bittium (Lirobittium) catalinense Bartsch, with two major spiral lirae, soon crossing over 1911:402-403, pi. 51, fig. 1. axial riblets (Fig. 15). Adult whorls sculptured with three major spiral cords crossed over by numerous thin axial ribs (24-26), forming Remarks cancellate appearance; small beads occur- ring at crossover points. Body whort (Fig. 15) The type species of this genus is a Pleis- sculptured with four major spiral cords and tocene fossil, but Bartsch (1911) described numerous axial ribs; moderately constricted many subspecies, some of which are Recent. at base. Shell base with about 7 spiral cords. Bittium cataliense is now regarded as a syn- Aperture ovate with oblique columella and onym of "Bittium"attenuatum Carpenter, 1864 curved, thin outer lip. Anterior canal moder- (Abbott, 1974: 106). ately developed; anal canal weak. Shell color Examination of a reconstituted, dried spec- white, covered with brown periostracum. imen of the type species of Lirobittium, Bittium catalinense (= Bittium attenuatum), showed Animal (Fig. 16A, B): Animal pure white with that the animal is basically the same as Liro- pink buccal mass showing through snout. bittium subplanatum. It is relatively unpig- Head large with very large, wide, extensible mented, has a large, broad snout, bilobed at snout, dorso-ventrally flattened, bilobed at tip; the anterior end and short cephalic tentacles, cephalic tentacles small, a little less than one- about half the snout length. The mantle edge third snout length, each with small black eye has many long papillae along its dorsal and adjacent to opaque white spot at tentacular lateral sides, while the mantle edge forming peduncular base. Snout ringed with many GENERIC REVIEW OF BITTIINAE 297 Stomach with central pad, gastric shield, short style sac and crystalline style; one opening to digestive gland. Nervous System: Cerebral ganglia joined by short connective. Pleural ganglia close to ce- rebral ganglia; left pleural ganglion connected to subesophageal by very short connective. Supraesophageal connective about two- thirds length of right pleural ganglion. Reproductive System (Fig. 16A); Testis white, producing dimorphic sperm; ovary cream-yellow containing large ova, 0.5 mm in diameter. Glandular portion of female palliai oviduct comprising many transverse folds, posterior opaque white portion comprising al- bumen gland (Fig. 16A, ag), and anterior, transparent greyish-white portion comprising capsule gland (Fig. 16A, eg). Anterior two- FIG. 15. SEM micrographs of shells of Lirobittium thirds of edge of medial lamina with large subplanatum from Palos Verdes, California (USNM sperm gutter (Fig. 16A, sg) leading into deep 881021). A, bar = 1.8 mm; B, detail of protoconch slit containing two openings: anterior opening and early teleoconch sculpture, bar = 0.6 mm; C, (Fig. 16A, osp) into large sperm bursa and bar = 1.8 mm. posterior opening (Fig. 16A, osr) into small deep, transverse epithelial folds (Fig. 16B). tubular sac-like seminal receptacle (Fig. 16A, Foot with very weak epipodial skirt and with- sr). Lateral lamina less glandular than medial out papillae or distinctive operculiferous lobe. lamina and with short ciliated ridge tract (Fig. No ciliated groove on right side of foot; no 16A, crt) leading into opening of spermato- ovipositor. Anterior of sole crescent shaped phore bursa (Fig. 16A, osb), adjacent to with deep transverse slit marking entrance to openings on medial lamina. Spermatophore anterior mucus gland. No metapodial mucus bursa (Fig. 16A, sb) small, elongate, sac-like. gland. Mantle edge bilobed, fringed with many papillae emerging from ventral side of Discussion mantle edge. Bartsch (1911) assigned this species to the Palliai Cavity: Osphradium brown, vermi- subgenus Semiblttium, and his assignment form, without pectins, wide, about one-third was followed by Dall (1921), Oldroyd (1927), the ctenidial width, nearly equaling ctenidial and Hertz (1981). Semiblttium is shown length. Ctenidium extending length of palliai herein to comprise a group of Eocene fossils cavity. Hypobranchial gland thick, comprising probably related to the extant Australian mo- transversely ridged glandular tissue. notypic genus Cacozellana, which differs con- Alimentary System: Mouth at tip of snout, de- siderably in anatomy from the California spe- fined by pair of fleshy pads. Buccal mass (Fig. cies. Abbott (1974) transferred this species, 16B, bm) pink, small, about one-third snout which he considered a synonym of BIttlum at- length. tenuatum Carpenter, 1864, to Lirobittium, but Radular ribbon (Fig. 17) small, about gave no reasons for doing so. one-ninth shell length. Rachidian tooth (Fig The shell is of moderate size (Table 3) and 17C) with large glabrella, long serrated cen- has a large protoconch sculptured with two tral cusp and 6 small denticles on each side. spiral lirae and lacking a sinusigeral notch. Lateral tooth (Fig. 17 B,C) with broad basal Although the shell of Lirobittium subplanatum plate; cutting edge has large denticle with 6 does not resemble that of Stylidlum es- inner denticles and 15-17 outer denticles. chrlchtil, the anatomical features of the two Marginal teeth (Fig. 17D) long, curving; inner species are quite similar. As far as can be marginal tooth with 15-19 inner denticles, seen in preserved material, Lirobittium sub- large central cusp and 5-6 outer denticles; planatum appears to have a very weak epi- outer marginal tooth same, but lacking outer podial skirt, but closer examination of living denticles. animals may show that this character is com- 298 HOUBRICK

FIG. 16. Lirobittium subplanatum. A, palliai oviduct, spread open to reveal details; B, head, showing broad snout, short cephalic tentacles and small buccal mass; C, dorsal view of attached spawn mass, showing individual capsules with enclosed embryos and attachment strands, ag = albumen gland; ant = anterior of palliai oviduct; bm = buccal mass; eg = capsule gland; cod = coelomic oviduct; crt = ciliated ridge tract; osp = opening to sperm pouch; osb = opening to spermatophore bursa; ovg = oviductal groove; sb = spermatophore bursa; sg = sperm groove; sp = sperm pouch; sr = seminal receptacle. pletely absent. The operculum also differs in Lirobittium subplanatum lives offshore on being more typically rounded than that of Sty- sandy-rubble bottoms. The shell is frequently lidium. severly eroded and abraided. The radula of Lirobittium subplanatum (Fig. Spawn morphology of Lirobittium is unique 16) is very similar to that of Lirobittium atten- among Bittiinae (Fig. 17C) and is deposited uatum, but differs in havjng many more den- on pieces of rubble or empty shells. It com- ticles on the teeth. The exact dentition for- prises clusters of large egg capsules, each mula is given in Table 2. about 0.5 mm in diameter and containing one There has apparently been some difficulty embryo. Each egg capsule is connected by a in identifying this species, as it has been con- strand to a central attachment point so that sidered synonymous with a number of other the spawn mass looks like a group of small sympatric species, but Hertz (1981) has balloons with their strings attached together. shown that it is a distinct, valid species. As Embryos revolve slowly with their capsules, mentioned above, the radula is distinct. where they pass through the veliger stage. GENERIC REVIEW OF BITTIINAE 299

FIG. 17. SEM micrographs of radula of LirobiWum subplanatum (USNM 881021). A, radular ribbon with marginal teeth spread open, bar = 35 |j.m; B, half row showing rachidian and lateral teeth, bar = 19 [¡.m; C, detail of dentition of rachidian and lateral teeth, bar = 10 |xm; D, detail of dentition of marginal teeth, bar = 12 (im. finally hatching out as small snails. Develop- nor Tuomey, 1848; nor J. de C. Sowerby, ment is direct (pers. obs.). in Dixon, 1850). Thiele, 1929: 211; Wenz, 1940: 756; Gründel, 1976: 56-57. CACOZEUANA STRAND, 1928 Cacozelia Iredale, 1924: 246 (Type species ? Semibittium Cossmann, 1896: 29 (Type by monotype: Cerittiium lacertinum species by original designation: Cerith- Gould, 1861); not Cacozelia Grots, 1878 ium cancellatum Lamarck, 1804; not [Lepidoptera]. Thiele, 1929: 211 ; Murray, Semibittium Bronn, 1831 ; nor Lea, 1842; 1969: 111. 300 HOUBRICK Cacozeliana Strand, 1928: 66 (new name for Cerithium granarium Kiener, the living type Cacozelia Iredale, 1924). Wenz, 1940: species of Cacozeliana from southern Austra- 756, lia redescribed herein; however, because the Lirobittium {Cacozeliana) Strand. Gründel, anatomy of the fossil is unknown, it is impos- 1976: 54-55. sible to declare with confidence that the two species are congeneric. Gründel (1976: 56) Diagnosis considered the Eocene genus Semiblttlum to be separate from Cacozeliana. He noted that Shell large, elongate with many weakly in- the shell of Semiblttlum species has a slight flated whorls, sculptured with four beaded varix on the lip of the protoconch followed by spiral cords per whorl and having overall pus- an almost simultaneous insertion of the three tuloso appearance. Protoconch unsculptured primary spiral cords. The name Cerithium except for microscopic subsutural pustules, cancellatum Lamarck is preoccupied, and but large sinusigeral notch present (Fig. 18F). needs a replacement name. Moreover, the Operculum circular-ovate, paucispiral with name Semiblttlum cannot be used because it subcentric nucleus and fringed edges. Epipo- is thrice preoccupied. The possibility that Ca- dial skirt with smooth edges. Snout short, nar- cozeliana granarla is a living survivor of the row. Opercular lobe lanceolate and with lon- Eocene genus Semiblttlum represented by gitudinal median groove. Large ovipositor Cerithium cancellatum should be considered, gland on right side of foot. Osphradium bipec- because several other Tethyan Eocene cer- tinate. Salivary glands anterior to nerve ring. ithioidean genera survive among the living Rachidian tooth without glabrella. Openings Australian molluscan fauna; e.g., Diastoma to sperm bursa and seminal receptacle well Deshayes, 1850; Gourmya Fischer, 1884; separated. Seminal receptacle comprising Campanile Fischer, 1884; and Plesiotrochus several grape-like lobes. Fischer, 1878 (Houbrick, 1981b, 1981c, 1981d, 1990b, respectively). It is also notable Remarks that Cacozeliana falls out at the base of the cladogram (Fig. 1) as the closest taxon to the The genus Cacozelia was proposed by Ire- outgroup. Moreover, Cacozeliana is sepa- dale for Cerithium lacertinum Gould, a sub- rated from all other Bittium-group genera by jective synonym of Cerithium granarium five non-homoplastic synapomorphies (Fig. Kiener. The living Australian species is 1), further demonstrating its distinctiveness. thought to be congeneric with the Paris Basin Gründers (1976: 56-57) separation of Semi- Eocene species Cerithium cancellatum La- bittium from Cacozeliana was based on the marck, which is the type species of Semibit- order of the insertion of spiral lirae on the tlum Cossmann; however, as Cacozelia is a early whorls, but this character has not been junior homonym, the name Cacozeliana was shown to be of generic weight, and therefore subsequently proposed by Strand (1928) as a is not seriously considered herein. If Cacoze- replacement. The allocation of Cacozeliana liana is truly congeneric with Semiblttlum, the as a subgenus of Liocerithium by Gründel genus would date from the Eocene, when the (1976) was made on the observation that In latter was common in the Paris Basin fauna Cacozeliana, the fourth primary spiral cord is (Cossmann, 1906:138). Cacozeliana is today initially weaker than the three formed earlier, monotypic and confined to the temperate wa- whereas in Liocerithium all four are equally ters of southern Australia. The type species, strong. Gründel (1976) also pointed out that Cacozeliana granaría (Kiener), undoubtedly varices are present in the subgenus, whereas has the largest shell of any representative of they are absent in Lirobittium. These minor the subfamily Bittiinae and differs from other sculptural differences hardly seem appropri- species of the group in several ways: ate as generic-level characters; furthermore, 1. The short narrow snout (Fig. 20A) is dis- radular and anatomical characters of Cacoz- tinctive, as is the fringed operculum (Fig. eliana show that it is far-removed from Liro- 1BG). bittium. 2. The rachidian tooth of Cacozeliana gra- The type species of Semiblttlum, which is narla is unique, differing from other Bittiinae placed into synonymy with Cacozeliana with a members in lacking a glabrella on the basal query, is an Eocene fossil from the Paris Ba- plate. Additionally, the rachidian tooth lacks sin, Cerithium cancellatum Lamarck. This fos- concave sides and a strong pair of basal but- sil species is conchologically very close to tresses (Fig. 19B). Moreover, the lateral basal GENERIC REVIEW OF BITTIINAE 301 extensions of the basal plate are nearly ab- weak, microscopic subsutural pustules, no sent. spiral lirae, and with deep sinusigeral notch. 3. The palliai oviduct of Cacozeliana grana- Early whorls (Fig. 18H) sculptured with 3 spi- rla (Fig. 20C), while having a typical layout, is nosely beaded spiral cords alined to form unique among known palliai oviducts in the about 12-13 axial riblets. Adult whorls slightly S/ïï/um-group in having the seminal recepta- beveled abapically, defining weak suture. cle divided into several grape-like lobes (Fig. Body whorl one-third shell length, having 6 20C, sr) and in having a highly developed, spiral beaded cords and weakly constricted swollen anterior capsule gland (Fig. 20C, eg). base. Aperture ovate, small, about one-fifth As pointed out earlier, a grape-like seminal shell length. Columella concave with weak receptacle also occurs in some species of columellar callus and smooth, rounded outer Cerithium Bruguière, 1789, Rhinoclavis lip. Anterior canal short, narrow, well defined. Swainson, 1840, and in Dialak. Adams, 1861 Shell color white to tan, blotched with pink to (Houbrick, 1971, 1978, 1992, pers. obser.; reddish brown and having brown spiral bands Ponder, 1991), although this structure in Diala with white flecks (Fig. 180, D). Beads some- is not proven to be a seminal receptacle. This times white (Fig. 18A B). Periostracum light kind of seminal receptacle does not necessar- tan, thin. ily indicate relatedness among these groups: the bulging, grape-like morphology may be Animal (Fig. 20): Head, snout and epipodium due to the swollen state of the filled seminal pigmented tan with chocolate blotches, tiny receptacle and may represent sexual "ripe- white spots, and irridescent green. Oephalic ness" rather than a distinct morphological tentacles darkly pigmented, having many character state of the seminal receptacle. black spots, slender, elongate, aboui twice snout length. Snout narrow, short (Fig. 20A, sn) with flared bilobed tip. Mantle edge Cacozeliana granaría (Kiener, 1842) fringed with very small papillae each bearing (Figs. 18-20) white spot. Pair of ciliated strips emerging Cerithium granarium Kiener, 1842: 72-73, pi. from mantle floor and running to base of 19, fig. 3 (Holotype MNHNP; type local- cephalic tentacles on each side of headfoot. ity, "les côtes de Timor," in error, here Deep ciliated groove running down right side corrected and restricted to Albany, West- of foot to edge, ending in small flap in males. ern Australia); G. B. Sowerby, 1855: 879, Ciliated groove in females having thick glan- pi. 184, figs. 225-227; 1865: pi. 19, fig. dular strips on each side of groove, compris- 135; Kobelt, 1898: 249, pi. 23, fig. 9. ing ovipositor. Epipodial skirt poorly devel- Ceritliium lacertinum Gould, 1861: 368 (Ho- oped, smooth along edge, forming short lotype USNM 16571; type locality Syd- lanceolate opercular lobe with dorsal longitu- ney Harbor, New South Wales, Austra- dinal furrow and without papillae along edge. lia); 1862: 141; G. B. Sowerby, 1866: pi. Orescent-shaped propodial slit at edge of an- 18, fig. 128; Tryon, 1884: 155, pi. 30, fig. terior foot leading into deep oval anterior mu- 100; R. Johnson, 1964: 96, pi. 11, fig. 4. cus gland (Fig. 20A, amg). Longitudinal fold in Bittium granarium (Kiener). Tryon, 1887: 155, middle of sole, but no metapodial mucus pi. 30, fig. 98; Wells, 1984: 30-31. gland present. Operculum (Fig. 18G) circular- ovate, paucispiral, with subcentral nucleus. Synonymic Remarks Opercular spiral fringed with thin lamella (Fig. 18G). Kiener's (1842) name, granarium, predates Paillai cavity: Osphradium bipectinate, with Gould's (1861) lacertinum. Examination of the weak pectins. Osphradium equaling ctenidial holotypes of both taxa leaves no doubt that length. Ctenidium comprising light tan elon- the two are conspecific. gate, triangular filaments. Hypobranchial gland thick, comprising irregular transverse Description glandular folds, secreting large amounts of mucus. Stielt (Fig. 18): Shell large, elongate, tur- reted, reaching 24 mm in length comprising Alimentary system (Fig. 19B): Buccal mass 12-13 nearly flat-sided whorls sculptured large, filling snout cavity, having small jaws with four beaded spiral cords. Protoconch and short radula (Fig. 19A). Rachidian tooth (Fig. 18F) comprising two smooth whorls with (Fig. 19B) with rectangular basal plate lacking 302 HOUBRICK

FIG. 18. Cacozeliana granaría ^rom King George Sound, Western Australia (USNM 858551). A-D, two shells showing variation in color pattern and sculpture, length 22.4 mm and 20.2 mm, respectively; E, SEM micrograph of immature shell, bar = 0.6 mm; F, SEM micrograph of protoconch, bar = 16 (xm; G, SEM micrograph of operculum, bar = 0.8 mm; H, SEM micrograph showing early sculpture, bar = 0.8 mm.

strong basal lateral buttresses, with straight many transverse internal epithelial folds com- base and equal in length to top of tooth; cut- prising esophageal gland. Stomach with one ting edge w/ith small central cusp flanked by digestive gland opening to left of large central two denticles on each side. Lateral tooth (Fig. pad dividing left sorting area from right gastric 19B) with one inner denticle and 3-4 outer shield complex. Style sac separated from in- denticles. Inner marginal tooth with 5-6 inner testinal opening by large typlosole fold. denticles and 3-4 outer denticles. Outer mar- ginal tooth (Fig. 19A) with 4 inner denticles. Nervous System (Fig. 20, B): Cerebral gan- Salivary glands (Fig. 208, Isg, rsg) paired, glia joined by short connective, one-third the vermiform, coiled, lying anterior to nerve ring. ganglion length. Subesophageal ganglion Midesophagus expanded laterally having very close to left pleural ganglion. GENERIC REVIEW OF BITTIINAE 303

FIG. 19. Radula of Cacozeliana granaría from King George Sound, Western Australia (USNM 858551). A, mid-section of radula, bar = 60 ^.m; B, details of racfiidian and lateral teeth, bar = 15 ^.m.

1mm osp

rpg 0.25mm

FIG. 20. Anatomical features of Cacozeliana granaría. A, head and foot anterior, showing narrow snout; B, position of salivary glands anterior to nerve ring; C, paillai oviduct, spread open to reveal interior details, a = anterior end of palliai oviduct; ag = albumen gland; eg = capsule gland; cod = coelomic oviduct; ctr = ciliated ridge tract; Isg = left salivary gland; osb = opening to spermatophore bursa; osp = opening to sperm pouch; osr = opening to seminal receptacle; rpg = right pleural ganglion; rsg = right salivary gland; sb = spermatophore bursa; sg = sperm groove; sp = sperm pouch; sr = seminal receptacle.

Reproductive System: Male palliai gonoduct half; anterior half of male palliai gonoduct less thick, glandular, having wide transverse folds glandular, white but not opaque. Female pal- forming spermatophore organ in posterior liai oviduct (Fig. 20C) having seminal recep- 304 HOUBRICK tacle comprising several grape-like lobes in medial lamina (Fig. 20C, sr). Openings to the sperm pouch (Fig. 20C, osp) and seminal re- ceptacle (Fig. 20C, osr) separated by long cil- iated groove. Ciliated ridge tract (Fig. 20C, ctr) beginning behind anterior capsule gland (Fig. 20C, eg) comprising many swollen trans- verse elements. Opening to spermatophore bursa (Fig. 200, osb) in lateral lamina adja- cent to opening of sperm pouch in medial lamina. Spawn mass comprising a jelly string containing many encapsulated eggs, 0.1-0.13 mm diameter, wound into flattened coil about 20 mm wide. Eggs opaque, white, each within hyaline capsule. Development in- direct with free swimming veliger stage.

Discussion

Although the shell of Cacozeliana granaría (Fig. 18) looks very much like those of some Centhium species, the weak epipodial skirt, palliai oviduct, and other anatomical features are very typical of members of the Bittiinae. The protoconch, as indicated by Gründel (1976), differs from those of most other gen- era in being nearly smooth, and in lacking any spiral threads (Fig. 18F; Table 3), but it does have a deep sinusigeral notch, indicative of planktotrophy. Stylidium species also have a smooth protoconch. The operculum of Caco- zeliana is unusual in having a thin lamellar- FIG. 21. SEM micrographs of shell of Argyropeza like fringe along its spiral (Fig. 186). The shell divina Melvill & Standen, from Refugio Id., Tanon of this species is undoubtedly the largest of Str., Philippines (USNM 302513); A, B, apertura! and dorsal views of adult shell, 6.3 mm length; C, any member of the S;Y//t/m-group (Table 3), protoconch showing sculpture and sinusigeral but the aperture is very small in relation to the notch, bar = 1 mm. shell length. There is much color variation within populations. The early life history of this species has similar habitats in Sydney Harbor and Botany been described by Murray (1969), who illus- Bay, New South Wales. trated the spawn (1969; pi. 17). The spawn comprises a coiled gelatinous thread contain- ARGYROPEZA MELVILL & ing encapsulated eggs that hatch as plank- STANDEN, 1901 totrophic veligers. Murray (1969) stated that 8-9 days after deposition, veliger-stage em- Argyropeza Melvill & Standen, 1901:371-372 bryos hatched out and were maintained in (Type species by original designation, Ar- sea water containers for up to 10 weeks. gyropeza divina Melvill & Standen, 1901 ). Cacozeliana granaría is found in the shal- Thiele, 1929: 212; Wenz, 1940: 757; low subtidal, temperate waters of southern Gründel, 1976: 44; Houbrick, 1980a: 2. Australia where it is common among Posi- donia, Zostera, and other sea grasses. It also Diagnosis occurs on moderately exposed and sheltered shores, on sandy-muddy bottoms, under Shell small, turreted, thin and vitreous, stones, and on rocky areas. I observed large sculptured with axial and spiral elements, va- populations of this species living on algal rices, and with many small nodules. Proto- mats and on Posidonia grass blades in King conch comprising three and a half whorls with George Sound, Western Australia, and in deep sinusigeral notch; sculptured with two GENERIC REVIEW OF BITTIINAE 305

FIG. 22. SEM micrographs of radula of Argyropeza divina (USNM 302513), A, radular ribbon with marginal teeth spread open, bar = 100 |xm; B, half row, bar = 50 \¡.m. spiral cords and many minute subsutural tuted, dried specimens. The shell and radula folds. Aperture ovate with well-developed, of the type species, Argyropeza divina Melvill short anterior canal. Operculum corneous, & Standen, 1901, are shown in Figures 21 subcircular, paucispiral, with subcentral nu- and 22. I do not agree with Powell's (1979) cleus. Snout broad with large cephalic tenta- suggestion that Tasmalira Dell, 1956, may be cles and large eyes. Foot with anterior mucus closely related to Argyropeza, because the gland. Mantle edge papillate. Palliai gono- shell morphology does not appear to fit the ducts open. Radula taenioglossate; rachidian limits of the genus. Argyropeza is tentatively tooth wider than tall; lateral tooth with trans- assigned to the Bittiinae until more complete verse ridge on basal plate; marginal teeth anatomical information is available. slender, scythe-shaped. VARICOPEZA GRÜNDEL, 1976 Remarks Varicopeza Gründel, 1976: 46 (Type species An alpha-level review of Argyropeza has by tautonomy, Varicopeza varicopeza been published by Houbrick (1980a), which Gründel, 1976). Houbrick, 1980b: 525; should be consulted for details about taxon- 1987: 85. omy, morphology and geographic distribution. The genus comprises five described species Diagnosis and several undescribed ones (pers. obser.). Members of this genus live on fine-grained Shell small, slender, turreted, vitreous, hav- substrates of deep water shelves and slopes, ing impressed suture, and sculptured with and not much is known about their biology. All strong spiral cords, weaker axial elements, examined species have small shells and pro- and many nodules. Protoconch having three toconchs sculptured with two spiral lirae, sub- and one-half smooth whorls, with weak, me- sutural pleats, and a deep sinusigeral notch dian spiral cord, minute subsutural pustules, (Fig. 21C; Table 3) indicative of a plank- and sinusigeral notch. Aperture ovate with totrophic larval stage. The anatomy of Argy- short, well-developed anal and anterior ca- ropeza species is virtually unknown except for nals. Operculum corneous, ovate, paucispi- superficial observations made from reconsti- ral, with subcentral nucleus. Radula taenio- 306 HOUBRICK 24) are well described, only a few external anatomical features are known. Varicopeza species occur at moderate subtidal depths on fine-grained substrates in the tropical Atlantic and Pacific. The shell sculpture of Varicopeza (Fig. 23A, B) is similar to that of Argyropeza species, differing chiefly in protoconch mor- phology. The aperture (Fig. 23A, B) is distinc- tive in having a large, flaring anal sinus. The radula (Fig. 24) has more denticles on the marginal teeth than in Argyropeza (Table 2). Grijndel (1976) suggested that Varicopeza was closely related to the extinct Jurassic ge- nus Cryptaulax and considered it to be a Re- cent representative of the of the extinct family Procerithiidae Cossmann, 1905. The shell and radula of Varicopeza pauxilla (A. Adams, 1854) is shown in Figures 23 and 24. This genus is tentatively assigned to the Bittium- group until more complete anatomical infor- mation is available.

ZEBITTIUM FmiAV, 1927 Zebittium Finlay, 1927; 381 (Type species by original designation, Cerithium exilis Hut- ton, 1873); Wenz, 1940; 756; fig. 2191; Powell, 1979: 132, fig. 32:1.

Diagnosis FIG. 23. SEM micrographs of shell of Varicopeza pauxilla (A. Adams, 1854) from Nagubat Id., E. Min- Shell very small, turreted, sculptured with danao, Philippines (USNM 276898). A, B, apertural beaded spiral cords, and weak axial riblets, and side views of adult shell, 8.1 mm length; C, having impressed suture. Aperture ovate with protoconch, bar = 100 Jí-HíI. weak notch-like anterior canal. Protoconch two and a half whorls, bluntly rounded, un- glossale with hourglass-shaped rachidian sculptured. tooth; lateral tooth with transverse ridge on basal plate; marginal teeth elongate, slender Remarks with denticulate sickle-shaped tips. Animal with large headfoot, elongate, wide snout, This genus was proposed without any de- long cephalic tentacles and very large eyes. fining characters, and was apparently intro- Deep ciliated groove on right side of foot. duced only to accomodate the New Zealand Mantle edge having short, thick papillae. species, Bittium exile Hutton and Bittium vit- reum Suter. The shell of Zebittium exile (Hut- Remarks ton, 1873) is shown in Figure 25. Zebittium was assigned as a subgenus of Bittium by The two known species of Varicopeza have Wenz (1940), who noted that the genus oc- been thoroughly described by Houbrick cured from the Miocene to the Recent of New (1980b, 1987a). These publications should be Zealand. The shell of the type species closely consulted for specific information about tax- resembles those of Bittium and Bittiolum spe- onomy and a detailed description of the type cies and does not appear to have any distin- species. The shell is of moderate length (Ta- guishing features of generic significance. The ble 3) and has a protoconch sculptured with unsculptured protoconch (Fig. 25D) appears one spiral lira and a shallow sinusigeral notch to indicate lecithotrophic development. No (Fig. 23C). Although the shell and radula (Fig. preserved material of this species was avail- GENERIC REVIEW OF BITTIINAE 307

FIG. 24. SEM micrographs of radula of Varicopeza pauxilla. A, section of ribbon with some marginal teeth spread open, bar = 50 i^m; B, detail of rachidian and lateral teeth, bar = 25 |j,m. able for study; therefore, the genus ZebiWum Remarks is included in this review only tentatively. This monotypic genus was recently pro- CASSIELLA GOFAS, 1976 posed and described by Gofas (1987), and his publication should be consulted for de- Cassiella Gofas, 1987: 109 (Type species by scriptive details of the genus and figures of original designation, Cassiella abylensis the type species. Cassiella abylensis does Gofas, 1987). not fit easily into the S/ff/um-group, although there are some resemblances. The shell of Diagnosis Cassiella abylensis (Fig. 26) varies highly in color pattern and in spiral sculpture (Gofas, Shell small, slender, turrited, sculptured 1987: 111). The shell morphology is unlike with spiral cords, without varices and with im- those of other members of the S/YI/um-group. pressed suture. Aperture ovate, without ante- No vestige of an anterior canal is present, and rior canal and simple outer lip. Operculum the shell morphology strongly resembles corneous, ovate, paucispiral, with subcentral those of some rissoids. The absence of an nucleus. Animal with bilobed snout and two anterior canal is also a feature of Cerithidium elongate cephalic tentacles. Foot short and Monerosato, a taxon I have excluded from broad without ovipositor or ciliated groove on Bittiinae. right side, and with large opercular lobe. Rad- The external anatomy of Cassiella abylen- ula taenioglossate; rachidian tooth with sis was depicted by Gofas (1987: figs. 10, 14, squarish basal plate, moderately concave on 15). The animal does not have epipodial ten- each side with small median glabrella, and tacles, although there is an inconspicuous having cutting edge with large central cusp groove around the foot, just above the edge of flanked by 3 smaller denticles on each side. the sole, which may be homologous with the Lateral tooth with large triangular cusp with epipodial skirt found in members of Bittiinae. one small inner denticle and 7-8 outer denti- The opercular lobes are said to be "massive" cles. Marginal teeth elongate, spatulate with (Gofas, 1987: 111 ), but they are not depicted curved tips; inner marginal teeth denticulate or labeled in the figures of the external anat- on both sides; outer marginal teeth lacking omy. The headfoot, operculum, and radula outer denticles. are not unlike those observed in other species 308 HOUBRICK

FIG. 26. SEM micrographs of shell of Cassiella abylensis Gofas, 1976, from Ceuta, Spain (USNM 869532); A, apertural view of shell, 2.3 mm length; B, dorsal view of shell, 2.5 mm length.

Estudos Açorianos "Alfonso Chaves." Work on the Western Atlantic species was done at the Smithsonian Marine Station, Link Port, Florida. I am grateful to Dr. Mary Rice and the staff of the marine station for their assistance throughout this project. This paper is Smith- FIG. 25. SEM micrographs of shell of Zebittium ex- sonian Marine Station contribution No. 272. ile (Hutton, 1873) from Long Bay, Auckland, New Work in Hawaii and Guam was supported by Zealand (USNM 681043); A, apertural view of adult two grants from the Smithsonian Secretary's shell, 4.7 mm length; B, dorsal view, 4.6 mm length; Research Opportunity Fund. I am grateful to C, immature shell, 4.4 mm length; D, protoconch, the University of Guam for laboratory space, bar = 0.25 mm. equipment and logistic support. I thank Dr. Michael Hadfield, of the University of Hawaii, of Bittiinae. There is no metapodial mucus for providing laboratory space at the Pacific gland, no ovipositor is indicated, and males Biomédical Research Laboratory, and tor his are aphallate (Gofas, 1987: 111). assistance with field work. A grant from the Pending further anatomical studies, the Smithsonian Secretary's Research Opportu- eastern Atlantic taxon Cassiella is tentatively nity Fund supported field and laboratory stud- assigned to Bittiinae with doubt. ies and attendance at the Workshop on Ma- rine Biology at Albany, Western Australia. I am Indebted to Dr. Fred Wells, Western Aus- ACKNOWLEDGEMENTS tralian Museum, Perth, for his assistance in the field. Dr. Henry Chaney, Mrs. Barbara This study was accomplished In many di- Chaney, and Mr. Paul Scott of the Santa Bar- verse places and with the help of many col- bara Museum of Natural History, provided lo- leagues and friends. I wish to thank Dr. Anto- gistic and field assistance In an heroic, alas nio Prias Martins, of the University of the unsuccessful, attempt to find living Lirobittium /Azores, for sponsoring me at the First Inter- specimens. I thank Don Cadlen for sending national Workshop of Malacology, held at Sao me live specimens of "Semibittium" sub- Miguel, Azores. This part of my study was planatum Bartsch from off Palos Verdes, Cal- supported by a grant of the Portuguese Uni- ifornia, and am grateful to Serge Gofas, Nat- \'ersity of the Azores and the Sociedade de ural History Museum, Paris, for sending shells GENERIC REVIEW OF BITTIINAE 309 of Cassiella abylensls. For technical assis- deren organische Einschlüsse, xii + 176 pp. tance (proofreading and SEM, and computer Heidelberg. macro design) I thank Shelley Greenhouse, BRUGUIÈRE, J. G., 1789-1792, Encyclopédie méthodique: Histoire naturelle des vers. Paris; National Museum of Natural History, Smith- Panckoucke. 1(1): 1-344 (1789); 1(2): 345-758 sonian Institution. Susanne Braden, National (1792). Museum of Natural History, Smithsonian In- CARPENTER, P. P., 1864, Supplementary report stitution, provided technical assistance with on the present state of our knowledge with regard SEM operation. John Wise provided valuable to the Mollusca of the west coast of North Amer- assistance in learning various aspects of the ica. Report of the British Association for the Ad- Hennig86 and GLADOS programs. Finally I vancement of Science, for 1863; 517-686. am grateful to Dr. Winston F. Ponder for crit- COSSMANN, M., 1884. Étude paléontologique et ically reading a draft of this paper and for stratigraphigue sur le terrain oligocène marin aux environs d'Étampes. Mémoires de la Société Gé- stimulating discussions and exchanges of ologique de France, (3)3(1): 1-187, pis. 1-6 data about anatomy and evolution of small- [With J. Lambert]. sized cerithioidean taxa. COSSMANN, M., 1889, Catalogue illustré des co- quilles fossiles de l'Éocène des environs de Paris. Quatrième fascicule. Annales de la So- ciété Royale Malacologique de Belgigue, 24: LITERATURE CITED 3-381, pis. 1-12. COSSMANN, M., 1896, Appendice No. 2 au Cata- ABBOTT, R. T., 1974, American seashells, 2ncl ed.: logue illustré des coquilles fossiles de l'Éocène 663 pp., illus. New York, Van Nostrand. des environs de Paris. Annales de la Société ADAMS, A., 1860. Mollusca Japónica: new species Royale Malacologique de Belgique, 31; 3-94, of Aclis, Ebala, DunlGastropoda and Scaphopoda. Bul- Sea. Journal of Conchology, 27: 221-223, pi. 7. letin of the Museum of Comparative Zoology, BLAINVILLE, H. M., 1816-1830. Vers et Zoo- Harvard ^8: 492 pp., 40 pis. phytes, in: Dictionnaire des sciences naturelles. DALL, W. H., 1892, Contributions to the Tertiary Pt. 2. Règne organisé. 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Paleontological Bul- senschaft, 2: 729-733. letin No. 15. Loney, Wellington. 140 pp., 17 pis. GRÜNDEL, J., 1976, ZurTaxonomie und Phyloge- FISCHER, P., 1878, Diagnosis molluscorum novo- nie der S/Yi/um-Gruppe. Malakologische Abhand- rum. Journal de Conchyliologie. 26: 211-212. lungen, 5: 33-59. FISCHER, P., 1880-1887 (1883-1884), Manuel de conchyliolgie et paléontologie conchyliologique HARDISON, L. K., & C. L. KITTING, 1985, Epi- (ou histoire naturelle des mollusques vivants et phytic algal browsing by Bittium varium (Gas- fossiles). Paris: Librairie F. Savy, xxiv + 1369 tropoda) among Thalassia testuidinum turtle- pp., 23 pis. (fascicule 6, pp. 513-608, 1883; fas- grass. Journal of Psychology, 21: 13. cicule 7, pp. 609-688, 1884). HASZPRUNAR, G., 1985. The fine morphology of FORBES, E. & S. HANLEY, 1850-1851. >4 history the esophageal sense organs of the Mollusca. 1. of British Mollusca and their shells, vol. 3. Van Gastropoda, Prosobranchia. Philosophical Voorst, London. 616 pp. Transactions of the Royal Society of London, B, FRETTER, V., 1948, The structure and life history 103: 457-496. of some minute prosobranchs of rock pools: Ske- HEALY, J. M., 1986. Ultrastructure of parasperma- neopsis planorbis (Fabricius), Omalogyra ato- tozoa of cerithiacean gastropods (Prosobran- mus (Philippi), Rissoella diaphana (Alder) and chia: Mesogastropoda). Helgoländer Meere- Rissoella opalina (Jeffreys). Journal of the Ma- suntersuchungen, 40: 177-199. rine Biological Association of the United King- HEDLEY, C, 1899, The Mollusca of Funafuti. Part dom, 27: 597-632. 1 .-Gastropoda. Memoirs of the Australian Mu- FRETTER, V., & A. GRAHAM, 1962, British proso- seum. 3: 397-567, 80 figs. branch molluscs. Ray Society, London. 755 pp.; HENNIG, W., 1966. Phylogeneticsystematics. Uni- 317 figs. versity of Illinois Press, Urbana. 263 pp. FRETTER, V. & M. PILKINGTON, 1970, Proso- HERTZ, J., 1981. A review of several eastern Pa- branchia. 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Cerithium nodulosum (Bruguière) [Gastropoda: HUTTON, F. W., 1873. Catalogue of the marine Prosobranchia]. Pacific Science, 25: 560-565. mollusks of New Zealand. Wellington, 1873. xx HOUBRICK, R. S., 1977, Réévaluation and new -I- 116 pp. description of the genus Bittium (Cerithiidae). IREDALE, T., 1924, Results from Roy Bell's mol- The Veliger, 20: 101-106. luscan collections. Proceedings of the Linnean HOUBRICK, R. S., 1978, Redescription of Bittium Society of New South Wales, 49: 179-278, pis. proteum (Jousseaume, 1930) witti comments on 33-36. its generic placement. The Nautilus, 92: 9-11. JEFFREYS, J. G., 1867-1869, British conchology, HOUBRICK, R. S., 1980a, Review of the deep-sea or an account of the Mollusca which now inhabit genus Argyropeza (Gastropoda: Prosobranchia: the British Isles and the surrounding seas. Lon- Cerithiidae). Smithsonian Contributions to Zool- don, van Voorst, vol. 4, 1867; vol. 5, 1869. ogy, 321: 30 pp., 11 figs. JEFFREYS, J. G., 1885, On the Mollusca procured HOUBRICK, R. S., 1980b, Reappraisal of the gas- during the Lightning and Porcupine expedition, tropod genus Varicopeza Gründel (Cerithiidae; 1868-70 (part 9). Proceedings of the Zoological Prosobranchia). Proceedings of the Biological Society of London, for 1885: 27-63, pis. 4-6. Society of Washington, 93'. 525-535. JOHANSSON, J., 1947, Über den offenen Uterus HOUBRICK, R. S., 1981a, Systematic position of bei einigen Monotocardiern ohne Kopulationsor- the genus Glyptozaria Iredale (Prosobranchia: gan. Zoologiska Bidrag fràn Uppsala, 25: 102- Gastropoda). Proceedings of the Biological Soci- 110. ety of Washington, 94: 838-847. JOHNSON, C. W., 1915, Fauna of New England, HOUBRICK, R. S., 1981b, Anatomy of Diastoma 13. List of the Mollusca. Occasional Papers of the melanoides (Reeve, 1849) with remarks on the Boston Society of Natural History, 7: 231 pp. systematic position of the family Diastomatidae JOHNSON, R., 1964. The Recent Mollusca of Au- (Prosobranchia: Gastropoda). Proceedings of the gustus Addison Gould. Bulletin of the United Biological Society of Washington, 94: 598-621. States National Museum, 239: 182 pp., 45 pis. HOUBRICK, R. S., 1981c, Anatomy and systemat- JOUSSEAUME, F., 1930, Cerithiidae de la Mer ics of Gourmya gourmyi (Prosobranchia: Cerithi- Rouge. Journal de Conchyliologie. 74: 270-296. idae), a Tethyan relict from the southwest Pacific. KAY, E. A., 1979, Hawaiian marine shells. Bishop The Nautilus, 95: 2-11. Museum Press, Special Publication 64(4), Hono- HOUBRICK, R. S., 1981d, Anatomy, biology and lulu, Hawaii. 653 pp. systematics of Campanile symbolicum with ref- KIENER, L C, 1841-1842, Spéies général et ico- erence to adaptive radiation of the Cerithiacea nographie des Coquilles vivantes . . . genre cé- (Gastropoda: Prosobranchia). Malacologia, 21 : rite. Paris: Rousseau, vol. 5: 1-104, pis. 1-32 263-289. [pis., 1841; text, 1841-1842]. HOUBRICK, R. S., 1987a, Transfer of Cerithiopsis KOBELT, W., 1888-1898, Die Gattung Cerithium, crystallina Dall to the genus Varicopeza Gründel, 297 pp., 47 pis., in F. H. W. MARTINI & J. H. CHEM- family Cerithiidae (Prosobranchia: Gastropoda). NITZ, Neues systematisches Conchylien- The Nautilus, 101: 80-85. Cabinet. . . . Nürntjerg, Bauer & Raspe, 1(26). HOUBRICK, R. S., 1987b, Anatomy of Alaba and KOSUGE, S., 1964. Anatomical study of Diala go- Litiopa (Prosobranchia: Litiopidae). The Nautilus, niochila (A. Adams) (Gastropoda). Bulletin of the 101: 9-18. National Science Museum, Tokyo, 7: 33-36. HOUBRICK, R. S., 1988, Cerithioidean phylogeny. LAMARCK, J. B. P. A. de, 1804, Suite des mé- in: W. F. PONDER, ed., Prosobranch phylogeny. moires sur les fossiles des environs de Paris. Proceedings of a symposium held at the 9th in- Annales du Muséum d'Histoire Naturelle, Paris. ternational malacological congress, Edinburgh, 3: 436-441. 1986. Malacological Review, Supplement 4: 88- LEA, H. 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